J. Jpn. Bot. 87: 169–186 (2012)

Delineation of Schoenoplectiella Lye (Cyperaceae), a Genus Newly Segregated from Schoenoplectus (Rchb.) Palla

Eisuke Hayasaka

Fukui Botanical Garden, Echizen, Fukui, 916-0146 JAPAN E-mail: [email protected]

(Accepted on March 25, 2012)

Schoenoplectiella Lye, a genus segregated from Schoenoplectus (Rchb.) Palla, is accepted with expanded circumscription to include all the species formerly placed in Schoenoplectus sectt. Actaeogeton (Rchb.) J. Rayn. and Supini (Cherm.) J. Rayn. A revised description of Schoenoplectiella is provided and all the 50 species included in the genus are listed with 34 new combinations for the species, varieties, and hybrids. Two sections, Schoenoplectiella sectt. Schoenoplectiella and Actaeogeton (Rchb.) Hayasaka are recognized with 25 species for each, and a key to and descriptions for the sections are given. Morphological characters that define Schoenoplectiella and its sections, as well as those that distinguish the genus from allied genera, are discussed.

Key words: Cyperaceae, generic circumscription, infrageneric classification, morphology, new combination, Schoenoplectiella, Schoenoplectus, Scirpus.

Schoenoplectus (Rchb.) Palla, one of the Ficinia Schrad. with 74 species and Isolepis R. segregates of Scirpus L. s. l., has been widely Br. with 74 species (Muasya and Simpson 2002, accepted since 1980’s in the floras of various Govaerts and Simpson 2007). regions of the world (Lye 1997, Kukkonen Species of Schoenoplectus show a 1998, Simpson and Koyama 1998, Smith considerable wide range of variation in their 2002, Liang et al. 2010), or in individual habit and morphology of both vegetative and taxonomic accounts (Wilson 1981, Strong 1994, reproductive organs. They can range from a Gordon-Gray 1995, Pignotti 2003, Egorova dwarf, tufted annual up to 5 cm high growing in 2005). Among the Scirpus segregates, either seasonally wet habitats, to a large, rhizomatous Actinoscirpus (Ohwi) R. W. Haines & Lye or perennial up to 350 cm high growing in Bolboschoenus (Asch.) Palla is merged into permanently wet, freshwater or brackish Schoenoplectus by some authors (Koyama 1978, habitats. Interspecific variation in morphological Lye 1997), but Schoenoplectus as excluding characters is remarkable especially in the shape these two genera is more frequently accepted of rhizomes, leaf blades, inflorescences, glumes, in recent years (Govaerts and Simpson 2007). perianth segments, nutlets, and in the presence Schoenoplectus in the latter sense comprises 77 or absence of nodes on a culm (Smith and species worldwide ranging from the subarctic to Hayasaka 2001). the tropics (Hayasaka 2002). The genus is now To grasp the species relationships within the largest of the Scirpus segregates, exceeding this large and polymorphic genus, attempts

