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Phylogenetic reconstruction for Carex and relatives in tribe Cariceae is complicated by species richness and nearly cosmopolitan distribution. In this investigation, our main objective was to estimate evolutionary relationships in tribe Cariceae using DNA sequence data from two spacer regions in nuclear ribosomal genes (ITS and ETS-1f) combined with noncoding chloroplast DNA (trnL intron, trnL-trnF intergenic spacer, and trnE-trnD intergenic spacers). Parsimony analyses of separate and combined data and Bayesian analysis of the combined data matrix revealed strong support for monophyly of tribe Cariceae and for monophyly of two major lineages, one comprising principally Carex subgen. Carex and Vigneastra, and the other representing subgen. Vignea. A third clade with representatives from Kobresia and Uncinia, along with Cymophyllus fraserianus, Carex curvula, and several unispicate Carex received weak-to-moderate support. A small clade comprising Schoenoxiphium and two unispicate carices was placed as sister to the clades comprising multispicate Carex species in the parsimony analysis, but sister to the clade of Kobresia, Uncinia, and unispicate Carex in the Bayesian analysis. Two large widespread groups within subgen. Carex, sect. Hymenochlaenae and sect. Physocarpae s.l. (''bladder sedges''), were highly polyphyletic, while ten clades that grouped species from two or more sections were each strongly supported as monophyletic. Within subgen. Vignea, three sections were strongly supported as monophyletic while sects. Phaestoglochin and Vulpinae were polyphyletic. Adding the variable ETS-1f region improved resolution and bootstrap support values over previous studies, but many of the characters supporting major branches came from the trnL region.

Cyperaceae tribe Cariceae is characterized by both species richness and habitat diversity, making it an ideal system to study ecological specialization and niche differentiation. We present a phylogenetic hypothesis for the tribe based on nuclear and chloroplast DNA sequence comparisons (ETS-1f, ITS, trnL intron, trnL-trnF intergenic spacer) for 140 representative species from five continents, and use this hypothesis to suggest patterns of both niche conservatism and niche differentiation, particularly within the large subgenus Carex. We identify a new major clade, comprising forest species of East Asian Carex section Siderostictae (subgenus Carex) as sister to the rest of tribe Cariceae. Within Carex subgenus Carex, species tolerant of water-saturated habitats occur in only a few, apparently derived groups, with varying species richness. Clades of predominantly wetland species tend to have broad geographic distribution, often with sister species on different continents, suggesting recent dispersal. In contrast, species within several clades are predominantly forest specialists with distinct Asian and North American lineages. Niche segregation along environmental gradients, such as soil moisture or acidity, is quite common among closely related wetland species, but more difficult to demonstrate within upland forest groups. More complete sampling of species within both wetland and forest groups, combined with comparable sampling of environmental preferences and testing against null models, will be needed for more rigorous exploration of the observed patterns.

We assessed the size and composition of the seed bank in 31 plots representing a range of habitats within an old-growth, temperate deciduous forest at Mont St. Hilaire, Québec, Canada. We identified 49 taxa in the seed bank, with an average of 40 species·m-2 and a median density of 1218 seeds·m-2. The most frequent seeds were species of Carex and Rubus, Diervilla lonicera, and Eupatorium rugosum, while seeds of Carex were the most numerous overall. Of the 12 species in the seed bank not found in the forest, 11 were found growing on the developed landscape surrounding this 10-km2 forest fragment. These nonforest species were numerically only a minor component of the forest seed bank. Vernal herbs were not in the seed bank, and there were only a few tree species. Variation in seed bank richness among habitats was correlated positively with canopy cover, soil moisture, and soil nutrients, but not with the seed bank density or total number of species in the aboveground vegetation. Seed bank density increased with plot soil moisture. Woody species predominated in the seed bank of plots with richer soils, deeper litter, and more closed canopies. Herbaceous species predominated in the seed bank of plots with more open canopies, more mesic water regimes, and greater species richness in the aboveground vegetation. Contrary to earlier results suggesting forest seed banks primarily include shade-intolerant species associated with canopy disturbance or secondary succession, the seed bank in this old-growth, primary forest contains many shade-tolerant forest species.Key words: seed bank, old-growth forest, primary forest, temperate deciduous forest, habitat diversity, seed dispersal.

