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The emergence of angiosperm-dominated tropical forests in the Cretaceous led to major shifts in the composition of biodiversity on Earth. Among these was the rise to prominence of epiphytic plant lineages, which today comprise an estimated one-quarter of tropical vascular plant diversity. Among the most successful epiphytic groups is the Polypodiaceae, which comprises an estimated 1500 species and displays a remarkable breadth of morphological and ecological diversity. Using a time-calibrated phylogeny for 417 species, we characterized macroevolutionary patterns in the family, identified shifts in diversification rate, and identified traits that are potential drivers of diversification. We find high diversification rates throughout the family, evidence for a radiation in a large clade of Paleotropical species, and support for increased rates of diversification associated with traits including chlorophyllous spores and noncordiform gametophytes. Contrary to previous hypotheses, our results indicate epiphytic species and groups with humus-collecting leaves diversify at lower rates than the family as a whole. We find that diversification rates in the Polypodiaceae are positively correlated with changes in elevation. Repeated successful exploration of novel habitat types, rather than morphological innovation, appears to be the primary driver of diversification in this group.

Homosporous vascular plants utilize three different mating systems, one of which, gametophytic selfing, is an extreme form of inbreeding only possible in homosporous groups. This mating system results in complete homozygosity in all progeny and has important evolutionary and ecological implications. Ferns are the largest group of homosporous land plants, and the significance of extreme inbreeding for fern evolution has been a subject of debate for decades. We cultured gametophytes in the laboratory and quantified the relative frequencies of sporophyte production from isolated and paired gametophytes, and examined associations between breeding systems and several ecological and evolutionary traits. The majority of fern species studied show a capacity for gametophytic selfing, producing sporophytes from both isolated and paired gametophytes. While we did not follow sporophytes to maturity to investigate potential detrimental effects of homozygosity at later developmental stages, our results suggest that gametophytic selfing may have greater significance for fern evolution and diversification than has previously been realized. We present evidence from the largest study of mating behavior in ferns to date that the capacity for extreme inbreeding is prevalent in this lineage, and we discuss its implications and relevance and make recommendations for future studies of fern mating systems.

Hybridization is an important evolutionary force in plants, but the mechanisms underlying it have not been well studied for many groups. In particular, the drivers of non‐random patterns of interspecific gene flow (asymmetrical hybridization) remain poorly understood, especially in the seed‐free vascular plants. Here, we examine patterns of asymmetrical hybridization in two widespread fern hybrids from eastern North America and study the role of gametophyte ecology in the determination of hybridization bias. We characterized the maternal parentage of > 140 hybrid sporophytes by sequencing a c. 350‐bp region of chloroplast DNA (cpDNA). To identify factors contributing to patterns of asymmetrical hybridization, we cultured gametophytes of the parental species and evaluated critical aspects of their reproductive biology. We found that asymmetrical hybridization was prevalent across the populations of both hybrids. Reproductive traits varied across species and suggest that selfing potential, antheridiogen responsiveness, sperm dispersal capacity and gamete size all contribute to the mediation of the direction of hybridization in this group. Our findings suggest that asymmetrical hybridization in ferns is driven by an array of reproductive traits. This study helps to sharpen and define a mechanistic understanding of patterns of hybridization in this group and demonstrates the importance of considering gametophyte biology when studying evolutionary processes in ferns.

• Tropical mountains are disproportionately biodiverse relative to their surface area, but the processes underlying their exceptional diversity require further study. Here, we use comparative phylogenetic methods to examine the impact of the Andean orogeny on the diversification of Neotropical Phlegmariurus, a species-rich lycophyte clade. • We generated a time-calibrated phylogeny of 105 species of Neotropical Phlegmariurus and estimated lineage diversification rates. We tested for correlations between lineage diversification rates and species range size, niche breadth, elevational range amplitude, and mean elevation of occurrence. A recently developed macroevolutionary model was used to incorporate geological data and test for an association between diversification rates and the Andean uplift. • Diversification rates of Neotropical Phlegmariurus are negatively correlated with species range size and positively correlated with mean elevation of species occurrence. The rise of the Andes is strongly associated with increased rates of diversification in Neotropical Phlegmariurus during the last 10 Myr. • Our study demonstrates the importance of mountain-building events and geographical isolation of alpine populations as drivers of rapid diversification, even in spore-dispersed plants. This work also highlights the usefulness of combined phylogenetic, geological and ecological datasets, and the promise of comparative environment-dependent diversification models in better understanding the evolutionary origins of biodiversity.

Parablechnum is the most diverse genus in the fern family Blechnaceae, with about 70 species, mainly from Central and South America, the Austropacific, and a few in Africa. Species delimitation in Parablechnum is challenging, and regional studies vary in species recognized. This genus is generally found in humid mid- to high-elevation forests, especially in the Andes. Ecuador is notable for its high species richness, particularly in the poorly explored Cordillera del Cóndor, a sub-Andean range with a distinctive geology contributing to high plant diversity and endemism. Since the early 2000s, botanical expeditions have revealed numerous endemic species, highlighting the region's significance. In 2006, an unusual Parablechnum species was collected in the Cordillera del Cóndor. Here, we describe it as a new species, Parablechnum shuariorum . It grows on sandstone cliffs along small rivers and can be distinguished by its fertile fronds, which are shorter than its sterile ones, and its densely scaly rachis. This species, endemic to the Cordillera del Cóndor, is found at elevations of 900–1,600 m. It is named after the Shuar people, whose lands include the collection sites. Preliminary conservation assessment suggests that P. shuariorum is endangered due to a limited area of occupancy and threats from human activities, such as mining.

