Thomas A. Püschel

Standing on the shoulders of apes: Analyzing the form and function of the hominoid scapula using geometric morphometrics and finite element analysis

OBJECTIVE The aim was to analyze the relationship between scapular form and function in hominoids by using geometric morphometrics (GM) and finite element analysis (FEA). METHODS FEA was used to analyze the biomechanical performance of different hominoid scapulae by simulating static postural scenarios. GM was used to quantify scapular shape differences and the relationship between form and function was analyzed by applying both multivariate-multiple regressions and phylogenetic generalized least-squares regressions (PGLS). RESULTS Although it has been suggested that primate scapular morphology is mainly a product of function rather than phylogeny, our results showed that shape has a significant phylogenetic signal. There was a significant relationship between scapular shape and its biomechanical performance; hence at least part of the scapular shape variation is due to non-phylogenetic factors, probably related to functional demands. DISCUSSION This study has shown that a combined approach using GM and FEA was able to cast some light regarding the functional and phylogenetic contributions in hominoid scapular morphology, thus contributing to a better insight of the association between scapular form and function.

Ecomorphological Variation of the Wireworm Cephalic Capsule: Studying the Interaction of Environment and Geometric Shape

Studying the association between organismal morphology and environmental conditions has been very useful to test hypothesis regarding the influence of climate on shape. It has been long recognized that different environments produce dissimilar stress levels in insects, which can be reflected on the ability of an individual to overcome these pressures and spread further. Agriotes (Coleoptera: Elateridae) species infest agricultural fields in different parts of Croatia, inhabiting different climatic conditions. Previous biological studies have indicated that there is a relationship between some Agriotes biological parameters such as density and climatic conditions such as soil moisture and temperature. However, it is still unknown how these environmental properties influence the wireworm morphological structure. This is highly relevant because the head of this species is directly involved in the mobility in the soil, thus affecting the invasive capacity of this insect. Therefore the aim of this study was to assess the association between different climatic conditions and the morphological variation of Agriotes cephalic capsule. Advanced multivariate analysis and geometric morphometric tool were applied to study the covariation between shape and environmental variables. Partial Least Squares methods were used in order to analyse the association between the wireworm head shape and three different climatic conditions: soil type, temperature and rainfall. Our results showed that there is a high covariation between the wireworm head shape and the climatic conditions. It was suggested that the observed shape–environment association could be result of the high plasticity of this species in relation to its invasive capacity.

A biomechanical approach to understand the ecomorphological relationship between primate mandibles and diet

The relationship between primate mandibular form and diet has been previously analysed by applying a wide array of techniques and approaches. Nonetheless, most of these studies compared few species and/or infrequently aimed to elucidate function based on an explicit biomechanical framework. In this study, we generated and analysed 31 Finite Element planar models of different primate jaws under different loading scenarios (incisive, canine, premolar and molar bites) to test the hypothesis that there are significant differences in mandibular biomechanical performance due to food categories and/or food hardness. The obtained stress values show that in primates, hard food eaters have stiffer mandibles when compared to those that rely on softer diets. In addition, we find that folivores species have the weakest jaws, whilst omnivores have the strongest mandibles within the order Primates. These results are highly relevant because they show that there is a strong association between mandibular biomechanical performance, mandibular form, food hardness and diet categories and that these associations can be studied using biomechanical techniques rather than focusing solely on morphology.

Erratum to: Left–right asymmetry and morphological consequences of a host shift in the oligophagous Neotropical moth Macaria mirthae (Lepidoptera: Geometridae)

