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Molecular Systematics of Cyperaceae Tribe Cariceae Based on Two Chloroplast DNA Regions: Ndhf and Trnl Intron-Intergenic Spacer

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Molecular Systematics of Cyperaceae Tribe Cariceae Based on Two Chloroplast DNA Regions: Ndhf and Trnl Intron-Intergenic Spacer Systematic Botany (2000), 25(3): pp. 479±494 q Copyright 2000 by the American Society of Plant Taxonomists Molecular Systematics of Cyperaceae Tribe Cariceae Based on Two Chloroplast DNA Regions: ndhF and trnL Intron-intergenic Spacer ALAN C. YEN1 and RICHARD G. OLMSTEAD Department of Botany, Box 355325, University of Washington, Seattle, Washington 98195-5325 1Present Address: The Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138 Communicating Editor: Jeff H. Rettig ABSTRACT. A phylogenetic analysis of Cyperaceae tribe Cariceae was conducted using chloroplast DNA sequences from the gene ndhF and trnL intron and trnL-trnF intergenic spacer. Twenty nine taxa within Cariceae, four outgroup genera, and approximately 3,000 bp of cpDNA were included in the study. Our analysis reveals a monophyletic Cariceae with 100% bootstrap support. Within Cariceae, the South African genus Schoenoxiphium forms a clade that is sister to the rest of the tribe. Our results indicate that genus Carex is paraphyletic with respect to Kobresia, Cymophyllus, and Uncinia. Cymophyllus and Uncinia are nested within an assemblage containing Kobresia, Cymophyllus, and several unispicate Carex species. At the subgeneric level within Carex, only Carex subgenus Vignea appears monophyletic. Several well supported clades were iden- ti®ed within the Cariceae, including the Schoenoxiphium clade, Uncinia clade, Carex subgenus Indocarex/Carex clade, and subgenus Vignea clade; however, relationships among some clades are only moderately supported. Interpretation of the phylogenetic patterns and an account of past phylogenetic hypotheses with respect to the new data are provided. The sedge tribe Cariceae, with over 2,000 species southern Africa and Madagascar (Kukkonen 1986). worldwide, is the largest tribe in the family Cyper- The monotypic Cymophyllus (Ker-Gawler) Kartesz & aceae. Five genera, including the large genus Carex Gandhi occurs in the southeastern United States. L., generally are included in this tribe. Several evo- Plants in Carex, Uncinia, and Cymophyllus have lutionary schemes have been proposed for Carex one-¯owered, unisexual spikelets with closed peri- and related genera based on in¯orescence mor- gynia, whereas plants in Kobresia and Schoenoxi- phology and cytology (KuÈ kenthal 1909; Heilborn phium usually have several-¯owered, bisexual 1924; Kreczetovicz 1936; Nelmes 1952; Savile and spikelets with open perigynia. In addition, Uncinia Calder 1953; Koyama 1961, 1962; Smith and Faulk- species are characterized by having a hook-like ner 1976; Kern and Nooteboom 1979; Reznicek structure, formed by the extension of the rachilla 1990); however, there is substantial disagreement and a leaf, that extends beyond the opening of the among various authors. perigynium. Kobresia has straight, terete rachillae, Members of tribe Cariceae are characterized by whereas Schoenoxiphium has straight, but ¯attened, having monoecious ¯owers and a sac-like structure rachillae. In both Kobresia and Schoenoxiphium, the termed a perigynium, that subtends the gynoeci- rachillae sometimes are well developed and bear um. In¯orescence morphology, the degree of clo- male ¯owers. Rachillae in Carex are of various sure of the perigynium, and the morphology of the forms including a reduced bud-like structure, a rachilla, are the key characters used in delimiting straight type, or a bent structure that is morpholog- genera. Carex L. (2,000 species) is found in all tem- ically similar to those in Uncinia (Snell 1936; Rez- perate regions of the world as well as montane ar- nicek 1990). eas in the tropics (Nelmes 1951). Kobresia Willd. (50 Because generic delimitation within Cariceae is species), is widely distributed at high altitudes in based largely on the morphology of the in¯ores- the Himalayas, China, and central Asia, with a few cence structure, blurring of the generic boundaries species found in the high mountains of Europe and becomes a problem for some taxa having morpho- North America (Dahlgren et al. 1985). Uncinia Pers. logical characters that are interpreted as interme- (50 species), is found widely in highlands of Central diate. Clarke (1908) placed a few species of Schoe- and South America, Australia, New Zealand, and noxiphium in Carex. Ivanova (1939) transferred sev- oceanic islands throughout the Southern hemi- eral Carex species to Kobresia. Kern (1958) consid- sphere (Kukkonen 1967). Schoenoxiphium Nees (17 ered the recognition of Kobresia and Schoenoxiphium species), is found in mountains in eastern and as two genera to be arti®cial. Koyama (1961) 479 480 SYSTEMATIC BOTANY [Volume 25 