Testing the Tropical Niche Conservatism Hypothesis: Climatic Niche Evolution of Escallonia Mutis ex L. F. (Escalloniaceae).

We assess the Tropical Niche Conservatism Hypothesis in the genus Escallonia in South America using phylogeny, paleoclimate estimation and current niche modelling. We tested four predictions: (1) the climatic condition where the ancestor of Escallonia grew is megathermal; (2) the temperate niche is a derived condition from tropical clades; (3) the most closely related species have a similar current climate niche (conservation of the phylogenetic niche); and (4) there is a range expansion from the northern Andes to high latitudes during warm times. Our phylogenetic hypothesis shows that Escallonia originated 52.17 ± 0.85 My, in the early Eocene, with an annual mean temperature of 13.8 °C and annual precipitation of 1081 mm, corresponding to a microthermal to mesothermal climate; the species of the northern and central tropical Andes would be the ancestral ones, and the temperate species evolved between 32 and 20 My in a microthermal climate. The predominant evolutionary models were Brownian and Ornstein-Uhlenbeck. There was phylogenetic signal in 7 of the 9 variables, indicating conservation of the climatic niche. Escallonia would have originated in the central and southern Andes and reached the other environments by dispersion.

Bizarre tail weaponry in a transitional ankylosaur from subantarctic Chile.

Armoured dinosaurs are well known for their evolution of specialized tail weapons-paired tail spikes in stegosaurs and heavy tail clubs in advanced ankylosaurs1. Armoured dinosaurs from southern Gondwana are rare and enigmatic, but probably include the earliest branches of Ankylosauria2-4. Here we describe a mostly complete, semi-articulated skeleton of a small (approximately 2 m) armoured dinosaur from the late Cretaceous period of Magallanes in southernmost Chile, a region that is biogeographically related to West Antarctica5. Stegouros elengassen gen. et sp. nov. evolved a large tail weapon unlike any dinosaur: a flat, frond-like structure formed by seven pairs of laterally projecting osteoderms encasing the distal half of the tail. Stegouros shows ankylosaurian cranial characters, but a largely ancestral postcranial skeleton, with some stegosaur-like characters. Phylogenetic analyses placed Stegouros in Ankylosauria; specifically, it is related to Kunbarrasaurus from Australia6 and Antarctopelta from Antarctica7, forming a clade of Gondwanan ankylosaurs that split earliest from all other ankylosaurs. The large osteoderms and specialized tail vertebrae in Antarctopelta suggest that it had a tail weapon similar to Stegouros. We propose a new clade, the Parankylosauria, to include the first ancestor of Stegouros-but not Ankylosaurus-and all descendants of that ancestor.

Biogeography of Argylia D. Don (Bignoniaceae): Diversification, Andean Uplift and Niche Conservatism.

Andean uplift and the concomitant formation of the Diagonal Arid of South America is expected to have promoted species diversification through range expansions into this novel environment. We evaluate the evolution of Argylia, a genus belonging to the Bignoniaceae family whose oldest fossil record dates back to 49.4 Ma. Today, Argylia is distributed along the Andean Cordillera, from 15°S to 38.5°S and from sea level up to 4,000 m.a.s.l. We ask whether Argylia's current distribution is a result of a range expansion along the Andes Cordillera (biological corridor) modulated by climatic niche conservatism, considering the timing of Andean uplift (30 Ma - 5 Ma). To answer this question, we reconstructed the phylogenetic relationships of Argylia species, estimated divergence times, estimated the realized climatic niche of the genus, reconstructed the ancestral climatic niche, evaluated its evolution, and finally, performed an ancestral range reconstruction. We found strong evidence for climatic niche conservatism for moisture variables, and an absence of niche conservatism for most of the temperature variables considered. Exceptions were temperature seasonality and winter temperature. Results imply that Argylia had the capacity to adapt to extreme temperature conditions associated with the Andean uplift and the new climatic corridor produced by uplift. Ancestral range reconstruction for the genus showed that Argylia first diversified in a region where subtropical conditions were already established, and that later episodes of diversification were coeval with the of Andean uplift. We detected a second climatic corridor along the coastal range of Chile-Peru, the coastal lomas, which allowed a northward range expansion of Argylia into the hyperarid Atacama Desert. Dating suggests the current distribution and diversity of Argylia would have been reached during the Late Neogene and Pleistocene.

