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Several studies describe the development of the chondrocranium of vertebrates. The details in these studies vary a lot, which makes it hard to compare developmental patterns and identify evolutionary trends. Therefore, we aim to close this gap for anurans, which is the largest order of amphibians. We present here a detailed description of the chondrocranium morphology and development of Smilisca phaeota, the New Granada cross-banded tree frog. The anatomy was described for the larvae at or older than Gossner stage 31 and before ossification starts. Following this, we describe the development of the chondrocranium from Gossner stages 19–26. Early in Gossner stage 19 no precursors of any cartilages are visible, while later in that stage the mesodermal Anlage of Meckel’s cartilage was observed. In the subsequent stages more and more mesodermal anlagen become identifiable, followed by chondrification, and final differentiation of the cartilage elements. We used serial sections to study all the developmental stages and additionally utilized cleared and stained specimens and CT scan data. The latter were also used for the 3D reconstruction of the chondrocranium. We previously studied several species and compared these developmental patterns with S. phaeota , revealing potentially characteristic patterns significant for Lalagobatrachia, a clade that includes over 7000 frog species. These include (1) the suprarostral alae develop before the suprarostral corpus, (2) the infrarostral cartilage chondrifies late, after the chondrification of ceratobranchial 1, and (3) the ceratohyal body is the first element to show chondrocytes and to chondrify. However, with only six species studied so far, our data only provide a basis for future studies and developing hypotheses about the ancestral developmental pattern in anurans.

Background The craniofacial skeleton is an evolutionary innovation of vertebrates. Due to its complexity and importance to protect the brain and aid in essential functions (e.g., feeding), its development requires a precisely tuned sequence of chondrification and/or ossification events. The comparison of sequential patterns of cartilage formation bears important insights into the evolution of development. Discoglossus scovazzi is a basal anuran species. The comparison of its chondrocranium (cartilaginous neuro- & viscerocranium) development with other basal anurans ( Xenopus laevis , Bombina orientalis ) will help establishing the ancestral pattern of chondrification sequences in anurans and will serve as basis for further studies to reconstruct ancestral conditions in amphibians, tetrapods, and vertebrates. Furthermore, evolutionary patterns in anurans can be studied in the light of adaptations once the ancestral sequence is established. Results We present a comprehensive overview on the chondrocranium development of D. scovazzi. With clearing and staining, histology and 3D reconstructions we tracked the chondrification of 44 elements from the first mesenchymal Anlagen to the premetamorphic cartilaginous head skeleton and illustrate the sequential changes of the skull. We identified several anuran and discoglossoid traits of cartilage development. In D. scovazzi the mandibular, hyoid, and first branchial arch Anlagen develop first followed by stepwise addition of the branchial arches II, III, and IV. Nonetheless, there is no strict anterior to posterior chondrification pattern within the viscerocranium of D. scovazzi . Single hyoid arch elements chondrify after elements of the branchial arch and mandibular arch elements chondrify after elements of the branchial arch I. Conclusions In Osteichthyes, neurocranial elements develop in anterior to posterior direction. In the anurans investigated so far, as well as in D. scovazzi , the posterior parts of the neurocranium extend anteriorly, while the anterior parts of the neurocranium, extend posteriorly until both parts meet and fuse. Anuran cartilaginous development differs in at least two crucial traits from other gnathostomes which further supports the urgent need for more developmental investigations among this clade to understand the evolution of cartilage development in vertebrates.

