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Evolution in Sedges (Carex, Cyperaceae)

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Evolution in Sedges (Carex, Cyperaceae) Evolution in sedges (Carex, Cyperaceae) A. A. REZNICEK University of Michigan Herbarium, North University Building, Ann Arbor, MI 48/09, U.S.A. Received January 2, 1990 REZNICEK,A. A. 1990. Evolution in sedges (Carex, Cyperaceae). Can. J. Bot. 68: 1409-1432. Carex is the largest and most widespread genus of Cyperaceae, but evolutionary relationships within it are poorly under- stood. Subgenus Primocarex was generally thought to be artificial and derived from diverse multispicate species. Relation- ships of rachilla-bearing species of subgenus Primocarex, however, were disputed, with some authors suggesting derivation from other genera, and others believing them to be primitive. Subgenus Indocarex, with compounded inflorescence units, was thought to be primitive, with subgenera Carex and Vignea reduced and derived. However, occurrence of rachillas is not confined to a few unispicate species, as previously thought, but is widespread. The often suggested connection between Uncinia and unispicate Carex is shown, based on rachilla morphology, to be founded on incorrect interpretation OF homology. Uncinia kingii, the alleged connecting link, is, in fact, a Carex. Unispicate Carex without close multispicate relatives probably originated from independent, ancient reductions of primitive, rachilla-bearing, multispicate Carex. The highly compounded inflorescences occumng in subgenus Vignea are hypothesized to represent a primitive state in Carex, and the more specialized inflorescences in subgenus Carex derived from inflorescences of this type. The relationships of subgenus Indocurex, with its unique perigynium-like inflorescence prophylls, remain unclear. REZNICEK,A. A. 1990. Evolution in sedges (Carex, Cyperaceae). Can. J. Bot. 68 : 1409-1432. Le Carex est le genre le plus irilportant et le plus rCpandu des Cyperaceae, mais les affinites Cvolutives a I'intCrieur de ce genre sont ma1 connues. Le sous-genre Primocarex Ctait gCnCralement considCrC commeztant artificiel et dCrivC d'une gamme d'espbces a plusieurs Cpis. Les affinitCs des espbces a rachColes du sous-genre Primocarex Ctaient, cependant, mises en question, certains auteurs invoquant la possibilitC d'une dirivation d'autres genres, d'autres les supposant primitives. Le sous-genre Indocarex, avec ses unitCs d'inflorescences glomCrulaires, Ctait considCrC comme primitif, et les sous-genres Carex et Vignea rkduits et dCrivCs. Cependant, la prCsence de rachColes ne se limite pas a quelques espbces a Cpi solitaire, comme on le croyait; elle est en fait trbs rkpandue. Le lien prCsumC entre le Uncinia et les Carex a Cpi solitaire est identifii, d'aprks la morphologie des rachkoles, comme Ctant fond6 sur une interpretation incorrecte de I'homologie. Le Uncinia-kingii, le chainon de liaison prCsumC, est en fait un Carex. Les Carex a Cpi solitaire sans proches parents a plusieurs Cpis sont probablement issus de forrnes rkduites, anciennes et indkpendantes de Carex primitifs a rachColes et a plusieurs Cpis. Les inflorescences hauternent glornCrulaires prCsentes dans le sous-genre Vignea representent, par hypothese, un Ctat primitif chez le Carex, et les inflorescences plus spCcialisCes dans le sous-genre Carex sont dCrivCes d'inflorescences de ce type. Les affinitCs du sous-genre Indocarex, avec ses prophylles d'inflorescences ressemblant a des pCrigynes, demeurent incertaines. [Traduit par la revue] Introduction Systematic position of Carex within the Cyperaceae Carex L., with about 2000 species, is by far the largest Although this paper is concerned with evolution within genus in the Cyperaceae and one of the most widespread and Carex, a brief diversion to establish the context within which ecologically important genera of vascular plants. Remarkably, Carex fits is necessary. Hypotheses of evolution within Carex evolutionary trends within the genus are poorly understood, invariably become entangled with the problems of generic lim- and published work on this topic is sketchy. This paper will its within the tribe Cariceae Kunth ex Dumort.' and frequently both briefly review past work on evolution in Carex and pre- also invoke the phytogeography of other genera in the tribe. sent new or overlooked information that may shed light on past As well, discussions of inflorescence structure in Carex rely problems or suggest alternative hypotheses for certain aspects heavily on the character states displayed by some of the other of the evolution of sedges. The review of past work will con- genera in the Cariceae. centrate primarily on more recent work that actually proposed With about 4500 to 5000 species in 100 to 105 genera (Goetghebeur 1987), the Cyperaceae are probably the seventh hypotheses rather than enumerate