Divergence in Plant Traits and Increased Modularity Underlie Repeated Transitions Between Low and High Elevations in the Andean Genus Leucheria.

Understanding why some plant lineages move from one climatic region to another is a mayor goal of evolutionary biology. In the southern Andes plant lineages that have migrated along mountain ranges tracking cold-humid climates coexist with lineages that have shifted repeatedly between warm-arid at low elevations and cold habitats at high elevations. Transitions between habitats might be facilitated by the acquisition of common traits favoring a resource-conservative strategy that copes with drought resulting from either low precipitation or extreme cold. Alternatively, transitions might be accompanied by phenotypic divergence and accelerated evolution of plant traits, which in turn may depend on the level of coordination among them. Reduced integration and evolution of traits in modules are expected to increase evolutionary rates of traits, allowing diversification in contrasting climates. To examine these hypotheses, we conducted a comparative study in the herbaceous genus Leucheria. We reconstructed ancestral habitat states using Maximum Likelihood and a previously published phylogeny. We performed a Phylogenetic Principal Components Analysis on traits, and then we tested the relationship between PC axes, habitat and climate using Phylogenetic Generalized Least Squares (PGLS). Finally, we compared the evolutionary rates of traits, and the levels of modularity among the three main Clades of Leucheria. Our results suggest that the genus originated at high elevations and later repeatedly colonized arid-semiarid shrublands and humid-forest at lower elevations. PGLS analysis suggested that transitions between habitats were accompanied by shifts in plant strategies: cold habitats at high elevations favored the evolution of traits related to a conservative-resource strategy (thicker and dissected leaves, with high mass per area, and high biomass allocation to roots), whereas warm-arid habitats at lower elevations favored traits related to an acquisitive-resource strategy. As expected, we detected higher levels of modularity in the clades that switched repeatedly between habitats, but higher modularity was not associated with accelerated rates of trait evolution.

Genetic and phenotypic diversity in 2000 years old maize (Zea mays L.) samples from the Tarapacá region, Atacama Desert, Chile.

The evolution of maize (Zea mays L.) is highly controversial given the discrepancies related to the phenotypic and genetic changes suffered by the species, the incidence of human groups and the times in which these changes occurred. Also, morphological and genetic traits of crops are difficult to evaluate in the absence of fossils macro-botanical remains. In contrast in the Tarapacá region (18-21° S), Atacama Desert of Chile, prehispanic settlements (ca. 2500-400 yr BP) displayed extensive maize agriculture. The presence of archaeological macro-botanical remains of maize provided a unique opportunity to study the evolution of this crop, covering a temporal sequence of at least 2000 years. Thus, in this study, we ask how the morphological and genetic diversity of maize has varied since its introduction during prehispanic times in the Tarapacá region. To answer this, we measured and compared morphological traits of size and shape between archaeological cobs and kernels and 95 ears from landraces. To established genetic diversity eight microsatellite markers (SSR) were analyzed in archaeological and modern kernels. Genetic diversity was estimated by allelic frequency rates, the average number of alleles per locus, observed heterozygosity (Ho) and expected heterozygosity (He). Differences between populations and genetic structure were estimated by fixation index FST and STRUCTURE analysis. Our results indicate significant phenotypic differences and genetic distance between archaeological maize and landraces. This result is suggestive of an introduction of new varieties or drastic selective changes in modern times in Tarapacá. Additionally, archaeological maize shows a low genetic diversity and a progressive increase in the size of ears and kernels. These results suggest a human selection during prehispanic times and establish that prehispanic farmers played an important role in maize development. They also provide new clues for understanding the evolutionary history of maize in hyperarid conditions.