Phytophagous insects are excellent model organisms to study the genetic and ecological components of adaptation and morphological divergence, because their host plants are one of the main environmental factors influencing their early life stages. Although many lepidopterans are highly specialized in their host use, shifts to exotic plants have been reported for some species. Macaria mirthae is a native moth from Northern Chile that feeds preferentially on the Fabacea species Acacia macracantha, however due to habitat loss a host shift has recently been observed to the introduced fabacean Leucaena leucocephala. We studied the impact that different host plants have on the developmental instability levels in the moth’s wing morphology evaluating both fluctuating asymmetry (FA) and directional asymmetry (DA). FA measures the small random deviations existing between the left and right sides of bilaterally symmetrical traits and it widely used as a biomonitor of environmental quality. DA refers to the tendency for a trait to be consistently developed in a different manner on the right and left sides of the body. It has been recently shown that subtle DA patterns seem to be a ubiquitous phenomenon among bilaterian animals. Our results confirmed the presence of FA in M. mirthae forewings by applying geometric morphometric techniques. Furthermore, it was found that the individuals feeding on the endemic tree (A. macracantha) showed marked DA levels, while the specimens inhabiting the exotic plant (L. leucocephala) did not. The absence of DA in the individuals occupying the exotic plant is striking, because it has been established that this asymmetry pattern is widespread among insect wings. This phenomenon could be related to the influence of L. leucocephala on normal wing development. Despite the reduced quality of L. leucocephala as host plant, its wider presence in the Azapa valley (Chile) could explain the host shift made by M. mirthae.

Analyzing the sclerocarpy adaptations of the Pitheciidae mandible

Primates are interpreted to be ancestrally adapted to frugivory, although some modern groups show clear adaptations to other diets. Among them, pitheciids stand out for specifically predating seeds. This dietary specialization is known as sclerocarpy and refers to the extraction of seeds from surrounding hard tissues using the anterior dentition followed by the mastication of seeds by the molars. It has been proposed that Callicebus‐Pithecia‐Chiropotes‐Cacajao represent a morphocline of increasingly specialized anatomical traits for sclerocarpic foraging. This study addresses whether there is a sclerocarpic specialization gradient in the mandibular morphology of pitheciids. Finite element analysis (FEA) was used to simulate two biting scenarios and the obtained stress values were compared between different pitheciids. Geometric morphometrics (GM) were used to display the morphological variation of this group. No support was found for the morphocline hypothesis from a biomechanical viewpoint since all pitheciins showed similar stress values and on average Chiropotes rather than Cacajao exhibited the strongest mandible. From a morphological perspective, it was found that there is indeed relative “robusticity” continuum in the pitheciid mandible for some aspects of shape as expected for the morphocline hypothesis, but this gradient could be related to other factors rather than sclerocarpic specialization. The present results are expected to contribute to a better insight regarding the ecomorphological relationship between mandibular morphology and mechanical performance among pitheciids.

The intervals method: a new approach to analyse finite element outputs using multivariate statistics

Background In this paper, we propose a new method, named the intervals’ method, to analyse data from finite element models in a comparative multivariate framework. As a case study, several armadillo mandibles are analysed, showing that the proposed method is useful to distinguish and characterise biomechanical differences related to diet/ecomorphology. Methods The intervals’ method consists of generating a set of variables, each one defined by an interval of stress values. Each variable is expressed as a percentage of the area of the mandible occupied by those stress values. Afterwards these newly generated variables can be analysed using multivariate methods. Results Applying this novel method to the biological case study of whether armadillo mandibles differ according to dietary groups, we show that the intervals’ method is a powerful tool to characterize biomechanical performance and how this relates to different diets. This allows us to positively discriminate between specialist and generalist species. Discussion We show that the proposed approach is a useful methodology not affected by the characteristics of the finite element mesh. Additionally, the positive discriminating results obtained when analysing a difficult case study suggest that the proposed method could be a very useful tool for comparative studies in finite element analysis using multivariate statistical approaches.

The evolution of the platyrrhine talus: a comparative analysis of the phenetic affinities of the Miocene platyrrhines with their modern relatives

Platyrrhines are a diverse group of primates that presently occupy a broad range of tropical-equatorial environments in the Americas. However, most of the fossil platyrrhine species of the early Miocene have been found at middle and high latitudes. Although the fossil record of New World monkeys has improved considerably over the past several years, it is still difficult to trace the origin of major modern clades. One of the most commonly preserved anatomical structures of early platyrrhines is the talus. This work provides an analysis of the phenetic affinities of extant platyrrhine tali and their Miocene counterparts through geometric morphometrics and a series of phylogenetic comparative analyses. Geometric morphometrics was used to quantify talar shape affinities, while locomotor mode percentages (LMPs) were used to test if talar shape is associated with locomotion. Comparative analyses were used to test if there was convergence in talar morphology, as well as different models that could explain the evolution of talar shape and size in platyrrhines. Body mass predictions for the fossil sample were also computed using the available articular surfaces. The results showed that most analyzed fossils exhibit a generalized morphology that is similar to some ‘generalist’ modern species. It was found that talar shape covaries with LMPs, thus allowing the inference of locomotion from talar morphology. The results further suggest that talar shape diversification can be explained by invoking a model of shifts in adaptive peak to three optima representing a phylogenetic hypothesis in which each platyrrhine family occupied a separate adaptive peak. The analyses indicate that platyrrhine talar centroid size diversification was characterized by an early differentiation related to a multidimensional niche model. Finally, the ancestral platyrrhine condition was reconstructed as a medium-sized, generalized, arboreal, quadruped.

