Trait evolution and historical biogeography shape assemblages of annual killifish

International audienceAim: Different species assemblages of annual killifish possess replicated body size distributions yet have unique sets of species in each area of endemism. Here, we use models of trait evolution and historical biogeography to discover how size variation originated and has been restructured.Location: South America.Taxon: Austrolebias (Cyprinodontiformes).Methods: We sampled 63 individuals from 26 Austrolebias species. Using phylogenetic trees (BEAST2), data on environmental variables at sampling locations and size data, we compare different models for trait evolution (SURFACE, l1OU) of body size and niche traits. We model the historical biogeography of the areas of endemism (BioGeoBEARS) and use both analyses in combination to reconstruct the history of four species assemblages.Results: We present new phylogenetic trees for Austrolebias and use them to show that large size principally arose within a single area driven by a shifted selection optimum for a subset of the species in that area. We suggest that ecological interactions triggered size divergence and that this large‐bodied lineage subsequently spread to two other areas. A second assemblage may have been shaped by adaptation to a new environment without an associated increase in size divergence. A third assemblage, which has the smallest size range and the most recent origin, is phylogenetically clustered, and we found no evidence of environmental filtering.Main conclusions: Assemblage similarity in Austrolebias is the result of contrasting ecological, evolutionary and historical processes. Modelling trait evolution together with historical biogeography can help to disentangle the complex histories of multispecies assemblages. This approach provides context to commonly used tests investigating the role of ecological processes from phylogenetic data and generates new testable hypotheses on the processes that generated trait diversity and assemblage similarit

Polyploidy on Islands: Its Emergence and Importance for Diversification.

Whole genome duplication or polyploidy is widespread among floras globally, but traditionally has been thought to have played a minor role in the evolution of island biodiversity, based on the low proportion of polyploid taxa present. We investigate five island systems (Juan Fernández, Galápagos, Canary Islands, Hawaiian Islands, and New Zealand) to test whether polyploidy (i) enhances or hinders diversification on islands and (ii) is an intrinsic feature of a lineage or an attribute that emerges in island environments. These island systems are diverse in their origins, geographic and latitudinal distributions, levels of plant species endemism (37% in the Galapagos to 88% in the Hawaiian Islands), and ploidy levels, and taken together are representative of islands more generally. We compiled data for vascular plants and summarized information for each genus on each island system, including the total number of species (native and endemic), generic endemicity, chromosome numbers, genome size, and ploidy levels. Dated phylogenies were used to infer lineage age, number of colonization events, and change in ploidy level relative to the non-island sister lineage. Using phylogenetic path analysis, we then tested how the diversification of endemic lineages varied with the direct and indirect effects of polyploidy (presence of polyploidy, time on island, polyploidization near colonization, colonizer pool size) and other lineage traits not associated with polyploidy (time on island, colonizer pool size, repeat colonization). Diploid and tetraploid were the most common ploidy levels across all islands, with the highest ploidy levels (>8x) recorded for the Canary Islands (12x) and New Zealand (20x). Overall, we found that endemic diversification of our focal island floras was shaped by polyploidy in many cases and certainly others still to be detected considering the lack of data in many lineages. Polyploid speciation on the islands was enhanced by a larger source of potential congeneric colonists and a change in ploidy level compared to overseas sister taxa

Polyploidy on Islands: Its Emergence and Importance for Diversification.

Whole genome duplication or polyploidy is widespread among floras globally, but traditionally has been thought to have played a minor role in the evolution of island biodiversity, based on the low proportion of polyploid taxa present. We investigate five island systems (Juan Fernández, Galápagos, Canary Islands, Hawaiian Islands, and New Zealand) to test whether polyploidy (i) enhances or hinders diversification on islands and (ii) is an intrinsic feature of a lineage or an attribute that emerges in island environments. These island systems are diverse in their origins, geographic and latitudinal distributions, levels of plant species endemism (37% in the Galapagos to 88% in the Hawaiian Islands), and ploidy levels, and taken together are representative of islands more generally. We compiled data for vascular plants and summarized information for each genus on each island system, including the total number of species (native and endemic), generic endemicity, chromosome numbers, genome size, and ploidy levels. Dated phylogenies were used to infer lineage age, number of colonization events, and change in ploidy level relative to the non-island sister lineage. Using phylogenetic path analysis, we then tested how the diversification of endemic lineages varied with the direct and indirect effects of polyploidy (presence of polyploidy, time on island, polyploidization near colonization, colonizer pool size) and other lineage traits not associated with polyploidy (time on island, colonizer pool size, repeat colonization). Diploid and tetraploid were the most common ploidy levels across all islands, with the highest ploidy levels (>8x) recorded for the Canary Islands (12x) and New Zealand (20x). Overall, we found that endemic diversification of our focal island floras was shaped by polyploidy in many cases and certainly others still to be detected considering the lack of data in many lineages. Polyploid speciation on the islands was enhanced by a larger source of potential congeneric colonists and a change in ploidy level compared to overseas sister taxa