—169— 170 植物研究雑誌 第 87 巻 第 3 号 2012 年 6 月 have been made to establish an infrageneric Young at University of Oklahoma (Unpublished system in Schoenoplectus. Oteng-Yeboah (1974) data, pers. comm. 2001–2002) used ITS and recognized and provided a key to the three trnL sequences for the infrageneric phylogeny subgenera Schoenoplectus, Actaeogeton (Rchb.) of Schoenoplectus, and indicated that the genus Oteng-Yeb., and Malacogeton (Ohwi) Oteng- is represented by two well-supported clades Yeb. His subgeneric system is recently adopted that are not sister to each other: one with the in the Schoenoplectus treatment of Pignotti species of the sections Schoenoplectus and (2003). Raynal (1976b) provided a monograph Malacogeton, the other with those of the sections of the section Supini (Cherm.) J. Rayn. with 21 Actaeogeton and Supini. Each of the above four species mainly from Africa and Madagascar, and sections recognized by Smith and Hayasaka he recognized the four sections Schoenoplectus, (2001, 2002) is supported as monophyletic in Actaeogeton (Rchb.) J. Rayn., Pterolepis her work. Molecular phylogeny within Japanese (Schrad.) J. Rayn., and Supini (Raynal 1976a, b, Schoenoplectus presented by Yano and Hoshino 1977). Raynal’s sectional treatment is followed (2005) also shows the monophyly of the two by Haines and Lye (1983) and Kukkonen sections Schoenoplectus and Actaeogeton. (1998). Smith and Hayasaka (2001, 2002) In their study, Japanese Schoenoplectus was partly accepted Raynal’s sectional treatment resolved into two clades, one with the species of and applied it also to the North American and the section Actaeogeton, the other with those of eastern Asian species, recognizing the four the sections Schoenoplectus and Malacogeton. sections Schoenoplectus (including Pterolepis), Deliberating on the results of these studies that Actaeogeton, Malacogeton (Ohwi) S. G. Smith indicate the polyphyly of Schoenoplectus as & Hayasaka, and Supini. Their infrageneric well as the monophyly of the four sections, the system is adopted in the Schoenoplectus genus should be divided into two genera, with treatments of Smith (2002) and Egorova (2005), the circumscription of the sections maintained as and is also used as a basis for discussion in the currently recognized under Schoenoplectus (s. molecular phylogenetic studies of the genus l. with 77 spp.). A solution could be delimiting (Yano and Hoshino 2005, Jung and Choi 2010). Schoenoplectus (s. s.) with 27 species classified Molecular phylogenetic studies in recent into the two sections Schoenoplectus and years show that Schoenoplectus as currently Malacogeton, and simultaneously establishing delimited is polyphyletic. In the suprageneric a new genus with 50 species classified into two phylogeny of Cyperaceae using rbcL sequences sections that correspond to Schoenoplectus sectt. (Muasya et al. 1998, Simpson et al. 2007), Actaeogeton and Supini. Schoenoplectus was resolved into two clades, Based on the rbcL suprageneric phylogeny one with S. lacustris (L.) Palla of the section of Muasya et al. (1998), Lye (2003) described Schoenoplectus and Actinoscirpus grossus (L. the new genus Schoenoplectiella Lye, which f.) Goetgh. & D. A. Simpson, the other with S. includes 26 species formerly placed in articulatus (L.) Palla and S. junceus (Willd.) Schoenoplectus. The greater part of these species J. Rayn. of the section Supini, as sister to are comparatively small, amphicarpous annuals Eleocharis. Muasya et al. (2009) analyzed eight of Africa and Madagascar. In the phylogenetic species of Schoenoplectus in their rbcL and tree presented by Muasya et al. (1998), only trnL-F phylogeny, showing the genus resolved two species of Schoenoplectiella, S. articulata into two strongly supported clades, one with and S. juncea are included, the former of which the species of the section Schoenoplectus as was designated as the type of the genus (Lye sister to Actinoscirpus, the other with those of 2003). Lye (2003) apparently intended to give the sections Actaeogeton and Supini. Laura A. a generic status to Schoenoplectus sect. Supini June 2012 Journal of Japanese Botany Vol. 87 No.3 171 because his key to distinguish Schoenoplectiella Schoenoplectus sect. Actaeogeton, which is from Schoenoplectus, as well as his description an acceptable solution to the taxonomy of of the new genus, refers to the morphology Schoenoplectus s. l. However, Schoenoplectiella and habit of the species in the section (Raynal would then be a comparatively large genus 1976b). Out of the 26 species that Lye (2003) widespread in the world that needs a taxonomic included in Schoenoplectiella, 23 are those of revision. The description of Schoenoplectiella Schoenoplectus sect. Supini (Raynal 1976b, should be revised with a close examination Hayasaka 2009). The remaining three species, of morphological characters that delimit the however, are those of Schoenoplectus sect. genus, and the generic circumscription should Actaeogeton (Smith and Hayasaka 2001, be clarified with an enumeration of the included 2002). They are Schoenoplectiella purshiana, species worldwide. In addition, an infrageneric non-amphicarpous annual of eastern North system is needed because Schoenoplectiella America, S. juncoides, non-amphicarpous is to be composed of all the species of perennial of Asia and the Pacific Islands, and S. Schoenoplectus sectt. Actaeogeton and Supini, wallichii, non-amphicarpous annual of eastern which are monophyletic groups respectively, as and southern Asia. Because the habit of these well as distinguishable from each other by their species is inconsistent with the definition of morphology and habit (Raynal 1976b, Smith Schoenoplectiella as amphicarpous annuals, Lye and Hayasaka 2001, 2002). This paper addresses (2003) put them into the new genus presumably these problems. being unaware of their habit. Jung and Choi (2010) presented molecular Morphological characters of Schoenoplectiella phylogeny of Scirpus s. l. of Korea using ITS and Morphological characters that define rbcL sequences. Nine species of Schoenoplectus Schoenoplectiella and distinguish it from s. l. were included in their phylogenetic analysis, Schoenoplectus s. s. are those shared between in which the genus was resolved into two clades Schoenoplectus (s. l.) sectt. Actaeogeton that are not sister to each other. The smaller and Supini. The distinguishing characters of of these clades consists of the representatives Schoenoplectiella include the shape of rhizomes, from Schoenoplectus sectt. Schoenoplectus and culm arrangement, presence or absence of nodes Malacogeton, as sister to Actinoscirpus, while on a culm, shape of glumes, presence or absence the larger one consists of the representatives of perianth segments, surface sculpturing of from Schoenoplectus sect. Actaeogeton, as sister nutlets, shape of epidermal cells of nutlets, and to Eleocharis. All of the nine species from Korea the habit whether annual or perennial, all of treated in Jung and Choi (2010) are distributed which are used in the key below. I examined also in Japan, and were previously included in specimens for these characters during the course the phylogenetic analysis of Yano and Hoshino of my revisional work of Schoenoplectus s. (2005). Results of the analyses performed by l. (Hayasaka 2002). For the purpose of this both of the researchers confirm that eastern paper, my earlier version of the description of Asian Schoenoplectus s. l. contains two remote Schoenoplectus is revised as well as the key clades. In accordance with their phylogenetic is reconstructed to explain the morphology tree obtained, Jung and Choi (2010) recognized of Schoenoplectiella. The descriptions of and Schoenoplectiella as a distinct genus, in the key to the two sections recognized under which they included six species of Korea Schoenoplectiella below are equivalent to that were formerly placed in Schoenoplectus those I have prepared for Schoenoplectus sectt. sect. Actaeogeton. This is an expansion of Actaeogeton and Supini. In the interest of further Schoenoplectiella sensu Lye (2003) to include studies on the morphology and taxonomy of 172 植物研究雑誌 第 87 巻 第 3 号 2012 年 6 月

Fig. 1. Nutlet epidermal cells (with the outer periclinal walls removed) of Schoenoplectiella and Schoenoplectus s. s. a. Schoenoplectiella articulata (L. J. Brass 19776, BRI). b. Schoenoplectiella mucronata (E. Hayasaka 2786, TUS). c. Schoenoplectus lacustris (E. Hayasaka 3282, TUS). d. Schoenoplectus nipponicus (E. Hayasaka 2969, TUS). Scale bars = 50 µm.