Analysis of the developmental sequence variability provides data linking the ontogeny and phylogeny of the investigated objects. Here, we present a study of cranial ossifications sequence variability in the polyphyletic group of Cyprininae Barbus sensu lato (Cypriniformes, Teleostei). We analyzed the intra- and interspecific variability, compared sequences of closely related and phylogenetically distant species, reconstructed an alleged ancestral sequence, and assessed the sequence variations’ evolutionary validity. As the results, we revealed that the observed sequence alterations occur due to intrinsic, likely genetic or epigenetic factors. The skull ossifications, as well as skull regions, differ in the variability level. The ossifications appearing at the intermediate phase and skull regions composing splanchnocranium are most variable. In contrast, the structures emerging at the early and late phases of skull development and regions of neurocranium are less variable. The majority of sequence alterations are non-adaptive and selectively neutral. Increase in their number accompanying the transition from intra- to interspecific and intergeneric levels resembles the accumulation of the selectively neutral genetic variations accompanying the increase of phylogenetic distance. Reconstructions of phylogenetic relationships between investigated species obtained with the analyses of sequence similarity and phylogenetic analyses using parsimony (PAUP) are consistent with the phylogenies based on molecular data and thus demonstrate the potential of sequence analysis for phylogenetic inference.

Developmental biology often yields data in a temporal context. Temporal data in phylogenetic systematics has important uses in the field of evolutionary developmental biology and, in general, comparative biology. The evolution of temporal sequences, specifically developmental sequences, has proven difficult to examine due to the highly variable temporal progression of development. Issues concerning the analysis of temporal sequences and problems with current methods of analysis are discussed. We present here an algorithm to infer ancestral temporal sequences, quantify sequence heterochronies, and estimate pseudoreplicate consensus support for sequence changes using Parsimov-based genetic inference [PGi]. Real temporal developmental sequence data sets are used to compare PGi with currently used approaches, and PGi is shown to be the most efficient, accurate, and practical method to examine biological data and infer ancestral states on a phylogeny. The method is also expandable to address further issues in developmental evolution, namely modularity.

• Premise of the study: Crested sepals, which have evolved at least five times in Iris, are adaxially elaborated with a sinuous and/or uneven median structure (crest) along the proximal-distal axis and sometimes with various lateral structures (ridges, crests, and linear protuberances) flanking the median crest. These structural elaborations are complex yet diverse in form, providing a good opportunity to investigate developmental mechanisms for the diversification of reproductive lateral organs. • Methods: Morphologies of the median and lateral structures at different developmental stages from selected crested sepals representing the major types of structural elaborations were recorded using scanning electron microscopy and light microscopy. Developmental (morphogenetic) events that contribute to changes in shape (eg., sinuousness, unevenness) between consecutive stages were recorded. Developmental sequences—trajectories that consist of a series of developmental events— were compared in a phylogenetic context. • Key results: Three developmental events (development of outgrowths, greater expansion of the upper zone, and greater expansion of the lower zone), are shared across lineages, occur in the same developmental sequences, and are responsible for the changes in shape during the development of diverse structural elaborations. In addition, two novel developmental events and the development of trichomes on elaborate structures were observed within the core-crested clade. • Conclusions: Developmental sequences are conserved across independently evolved crested lineages. Heterochronic and heterotopic shifts of developmental events play the major role in the diversification of elaborations of crested sepals in Iris. The evolution of novel developmental events and the development of trichomes also contribute to the diversity.

The current study proposes a Developmental Sequence Model to University Adjustment and uses a multifaceted measure, including academic, social and psychological adjustment, to examine factors predictive of undergraduate international student adjustment. A hierarchic regression model is carried out on the Student Adaptation to College Questionnaire to examine theoretically pertinent predictors arranged in a developmental sequence in determining adjustment outcomes. This model accounted for over 60% of the variance in adjustment scores, and highlighted the importance of ecological factors in relation to student adjustment, such as social support, relationship with parents, and perceived institutional support as more useful than static measures of achievement such as high school GPA. Implications of these findings, including the important role of campus services, are discussed.

