Community assembly of the ferns of Florida.

PREMISE OF THE STUDY: Many ecological and evolutionary processes shape the assembly of organisms into local communities from a regional pool of species. We analyzed phylogenetic and functional diversity to understand community assembly of the ferns of Florida at two spatial scales. METHODS: We built a phylogeny for 125 of the 141 species of ferns in Florida using five chloroplast markers. We calculated mean pairwise dissimilarity (MPD) and mean nearest taxon distance (MNTD) from phylogenetic distances and functional trait data for both spatial scales and compared the results to null models to assess significance. KEY RESULTS: Our results for over vs. underdispersion in functional and phylogenetic diversity differed depending on spatial scale and metric considered. At the county scale, MPD revealed evidence for phylogenetic overdispersion, while MNTD revealed phylogenetic and functional underdispersion, and at the conservation area scale, MPD revealed phylogenetic and functional underdispersion while MNTD revealed evidence only of functional underdispersion. CONCLUSIONS: Our results are consistent with environmental filtering playing a larger role at the smaller, conservation area scale. The smaller spatial units are likely composed of fewer local habitat types that are selecting for closely related species, with the larger-scale units more likely to be composed of multiple habitat types that bring together a larger pool of species from across the phylogeny. Several aspects of fern biology, including their unique physiology and water relations and the importance of the independent gametophyte stage of the life cycle, make ferns highly sensitive to local, microhabitat conditions.

Into Africa: Molecular phylogenetics and historical biogeography of sub-Saharan African woodferns (Dryopteris).

PREMISE OF THE STUDY: Our goal was to infer the phylogenetic relationships and historical biogeography of the genus Dryopteris with a focus on taxa in sub-Saharan Africa and neighboring islands. In general, little is known about the relationships between African fern species and their congeners in other geographic regions, and our aim was to determine whether the sub-Saharan African species of Dryopteris are monophyletic and evolved within Africa or arrived there via repeated dispersals into Africa from other regions. METHODS: We obtained sequence data for five chloroplast markers from 214 species of Dryopteris and 18 outgroups. We performed phylogenetic and molecular dating analyses using a Bayesian relaxed clock method in BEAST with fossil and secondary calibration points and estimated ancestral ranges for the genus globally by comparing multiple models in BioGeoBEARS. KEY RESULTS: We found that 22 of 27 accessions of sub-Saharan African Dryopteris belong to a large clade of 31 accessions that also includes taxa from Indian and Atlantic Ocean islands. Additional accessions of taxa from our regions of interest have Asian, Hawaiian, European, or North American species as their closest relatives. CONCLUSIONS: The majority of sub-Saharan African Dryopteris species are descended from a shared common ancestor that dispersed to Africa from Asia approximately 10 Ma. There have been subsequent dispersal events from the African mainland to islands in the Atlantic and Indian Oceans, including Madagascar. Several additional species are estimated to have descended from ancestors that reached Africa via separate events over the last roughly 20 million years.

Differences in dehydration tolerance among populations of a gametophyte-only fern.

PREMISE OF THE STUDY: For many plant species, historical climatic conditions may have left lasting imprints that are detectable in contemporary populations. Additionally, if these historical conditions also prevented gene flow among populations, these populations may be differentiated with respect to one another and their contemporary environmental conditions. For the fern, Vittaria appalachiana, one theory is that historical conditions during the Pleistocene largely shaped both the distribution and lack of sporophyte production. Our goals-based on this theory-were to examine physiological differences among and within populations spanning the species' geographic range, and the contribution of historical climatic conditions to this differentiation. METHODS: We exposed explants from five populations to four drying treatments and examined differences in physiological response. Additionally, we examined the role of historical and current climatic conditions in driving the observed population differentiation. KEY RESULTS: Populations differ in their ability to tolerate varying levels of dehydration, displaying a pattern of countergradient selection. Exposure to historical and contemporary climatic conditions, specifically variation in temperature and precipitation regimes, resulted in population divergence observed among contemporary populations. CONCLUSIONS: Historical conditions have shaped not only the distribution of V. appalachiana, but also its current physiological limitations. Results from this study support the hypothesis that climatic conditions during the Pleistocene are responsible for the distribution of this species, and may be responsible for the observed differences in dehydration tolerance. Additionally, dehydration tolerance may be the driving factor for previously reported patterns of countergradient selection in this species.

Population differentiation and countergradient variation throughout the geographic range in the fern gametophyte Vittaria appalachiana.