the schemes of the numerous largest family of vascular plants. The subfamily Caricoideae authors that merely developed an arrangement convenient for Pax, with one tribe, Cariceae, to which Carex belongs, com- listing the species in their local region. Since this review is prises five genera and about 2100 species, thus constituting concerned with evolution within Carex, it will also not touch nearly half the family. Almost all the species in the subfamily on the extensive literature concerning topics such as the der- belong to the enormous genus Carex. ivation of the perigynium and the controversy over whether or The tribe Cariceae is characterized by uniformly unisexual not male flowers are true flowers or synanthia. However, a flowers with the female subtended by a partially or wholly brief discussion of terminology, especially as it relates to Carex closed perigynium of prophyllar origin (Blaser 1944). The five inflorescences, is included, since varied and conflicting ter- genera here recognized are Carex, Cymophyllus Mackenzie, minology is in use in recent literature. While developing a Kobresia Willd., Schoenoxiphium Nees, and Utzcinia Pers. AS natural classification of Carex reaching to the species or sec- noted by Mora-Osejo (1982) and Wheeler (1989), the South tional level is not yet feasible, comments are brought to bear American segregate genus Vesicarex (Steyermark 1951; Cleef on the classification at the subgeneric level. A few previously 1982) is referable to Carex sect. Abditispicae G. Wheeler. The proposed hypotheses are discussed and rejected, but unfortu- nately, more questions are raised than are answered. 'Authors for suprageneric names follow Goetghebeur (1985). Printed in Canada i Imprim6 au Canada 1410 CAN. J. BOT. VOL. 68, 1990 peculiar collarlike appendage noted by Steyermark (195 1) may tiated inflorescences, Schoenoxiphium has often been regarded be an artifact caused by separation of the pericarp and seed as the most primitive genus in the Cariceae. coat, which sometimes occurs when immature material of A fundamental blurring of generic limits may occur between Carex is softened too long in water before dissection. The Carex aand Kobresia, with certain species traditional!^ put in small, South American genus Bisboeckelera Kuntze (Hoppia Carex, but very similar to Kobresia except for the more Nees non Sprengel), sometimes cited as belonging in the tribe reduced rachilla (Nelmes 1952). Some of these, such as Cariceae (Jermy et. al. 1982), is here considered to belong in C. bucharica Kiikenth., C. hepburnii Boott, and C. nardina the subfamily Sclerioideae C. B. Clarke, tribe Bisboeckeler- Fries were, in fact, transferred to Kobresia by Ivanova (1939). eae Pax ex L. Eiten (Koyama 1965; Smith and Faulkner 1976; In addition, the considerable similarity of Schoenoxiphium to Meert and Goetghebeur 1979). members of Carex subgenus Indocarex has not gone unno- Except for Cymophyllus, which has unique, flat, sheathless ticed, and may indicate a close relationship (Haines and Lye leaves without a midvein or ligule, the genera are all essen- 1983). Some species of Schoenoxiphium have also been placed tially indistinguishable vegetatively (Metcalfe 1969, 1971). in Carex in the past by Clarke (1898, 1908). The delimitation of the genera based on reproductive charac- Carex is cosmopolitan in distribution, but with the great ters is also difficult, with apparently transitional species link- majority of the species in the north and south temperate re,'OIO~S ing all the genera except Cymophyllus. The brief outline fol- and the montane tropics. Uncinia, with perhaps 50 species lowing is, of necessity, inadequate since the topic of generic (Kukkonen 1967a), is predominantly Southern Hemisphere, limits is beyond the scope of this work. with occurrences north of the Equator from northern South Carex, Cymophyllus, and Uncinia are characterized by a America to Mexico and the West Indies, and in Hawaii. completely closed perigynium. Only in Carex subgenus Vig- Kobresia, with about 40 species (Ivanova 1939), is best devel- nea (P. Beauv. ex Lestib. f.) Peterm. does the perigynium oped in the Himalayas and adjacent regions of central Asia, have an abaxial false suture (Mackenzie 193 1; Nannfeldt 1977) with a few boreal species and one in alpine Sumatra (Kern that may be a vestige of a partially open perigynium. As well, 1958). Schoenoxip&um, with 17 species now known in all three genera, there is a distinction between spikes and (Kukkonen 1983, 1986a), is restricted to southern and eastern one-flowered spikelets, as defined by Smith and Faulkner Africa and Madagascar. 'The monotypic Cymophyllus is a (1976). In Uncitzia, the rachilla (the vestigial continuation of southern Appalachian Mountain endemic, occurring from the axis that bears the female flower laterally) is smooth,
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