Analysing the Floral Elements of the Lost Tree of Easter Island: A Morphometric Comparison between the Remaining Ex- Situ Lines of the Endemic Extinct Species Sophora toromiro

Sophora toromiro (Phil) Skottsb. is a species that has been extinct in its natural habitat Easter Island (Rapa Nui) for over 50 years. However, seed collections carried out before its extinction have allowed its persistence ex-situ in different botanical gardens and private collections around the world. The progenies of these diverse collections have been classified in different lines, most of them exhibiting high similarity as corroborated by molecular markers. In spite of this resemblance observed between the different lines, one of them (Titze) has dissimilar floral elements, thus generating doubts regarding its species classification. The floral elements (wing, standard and keel) belonging to three different S. toromiro lines and two related species were analyzed using geometric morphometrics. This method was applied in order to quantify the floral shape variation of the standard, wing, and keel between the different lines and control species. Geometric morphometrics analyses were able to distinguish the floral elements at both intra (lines) and inter-specific levels. The present results are on line with the cumulative evidence that supports the Titze line as not being a proper member of the S. toromiro species, but probably a hybridization product or even another species of the Edwardsia section. The reintroduction programs of S. toromiro should consider this information when assessing the authenticity and origin of the lines that will be used to repopulate the island.

Inferring locomotor behaviours in Miocene New World monkeys using finite element analysis, geometric morphometrics and machine-learning classification techniques applied to talar morphology

The talus is one of the most commonly preserved post-cranial elements in the platyrrhine fossil record. Talar morphology can provide information about postural adaptations because it is the anatomical structure responsible for transmitting body mass forces from the leg to the foot. The aim of this study is to test whether the locomotor behaviour of fossil Miocene platyrrhines could be inferred from their talus morphology. The extant sample was classified into three different locomotor categories and then talar strength was compared using finite-element analysis. Geometric morphometrics were used to quantify talar shape and to assess its association with biomechanical strength. Finally, several machine-learning (ML) algorithms were trained using both the biomechanical and morphometric data from the extant taxa to infer the possible locomotor behaviour of the Miocene fossil sample. The obtained results show that the different locomotor categories are distinguishable using either biomechanical or morphometric data. The ML algorithms categorized most of the fossil sample as arboreal quadrupeds. This study has shown that a combined approach can contribute to the understanding of platyrrhine talar morphology and its relationship with locomotion. This approach is likely to be beneficial for determining the locomotor habits in other fossil taxa.

Modificación Intencional del Cráneo en San Pedro de Atacama y la Cuenca del Loa: Un Análisis Cuantitativo de su Rol como Marcador de Identidad Social

Diverse hypotheses have been proposed to explain artificial cranial modification (ACM) in South America. In the Atacama area (Northern Chile), some studies have concluded that it was used to create a communal identity that could serve to resist, or to form alliances with, different external groups (Inter Site Distinction hypothesis, this work). On the other hand, other studies have suggested that there was a relationship between ACM and the social status and gender of the individuals within the community (Intra Site Distinction hypothesis, this work). These studies, however, have relied mainly on typological methods and the archaeological contexts to which these modification categories are associated have been simplified as well. In this work we use a quantitative multivariate approach to assess the relationship between cranial morphology and funerary context. The modification patterns of populations inhabiting Northern Chile during the Formative (3500-1600 B.P) and Late Intermediate (950-500 B.P.) periods were studied. We analyzed the X-rays of 203 individuals belonging to 7 archaeological sites and, when possible, they were correlated with the corresponding funerary context. The results indicate that cranial morphology correlates with the interaction networks among sites, therefore these results support the Inter Site hypothesis.