Evolutionary persistence in Gunnera and the contribution of southern plant groups to the tropical Andes biodiversity hotspot.

Several studies have demonstrated the contribution of northern immigrants to the flora of the tropical Andes-the world's richest and most diverse biodiversity hotspot. However, much less is known about the biogeographic history and diversification of Andean groups with southern origins, although it has been suggested that northern and southern groups have contributed roughly equally to the high Andean (i.e., páramo) flora. Here we infer the evolutionary history of the southern hemisphere plant genus Gunnera, a lineage with a rich fossil history and an important ecological role as an early colonising species characteristic of wet, montane environments. Our results show striking contrasts in species diversification, where some species may have persisted for some 90 million years, and whereas others date to less than 2 Ma since origination. The outstanding longevity of the group is likely linked to a high degree of niche conservatism across its highly disjunct range, whereby Gunnera tracks damp and boggy soils in cool habitats. Colonisation of the northern Andes is related to Quaternary climate change, with subsequent rapid diversification appearing to be driven by their ability to take advantage of environmental opportunities. This study demonstrates the composite origin of a mega-diverse biota.

Sensitivity of leaf size and shape to climate: global patterns and paleoclimatic applications.

• Paleobotanists have long used models based on leaf size and shape to reconstruct paleoclimate. However, most models incorporate a single variable or use traits that are not physiologically or functionally linked to climate, limiting their predictive power. Further, they often underestimate paleotemperature relative to other proxies. • Here we quantify leaf-climate correlations from 92 globally distributed, climatically diverse sites, and explore potential confounding factors. Multiple linear regression models for mean annual temperature (MAT) and mean annual precipitation (MAP) are developed and applied to nine well-studied fossil floras. • We find that leaves in cold climates typically have larger, more numerous teeth, and are more highly dissected. Leaf habit (deciduous vs evergreen), local water availability, and phylogenetic history all affect these relationships. Leaves in wet climates are larger and have fewer, smaller teeth. Our multivariate MAT and MAP models offer moderate improvements in precision over univariate approaches (± 4.0 vs 4.8°C for MAT) and strong improvements in accuracy. For example, our provisional MAT estimates for most North American fossil floras are considerably warmer and in better agreement with independent paleoclimate evidence. • Our study demonstrates that the inclusion of additional leaf traits that are functionally linked to climate improves paleoclimate reconstructions. This work also illustrates the need for better understanding of the impact of phylogeny and leaf habit on leaf-climate relationships.

A climatic and taxonomic comparison between leaf litter and standing vegetation from a Florida swamp woodland.

One method to determine past climate has been the use of leaf morphological characteristics of fossil leaves quantified using modern climate and canopy leaf characteristics. Fossil assemblages are composed of abscised leaves, and climate may be more accurately determined by using leaves from leaf litter instead of the canopy. To better understand whether taphonomic processes make a difference in this relationship, a north-central Florida woodland was sampled to determine the morphologically based climate estimates from these leaves. Leaves from woody, dicotyledonous plants were collected and identified, then compared using presence/absence data and analyzed using several linear regression equations and the CLAMP data set. Although the majority of standing vegetation was reflected in leaf litter, some inconsistencies were observed, which may reflect plant community structure or sampling technique. Mean annual temperature (MAT) and growing season precipitation (GSP) were estimated from leaf litter morphological characters and living leaves. Overall, values for MAT estimated from litter and living leaves were cooler than actual MATs, although several accurate and high estimates were obtained depending on the predictive method used. Estimated GSP values were higher than actual GSPs. Statistically, no difference was observed between MAT and GSP estimates derived from leaf litter vs. estimates derived from living leaves, with one exception.