Infraspecific diversification of the star cloak fern (Notholaena standleyi) in the deserts of the United States and Mexico.

PREMISE: Not all ferns grow in moist and shaded habitats. One well-known example is Notholaena standleyi, a species that thrives in deserts of the southwestern United States and Mexico. This species exhibits several "chemotypes" that differ in farina (flavonoid exudates) color and chemistry. By integrating data from molecular phylogenetics, cytology, biochemistry, and biogeography, we circumscribed the major evolutionary lineages within N. standleyi and reconstructed their diversification histories. METHODS: Forty-eight samples were selected from across the geographic distribution of N. standleyi. Phylogenetic relationships were inferred using four plastid and five nuclear markers. Ploidy levels were inferred using spore sizes calibrated by chromosome counts, and farina chemistry was compared using thin-layer chromatography. RESULTS: Four clades are recognized, three of which roughly correspond to previously recognized chemotypes. The diploid clades G and Y are found in the Sonoran and Chihuahuan deserts, respectively; they are estimated to have diverged in the Pleistocene, congruent with the postulated timing of climatological events separating these two deserts. Clade P/YG is tetraploid and partially overlaps the distribution of clade Y in the eastern Chihuahuan Desert. It is apparently confined to limestone, a geologic substrate rarely occupied by members of the other clades. The cryptic (C) clade, a diploid group known only from southern Mexico and highly disjunct from the other three clades, is newly recognized here. CONCLUSIONS: Our results reveal a complex intraspecific diversification history of N. standleyi, traceable to a variety of evolutionary drivers including classic allopatry, parapatry with or without changes in geologic substrate, and sympatric divergence through polyploidization.

Regional signatures of plant response to drought and elevated temperature across a desert ecosystem.

The performance of many desert plant species in North America may decline with the warmer and drier conditions predicted by climate change models, thereby accelerating land degradation and reducing ecosystem productivity. We paired repeat measurements of plant canopy cover with climate at multiple sites across the Chihuahuan Desert over the last century to determine which plant species and functional types may be the most sensitive to climate change. We found that the dominant perennial grass, Bouteloua eriopoda, and species richness had nonlinear responses to summer precipitation, decreasing more in dry summers than increasing with wet summers. Dominant shrub species responded differently to the seasonality of precipitation and drought, but winter precipitation best explained changes in the cover of woody vegetation in upland grasslands and may contribute to woody-plant encroachment that is widespread throughout the southwestern United States and northern Mexico. Temperature explained additional variability of changes in cover of dominant and subdominant plant species. Using a novel empirically based approach we identified "climate pivot points" that were indicative of shifts from increasing to decreasing plant cover over a range of climatic conditions. Reductions in cover of annual and several perennial plant species, in addition to declines in species richness below the long-term summer precipitation mean across plant communities, indicate a decrease in the productivity for all but the most drought-tolerant perennial grasses and shrubs in the Chihuahuan Desert. Overall, our regional synthesis of long-term data provides a robust foundation for forecasting future shifts in the composition and structure of plant assemblages in the largest North American warm desert.

Molecular analysis of stomach contents reveals important grass seeds in the winter diet of Baird's and Grasshopper sparrows, two declining grassland bird species.

We analyzed the diet of Baird's Sparrow (Ammodramus bairdii) and Grasshopper Sparrow (A. savannarum) in three different sites and sampling periods across the Chihuahuan Desert in northern Mexico. DNA from seeds in regurgitated stomach contents was sequenced using NGS technology and identified with a barcoding approach using the P6 loop of the trnL intron as genetic marker. During each sampling period, we collected random soil samples to estimate seed availability in the soil seed bank. Due to the variability and size of the genetic marker, the resolution was limited to a family level resolution for taxonomic classification of seeds, but in several cases a genus level was achieved. Diets contained a high diversity of seeds but were dominated by a limited number of genera/families. Seeds from Panicoideae (from the genera Panicum, Setaria, Eriochloa, Botriochloa, and Hackelochloa) contributed for the largest part to the diets (53 ± 19%), followed by Bouteloua (10 ± 12%). Depending on the site and sampling period, other important seeds in the diets were Eragrostideae, Pleuraphis, Asteraceae, Verbena, and Amaranthus. The most abundant seeds were not always preferred. Aristida and Chloris were common in the soil seed bank but these seeds were avoided by both bird species. Baird's and Grasshopper sparrows did not differ in seed preferences. This work highlights the importance of range management practices that favor seed production of Panicoideae and Bouteloua grasses to enhance winter habitat use and survival of Baird's and Grasshopper sparrows in the Chihuahuan Desert.