Phylogenetic studies of Carex L. (Cyperaceae) have consistently demonstrated that most subgenera and sections are para-or polyphyletic. Yet, taxonomists continue to use subgenera and sections in Carex classification. Why? The Global Carex Group (GCG) here takes the position that the historical and continued use of subgenera and sections serves to (i) organize our understanding of lineages in Carex, (ii) create an identification mechanism to break the~2000 species of Carex into manageable groups and stimulate its study, and (iii) provide a

The field of systematics is experiencing a new molecular revolution driven by the increased availability of high-throughput sequencing technologies. As these techniques become more affordable, the increased genomic resources have increasingly far-reaching implications for our understanding of the Tree of Life. With c. 2000 species, Carex (Cyperaceae) is one of the five largest genera of angiosperms and one of the two largest among monocots, but the phylogenetic relationships between the main lineages are still poorly understood. We designed a Cyperaceae-specific HybSeq bait kit using transcriptomic data of Carex siderosticta and Cyperus papyrus. We identified 554 low-copy nuclear orthologous loci, targeting a total length of c. 1 Mbp. Our Cyperaceae-specific kit shared loci with a recently published angiosperm-specific Anchored Hybrid Enrichment kit, which enabled us to include and compile data from different sources. We used our Cyperaceae kit to sequence 88 Carex spp., including samples of all the five major clades in the genus. For the first time, we present a phylogenetic tree of Carex based on hundreds of loci (308 nuclear exon matrices, 543 nuclear intron matrices and 66 plastid exon matrices), demonstrating that there are six strongly supported main lineages in Carex: the Siderostictae, Schoenoxiphium, Unispicate, Uncinia, Vignea and Core Carex clades. Based on our results, we suggest a revised subgeneric treatment and provide lists of the species belonging to each of the subgenera. Our results will inform future biogeographic, taxonomic, molecular dating and evolutionary studies in Carex and provide the step towards a revised classification that seems likely to stand the test of time.

Summary 1 A ®eld experiment was designed to investigate the relationship between environmental heterogeneity and species diversity in a group of sedges (Cyperaceae: Carex) growing in old-growth forest. 2 A measure of environmental quality, as perceived by the sedges, was obtained from the survival of clonal ramets of 11 species of Carex planted at 10-m intervals along each of three 1-km transect lines. 3 The resident assemblage of sedges was censused along the same three transect lines and along a further 24 km of survey lines in the same forest. 4 The general state of a site was represented by the overall survival of the experimental implants at that site. The general environmental variance between sites provided a measure of environmental heterogeneity. This could be partitioned into a speci®c variance (mean environmental variance of species) and an environmental covariance. The rate of increase of the general and speci®c variances with distance between sites re¯ected environmental structure. 5 The three transects diered in scale. The species diversity of the resident Carex assemblage was correlated with general environmental quality both among and within transects. 6 The three transects diered in structure. The number of resident species, relative to the number expected from the number of individuals sampled, was greatest on the most coarse-grained transect (steepest increase in general environmental variance with distance). 7 Within each transect, species diversity increased with general environmental variance because the speci®c correlation of performance (correlation among species of survival in pair-wise combinations of sites) decreased as the general environmental variance increased. 8 The eect of speci®c environmental variance was weaker. Overall survival of a species on the transects was not correlated with its abundance in the forest. Neither the transects nor a targeted implant experiment provided evidence for a close relationship between the distribution of species and the state of the environment. 9 As a general explanation of our results, we propose a`marginal-specialist' model in which the species that dominate the most productive sites also have the broadest ranges, whereas other species are superior in a more restricted range of less productive sites.

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