PREMISE OF THE STUDY: Theory predicts that limited gene flow between populations will promote population differentiation, and experimental studies have found that differentiation is often explained by local adaptation in sexually reproducing angiosperms. However, few experiments have examined the drivers of differentiation among populations in asexual land plants with limited dispersal potential. Here, we evaluated the role of temperature in driving population differentiation in an asexual, obligate gametophyte fern species. METHODS: We reciprocally transplanted Vittaria appalachiana gametophytes among six populations that spanned the species' geographic range in the Appalachian Mountains and Plateau. Temperature, survival, and senescence rates were measured for 1 year. KEY RESULTS: Populations had significantly different fitness responses to different sites, consistent with the hypothesis that populations have differentiated across the species' range. There was some evidence for local adaptation in marginal populations and for countergradient selection favoring particularly robust genotypes at the northern range edge. Most populations had relatively high fitness at the site with the most stable temperature conditions and were negatively affected by decreasing minimum temperatures. CONCLUSIONS: Populations of Vittaria appalachiana exhibit highly variable responses to transplantation across the species' range, and only a small subset of these responses are due to local adaptation. Differences in daily minimum temperature explain some variation in fitness, but other site-specific factors also have significant impacts on transplant fitness. These results indicate that asexual, patchily distributed species with limited dispersal may exhibit population-specific responses to global climate change that have not been elucidated by empirical work focused on sexually reproducing angiosperms.

Identifying cryptic fern gametophytes using DNA barcoding: A review.

Ferns and lycophytes are unique among land plants in having sporophyte (diploid) and gametophyte (haploid) generations that can grow independently of each other. While most studies of fern ecology focus on the more visible sporophytic stage, the gametophyte is critically important, as it is the sexual phase of the life cycle. Yet, fern gametophytes have long been neglected in field studies due to their small size and cryptic morphology. DNA barcoding is a powerful method that can be used to identify field-collected gametophytes to species and allow for detailed study of their ecology. Here, we review the state of DNA barcoding as applied to fern gametophytes. First, we trace the history of DNA barcoding and how it has come to be applied to fern gametophytes. Next, we summarize case studies that show how DNA barcoding has been used to better understand fern species distributions, gametophyte ecology, and community ecology. Finally, we propose avenues for future research using this powerful tool, including next-generation DNA sequencing for in-field identification of cryptic gametophytes.

The ecology and physiology of fern gametophytes: A methodological synthesis.

All green plants alternate between the gametophyte and sporophyte life stages, but only seed-free vascular plants (ferns and lycophytes) have independent, free-living gametophytes. Fern and lycophyte gametophytes are significantly reduced in size and morphological complexity relative to their sporophytic counterparts and have often been overlooked in ecological and physiological studies. Understanding the ecological and physiological factors that directly impact this life stage is of critical importance because the ultimate existence of a sporophyte is dependent upon successful fertilization in the gametophyte generation. Furthermore, previous research has shown that the dual nature of the life cycle and the high dispersibility of spores can result in different geographic patterns between gametophytes and their respective sporophytes. This variation in distribution patterns likely exacerbates the separation of selective pressures acting on gametophyte and sporophyte generations, and can uniquely impact a species' ecology and physiology. Here, we provide a review of historical and contemporary methodologies used to examine ecological and physiological aspects of fern gametophytes, as well as those that allow for comparisons between the two generations. We conclude by suggesting methodological approaches to answer currently outstanding questions. We hope that the information covered herein will serve as a guide to current researchers and stimulate future discoveries in fern gametophyte ecology and physiology.

Conserved thermal performance curves across the geographic range of a gametophytic fern.

Species-level responses to environmental change depend on the collective responses of their constituent populations and the degree to which populations are specialized to local conditions. Manipulative experiments in common-garden settings make it possible to test for population variation in species' responses to specific climate variables, including those projected to shift as the climate changes in the future. While this approach is being applied to a variety of plant taxa to evaluate their responses to climate change, these studies are heavily biased towards seed-bearing plant species. Given several unique morphological and physiological traits, fern species may exhibit very different responses from angiosperms and gymnosperms. Here, we tested the hypothesis that previously detected population differentiation in a fern species is due to differentiation in thermal performance curves among populations. We collected explants from six populations spanning the species' geographic range and exposed them to 10 temperature treatments. Explant survival, lifespan and the change in photosynthetic area were analysed as a function of temperature, source population and their interaction. Overall results indicated that explants performed better at the lowest temperature examined, and the threshold for explant performance reflects maximum temperatures likely to be experienced in the field. Surprisingly, explant fitness did not differ among source populations, suggesting that temperature is not the driver behind previously detected patterns of population differentiation. These results highlight the importance of other environmental axes in driving population differentiation across a species range, and suggest that the perennial life history strategy, asexual mating system and limited dispersal potential of Vittaria appalachiana may restrict the rise and differentiation of adaptive genetic variation in thermal performance traits among populations.