Polyploid goldback and silverback ferns (Pentagramma) occupy a wider, colder, and wetter bioclimatic niche than diploid counterparts.

PREMISE: The western North American fern genus Pentagramma (Pteridaceae) is characterized by complex patterns of ploidy variation, an understanding of which is critical to comprehending both the evolutionary processes within the genus and its current diversity. METHODS: We undertook a cytogeographic study across the range of the genus, using a combination of chromosome counts and flow cytometry to infer ploidy level. Bioclimatic variables and elevation were used to compare niches. RESULTS: We found that diploids and tetraploids are common and widespread, and triploids are rare and sporadic; in contrast with genome size inferences in earlier studies, no hexaploids were found. Diploids and tetraploids show different geographic ranges: only tetraploids were found in the northernmost portion of the range (Washington, Oregon, and British Columbia) and only diploids were found in the Sierra Nevada of California. Diploid, triploid, and tetraploid cytotypes were found to co-occur in relatively few localities: in the southern (San Diego County, California) and desert Southwest (Arizona) parts of the range, and along the Pacific Coast of California. CONCLUSIONS: Tetraploids occupy a wider bioclimatic niche than diploids both within P. triangularis and at the genus-wide scale. It is unknown whether the wider niche of tetraploids is due to their expansion upon the diploid niche, if diploids have contracted their niche due to competition or changing abiotic conditions, or if this wider niche occupancy is due to multiple origins of tetraploids.

Cytogeography of the subalpine marsh marigold polyploid complex (Caltha leptosepala s.l., Ranunculaceae).

PREMISE OF THE STUDY: Unrecognized variation in ploidy level can lead to an underestimation of species richness and a misleading delineation of geographic range. Caltha leptosepala (Ranunculaceae) comprises a complex of hexaploids (6x), rare nonaploids (9x), and dodecaploids (12x), all with unknown distributions. We delineate the geographic distribution and contact zones of the cytotypes, investigate morphologies of cytotypes and subspecies, and discuss the biogeography and evolutionary history of the polyploid complex. METHODS: Using cytologically determined specimens as reference, propidium iodide flow cytometry was performed on silica-dried samples and herbarium specimens from across the range of C. leptosepala s.l. Genome size estimates from flow cytometry were used to infer cytotypes. A key morphological character, leaf length-to-width ratio, was measured to evaluate whether these dimensions are informative for taxon and/or cytotype delimitation. KEY RESULTS: Dodecaploids were more northerly in distribution than hexaploids, and a single midlatitude population in the Northern Rockies yielded nonaploids. Genome size estimates were significantly different between all cytotypes and between hexaploid subspecies. Leaf length-to-width ratios were significantly different between subspecies and some cytotypes. CONCLUSIONS: Caltha leptosepala presents clear patterns of cytotype distribution at the large scale. Marked differences in morphology, range, and genome size were detected between the hexaploid subspecies, C. leptosepala subsp. howellii in the Cascade-Sierra axis and C. leptosepala subsp. leptosepala in the Rockies. Sympatry between cytotypes in the Cascades and a parapatric distribution in the Northern Rockies suggest unique origins and separate lineages in the respective contact zones.

Phylogeny and fruit evolution in Menispermaceae.

PREMISE OF THE STUDY: This work surveys endocarp morphology of Menispermaceae in the context of a well-supported molecular phylogeny. The study is important since menispermaceous endocarps appear often in the fossil record and indicate the presence of a wet forest ecosystem. • METHODS: Three chloroplast regions were used to derive phylogenies for 53 genera and 60 species. Endocarps of 47 genera and 92 species were dissected and morphological characters scored. Photographs of key features are presented. We superimposed our morphological matrix onto the phylogeny to explore character evolution. A detailed key to fruits is presented, allowing identification of extant and fossil specimens to the level of clade or genus. • KEY RESULTS: Menispermaceae consists of two major subfamilies: Tinosporoideae and Menispermoideae. Within Tinosporoideae, tribe Coscineae is basal. Within Menispermoideae, tribe Menispermeae is basal. Tinosporoideae consists mainly of taxa with apical style scars, bilateral curvature, subhemispherical condyles, and foliaceous cotyledons with divaricate or imbricate orientation. Menispermoideae consists almost entirely of taxa with basal or subbasal style scars, dorsoventral curvature, bilaterally and/or dorsoventrally compressed condyles, and subterete or fleshy cotyledons oriented dorsoventrally or laterally. • CONCLUSIONS: Several fruit characters differentiate major clades, and further synapomorphies are diagnostic of various subclades. Fruit characters that can be inferred as ancestral in the family are basal or subbasal stylar scars, endocarps with dorsoventral curvature, endocarp walls woody or bony, presence of a condyle, locule without ribs, sublateral vascular traces, presence of endosperm, and foliaceous or subterete cotyledons.

Phytogeographic implications of fossil endocarps of Menispermaceae from the Paleocene of Colombia.

PREMISE OF THE STUDY: Fossil leaves of Menispermaceae were previously described from the Paleocene of Colombia. Because of strong homoplasy of leaf characters, the fossils could not be placed more specifically within recognized clades, and additional data were needed to specify intrafamilial and paleogeographic relationships during the Paleocene. METHODS: Fossil endocarps of Menispermaceae were collected from the Cerrejón Formation, the recently discovered Bogotá flora, and Wyoming (∼60 Ma). We surveyed the endocarp morphology of almost all extant genera, conducted character optimization, a molecular scaffold analysis, and critically reviewed the related fossil genera. KEY RESULTS: Parallel syndromes of fruit characters have appeared in unrelated clades of the family according to current phylogenetic reconstructions. However, mapping selected endocarp characters across those clades that contain horseshoe-shaped endocarps facilitates identification and phylogenetic assessment of the fossils. Three fossil species are recognized. One of them belongs to the extant genus Stephania, which today grows only in Africa and Australasia. Palaeoluna gen. nov. is placed within the pantropical clade composed of extant Stephania, Cissampelos, and Cyclea; this morphogenus is also recognized from the Paleocene of Wyoming. Menispina gen. nov. shows similarity with several unrelated clades. CONCLUSIONS: The new fossils from Colombia reveal a complex paleobiogeographic history of the recognized clades within Menispermaceae, suggesting a more active exchange among neotropical, paleotropical, North American, and European paleoforests than previously recognized. In addition, the new fossils indicate that neotropical forests were an important biome for the radiation and dispersal of derived lineages in Menispermaceae after the Cretaceous-Paleogene boundary.