Alexander O. Vargas

Development of neural crest in Xenopus.

The neural crest is a unique cell population among embryonic cell types, displaying properties of both ectodermal and mesodermal lineages. Most of the recent studies examining the neural crest have been performed in avian embryos. Only in the first half of this century were amphibians extensively used. We first summarize this important older source of information, reviewing studies made since the turn of the century. Due to the increasingly detailed in cellular and molecular knowledge of the early development of Xenopus laevis, the remainder of the review focuses on this species. We describe the route of migration and fate of the neural crest and propose a new model of neural crest induction in which prospective cells are induced independently of the neural plate by a double gradient of a morphogen that patterns the entire ectoderm. This model is also discussed in a more general context in connection with the dorsoventral patterning of the neural tube. Finally, we discuss some ideas concerning neural crest evolution and propose a novel hypothesis about its phylogenetic origin.

An enigmatic plant-eating theropod from the Late Jurassic period of Chile

Theropod dinosaurs were the dominant predators in most Mesozoic era terrestrial ecosystems. Early theropod evolution is currently interpreted as the diversification of various carnivorous and cursorial taxa, whereas the acquisition of herbivorism, together with the secondary loss of cursorial adaptations, occurred much later among advanced coelurosaurian theropods. A new, bizarre herbivorous basal tetanuran from the Upper Jurassic of Chile challenges this conception. The new dinosaur was discovered at Aysén, a fossil locality in the Upper Jurassic Toqui Formation of southern Chile (General Carrera Lake). The site yielded abundant and exquisitely preserved three-dimensional skeletons of small archosaurs. Several articulated individuals of Chilesaurus at different ontogenetic stages have been collected, as well as less abundant basal crocodyliforms, and fragmentary remains of sauropod dinosaurs (diplodocids and titanosaurians).

New developmental evidence clarifies the evolution of wrist bones in the dinosaur-bird transition.

A new study that integrates developmental and paleontological data reveals previously unsuspected evolutionary transformations during the emergence of the bird wrist, consistent with its derivation from non-avian dinosaurs.

Skeletal plasticity in response to embryonic muscular activity underlies the development and evolution of the perching digit of birds

Most birds have an opposable digit 1 (hallux) allowing the foot to grasp, which evolved from the non-opposable hallux of early theropod dinosaurs. An important morphological difference with early theropods is the twisting of the long axis of its metatarsal. Here, we show how embryonic musculature and the onset of its activity are required for twisting of metatarsal 1 (Mt1) and retroversion of the hallux. Pharmacologically paralyzed embryos do not fully retrovert the hallux and have a straight Mt1 shaft, phenocopying the morphology of early tetanuran dinosaurs. Molecular markers of cartilage maturation and ossification show that differentiation of Mt1 is significantly delayed compared to Mt2-4. We hypothesize on how delayed maturation may have increased plasticity, facilitating muscular twisting. Our experimental results emphasize the importance of embryonic muscular activity in the evolutionary origin of a crucial adaptation.

The developmental origin of zygodactyl feet and its possible loss in the evolution of Passeriformes

The zygodactyl orientation of toes (digits II and III pointing forwards, digits I and IV pointing backwards) evolved independently in different extant bird taxa. To understand the origin of this trait in modern birds, we investigated the development of the zygodactyl foot of the budgerigar (Psittaciformes). We compared its muscular development with that of the anisodactyl quail (Galliformes) and show that while the musculus abductor digiti IV (ABDIV) becomes strongly developed at HH36 in both species, the musculus extensor brevis digiti IV (EBDIV) degenerates and almost disappears only in the budgerigar. The asymmetric action of those muscles early in the development of the budgerigar foot causes retroversion of digit IV (dIV). Paralysed budgerigar embryos do not revert dIV and are anisodactyl. Both molecular phylogenetic analysis and palaeontological information suggest that the ancestor of passerines could have been zygodactyl. We followed the development of the zebra finch (Passeriformes) foot muscles and found that in this species, both the primordia of the ABDIV and of the EBDIV fail to develop. These data suggest that loss of asymmetric forces of muscular activity exerted on dIV, caused by the absence of the ABDIV, could have resulted in secondary anisodactyly in Passeriformes.

New developmental evidence supports a homeotic frameshift of digit identity in the evolution of the bird wing