Hard X-ray flux from low-mass stars in the Cygnus OB2 Association

Context. The Cygnus OB2 association, the central engine of the Cygnus X star-forming region, is the subject of an extensive INTEGRAL Key Project that will accumulate 6Ms of observations. Analysis of 2Ms of observations by De Becker and co-workers provides the most sensitive limit yet obtained on hard X-ray emission from the cluster. Aims. We investigate the X-ray emission in the 20-40 keV band expected from the flaring low-mass stellar population in Cygnus OB2. We discuss whether such emission needs to be considered in the interpretation of existing and future X-ray observations of the region, and whether such observations might provide insight into the high-energy processes on low-mass pre-main sequence stars. Methods. The total hard X-ray flux from low-mass stars is estimated by assuming the observed soft X-ray emission stems from a superposition of flares. We further assume the ratio of hard X-ray to soft X-ray emission is described by a scaling found for solar flares by Isola and co-workers. Results. We estimate the low-mass stellar hard X-ray flux in the 20-40 keV band to lie in the range ~2x10^31-6x10^32 erg/s and discuss some potential biases that might affect this result. Conclusions. Hard X-ray emission could lie at a level not much below the current observed flux upper limits for Cygnus OB2. If this emission could be detected, it would provide insight into the hard X-ray production of large flares on pre-main sequence stars. We highlight the penetrating power of hard X-rays from low-mass stellar populations as a possible pointer to our Galaxy's hidden star-forming clusters and super-clusters using more sensitive observations from future missions.Comment: 5 page

Inside-Out Evacuation of Transitional Protoplanetary Disks by the Magneto-Rotational Instability

How do T Tauri disks accrete? The magneto-rotational instability (MRI) supplies one means, but protoplanetary disk gas is typically too poorly ionized to be magnetically active. Here we show that the MRI can, in fact, explain observed accretion rates for the sub-class of T Tauri disks known as transitional systems. Transitional disks are swept clean of dust inside rim radii of ~10 AU. Stellar coronal X-rays ionize material in the disk rim, activating the MRI there. Gas flows from the rim to the star, at a rate limited by the depth to which X-rays ionize the rim wall. The wider the rim, the larger the surface area that the rim wall exposes to X-rays, and the greater the accretion rate. Interior to the rim, the MRI continues to transport gas; the MRI is sustained even at the disk midplane by super-keV X-rays that Compton scatter down from the disk surface. Accretion is therefore steady inside the rim. Blown out by radiation pressure, dust largely fails to accrete with gas. Contrary to what is usually assumed, ambipolar diffusion, not Ohmic dissipation, limits how much gas is MRI-active. We infer values for the transport parameter alpha on the order of 0.01 for GM Aur, TW Hyd, and DM Tau. Because the MRI can only afflict a finite radial column of gas at the rim, disk properties inside the rim are insensitive to those outside. Thus our picture provides one robust setting for planet-disk interaction: a protoplanet interior to the rim will interact with gas whose density, temperature, and transport properties are definite and decoupled from uncertain initial conditions. Our study also supplies half the answer to how disks dissipate: the inner disk drains from the inside out by the MRI, while the outer disk photoevaporates by stellar ultraviolet radiation.Comment: Accepted to Nature Physics June 7, 2007. The manuscript for publication is embargoed per Nature policy. This arxiv.org version contains more technical details and discussion, and is distributed with permission from the editors. 10 pages, 4 figure

Empirical Constraints on Turbulence in Protoplanetary Accretion Disks

We present arcsecond-scale Submillimeter Array observations of the CO(3-2) line emission from the disks around the young stars HD 163296 and TW Hya at a spectral resolution of 44 m/s. These observations probe below the ~100 m/s turbulent linewidth inferred from lower-resolution observations, and allow us to place constraints on the turbulent linewidth in the disk atmospheres. We reproduce the observed CO(3-2) emission using two physical models of disk structure: (1) a power-law temperature distribution with a tapered density distribution following a simple functional form for an evolving accretion disk, and (2) the radiative transfer models developed by D'Alessio et al. that can reproduce the dust emission probed by the spectral energy distribution. Both types of models yield a low upper limit on the turbulent linewidth (Doppler b-parameter) in the TW Hya system (<40 m/s), and a tentative (3-sigma) detection of a ~300 m/s turbulent linewidth in the upper layers of the HD 163296 disk. These correspond to roughly <10% and 40% of the sound speed at size scales commensurate with the resolution of the data. The derived linewidths imply a turbulent viscosity coefficient, alpha, of order 0.01 and provide observational support for theoretical predictions of subsonic turbulence in protoplanetary accretion disks.Comment: 18 pages, 9 figures, accepted for publication in Ap

