Stable Heliconius butterfly hybrid zones are correlated with a local rainfall peak at the edge of the Amazon basin

Multilocus clines between Müllerian mimetic races of Heliconius butterflies provide a classic example of the maintenance of hybrid zones and their importance in speciation. Concordant hybrid zones in the mimics Heliconius erato and H. melpomene in northern Peru were carefully documented in the 1980s, and this prior work now permits a historical analysis of the movement or stasis of the zones. Previous work predicted that these zones might be moving toward the Andes due to selective asymmetry. Extensive deforestation and climate change might also be expected to affect the positions and widths of the hybrid zones. We show that the positions and shapes of these hybrid zones have instead remained remarkably stable between 1985 and 2012. The stability of this interaction strongly implicates continued selection, rather than neutral mixing following secondary contact. The stability of cline widths and strong linkage disequilibria (gametic correlation coefficients R_max = 0.35–0.56 among unlinked loci) over 25 years suggest that mimetic selection pressures on each color pattern locus have remained approximately constant (s ≈ 0.13–0.40 per locus in both species). Exceptionally high levels of precipitation at the edge of the easternmost Andes may act as a population density trough for butterflies, trapping the hybrid zones at the foot of the mountains, and preventing movement. As such, our results falsify one prediction of the Pleistocene Refugium theory: That the ranges of divergent species or subspecies should be centered on regions characterized by maxima of rainfall, with hybrid zones falling in more arid regions between them.     

The east‐west‐north colonization history of the Mediterranean and Europe by the coastal plant Carex extensa (Cyperaceae)

Coastal plants are ideal models for studying the colonization routes of species because of the simple linear distributions of these species. Carex extensa occurs mainly in salt marshes along the Mediterranean and European coasts. Variation in cpDNA sequences, amplified fragment length polymorphisms (AFLPs) and simple sequence repeats (SSRs) of 24 populations were analysed to reconstruct its colonization history. Phylogenetic relationships indicate that C. extensa together with the South American Carex vixdentata and the southern African Carex ecklonii form a monophyletic group of halophilic species. Analyses of divergence times suggest that early lineage diversification may have occurred between the late Miocene and the late Pliocene (Messinian crisis). Phylogenetic and network analyses of cpDNA variation revealed the monophyly of the species and an ancestral haplotype contained in populations of the eastern Mediterranean. The AFLP and SSR analyses support a pattern of variation compatible with these two lineages. These analyses also show higher levels of genetic diversity and differentiation in the eastern population group, which underwent an east‐to‐west Mediterranean colonization. Quaternary climatic oscillations appear to have been responsible for the split between these two lineages. Secondary contacts may have taken place in areas near the Ligurian Sea in agreement with the gene flow detected in Corsican populations. The AFLP and SSR data accord with the ‘tabula rasa’ hypothesis in which a recent and rapid colonization of northern Europe took place from the western Mediterranean after the Last Glacial Maximum. The unbalanced west‐east vs. west‐north colonization may be as a result of ‘high density blocking’ effect.     

The costal skeleton of Homo antecessor: preliminary results

The Lower Pleistocene TD6 level at the Gran Dolina site in the Sierra de Atapuerca (Burgos, Spain) has yielded nine ribs that represent a minimum of three individuals of the species, Homo antecessor. We present a detailed morphological and metric study of these costal elements, including the siding and anatomical position of all of the rib remains. The adult or nearly adult ribs are also metrically compared with other fossil hominins and with modern comparative samples. The costal elements recovered to date from the TD6 level at Gran Dolina can neither confirm nor reject the hypothesis that H. antecessor had a large thorax, similar to that of Neandertals. However, the fragmentary evidence of the H. antecessor thoracic skeleton is not inconsistent with this suggestion based on other skeletal elements, such as clavicles.

Resumen

En el nivel TD6 del Pleistoceno inferior del yacimiento de Gran Dolina, en la Sierra de Atapuerca (Burgos, España) se han recuperado nueve costillas que pertenecen a un mínimo de tres individuos de la especie Homo antecessor. Presentamos un detallado estudio métrico y morfológico incluyendo el lado y la determinación anatómica. Las costillas pertenecientes a individuos adultos o casi adultos también son comparadas métricamente a muestras modernas de comparación y otros homininos fósiles. Basándonos en el registro de costillas de Homo antecessor recuperado hasta el momento no podemos probar ni refutar la hipótesis de que esta especie presentaba un tórax grande similar al de los Neandertales. Sin embargo, el registro de costillas no es inconsistente con la hipótesis de un tórax grande como sugiere la gran longitud de sus clavículas.     