Schoenoplectiella, I here discuss the morphology (Ragonese et al. 1984), Schoenus (Liu and Lin of nutlet epidermal cells and inflorescences, 1999), and Scirpus s. l. (Schuyler 1971). As which I regard as important for recognizing the confirmed by these authors, morphology of genus. nutlet epidermal cells provides useful characters for the taxonomy of various genera because Morphology of nutlet epidermal cells in of the infraspecific stability of the shape, size, Schoenoplectiella and Schoenoplectus s. s. and arrangement of the cells as well as their The nutlet epidermal cells of Cyperaceae interspecific similarities or differences that can be observed to the detail with the scanning correlate with other morphological characters. electron microscope after removing the outer Schuyler (1971) examined the nutlet periclinal walls by sonicating the nutlets with an epidermal cells of nine species of Eriophorum ultrasonic cleaner (Schuyler 1971). This method and 49 species of Scirpus s. l. In his paper, he has been applied to the observations of the cells provided scanning electron micrographs for in various genera including Carex (Standley five species of Schoenoplectiella and 11 species 1985), Cyperus (Wilson 1991), Fimbristylis (Oh of Schoenoplectus s. s. (treated under Scirpus), and Park 1997), Gahnia (Liu and Lin 1999), of which four in Schoenoplectiella and nine in Eleocharis (Menapace 1991), Eriophorum Schoenoplectus s. s. are distinct species in my (Tucker and Miller 1990), Rhynchospora treatment (Hayasaka 2002). For the purpose June 2012 Journal of Japanese Botany Vol. 87 No.3 173 of disccusion, Schuyler (1971) organized the absent or often one to several per cell. In all the examined species of Scirpus s. l. into five groups, species examined in Schoenoplectiella sect. which correspond roughly to the segregate Actaeogeton, the cells were uniformly narrowly genera. The four species of Schoenoplectiella he oblong to linear with the inner periclinal walls examined were arranged in the group ‘Scirpus lacking pits (Fig. 1b). In this section, silica supinus and related species’, and the nine of bodies were absent or often 1–2 that were less Schoenoplectus s. s. in ‘Scirpus lacustris and developed than in the section Schoenoplectiella. related species’. Based on his observations on The cells observed in Schoenoplectus sect. the nutlet epidermal cells, he mentioned that the Schoenoplectus were somewhat various in the cells much longer than wide shared by ‘Scirpus outline shape and cellular details. The majority supinus and related species’ indicate the close of the species in the section had cells that were relationship within the group although many isodiametric to oblong with 1(–2) silica bodies more species should be examined. He also and without pits on the inner periclinal walls pointed out that the cells of ‘Scirpus lacustris (Fig. 1c). In one species of the section, however, and related species’ are much shorter than those the cells were narrowly oblong without silica of ‘Scirpus supinus and related species’, which bodies, and the inner walls were pitted; and distinguish between the two groups. in another species, the cells were oblong to By Schuyler’s (1971) method, I examined narrowly oblong with 1–5 bodies, and the the nutlet epidermal cells of 17 species inner walls were not pitted. The species of of Schoenoplectiella and 14 species of Schoenoplectus sect. Malacogeton had cells Schoenoplectus s. s. (Hayasaka unpublished oblong to narrowly oblong without silica bodies data). For the purpose of this paper, I here (Fig. 1d). Pits were present or absent on the inner present the minimum data on the morphology periclinal walls in this section. of the cells that represent the character states in The above observations suggest that Schoenoplectiella, Schoenoplectus s. s., and their Schoenoplectiella and Schoenoplectus s. s. can sections. The rest of the data will be published be distinguished from each other by the outline elsewhere. shape of the nutlet epidermal cells, which are Interspecific variation in the morphology narrowly oblong to linear in the former and of the cells was detected in the outline shape, isodiametric to narrowly oblong in the latter, as the presence or absence of silica bodies in indicated in the key below in combination with the lumina, and the presence or absence of other characters. The shape of the cells together pits on the inner periclinal walls. Each of with other cellular details further provides the the species whose nutlet epidermal cells are distinguishing characters for the sections of both shown in Fig. 1 is the type of the genera or genera. sections, i. e., Schoenoplectiella articulata is the type of Schoenoplectiella, S. mucronata Interspecific variation in inflorescence gross of Schoenoplectiella sect. Actaeogeton, morphology within Schoenoplectiella Schoenoplectus lacustris of Schoenoplectus, The inflorescences most commonly observed and S. nipponicus of Schoenoplectus sect. in Schoenoplectiella are capitate with a few to Malacogeton. numerous sessile spikelets densely crowded (Fig. The cells in the outline shape observed 2b). In contrast, most species of Schoenoplectus in Schoenoplectiella sect. Schoenoplectiella s. s. have anthelate inflorescences with were uniformly narrowly oblong to linear, branched or unbranched peduncles, with the with the inner periclinal walls lacking pits exceptions of S. subterminalis (Torr.) Soják (Fig. 1a). In this section, silica bodies were having inflorescences of solitary spikelet, 174 植物研究雑誌 第 87 巻 第 3 号 2012 年 6 月

Fig. 2. Inflorescences ofSchoenoplectiella . a. S. lineolata (S. Iwano 1028, TUS). b. S. triangulata (E. Hayasaka 2272, TUS). c. S. komarovii (Y. Murakami s. n., TUS 183236). d. S. lateriflora (S. T. Blake 12626, BRI). Scale bars = 10 mm. and also S. pungens (Vahl) Palla with several Pedunculate inflorescences are also observed other species having capitate or subcapitate in Schoenoplectiella. In S. komarovii, inflorescences. A close examination of the inflorescences are capitate or often shortly inflorescences in Schoenoplectiella also reveals pedunculate on a well-developed tussock, with the interspecific variation within the genus. peduncles terminated by a cluster of several Schoenoplectiella lineolata has inflorescences of sessile spikelets (Fig. 2c). The occurrence of solitary spikelet (Fig. 2a). The reduced structure pedunculate inflorescences in this species has of inflorescences in this species is considered been overlooked in the previous taxonomic as the adaptation to its highly aquatic habit, as accounts (Ohwi 1944: 116, Koyama 1958: in the case of Schoenoplectus subterminalis. 307). Another example is the inflorescences June 2012 Journal of Japanese Botany Vol. 87 No.3 175 of S. lateriflora, which are capitate or often was rooting and vegetatively propagating. lax with peduncles to ca. 35 mm long that are The occurrence of proliferous inflorescences terminated by solitary or often a few-clustered is known in various genera of Cyperaceae, spikelets (Fig. 2d). Jung and Choi (2010) including Cyperus (Bruhl 1995: 276), Eleocharis gave ‘unbranched inflorescence’ as one of the (Ueno et al. 1988), Isolepis (Muasya and distinguishing characters of Schoenoplectiella. Simpson 2002: 259), Scirpus (Strong 1994: 41), The above observations, however, suggest and Trichophorum (Swan 1999: 217). In contrast that the shape of inflorescences is variable to Schoenoplectiella, however, I have not seen within the genus, and thus should be used in proliferous inflorescences in Schoenoplectus s. combination with other characters for defining s. nor any pieces of published information about Schoenoplectiella. them. Although the ability to produce proliferous Occurrence of proliferous inflorescences in inflorescences seems species-specific in Schoenoplectiella Cyperaceae, proliferation has rarely been used In some species of Schoenoplectiella, as a character of taxonomic utility probably inflorescences are sometimes proliferating, i. e. because it is only occasionally observed in most bearing plantlets at the apex of a culm where an of the species. Therefore, the following cases inflorescence develops. I examined a specimen are exceptional. Based on the proliferating of S. clemensiae that bears vegetative plantlets inflorescences and the morphology of at the culm apex together with the ordinary nutlets, Muasya and Simpson (2002, 2005) inflorescence (Fig. 3a). Flowering plantlets recognized Isolepis sect. Proliferae Muasya, growing at the apex of a culm were observed which includes 25 species that are ‘usually in S. hotarui (Fig. 3b), which bear secondary proliferating’. Proliferation is frequent in spikelets normally developed. Similar plantlets Schoenoplectiella gemmifera, a submerged with secondary inflorescences were also found aquatic of Japan, which grows in streams where in S. triangulata, which have immature spikelets the water temperature is stable throughout a (Fig. 3d). Submerged plants of S. gemmifera year. This species vegetatively propagates by growing in running water usually produce means of the proliferous plantlets that grow vegetative, rooting plantlets at the culm apex, at the apex of a submerged culm in running which lack secondary culms but have linear, water (Fig. 3c). The proliferating habit of S. flaccid leaf blades (Fig. 3c). In addition to the gemmifera was used as one of the diagnostic observations above, I have seen proliferating characters of the species (Sato et al. 2004). inflorescences on some specimens of S. erecta Based on the observations above, I stated and S. lateriflora. Furthermore a few species ‘Inflorescences...sometimes proliferous’ in of Schoenoplectiella have been reported to be the description of Schoenoplectiella below. proliferating. Schuyler (1970), in the original The most notable finding about proliferation description of Scirpus heterophyllus Schuyler in this study, however, is that proliferation is (= Schoenoplectiella heterophylla), noted sometimes observed in Schoenoplectiella but not ‘Inflorescentia...saepe prolifera’. This species is in Schoenoplectus s. s., which might imply the a submerged aquatic having ribbon-shaped leaf remote relationship between the two. blades. Schoenoplectiella hondoensis was also observed proliferating (Hayasaka and Ohashi Taxonomic treatment 2000). In the case of this species, the culm Although the remote relationship between bearing a plantlet was the previous year’s that Schoenoplectiella and Schoenoplectus s. s. is was submerged and withered, and the plantlet now evident from the molecular phylogeny, 176 植物研究雑誌 第 87 巻 第 3 号 2012 年 6 月