BackgroundThe homology of the digits in the bird wing is a high-profile controversy in developmental and evolutionary biology. The embryonic position of the digits cartilages with respect to the primary axis (ulnare and ulna) corresponds to 2, 3, 4, but comparative-evolutionary morphology supports 1, 2, 3. A homeotic frameshift of digit identity in evolution could explain how cells in embryonic positions 2, 3, 4 began developing morphologies 1, 2, 3. Another alternative is that no re-patterning of cell fates occurred, and the primary axis shifted its position by some other mechanism. In the wing, only the anterior digit lacks expression of HoxD10 and HoxD12, resembling digit 1 of other limbs, as predicted by 1, 2, 3. However, upon loss of digit 1 in evolution, the most anterior digit 2 could have lost their expression, deceitfully resembling a digit 1. To test this notion, we observed HoxD10 and HoxD12 in a limb where digit 2 is the most anterior digit: The rabbit foot. We also explored whether early inhibition of Shh signalling in the embryonic wing bud induces an experimental homeotic frameshift, or an experimental axis shift. We tested these hypotheses using DiI injections to study the fate of cells in these experimental wings.ResultsWe found strong transcription of HoxD10 and HoxD12 was present in the most anterior digit 2 of the rabbit foot. Thus, we found no evidence to question the use of HoxD expression as support for 1, 2, 3. When Shh signalling in early wing buds is inhibited, our fate maps demonstrate that an experimental homeotic frameshift is induced.ConclusionAlong with comparative morphology, HoxD expression provides strong support for 1, 2, 3 identity of wing digits. As an explanation for the offset 2, 3, 4 embryological position, the homeotic frameshift hypothesis is consistent with known mechanisms of limb development, and further proven to be experimentally possible. In contrast, the underlying mechanisms and experimental plausibility of an axis shift remain unclear.

A new species of chimaeriform (Chondrichthyes, Holocephali) from the uppermost Cretaceous of the Lopez de Bertodano Formation, Isla Marambio (Seymour Island), Antarctica

Abstract We describe a new chimaeriform fish, Callorhinchus torresi sp. nov., from the uppermost Cretaceous (late Maastrichtian) of the López de Bertodano Formation, Isla Marambio (Seymour Island), Antarctica. The material shows it is distinct from currently known fossil and extant species of the genus, whereas the outline of the tritors (abrasive surfaces of each dental plate) shows an intermediate morphology between earlier records from the Cenomanian of New Zealand and those from the Eocene of Isla Marambio. This suggests an evolutionary trend in tritor morphology in the lineage leading to modern callorhynchids, during the Late Cretaceous-Palaeogene interval.

A new titanosaur sauropod from the Atacama Desert, Chile

Partial remains of a titanosaur sauropod collected in the Tolar Formation (Upper Cretaceous) at the Atacama Desert (Antofagasta Region), northern Chile, is described, and a new species, Atacamatitan chilensis gen. et sp. nov., is erected. The material consists mainly of dorsal and caudal vertebrae, part of a humerus and a femur. The presence of a titanosaur confirms the Cretaceous age for the outcrops of red sandstone of the Tolar Formation whose age was previously uncertain, ranging from the Upper Cretaceous to the Paleocene. The new specimen represents the most complete dinosaur reported for this region and one of the most complete titanosaur known from Chile and the pacific margin of South America so far.

Altriciality and the Evolution of Toe Orientation in Birds

Specialized morphologies of bird feet have evolved several times independently as different groups have become zygodactyl, semi-zygodactyl, heterodactyl, pamprodactyl or syndactyl. Birds have also convergently evolved similar modes of development, in a spectrum that goes from precocial to altricial. Using the new context provided by recent molecular phylogenies, we compared the evolution of foot morphology and modes of development among extant avian families. Variations in the arrangement of toes with respect to the anisodactyl ancestral condition have occurred only in altricial groups. Those groups represent four independent events of super-altriciality and many independent transformations of toe arrangements (at least four zygodactyl, three semi-zygodactyl, one heterodactyl, one pamprodactyl group, and several syndactyl). We propose that delayed skeletal maturation due to altriciality facilitates the epigenetic influence of embryonic muscular activity over developing toes, allowing for repeated evolution of innovations in their morphology.

NEW CHONDRICHTHYANS FROM THE UPPER CRETACEOUS (CAMPANIAN-MAASTRICHTIAN) OF SEYMOUR AND JAMES ROSS ISLANDS, ANTARCTICA

Abstract We present new records of chondrichthyans recovered from strata of Maastrichtian age of the López de Bertodano Formation, Seymour (=Marambio) Island, and from levels of latest Campanian age of the Santa Marta Formation, James Ross Island, both located in the eastern Antarctic Peninsula. The material from Marambio Island comprises an associated assemblage with the first records of an indeterminate odontaspidid different from Odontaspis, as well as the genera Pristiophorus, Squatina, Paraorthacodus, and the species Chlamydoselachus tatere from the López de Bertodano Formation. Also, the studied section provides a well-constrained age for several taxa already recognized in the López de Bertodano Formation only by scattered samples of Maastrichtian age for the first time. The assemblage from Marambio Island is representative of one of the latest environmental conditions during the end of the Cretaceous in the coastal seas of the Larsen Basin before major changes that began after the K/P boundary. In addition, the finds from James Ross Island comprise the southernmost records of the neoselachians Cretalamna sp., Centrophoroides sp., as well as the holocephalans Callorhinchus sp. and an indeterminate rhinochimaerid, extending the occurrence of some of these taxa into the late Campanian, being their oldest record of the Weddellian Biogeographic Province.