Phylogeography of Podocarpus matudae (Podocarpaceae): pre‐Quaternary relicts in northern Mesoamerican cloud forests

Aim Cloud forests of northern Mesoamerica represent the northern and southern limit of the contact zone between species otherwise characteristic of North or South America. Several phylogeographic studies featuring temperate conifer species have improved our understanding of species responses to environmental changes. In contrast, conifer species that presumably colonized northern Mesoamerica from South America are far less studied. A phylogeographic study of Podocarpus matudae (Podocarpaceae) was conducted to identify any major evolutionary divergences or disjunctions across its range and to determine if its current distribution is associated with pre‐Quaternary climatic and/or long‐distance dispersal events. Location Northern Mesoamerica (Mexico and Guatemala). Methods Sixteen populations (157 individuals) of P. matudae were screened for variation at two plastid DNA markers. The intra‐specific phylogenetic relationships among haplotypes were reconstructed using Bayesian inference. Population genetic analyses were undertaken to gain insight into the evolutionary history of these populations. To test whether genetic divergence among populations occurred at different time‐scales plastid DNA sequence data and fossil‐ and coalescent‐based calibrations were integrated. Results The combination of plastid markers yielded 11 haplotypes. Differentiation among populations based on DNA variation (G_ST) (0.707, SE 0.0807) indicated a clear population structure in P. matudae. Differentiation for ordered alleles (N_ST) (0.811, SE 0.0732) was higher than that for G_ST, indicating phylogeographical structure in P. matudae. Most of the total variation (81.3%, P < 0.0001) was explained by differences among populations. The estimated divergence time between the unique haplotypes from a Guatemalan population and the two most common haplotypes from the Sierra Madre Oriental in Mexico was between 10 and 20 Ma, and further haplotype divergence in the poorly resolved clade of the Sierra Madre Oriental occurred between 3 and 0.5 Ma. Main conclusions Divergence estimations support the hypothesis that extant Podocarpus matudae populations are pre‐Quaternary relicts. This finding is consistent with fossil and pollen data that support a Miocene age for temperate floristic elements in Mesoamerican cloud forests, whereas further haplotype divergence within the Sierra Madre Oriental, Chiapas and Guatemala occurred more recently, coinciding with Pleistocene cloud forest refugia.     

Glacial and interglacial refugia within a long-term rainforest refugium: The Wet Tropics Bioregion of NE Queensland, Australia

An artificial neural network is used to classify environments, including climate, terrain and soil variables, according to their suitability for fifteen structural/environmental forest classes in the Wet Tropics Bioregion of north-east Queensland. We map the environments characteristic of these forest classes in four climate regimes (the present and three past climate scenarios), quantify the changes in area of these environments in response to past regional changes in climate and identify areas that would have been environmentally suitable for rainforests at last glacial maximum (glacial refugia). We also identify areas that would have been suitable for upland and highland rainforest classes during the warmest parts of the interglacial (interglacial refugia) and map locations that consistently remain favourable to specific forest classes despite large changes in climate.In the climate of the last glacial maximum (LGM), rainforest environments are predicted in three relatively distinct refugia in the northern, central and southern Wet Tropics. Only three percent of the total area contains lowland, Mesophyll Vine Forest and the majority of the area of the rainforest refugia supports upland rainforest classes. In the cool, wet climate of the Pleistocene/Holocene transition (PHT), rainforest environments expand to form a more or less continuous block from the northern limits of the region to the Walter Hill Range, except for discontinuous patches extending through the Seaview and Paluma Ranges in the south. During the Holocene climatic optimum (HCO), rainforest environments become more fragmented, especially in the south. Lowland rainforest environments are very extensive in this climate while upland rainforest classes are restricted to what we term “interglacial refugia”.Estimated distributions and stable locations (consistently predicted in all four climate scenarios) for the various rainforest environment classes are our main, novel contribution. Each forest environment responds individualistically to climate change. Our results confirm the highly dynamic nature of the Wet Tropics landscape and present a much more detailed picture of landscape change since the late Pleistocene than previously has been available. This mapping exercise should be useful in the future for analyses of present-day biogeographic patterns. We argue that empirical modelling approaches have an important role in palaeoecology and global change research that is complementary to the developing mechanistic methods.