Fig. 3. Proliferous inflorescences of Schoenoplectiella. a. S. clemensiae (R. Schodde 1758, L). b. S. hotarui (C. Sato 3371, TUS). c. S. gemmifera (C. Sato 3384, TUS). d. S. triangulata (E. Hayasaka 2985, TUS). Arrow-heads indicate the apex of a culm. Scale bars = 20 mm. the two genera should still be compared used to define Schoenoplectus s. l. as well as to morphologically with each other, especially for exclude Actinoscirpus or Bolboschoenus from the purpose of identifying specimens, because it. Such characters include (1) the pseudolateral they share a few characters that have been inflorescences with involucral bract solitary June 2012 Journal of Japanese Botany Vol. 87 No.3 177

Fig. 4. Inner surface of the leaf sheath at the junction with the leaf blade (cut-open at the contralaminar side and flattened) showing the presence or absence of a ligule in Schoenoplectiella and allied genera. a. Actinoscirpus grossus (E. E. Henty NGF49262, L). b. Bolboschoenus fluviatilis subsp. yagara (E. Hayasaka 3279, TUS). c. Schoenoplectiella triangulata (E. Hayasaka 2272, TUS). d. Schoenoplectus triqueter (E. Hayasaka 2143, TUS). l. Ligule. bl. Leaf blade. s. Inner surface of the leaf sheath. h. Hyaline margin of the leaf sheath. Scale bars = 5 mm. and culm-like (vs. inflorescences terminal Actinoscirpus but absent in Bolboschoenus). with involucral bracts several and leaf-like Among these characters, morphology of the in Actinoscirpus and Bolboschoenus, with ligules is discussed below as their presence in the exception of B. planiculmis (F. Schmidt) Schoenoplectiella and Schoenoplectus s. s. needs T. V. Egor. having Schoenoplectus-like to be clearly explained. inflorescences), (2) the glabrous glumes (vs. Ligules are present in the monotypic puberulent in Actinoscirpus and Bolboschoenus), Actinoscirpus, which consists of A. grossus and (3) the presence of ligules (present in with well-developed leaf blades (Fig. 4a). In 178 植物研究雑誌 第 87 巻 第 3 号 2012 年 6 月