Purpose: The purposes of this study were to examine age and gender differences in throwing performance across an underexplored portion of the lifespan: middle adolescents (14-17 years old), young adults (18-25 years old), and adults (35-55 years old). Method: Throwing performance was assessed using the body component levels from Roberton's developmental sequences for force and ball velocity that were recorded by a radar gun. Participants in each age group performed between 5 to 10 forceful overhand throws toward a target approximately 15 m to 20 m from the thrower. A Wilcoxon-Mann-Whitney Test was used to determine gender differences and a Wilcoxon-Signed Ranks Test was used to determine age-group differences for each component. Gender and age-group differences in ball speed were determined by a 3 (age group) × 2 (gender) factorial analysis of variance with follow-up post-hoc tests. Results: Young-adult men had higher body component levels and ball speed compared with the adolescent boys and adult men. Female age-group differences existed only for humerus action between young-adult and adult groups and for ball speed between young-adult and adolescent groups. Gender differences (p < .01) existed in component levels for the adolescent and young-adult groups, but not the adult groups. Gender differences in ball speed (p < .001) existed within each age group. Conclusion: Although these data were cross-sectional, the regressive developmental changes observed and the narrowing gender gap may eventually provide insight related to the relationships among motor skill competence, physical fitness, and physical activity across the lifespan.

Sequence heterochrony (changes in the order in which events occur) is a potentially important, but relatively poorly explored, mechanism for the evolution of development. In part, this is because of the inherent difficulties in inferring sequence heterochrony across species. The event-pairing method, developed independently by several workers in the mid-1990s, encodes sequences in a way that allows them to be examined in a phylogenetic framework, but the results can be difficult to interpret in terms of actual heterochronic changes. Here, we describe a new, parsimony-based method to interpret such results. For each branch of the tree, it identifies the least number of event movements (heterochronies) that will explain all the observed event-pair changes. It has the potential to find all alternative, equally parsimonious explanations, and generate a consensus, containing the movements that form part of every equally most parsimonious explanation. This new technique, which we call Parsimov, greatly increases the utility of the event-pair method for inferring instances of sequence heterochrony.

INTRODUCTION: One of the most interesting riddles within crustaceans is the origin of Cladocera (water fleas). Cladocerans are morphologically diverse and in terms of size and body segmentation differ considerably from other branchiopod taxa (Anostraca, Notostraca, Laevicaudata, Spinicaudata and Cyclestherida). In 1876, the famous zoologist Carl Claus proposed with regard to their origin that cladocerans might have evolved from a precociously maturing larva of a clam shrimp-like ancestor which was able to reproduce at this early stage of development. In order to shed light on this shift in organogenesis and to identify (potential) changes in the chronology of development (heterochrony), we investigated the external and internal development of the ctenopod Penilia avirostris and compared it to development in representatives of Anostraca, Notostraca, Laevicaudata, Spinicaudata and Cyclestherida. The development of the nervous system was investigated using immunohistochemical labeling and confocal microscopy. External morphological development was followed using a scanning electron microscope and confocal microscopy to detect the autofluorescence of the external cuticle. RESULTS: In Anostraca, Notostraca, Laevicaudata and Spinicaudata development is indirect and a free-swimming nauplius hatches from resting eggs. In contrast, development in Cyclestherida and Cladocera, in which non-swimming embryo-like larvae hatch from subitaneous eggs (without a resting phase) is defined herein as pseudo-direct and differs considerably from that of the other groups. Both external and internal development in Anostraca, Notostraca, Laevicaudata and Spinicaudata is directed from anterior to posterior, whereas in Cyclestherida and Cladocera differentiation is more synchronous. CONCLUSIONS: In this study, developmental sequences from representatives of all branchiopod taxa are compared and analyzed using a Parsimov event-pairing approach. The analysis reveals clear evolutionary transformations towards Cladocera and the node of Cladoceromorpha which correspond to distinct heterochronous signals and indicate that the evolution of Cladocera was a stepwise process. A switch from a strategy of indirect development to one of pseudo-direct development was followed by a shift in a number of morphological events to an earlier point in ontogenesis and simultaneously by a reduction in the number of pre-metamorphosis molts. A compression of the larval phase as well as a shortening of the juvenile phase finally leads to a precocious maturation and is considered as a gradual progenetic process.