Marina Alcobendas

Initial Diversification of Living Amphibians Predated the Breakup of Pangaea

The origin and divergence of the three living orders of amphibians (Anura, Caudata, Gymnophiona) and their main lineages are one of the most hotly debated topics in vertebrate evolution. Here, we present a robust molecular phylogeny based on the nuclear RAG1 gene as well as results from a variety of alternative independent molecular clock calibrations. Our analyses suggest that the origin and early divergence of the three living amphibian orders dates back to the Palaeozoic or early Mesozoic, before the breakup of Pangaea, and soon after the divergence from lobe‐finned fishes. The resulting new biogeographic scenario, age estimate, and the inferred rapid divergence of the three lissamphibian orders may account for the lack of fossils that represent plausible ancestors or immediate sister taxa of all three orders and the heretofore paradoxical distribution of some amphibian fossil taxa. Furthermore, the ancient and rapid radiation of the three lissamphibian orders likely explains why branch lengths connecting their early nodes are particularly short, thus rendering phylogenetic inference of implicated relationships especially difficult.

DISPERSAL OF VIVIPARITY ACROSS CONTACT ZONES IN IBERIAN POPULATIONS OF FIRE SALAMANDERS (SALAMANDRA) INFERRED FROM DISCORDANCE OF GENETIC AND MORPHOLOGICAL TRAITS

Abstract.— We used partial sequences of the cytochrome b mitochondrial DNA (mtDNA) gene, obtained from 76 individuals representing 45 populations of Iberian Salamandra salamandra plus 15 sequences of additional species of Salamandra and related genera, to investigate contact zones. These zones, identified by earlier allozymic and morphological analyses, are between populations of viviparous (S. s. bernardezi and S. s. fastuosa) and ovoviviparous (S. s. gallaica and S. s. terrestris) salamanders. The distribution of mtDNA and nuclear markers is mostly concordant at one contact zone (between S. s. gallaica and S. s. bernardezi), but at another (between S. s. fastuosa and S. s. terrestris) the markers are offset by about 250 km. The observed geographic variation fits a model of mtDNA capture. Among the potential mechanisms responsible for such discordance we favor a combination of range shifts due to climatic fluctuations and biased genetic admixture across moving contact zones. We apply our findings to the issue of possible homoplasy in the evolution of viviparity and conclude that viviparity likely arose only once within S. salamandra.

Heterochrony, cannibalism, and the evolution of viviparity in Salamandra salamandra

SUMMARY The way in which novelties that lead to macroevolutionary events originate is a major question in evolutionary biology, and one that can be addressed using the fire salamander (Salamandra salamandra) as a model system. It is exceptional among amphibians in displaying intraspecific diversity of reproductive strategies. In S. salamandra, two distinct modes of reproduction co‐occur: the common mode, ovoviviparity (females giving birth to many small larvae), and a phylogenetically derived reproductive strategy, viviparity (females producing only a few large, fully metamorphosed juveniles, which are nourished maternally). We examine the relationship between heterochronic modifications of the ontogeny and the evolution of the new reproductive mode in the fire salamander. The in vitro development of embryos of ovoviviparous and viviparous salamanders from fertilization to metamorphosis is compared, highlighting the key events that distinguish the two modes of reproduction. We identify the heterochronic events that, together with the intrauterine cannibalistic behavior, characterize the derived viviparous reproductive strategy. The ways in which evolutionary novelties can arise by modification of developmental programs can be studied in S. salamandra. Moreover, the variation in reproductive modes and the associated variation of sequences of development occur in neighboring, conspecific populations. Thus, S. salamandra is a unique biological system in which evolutionary developmental research questions can be addressed at the level of populations.

VARIABILITY IN SURVIVAL, GROWTH AND METAMORPHOSIS IN THE LARVAL FIRE SALAMANDER (SALAMANDRA SALAMANDRA): EFFECTS OF LARVAL BIRTH SIZE, SIBSHIP AND ENVIRONMENT

Salamandra salamandra has an unusual mode of reproduction. Females retain eggs in their oviducts and give birth to aquatic larvae or, in some populations, to fully metamorphosed juveniles. We investigated how variation in the birth size of larvae of five different sibships of S. salamandra affected larval survival, larval growth rates, time to metamorphosis, and size at metamorphosis under different food and temperature conditions. Maternal effects, via the effect of larval birth size, attenuated throughout development in all environments but still affected metamorphic traits at lower temperatures. Larval developmental time was negatively correlated with larval birth size and positively correlated with size at metamorphosis. After controlling for the effect of larval birth size, broad-sense heritabilities were obtained for residual values of metamorphic traits, indicating that enough genetic variability exists at lower temperature environments and that responses to selection of metamorphic traits are possible. The size at and the time to metamorphosis were phenotypically and genetically correlated. Since both traits presumably affect fitness, genetic dependence between size and time to metamorphosis may act as a constraint on adaptive evolution through antagonistic pleiotropy. Mass-specific growth, time to metamorphosis, and size at metamorphosis were plastic under different temperature and food conditions, but no significant sibship × environment interaction was found for any trait. In addition, significant cross-environment correlation for size at metamorphosis with respect to the food gradient indicates low potential for the adaptive evolution of reaction norms to heterogeneous food environments.