Bolboschoenus, in which all the species have the culm-bases, or shortly lying, or elongate and well-developed leaf blades, ligules are always creeping, with or without small fleshy tubers. absent (Fig. 4b). Species of Schoenoplectiella Roots fibrous. Culms erect or spreading, or and Schoenoplectus s. s. have ligules in common submerged with the upper part floating on the when leaf blades are developed at least into water surface, tufted or solitary and distantly short setiform ones (Fig. 4d). When leaf blades arranged in a row, (sub)terete or trigonous, or are absent, there is no structure that could be several-angled, to 138 cm long, 0.2–10 mm wide referred to as a ligule because it is defined as ‘the in the middle, glabrous, rigid or flaccid, nodeless membranous appendage arising from the inner or 1(–3)-noded above the base. Leaves all surface of the leaf at the junction with the leaf basal or upper 1(–3) cauline; sheaths glabrous, sheath’ (Harris and Harris 2000). In that case, lower ones tubular or scale-like, the uppermost however, the apex of a leaf sheath has a hyaline tubular, contralaminar side often hyaline- margin, which I consider is homologous with banded, orifice oblique and hyaline-margined; a ligule (Fig. 4c). Based on these observations, blades reduced to mucro or elongate to ca. ligules are described simply as ‘present’ in the 23 cm, shorter than the culm; ligules present. description of Schoenoplectiella below. Inflorescences pseudolateral with involucral bract continuous from the culm, of solitary Key to distinguish Schoenoplectiella from spikelet or capitate, or shortly pedunculate to Schoenoplectus s. s. lax, sometimes proliferous; involucral bract 1. Nutlets smooth, epidermal cells isodiametric culm-like, (sub)terete or trigonous, or several- to narrowly oblong; rhizome elongate, angled, canaliculate on the adaxial side, erect or creeping or ascending; glumes entire or patent while fruiting, to 75 cm long, transversely minutely notched, or emarginate to bifid at septate or aseptate. Spikelets 1 to more than 300, the apex; perianth segments present; culms sessile or pedunculate, (sub)terete or several- solitary or a few-fascicled, nodeless above the angled; rachilla persistent, not winged; flowers base; perennial ...... Schoenoplectus s. s. hermaphroditic. Glumes spirally arranged 1. Nutlets smooth or transversely rugulose to around the rachilla, cymbiform, membranous to sharply ridged, epidermal cells narrowly chartaceous, each subtending a flower, caducous oblong to linear; rhizome very short and or persistent, 1.2–5.5 mm long, mostly equally hidden among the culm-bases, or shortly sized and shaped within a spikelet, margin lying, or elongate and creeping; glumes smooth or ciliolate, abaxial surface glabrous, entire at the apex; perianth segments present apex entire. Stamens 2–3; filaments glabrous; or absent; culms tufted or solitary, nodeless anthers 0.1–3 mm long. Pistil bicarpellate or or 1(–3)-noded above the base; annual or tricarpellate; styles bifid or trifid, glabrous or perennial ...... Schoenoplectiella minutely papillate, continuous with the ovary, without tubercle at the base. Perianth segments Schoenoplectiella Lye in Lidia 6: 20 (2003); 0–10, setiform, smooth or spinulose, not Beentje, Fl. Trop. E. Africa, Cyperaceae: 24 lengthening after flowering. Nutlets trigonous (2010); J. Jung & H. K. Choi in J. Plant Biol. 53: or plano-convex, or biconvex, 0.7–3 mm long, 229 (2010). smooth or transversely rugulose to sharply Type: Scirpus articulatus L. ridged, apex beaked; epidermal cells narrowly Tufted, amphicarpous or non-amphicarpous oblong to linear, longitudinally elongate. Basal annual, or rhizomatous, non-amphicarpous pistillate flower absent or present at the culm perennial; emergent or submerged, or on the wet bases in axil of leaf sheath: basal flower sessile; ground. Rhizome very short and hidden among style of basal flower bifid or trifid, elongate June 2012 Journal of Japanese Botany Vol. 87 No.3 179 and projecting from the orifice of leaf sheath, cm long, transversely septate (in S. articulata glabrous; basal nutlets inflated, larger than and aliies) or aseptate. Spikelets 1 to more than those from spikelets, surface sculpturing mostly 300, sessile or pedunculate. Glumes 1.2–5.5 similar to that of spikelet nutlet within a species, mm long, margin smooth or ciliolate. Anthers apex beaked; perianth segments of basal flower 0.1–2.4 mm long. Perianth segments absent or absent or present, setiform or scale-like. 1–6 (in S. blakei, S. dissachantha, S. hooperiae, Fifty species worldwide in temperate to and S. supina), setiform. Nutlets 0.7–2.4 mm tropical regions. long, smooth or transversely rugulose to sharply ridged. Basal pistillate flower present or only Key to the sections of Schoenoplectiella rarely present (in S. supina) at the culm bases in 1. Basal pistillate flower often present at the axil of leaf sheath. culm-base in axil of leaf sheath; culms tufted, Twenty-five species worldwide, diversified nodeless or often 1(–3)-noded above the base; especially in Africa, Madagascar, and Australia. leaves all basal or often upper 1(–3) cauline; Growing on freshwater, seasonally wet marshes rhizome very short and hidden among or grasslands, ditches, shore of lakes or the culm-bases; nutlets 0.7–2.4 mm long; temporary ponds, or paddy fields. involucral bract erect ...... Endemism of species in Schoenoplectiella ...... Sect. Schoenoplectiella sect. Schoenoplectiella is particularly notable 1. Basal pistillate flower absent; culms tufted in Africa and Madagascar. Seven species are or solitary (in S. lineolata and S. multiseta), endemic to Africa (S. hooperiae, S. juncea, S. nodeless above the base; leaves all basal; microglumis, S. oxyjulos, S. patentiglumis, S. rhizome very short and hidden among the proxima, and S. raynaliana), five to Madagascar culm-bases, or shortly lying, or elongate and (S. aberrans, S. heterophylla, S. perrieri, S. creeping (in S. lineolata and S. multiseta); reducta, and S. vohemarensis), three to Australia nutlets 1.2–3 mm long; involucral bract erect (S. blakei, S. dissachantha, and S. laevis), two or patent while fruiting ...... Sect. Actaeogeton to North America (S. hallii and S. saximontana), and one to India (S. naikiana). There are Schoenoplectiella Lye sect. Schoenoplectiella also several species widespread in the world. Scirpus L. sect. Supini Cherm. in Arch. Bot. Schoenoplectiella articulata and S. lateriflora are Bull. Men. 3: 193 (1929), syn. nov. widely distributed in Africa, Madagascar, Asia, Type: Scirpus supinus L. and Australia; S. erecta in Africa, Madagascar, Schoenoplectus (Rchb.) Palla sect. Supini Asia, Australia, North America, and South (Cherm.) J. Rayn. in Adansonia, ser. 2, 16: 130 America; S. praelongata in Asia and Australia; (1976). S. roylei and S. senegalensis in Africa and Asia; Tufted amphicarpous annual or short- and S. supina in Africa and Eurasia. lived perennial, emergent or submerged (S. The type of the name Schoenoplectiella heterophylla), or on the wet ground. Rhizome is Scirpus articulatus L. (= Schoenoplectiella very short and hidden among the culm-bases, articulata), while that of the synonym Scirpus without tubers. Culms tufted, to 90 cm long, sect. Supini (= Schoenoplectus sect. Supini) 0.2–8 mm wide in the middle, nodeless or often is Scirpus supinus L. (= Schoenoplectiella 1(–3)-noded above the base. Leaves all basal supina). Comparison in the gross morphology or often upper 1(–3) cauline; blades reduced of S. articulata with S. supina, however, implies to mucro or elongate to ca. 23 cm, shorter than that these two species have a rather remote the culm. Inflorescences of solitary spikelet or relationship with each other. Schoenoplectiella capitate, or lax; involucral bract erect, 0.7–75 articulata is a robust plant with culms to ca. 180 植物研究雑誌 第 87 巻 第 3 号 2012 年 6 月

