Edaphic adaptation maintains the coexistence of two cryptic species on serpentine soil

• Premise of the study: Divergent edaphic adaptation can contribute to reproductive isolation and coexistence between closely related species, yet we know little about how small-scale continuous edaphic gradients contribute to this phenomenon. We investigated edaphic adaptation between two cryptic species of California wildflower, Lasthenia californica and L. gracilis (Asteraceae), which grow in close parapatry on serpentine soil. • Methods: We reciprocally transplanted both species into the center of each species’ habitat and the transition zone between species. We quantified multiple components of fitness and used aster models to predict fitness based on environmental variables. We sampled soil across the ridge throughout the growing season to document edaphic changes through time. We sampled naturally germinating seedlings to determine whether there was dispersal into the adjacent habitat and to help pinpoint the timing of any selection against migrants. • Key results: We documented within-serpentine adaptation contributing to habitat isolation between close relatives. Both species were adapted to the edaphic conditions in their native region and suffered fitness trade-offs when moved outside that region. However, observed fitness values did not perfectly match those predicted by edaphic variables alone, indicating that other factors, such as competition, also contributed to plant fitness. Soil water content and concentrations of calcium, magnesium, sodium, and potassium were likely drivers of differential fitness. Plants either had limited dispersal ability or migrants experienced early-season mortality outside their native region. • Conclusions: Demonstrating that continuous habitats can support differently adapted, yet closely related, taxa is important to a broader understanding of how species are generated and maintained in nature

Early Developmental Responses to Seedling Environment Modulate Later Plasticity to Light Spectral Quality

Correlations between developmentally plastic traits may constrain the joint evolution of traits. In plants, both seedling de-etiolation and shade avoidance elongation responses to crowding and foliage shade are mediated by partially overlapping developmental pathways, suggesting the possibility of pleiotropic constraints. To test for such constraints, we exposed inbred lines of Impatiens capensis to factorial combinations of leaf litter (which affects de-etiolation) and simulated foliage shade (which affects phytochrome-mediated shade avoidance). Increased elongation of hypocotyls caused by leaf litter phenotypically enhanced subsequent elongation of the first internode in response to low red∶far red (R∶FR). Trait expression was correlated across litter and shade conditions, suggesting that phenotypic effects of early plasticity on later plasticity may affect variation in elongation traits available to selection in different light environments

Mutualism and Adaptive Divergence: Co-Invasion of a Heterogeneous Grassland by an Exotic Legume-Rhizobium Symbiosis

Species interactions play a critical role in biological invasions. For example, exotic plant and microbe mutualists can facilitate each other's spread as they co-invade novel ranges. Environmental context may influence the effect of mutualisms on invasions in heterogeneous environments, however these effects are poorly understood. We examined the mutualism between the legume, Medicago polymorpha, and the rhizobium, Ensifer medicae, which have both invaded California grasslands. Many of these invaded grasslands are composed of a patchwork of harsh serpentine and relatively benign non-serpentine soils. We grew legume genotypes collected from serpentine or non-serpentine soil in both types of soil in combination with rhizobium genotypes from serpentine or non-serpentine soils and in the absence of rhizobia. Legumes invested more strongly in the mutualism in the home soil type and trends in fitness suggested that this ecotypic divergence was adaptive. Serpentine legumes had greater allocation to symbiotic root nodules in serpentine soil than did non-serpentine legumes and non-serpentine legumes had greater allocation to nodules in non-serpentine soil than did serpentine legumes. Therefore, this invasive legume has undergone the rapid evolution of divergence for soil-specific investment in the mutualism. Contrary to theoretical expectations, the mutualism was less beneficial for legumes grown on the stressful serpentine soil than on the non-serpentine soil, possibly due to the inhibitory effects of serpentine on the benefits derived from the interaction. The soil-specific ability to allocate to a robust microbial mutualism may be a critical, and previously overlooked, adaptation for plants adapting to heterogeneous environments during invasion

Appendix A. Aerial photograph of “the grid” at the University of California McLaughlin Reserve with grid points overlaid onto the landscape.