Premise of the Study Understanding the relationship between phenotypic evolution and lineage diversification is a central goal of evolutionary biology. To extend our understanding of the role morphological evolution plays in the diversification of plants, we examined the relationship between leaf size evolution and lineage diversification across ferns. Methods We tested for an association between body size evolution and lineage diversification using a comparative phylogenetic approach that combined a time‐calibrated phylogeny and leaf size data set for 2654 fern species. Rates of leaf size change and lineage diversification were estimated using BAMM, and rate correlations were performed for rates obtained for all families and individual species. Rates and patterns of rate–rate correlation were also analyzed separately for terrestrial and epiphytic taxa. Key Results We find no significant correlation between rates of leaf area change and lineage diversification, nor was there a difference in this pattern when growth habit is considered. Our results are consistent with the findings of an earlier study that reported decoupled rates of body size evolution and diversification in the Polypodiaceae, but conflict with a recent study that reported a positive correlation between body size evolution and lineage diversification rates in the tree fern family Cyatheaceae. Conclusions Our findings indicate that lineage diversification in ferns is largely decoupled from shifts in body size, in contrast to several other groups of organisms. Speciation in ferns appears to be primarily driven by hybridization and isolation along elevational gradients, rather than adaptive radiations featuring prominent morphological restructuring. The exceptional diversity of leaf morphologies in ferns appears to reflect a combination of ecophysiological constraints and adaptations that are not key innovations.

Key to Dave's long and active career has been his enthusiasm for applying contemporary techniques and approaches in pursuit of the principal goals of his research: resolving reticulate evolutionary histories and understanding plant biogeography. The goal of that visit was to learn isozyme electrophoresis, a technique that by that time was widely used in angiosperms but was just gaining traction in the fern and lycophyte community as a powerful way to infer the relationships of taxa in groups with complex histories of hybridization and polyploidy. Two other papers, each published in the American Fern Journal and with Dave as the lead author, stand out to me as being particularly influential contributions to the study of reticulate evolution in ferns and lycophytes. The second, \"Hybridization, reticulation, and species concepts in the ferns\" by Barrington, Haufler, and Werth (1989), carefully examines how hybridization and polyploidy complicate the application of species concepts in ferns and provides thoughtful suggestions for how fern and lycophyte biologists can make sense of extant fern diversity (namely by recognizing evolutionary cohesive lineages of hybrid origin as species, in most cases).

The taxonomy of the Polypodiaceae subfamily Microsoroideae is highly problematic, especially with respect to the circumscription of the highly variable and non-monophyletic genus Microsorum. Using phylogenetic analyses and morphological evidence, we demonstrate that sixteen taxa typically treated in the genera Microsorum and Colysis are not closely related to those groups and instead belong to three clades that are successive sister groups to the Old-World ant-fern genus, Lecanopteris. We use the available genus name Dendroconche for one of these clades and propose the new genera Bosmania and Zealandia to accommodate the remaining two groups. We provide a description and identify morphological synapomorphies for each of the genera, make new combinations and designate lectotypes where necessary, and present keys and descriptions for all relevant species. We also discuss the evolution of ant-fern associations in the lecanopteroid ferns and highlight the need for additional taxonomic work in the subfamily. The following new combinations are provided: Bosmania lastii, B. leandriana, B. membranacea, Dendroconche ampla, D. latilobata, D. linguiforme, D. sayeri, D. scandens, D. varians, Zealandia novae-zealandiae, Z. powellii, Z. pustulata, Z. pustulata subsp. howensis, and Z. vieillardii.

• Sex expression of homosporous ferns is controlled by multiple factors, one being the antheridiogen system. Antheridiogens are pheromones released by sexually mature female fern gametophytes, turning nearby asexual gametophytes precociously male. Nevertheless, not all species respond. It is still unknown how many fern species use antheridiogens, how the antheridiogen system evolved, and whether it is affected by polyploidy and/or apomixis. • We tested the response of 68 fern species to antheridiogens in cultivation. These results were combined with a comprehensive review of literature to form the largest dataset of antheridiogen interactions to date. Analyzed species also were coded as apomictic or sexual and diploid or polyploid. • Our final dataset contains a total of 498 interactions involving 208 species (c. 2% of all ferns). About 65% of studied species respond to antheridiogen. Multiple antheridiogen types were delimited and their evolution is discussed. Antheridiogen responsiveness was not significantly affected by apomixis or polyploidy. • Antheridiogens are widely used by ferns to direct sex expression. The antheridiogen system likely evolved multiple times and provides homosporous ferns with the benefits often associated with heterospory, such as increased rates of outcrossing. Despite expectations, antheridiogens may be beneficial to polyploids and apomicts.

• Lycopodiaceae are one of three surviving families of lycopsids, a lineage of vascular plants with a fossil history dating to at least the Early Devonian or perhaps the Late Silurian (c. 415 Ma). Many fossils have been linked to crown Lycopodiaceae, but the lack of well-preserved material has hindered definitive recognition of this group in the paleobotanical record. • New, exceptionally well-preserved permineralized lycopsid fossils from the Early Cretaceous (125.6 ± 1.0 Ma) of Inner Mongolia, China, were examined in detail using acetate peel and micro-computed tomography techniques. The anatomy of extant Lycopodiaceae was analyzed for comparison using fluorescence microscopy. Phylogenetic relationships of the new fossil to extant Lycopodiaceae were evaluated using parsimony and maximum likelihood analyses. • Lycopodicaulis oellgaardii gen. et sp. nov. provides the earliest unequivocal and best-documented evidence of crown Lycopodiaceae and Lycopodioideae, based on anatomically-preserved fossil material. • Recognition of Lycopodicaulis in Asia during the Early Cretaceous indicates the presence of crown Lycopodiaceae at this time, and striking similarities of stem anatomy with extant species provide a framework for the understanding of the interaction of branching and vascular anatomy in crown-group lycopsids.