90 cm long and 3–8 mm wide in the middle, 2, 16: 130 (1976). and involucral bracts transversely septate. Tufted annual or rhizomatous perennial, This species with other five species forms emergent or submerged (S. gemmifera), or the S. articulata complex characterized by on the wet ground. Rhizome very short and the septate involucral bract (Hayasaka 2003). hidden among the culm-bases, or shortly lying, Schoenoplectiella supina on the other hand is or elongate and creeping (in S. lineolata and S. a small and slender plant with culms to ca. 20 multiseta), with (in S. lineolata) or without small cm long and 0.4–2 mm wide in the middle, fleshy tubers. Culms tufted or solitary (in S. and aseptate involucral bracts. This species is lineolata and S. multiseta), to 138 cm long, 0.5– closely related to S. juncea, S. lateriflora, S. 10 mm wide in the middle, nodeless above the microglumis, S. proxima, S. raynaliana, and S. base. Leaves all basal; blades reduced to mucro saximontana, with which it shares trigonous or elongate to ca. 15 cm, much shorter than nutlets rugulose to rugose on the sides, and the culm. Inflorescences of solitary spikelet or aseptate involucral bracts. capitate, or shortly pedunculate (in S. komarovii); After the revision of Schoenoplectus sect. involucral bract erect or patent while fruiting, Supini by Raynal (1976b), two new species from 0.5–28 cm long, aseptate. Spikelets 1 to ca. Africa and one from Australia were described 100, sessile. Glumes 1.8–5.5 mm long, margin and placed in the section (Scholz 1981, smooth or distally minutely ciliolate. Anthers Hayasaka 2003, 2009). North American Scirpus 0.3–3 mm long. Perianth segments 0–10, hallii A. Gray also has become recognized setiform. Nutlets 1.2–3 mm long, smooth or under the section (Smith 1995, 2002). These transversely rugulose to rugose. Basal pistillate species are included in Schoenoplectiella sect. flower absent. Schoenoplectiella in the list below. In addition, Twenty-five species worldwide except S. Scirpus naikianus Wad. Khan of India is America, diversified especially in eastern Asia. transferred to Schoenoplectiella in this paper. I Growing on freshwater marshes, river banks, examined an isotype of it and confirmed that it ditches, paddy fields, sandy to muddy shore of is an amphicarpous species to be included in the lakes, ponds, or reservoirs. Some species are section Schoenoplectiella. troublesome weeds of paddy fields. Species diversity in Schoenoplectiella sect. Schoenoplectiella Lye sect. Actaeogeton Actaeogeton is especially remarkable in China (Rchb.) Hayasaka, comb. nov. and Japan. There are six species endemic to Scirpus L. sect. Actaeogeton Rchb., Fl. China (S. chen-moui, S. chuana, S. fohaiensis, Germ. Excurs. 1: 78 (1830). S. jingmenensis, S. schoofii, and S. subbisetosa), Type: Scirpus mucronatus L. four to Japan (S. gemmifera, S. hondoensis, S. Scirpus L. subgen. Actaeogeton (Rchb.) multiseta, and S. orthorhizomata), and also two Börner in Abh. Naturw. Ver. Bremen 21: 261 to North America (S. purshiana and S. smithii). (1913). A few species have a wide range of distribution. Scirpus L. ser. Actaeogeton (Rchb.) T. Schoenoplectiella juncoides and S. triangulata Koyama in J. Fac. Sci. Univ. Tokyo, sect. 3, 7: are widespread in Asia and the Pacific Islands, 284 (1958). and S. mucronata is widespread in Eurasia. Schoenoplectus (Rchb.) Palla subgen. The recent updates in the taxonomy of the Actaeogeton (Rchb.) Oteng-Yeb. in Notes Roy. section Actaeogeton include the additions of Bot. Gard. Edinb. 33: 315 (1974). two new species and two new varieties to the Schoenoplectus (Rchb.) Palla sect. Schoenoplectiella mucronata complex from Actaeogeton (Rchb.) J. Rayn. in Adansonia, ser. Japan (Kohno et al. 2001, Iokawa et al. 2004, June 2012 Journal of Japanese Botany Vol. 87 No.3 181

Sato et al. 2004, Hayasaka and Sato 2004), and Hayasaka, comb. nov. a new species from Myanmar that is comparable Scirpus naikianus Wad. Khan in Rheedea 8: with S. fuscorubens or S. juncoides (Koyama 71 (1998). 2008). Further study on the Asian species of the Schoenoplectus naikianus (Wad. Khan) M. section is needed because of their great diversity R. Almeida, Fl. Maharashtra 5B: 386 (2009). there. 14. Schoenoplectiella oxyjulos (S. S. Hooper) Lye in Lidia 6: 26 (2003). List of the species included in Schoenoplectiella 15. Schoenoplectiella patentiglumis (Hayasaka) and new combinations Hayasaka, comb. nov. Schoenoplectus patentiglumis Hayasaka in J. (A) Sect. Schoenoplectiella Jpn. Bot. 78: 65 (2003). 1. Schoenoplectiella aberrans (Cherm.) Lye in 16. Schoenoplectiella perrieri (Cherm.) Lye in Lidia 6: 24 (2003). Lidia 6: 26 (2003). 2. Schoenoplectiella articulata (L.) Lye in Lidia 17. Schoenoplectiella praelongata (Poir.) Lye in 6: 20 (2003). Lidia 6: 26 (2003). 3. Schoenoplectiella blakei (Hayasaka) 18. Schoenoplectiella proxima (Steud.) Lye in Hayasaka, comb. nov. Lidia 6: 26 (2003). Schoenoplectus blakei Hayasaka in J. Jpn. var. botswanensis (Hayasaka) Hayasaka, Bot. 84: 14 (2009). comb. nov. 4. Schoenoplectiella dissachantha (S. T. Blake) Schoenoplectus proximus (Steud.) J. Rayn. Lye in Lidia 6: 24 (2003). var. botswanensis Hayasaka in J. Jpn. Bot. 80: 5. Schoenoplectiella erecta (Poir.) Lye in Lidia 161 (2005). 6: 25 (2003). 19. Schoenoplectiella raynaliana (U. Scholz) 6. Schoenoplectiella hallii (A. Gray) Lye in Lye in Lidia 6: 26 (2003). Lidia 6: 25 (2003). 20. Schoenoplectiella reducta (Cherm.) Lye in 7. Schoenoplectiella heterophylla (Schuyler) Lidia 6: 26 (2003). Lye in Lidia 6: 25 (2003). 21. Schoenoplectiella roylei (Nees) Lye in Lidia 8. Schoenoplectiella hooperiae (J. Rayn.) Lye in 6: 26 (2003). Lidia 6: 25 (2003). 22. Schoenoplectiella saximontana (Fern.) Lye 9. Schoenoplectiella juncea (Willd.) Lye in in Lidia 6: 27 (2003). Lidia 6: 25 (2003). 23. Schoenoplectiella senegalensis (Steud.) Lye 10. Schoenoplectiella laevis (S. T. Blake) Lye in in Lidia 6: 27 (2003). Lidia 6: 25 (2003). 24. Schoenoplectiella supina (L.) Lye in Lidia 11. Schoenoplectiella lateriflora (J. F. Gmel.) 6: 27 (2003). Lye in Lidia 6: 25 (2003). 25. Schoenoplectiella vohemarensis (Cherm.) var. laevinux (Lye) Hayasaka, comb. nov. Lye in Lidia 6: 27 (2003). Schoenoplectus lateriflorus (J. F. Gmel.) Lye subsp. laevinux Lye in Nord. J. Bot. 3: 242 (B) Sect. Actaeogeton (1983). 26. Schoenoplectiella bucharica (Roshev.) Schoenoplectiella lateriflora (J. F. Gmel.) Hayasaka, comb. nov. Lye subsp. laevinux (Lye) Beentje, Fl. Trop. E. Scirpus bucharicus Roshev., Fl. SSSR 3: 55, Africa, Cyperaceae: 36 (2010). 457 (1935). 12. Schoenoplectiella microglumis (Lye) Lye in Schoenoplectus bucharicus (Roshev.) Lidia 6: 26 (2003). Grossh., Oraspr. odnodol’n. prishel’chev: 47 13. Schoenoplectiella naikiana (Wad. Khan) (1939). 182 植物研究雑誌 第 87 巻 第 3 号 2012 年 6 月