Aerial photograph of “the grid” at the University of California McLaughlin Reserve with grid points overlaid onto the landscape

Flowering phenology shifts in response to biodiversity loss

Observational studies and experimental evidence agree that rising global temperatures have altered plant phenology—the timing of life events, such as flowering, germination, and leaf-out. Other large-scale global environmental changes, such as nitrogen deposition and altered precipitation regimes, have also been linked to changes in flowering times. Despite our increased understanding of how abiotic factors influence plant phenology, we know very little about how biotic interactions can affect flowering times, a significant knowledge gap given ongoing human-caused alteration of biodiversity and plant community structure at the global scale. We experimentally manipulated plant diversity in a California serpentine grassland and found that many plant species flowered earlier in response to reductions in diversity, with peak flowering date advancing an average of 0.6 days per species lost. These changes in phenology were mediated by the effects of plant diversity on soil surface temperature, available soil N, and soil moisture. Peak flowering dates were also more dispersed among species in high-diversity plots than expected based on monocultures. Our findings illustrate that shifts in plant species composition and diversity can alter the timing and distribution of flowering events, and that these changes to phenology are similar in magnitude to effects induced by climate change. Declining diversity could thus contribute to or exacerbate phenological changes attributed to rising global temperatures

Potential for adaptive evolution at species range margins: Contrasting interactions between red coral populations and their environment in a changing ocean

15 pages, 4 figures, 2 tables, supporting information http://onlinelibrary.wiley.com/doi/10.1002/ece3.1324/suppinfoStudying population-by-environment interactions (PEIs) at species range margins offers the opportunity to characterize the responses of populations facing an extreme regime of selection, as expected due to global change. Nevertheless, the importance of these marginal populations as putative reservoirs of adaptive genetic variation has scarcely been considered in conservation biology. This is particularly true in marine ecosystems for which the deep refugia hypothesis proposes that disturbed shallow and marginal populations of a given species can be replenished by mesophotic ones. This hypothesis therefore assumes that identical PEIs exist between populations, neglecting the potential for adaptation at species range margins. Here, we combine reciprocal transplant and common garden experiments with population genetics analyses to decipher the PEIs in the red coral, Corallium rubrum. Our analyses reveal partially contrasting PEIs between shallow and mesophotic populations separated by approximately one hundred meters, suggesting that red coral populations may potentially be locally adapted to their environment. Based on the effective population size and connectivity analyses, we posit that genetic drift may be more important than gene flow in the adaptation of the red coral. We further investigate how adaptive divergence could impact population viability in the context of warming and demonstrate differential phenotypic buffering capacities against thermal stress. Our study questions the relevance of the deep refugia hypothesis and highlights the conservation value of marginal populations as a putative reservoir of adaptive genetic polymorphism. In marine conservation, the deep-refugia hypothesis proposes that disturbed shallow and marginal populations of a given species can be replenished by mesophotic populations. Combining reciprocal transplant and common garden experiments with population genetics analyses we question the relevance of this hypothesis in the red coral, Corallium rubrum. Our study highlights the conservation value of marginal populations as a putative reservoir of adaptive genetic polymorphism. © 2015 Published by John Wiley & Sons Ltd.This study was funded by the “Provence–Alpes–Côte d'Azur” region (J-B. Ledoux Doctoral Fellowship) and by the IFB Foundation (Institut Français de la Biodiversité), the environmental department of TOTAL-FINA-ELF group, an “Action Concertée Initiative” (ACI) program “Jeunes chercheurs 2003” (number JC5360) from the French Research Ministry and the MEDCHANGE project funded by the Agence Nationale pour la Recherche (ANR) J-B. L. is supported by a post-doctoral grant SFRH/BPD/74400/2010 from Fundacão para a Ciência e Tecnologia (FCT; Portugal)Peer Reviewe