Morphological variation, modularity and integration in the scapula and humerus of Lissotriton newts.

The modular organization of tetrapod paired limbs and girdles, influenced by the expression of Hox genes is one of the primary driving forces of the evolution of animal locomotion. The increased morphological diversification of the paired limbs is correlated with reduced between-limb covariation, while correlation within the elements is usually higher than between the elements. The tailed amphibians, such as Lissotriton newts, have a biphasic lifestyle with aquatic and terrestrial environments imposing different constraints on limb skeleton. By employing the methods of computerized microtomography and 3D geometric morphometrics, we explored the pattern of morphological variation, disparity, modularity and morphological integration in the proximal parts of the anterior limbs of six species of Eurasian small bodied newts. Although the species significantly differ in limb shape, there is a great overlap in morphology of scapula and humerus, and there are no differences in morphological disparity. For the scapula, the shape differences related to the duration of the aquatic period are in length, depth and curvature. The shape of the humerus is not affected by the length of aquatic period, and shape differences between the species are related to robustness of the body. The length of aquatic period has statistically supported phylogenetic signal. The scapula and humerus are structures of varying modularity. For the humerus, the strongest support on the phylogenetic level was for the capitulum/shaft hypothesis, which can also be interpreted as functional modularity. For the scapula, the greatest support was for the antero-posterior hypothesis of modularity in case of Lissotriton vulgaris, which can be explained by different functional roles and muscle insertion patterns, while there was no phylogenetic modularity. The modularity patterns seem to correspond with the general tetrapod pattern, with modularity being more pronounced in the distal structure. The future research should include more salamandrid taxa with different habitat preferences and both adult and larval stages, in order to explore how size, phylogeny and ecology affect the morphology and covariation patterns of limbs.

Geological Substrate Effects on Teucrium montanum L. (Lamiaceae) Morphological Traits: Geometric Morphometrics Approach.

The shape-environment relationship in plants refers to the ways in which the physical characteristics and structures of plants are influenced by their environment. Plants have evolved a remarkable ability to adapt to their specific habitats, and their shape and form play a crucial role in determining their survival and reproductive success. This study aimed to examine differences in size and shape between morphological traits in mountain germander (Teucrium montanum L.) from different geological substrates (calcareous and serpentinite). For this study, 400 individuals of T. montanum from 20 populations (ten populations from the serpentinite and ten from the calcareous substrate) were selected. Using the geometric morphometrics approach, it was shown that the degree of phenotypic variation in the size and shape of the corolla, leaf, and stem of T. montanum depends on the type of substrate. The main differences between the populations are the narrower part of the lower lip of the corolla, the narrower leaf, and the wider central part of the vascular system stem from serpentinite populations. The results of this study will contribute to a better understanding of the morphological variability of T. montanum in relation to edaphic conditions. In addition, the results confirm that certain morphological differences play an important role in the adaptive response in relation to substrate composition, especially for substrates with increased metal content, such as serpentinite. The shape-environment relationship in plants could define diversity and complexity in plant life, and underscores the importance of shape as a key factor in their survival and success in different habitats.

Directional asymmetry and direction-giving factors: Lessons from flowers with complex symmetry.

Directional asymmetry is a systematic difference between the left and right sides for structures with bilateral symmetry or a systematic differentiation among repeated parts for complex symmetry. This study explores factors that produce directional asymmetry in the flower of Iris pumila, a structure with complex symmetry that makes it possible to investigate multiple such factors simultaneously. The shapes and sizes of three types of floral organs, the falls, standards, and style branches, were quantified using the methods of geometric morphometrics. For each flower, this study recorded the compass orientations of floral organs as well as their anatomical orientations relative to the two spathes subtending each flower. To characterize directional asymmetry at the whole-flower level, differences in the average sizes and shapes according to compass orientation and relative orientation were computed, and the left-right asymmetry was also evaluated for each individual organ. No size or shape differences within flowers were found in relation to anatomical position; this may relate to the terminal position of flowers in Iris pumila, suggesting that there may be no adaxial-abaxial polarity, which is very prominent in many other taxa. There was clear directional asymmetry of shape in relation to compass orientation, presumably driven by a consistent environmental gradient such as solar irradiance. There was also clear directional asymmetry between left and right halves of every floral organ, most likely related to the arrangement of organs in the bud. These findings indicate that different factors are acting to produce directional asymmetry at different levels. In conventional analyses not recording flower orientations, these effects would be impossible to disentangle from each other and would probably be included as part of fluctuating asymmetry.

Sexual Dimorphism and Morphological Modularity in Acanthoscelides obtectus (Say, 1831) (Coleoptera: Chrysomelidae): A Geometric Morphometric Approach.

Sexual dimorphism and specific patterns of development contribute in a great manner to the direction and degree of the sexual differences in body size and shape in many insects. Using a landmark-based geometric morpohometrics approach, we investigated sex-specific morphological size and shape variation in the seed beetle, Acanthoscelides obtectus. We also tested the functional hypothesis of the two morphological modules-thorax and abdomen in both sexes. Female-biased sexual dimorphism in size was shown, while differences in shape were reflected in the wider thorax and abdomen and shorter abdomen in females in comparison to males. The functional hypothesis of a two-module body was confirmed only in females before correction for size, and in both sexes after the allometry correction. Our results indicate that reproductive function has the central role in forming the patterns of modularity. We hypothesize that high morphological integration of the abdomen in females results from intense stabilizing selection, while the more relaxed integration in males is driven by the higher intensity of sexual selection.

Phenotypic plasticity in response to environmental heterogeneity contributes to fluctuating asymmetry in plants: first empirical evidence.

Fluctuating asymmetry (FA) is widely used to quantify developmental instability (DI) in ecological and evolutionary studies. It has long been recognized that FA may not exclusively originate from DI for sessile organisms such as plants, because phenotypic plasticity in response to heterogeneities in the environment might also produce FA. This study provides the first empirical evidence for this hypothesis. We reasoned that solar irradiance, which is greater on the southern side than on the northern side of plants growing in the temperate zone of the Northern Hemisphere, would cause systematic morphological differences and asymmetry associated with the orientation of plant parts. We used geometric morphometrics to characterize the size and shape of flower parts in Iris pumila grown in a common garden. The size of floral organs was not significantly affected by orientation. Shape and particularly its asymmetric component differed significantly according to orientation for three different floral parts. Orientation accounted for 10.4% of the total shape asymmetry within flowers in the falls, for 11.4% in the standards and for 2.2% in the style branches. This indicates that phenotypic plasticity in response to a directed environmental factor, most likely solar irradiance, contributes to FA of flowers under natural conditions. That FA partly results from phenotypic plasticity and not just from DI needs to be considered by studies of FA in plants and other sessile organisms.