Schoenoplectus bucharicus (Roshev.) V. Schoenoplectus gemmifer C. Sato & al. in J. I. Krecz., Fl. Uzbekist. 1: 328 (1941), comb. Jpn. Bot. 79: 23 (2004). superfl. 33. Schoenoplectiella hondoensis (Ohwi) 27. Schoenoplectiella chen-moui (Tang & F. T. Hayasaka, comb. nov. Wang) Hayasaka, comb. nov. Scirpus hondoensis Ohwi in Bot. Mag. Scirpus chen-moui Tang & F. T. Wang, Fl. (Tokyo) 45: 189 (1931). Reip. Pop. Sin. 11: 26 & 223 (1961). Schoenoplectus hondoensis (Ohwi) Soják in Schoenoplectus chen-moui (Tang & F. T. Čas. Nár. Muz. Odd. Přír. 141: 62 (1972). Wang) Hayasaka in J. Jpn. Bot. 84: 49 (2009). 34. Schoenoplectiella hotarui (Ohwi) J. Jung & 28. Schoenoplectiella chuana (Tang & F. T. H. K. Choi in J. Plant Biol. 53: 230 (2010). Wang) Hayasaka, comb. nov. 35. Schoenoplectiella jingmenensis (Tang & F. Scirpus chuanus Tang & F. T. Wang, Fl. T. Wang) Hayasaka, comb. nov. Reip. Pop. Sin. 11: 22, 222 (1961). Scirpus jingmenensis Tang & F. T. Wang, Fl. Schoenoplectus chuanus (Tang & F. T. Reip. Pop. Sin. 11: 25, 222 (1961). Wang) S. Yun Liang & S. R. Zhang in Novon Schoenoplectus jingmenensis (Tang & F. T. 20: 170 (2010). Wang) S. Yun Liang & S. R. Zhang in Novon 29. Schoenoplectiella clemensiae (Kük.) 20: 170 (2010). Hayasaka, comb. nov. 36. Schoenoplectiella juncoides (Roxb.) Lye in Scirpus mucronatus L. subsp. clemensiae Lidia 6: 25 (2003). Kük. in Bot. Jahrb. Syst. 69: 259 (1938). var. rockii (Kük.) Hayasaka, comb. nov. Scirpus clemensiae (Kük.) Ohwi in Bot. Scirpus rockii Kük. in Feddes Repert. Sp. Mag. (Tokyo) 56: 203 (1942). Nov. Regni Veg. 16: 432 (1920). Scirpus clemensiae (Kük.) Kük. in Mitteil. Scirpus juncoides Roxb. var. rockii (Kük.) Thuring. Bot. Ver. N. F. 50: 13 (1943), comb. T. Koyama & B. C. Stone in Bot. Mag. (Tokyo) superfl. 73: 290 (1960). Schoenoplectus mucronatus (L.) Palla subsp. Schoenoplectus juncoides (Roxb.) Palla clemensiae (Kük.) Soják in Čas. Nár. Muz. Odd. subsp. rockii (Kük.) Soják in Čas. Nár. Muz. Přír. 148: 194 (1980). Odd. Přír. 141: 62 (1972). Schoenoplectus clemensiae (Kük.) G. C. 37. Schoenoplectiella kandawlayensis (T. Tucker, Fl. China 23: 185 (2010). Koyama) Hayasaka, comb. nov. 30. Schoenoplectiella fohaiensis (Tang & F. T. Schoenoplectus kandawlayensis T. Koyama Wang) Hayasaka, comb. nov. in Makinoa N. S. 7: 55 (2008). Scirpus fohaiensis Tang & F. T. Wang, Fl. 38. Schoenoplectiella komarovii (Roshev.) Reip. Pop. Sin. 11: 23, 222 (1961). J. Jung & H. K. Choi in J. Plant Biol. 53: 230 Schoenoplectus fohaiensis (Tang & F. T. (2010). Wang) Hayasaka in J. Jpn. Bot. 84: 49 (2009). 39. Schoenoplectiella lineolata (Franch. & Sav.) 31. Schoenoplectiella fuscorubens (T. Koyama) J. Jung & H. K. Choi in J. Plant Biol. 53: 230 Hayasaka, comb. nov. (2010). Scirpus fuscorubens T. Koyama in 40. Schoenoplectiella mucronata (L.) J. Jung & Willdenowia 5: 491 (1969). H. K. Choi in J. Plant Biol. 53: 230 (2010). Schoenoplectus fuscorubens (T. Koyama) var. antrorsispinulosa (Iokawa & al.) T. Koyama in H. Hara & al., Enum. Flow. Pl. Hayasaka, comb. nov. Nepal 1: 118 (1978). Schoenoplectus mucronatus (L.) Palla var. 32. Schoenoplectiella gemmifera (C. Sato & al.) antrorsispinulosus Iokawa & al. in J. Jpn. Bot. Hayasaka, comb. nov. 79: 1 (2004). June 2012 Journal of Japanese Botany Vol. 87 No.3 183