merged Uncinia with Carex, and Schoenoxiphium related to Kobresia or Schoenoxiphium than they are with Kobresia. to Carex. Savile and Calder (1953), in their studies At the subgeneric level in Carex, the present clas- of smut fungi that infect Carex, suggested that Carex si®cation largely follows the system of KuÈ kenthal is a natural group. Smith and Faulkner (1976) sug- (1909), who organized Carex into four subgenera gested that Kobresia and Schoenoxiphium are the based on in¯orescence structure. Recognition of most primitive genera of the Cariceae and pro- subgenera within Carex is based entirely on phe- posed an evolutionary link between Schoenoxiphium, netic similarity in the in¯orescence structure and Kobresia, and Carex subgenus Indocarex. Reznicek probably does not re¯ect phylogenetic relation- (1990) suggested that Uncinia and Carex may not be ships. Subgenus Primocarex KuÈ kenth. (60 species), closely related, and disputed the importance of the characterized by a single terminal in¯orescence, rachilla as a systematic character. generally is considered to be arti®cial (Kreczetovicz Reznicek (1990) presented the most recent hy- 1936; Nelmes 1952; Smith and Faulkner 1976; Rez- potheses on the evolution of Carex, which he con- nicek 1990). To reduce confusion, instead of using sidered to be derived from Schoenoxiphium-orKo- the designation ``subgenus Primocarex,'' we refer to bresia-like ancestors. He suggested that the most these taxa simply as ``unispicate species'' in this pa- primitive subgenus in Carex is subgenus Vignea, per. Subgenus Vignea (P. Beauv.) Nees (400 to 500 which he considered to contain the most complex species), is characterized by having bisexual spikes in¯orescence structures in the genus, and that sub- and two stigmas, and generally is considered a nat- genus Carex could have evolved through a reduc- ural group (Reznicek 1990), although this hypoth- tion in in¯orescence structure and branching. How- esis has not been veri®ed cladistically. Subgenus ever, Reznicek (1990) considered the origin and Carex ( 5 Eucarex Coss. et Germ.) (1,400 species), is evolution of subgenus Indocarex to be unclear. He a morphologically diverse group characterized by regarded the unispicate species to be polyphyletic having a cladoprophyll (a tubular or utriculiform and derived by reduction in in¯orescence complex- structure found at the base of the in¯orescence) and ity on multiple occasions from the other three sub- usually unisexual spikes with ¯owers having two genera and possibly from other genera in the Car- or three stigmas. It is unclear from the taxonomic iceae (Nelmes 1952). Reznicek's (1990) views on the literature whether subgenus Carex is a natural evolution and the phylogenetic position of Carex group or what the phylogenetic relationships be- subgenus Vignea were quite different from those of tween it and the other subgenera may be. Subgenus most authors, who generally consider subgenus In- Indocarex Baill. (100 mostly tropical species), is char- docarex to be most primitive and subgenus Vignea acterized by the presence of a cladoprophyll and to be derived (KuÈ kenthal 1909; Kreczetovicz 1936; highly branched, bisexual spikes with tristigmatic Nelmes 1952; Savile and Calder 1953; Koyama 1961; ¯owers, and is considered the most primitive sub- Kern and Nooteboom 1979). genus in Carex by many (Kreczetovicz 1936; Nelmes Molecular data can provide an evaluation of clas- 1952; Koyama 1962; Smith and Faulkner 1976; Kern si®cations based on morphological characters, the and Nooteboom 1979), but not all authors (KuÈken- interpretation of which have led to confusing and thal 1909; Reznicek 1990). contradictory hypotheses of the phylogenetic rela- Many hypotheses on the evolution of Carex and tionships in the Cariceae. Because of the reduced Cariceae have been proposed. Heilborn (1924) pro- ¯oral structures, the uniform vegetative morphol- duced the ®rst ``phylogenetic tree'' of the genus ogy, and traits unique to the Cariceae (e.g., peri- based largely on chromosome numbers. KuÈ kenthal gynium), polarization of characters based on out- (1909) and Heilborn (1924) considered the unispi- group comparison is dif®cult (Crins 1990; Bruhl cate species and species with low chromosome 1995). Recent phylogenetic studies based on mor- numbers to be primitive in the genus. However, phology (Goetghebeur 1986; Bruhl 1995; Simpson Kreczetovicz (1936) suggested that unispicate spe- 1995) and molecular data (Plunkett 1995; Muasya et cies are derived from other subgenera within Carex al. 1998) on Cyperaceae and Cyperales (sensu through the reduction in in¯orescence complexity. Dalghren et al. 1985) indicate a monophyletic Cy- Kreczetovicz (1936) also suggested that some un- peraceae and tribe Cariceae sensu KuÈ kenthal and ispicate species may have had their origins outside suggest possible sister groups of tribe Cariceae, in- the genus. Nelmes (1952) elaborated
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