Surface magnetic fields on two accreting T Tauri stars: CV Cha and CR Cha

We have produced brightness and magnetic field maps of the surfaces of CV Cha and CR Cha: two actively accreting G and K-type T Tauri stars in the Chamaeleon I star-forming cloud with ages of 3-5 Myr. Our magnetic field maps show evidence for strong, complex multi-polar fields similar to those obtained for young rapidly rotating main sequence stars. Brightness maps indicate the presence of dark polar caps and low latitude spots -- these brightness maps are very similar to those obtained for other pre-main sequence and rapidly rotating main sequence stars. Only two other classical T Tauri stars have been studied using similar techniques so far: V2129 Oph and BP Tau. CV Cha and CR Cha show magnetic field patterns that are significantly more complex than those recovered for BP Tau, a fully convective T Tauri star. We discuss possible reasons for this difference and suggest that the complexity of the stellar magnetic field is related to the convection zone; with more complex fields being found in T Tauri stars with radiative cores (V2129 Oph, CV Cha and CR Cha). However, it is clearly necessary to conduct magnetic field studies of T Tauri star systems, exploring a wide range of stellar parameters in order to establish how they affect magnetic field generation, and thus how these magnetic fields are likely to affect the evolution of T Tauri star systems as they approach the main sequence.Comment: Accepted for publication by MNRAS: 15 pages, 11 figure

Magnetic fields in protoplanetary disks

Magnetic fields likely play a key role in the dynamics and evolution of protoplanetary discs. They have the potential to efficiently transport angular momentum by MHD turbulence or via the magnetocentrifugal acceleration of outflows from the disk surface, and magnetically-driven mixing has implications for disk chemistry and evolution of the grain population. However, the weak ionisation of protoplanetary discs means that magnetic fields may not be able to effectively couple to the matter. I present calculations of the ionisation equilibrium and magnetic diffusivity as a function of height from the disk midplane at radii of 1 and 5 AU. Dust grains tend to suppress magnetic coupling by soaking up electrons and ions from the gas phase and reducing the conductivity of the gas by many orders of magnitude. However, once grains have grown to a few microns in size their effect starts to wane and magnetic fields can begin to couple to the gas even at the disk midplane. Because ions are generally decoupled from the magnetic field by neutral collisions while electrons are not, the Hall effect tends to dominate the diffusion of the magnetic field when it is able to partially couple to the gas. For a standard population of 0.1 micron grains the active surface layers have a combined column of about 2 g/cm^2 at 1 AU; by the time grains have aggregated to 3 microns the active surface density is 80 g/cm^2. In the absence of grains, x-rays maintain magnetic coupling to 10% of the disk material at 1 AU (150 g/cm^2). At 5 AU the entire disk thickness becomes active once grains have aggregated to 1 micron in size.Comment: 11 pages, 11 figs, aastex.cls. Accepted for publication in Astrophysics & Space Science. v3 corrects bibliograph

Trapping Solids at the Inner Edge of the Dead Zone: 3-D Global MHD Simulations

The poorly-ionized interior of the protoplanetary disk is the location where dust coagulation processes may be most efficient. However even here, planetesimal formation may be limited by the loss of solid material through radial drift, and by collisional fragmentation of the particles. Our aim is to investigate the possibility that solid particles are trapped at local pressure maxima in the dynamically evolving disk. We perform the first 3-D global non-ideal MHD calculations of the disk treating the turbulence driven by the magneto-rotational instability. The domain contains an inner MRI-active region near the young star and an outer midplane dead zone, with the transition between the two modeled by a sharp increase in the magnetic diffusivity. The azimuthal magnetic fields generated in the active zone oscillate over time, changing sign about every 150 years. We thus observe the radial structure of the `butterfly pattern' seen previously in local shearing-box simulations. The mean magnetic field diffuses from the active zone into the dead zone, where the Reynolds stress nevertheless dominates. The greater total accretion stress in the active zone leads to a net reduction in the surface density, so that after 800 years an approximate steady state is reached in which a local radial maximum in the midplane pressure lies near the transition radius. We also observe the formation of density ridges within the active zone. The dead zone in our models possesses a mean magnetic field, significant Reynolds stresses and a steady local pressure maximum at the inner edge, where the outward migration of planetary embryos and the efficient trapping of solid material are possible.Comment: 17 pages, 30 *.ps files for figures. Accepted 16 November 2009 in A&