var. ishizawae (K. Kohno & al.) Hayasaka, 46. Schoenoplectiella schoofii (Beetle) comb. nov. Hayasaka, comb. nov. Schoenoplectus mucronatus (L.) Palla var. Scirpus shoofii Beetle in Am. J. Bot. 29: 654 ishizawae K. Kohno & al. in J. Jpn. Bot. 76: 228 (1942). (2001). Schoenoplectus schoofii (Beetle) Soják in 41. Schoenoplectiella multiseta (Hayasaka & C. Čas. Nár. Muz. Odd. Přír. 148: 194 (1980). Sato) Hayasaka, comb. nov. 47. Schoenoplectiella smithii (A. Gray) Schoenoplectus multisetus Hayasaka & C. Hayasaka, comb. nov. Sato in J. Jpn. Bot. 79: 322 (2004). Scirpus smithii A. Gray, Man. Bot. ed. 5.: 42. Schoenoplectiella oligoseta (A. E. 563 (1867). Kozhevn.) Hayasaka, comb. nov. Schoenoplectus smithii (A. Gray) Soják in Scirpus oligosetus A. E. Kozhevn. in Bot. Čas. Nár. Muz. Odd. Přír. 141: 62 (1972). Zhurn. 72: 1255 (1987). Schoenoplectus smithii (A. Gray) J. Rayn. in Schoenoplectus oligosetus (A. E. Kozhevn.) Adansonia, ser. 2, 16: 530 (1977), comb. superfl. T. V. Egor., Novosti Sist. Vyssh. Rast. 37: 71 var. leviseta (Fassett) Hayasaka, comb. nov. (2005). Scirpus smithii A. Gray var. levisetus Fassett 43. Schoenoplectiella orthorhizomata (Kats. in Rhodora 23: 42 (1921). Arai & Miyam.) Hayasaka, comb. nov. Scirpus smithii A. Gray f. levisetus (Fassett) Scirpus orthorhizomatus Kats. Arai & Fern. in Rhodora 44: 479 (1942). Miyam. in J. Jpn. Bot. 72: 299 (1997). Schoenoplectus smithii (A. Gray) Soják var. Schoenoplectus orthorhizomatus (Kats. Arai levisetus (Fassett) S. G. Smith in Novon 12: 108 & Miyam.) Hayasaka & H. Ohashi in J. Jpn. (2002). Bot. 75: 224 (2000). var. setosa (Fern.) Hayasaka, comb. nov. 44. Schoenoplectiella purshiana (Fern.) Lye in Scirpus smithii A. Gray var. setosus Fern. in Lidia 6: 26 (2003). Rhodora 3: 252 (1901). var. williamsii (Fern.) Hayasaka, comb. nov. Scirpus smithii A. Gray f. setosus (Fern.) Scirpus debilis Pursh var. williamsii Fern. in Fern. in Rhodora 44: 479 (1942). Rhodora 3: 252 (1901). Schoenoplectus smithii (A. Gray) Soják var. Scirpus purshianus Fern. f. williamsii (Fern.) setosus (Fern.) S. G. Smith in Novon 12: 107 Fern. in Rhodora 44: 479 (1942). (2002). Scirpus smithii A. Gray var. williamsii (Fern.) 48. Schoenoplectiella subbisetosa (T. Koyama) Beetle in Am. J. Bot. 29: 655 (1942). Hayasaka, comb. nov. Scirpus juncoides Roxb. var. williamsii Scirpus subbisetosus T. Koyama in Bot. (Fern.) T. Koyama in Can. J. Bot. 40: 914 Mag. (Tokyo) 86: 281 (1973). (1962). Schoenoplectus subbisetosus (T. Koyama) Schoenoplectus smithii (A. Gray) Soják Soják in Čas. Nár. Muz. Odd. Přír. 148: 194 subsp. williamsii (Fern.) Soják in Čas. Nár. Muz. (1980). Odd. Přír. 148: 194 (1980). 49. Schoenoplectiella triangulata (Roxb.) J. Schoenoplectus purshianus (Fern.) M. T. Jung & H. K. Choi in J. Plant Biol. 53: 230 Strong var. williamsii (Fern.) S. G. Smith in (2010). Novon 12: 107 (2002). 50. Schoenoplectiella wallichii (Nees) Lye in 45. Schoenoplectiella rechingeri (I. Kukkonen) Lidia 6: 27 (2003). Hayasaka, comb. nov. Schoenoplectus rechingeri I. Kukkonen in (C) Hybrids Ann. Bot. Fennici 32: 154 (1995). Schoenoplectiella ×igaensis (T. Koyama) 184 植物研究雑誌 第 87 巻 第 3 号 2012 年 6 月

Hayasaka, comb. nov. Smith at University of Wisconsin-Whitewater Scirpus ×igaensis T. Koyama in J. Fac. Sci. for our collaboration in North American and Univ. Tokyo, sect. 3, 7: 362 (1958). eastern Asian Schoenoplectus. Koji Yonekura at Schoenoplectus ×igaensis (T. Koyama) Tohoku University kindly read the manuscript Hayasaka & H. Ohashi in J. Jpn. Bot. 75: 224 and provided useful comments on it. I thank the (2000). curators of the following herbaria for the loan Schoenoplectiella ×intermedia (Tang & F. T. of specimens or accommodating my visits: A, Wang) Hayasaka, comb. nov. BISH, BM, BRI, E, FLAS, FUK, GH, IBSC, Scirpus ×intermedius Tang & F. T. Wang, Fl. K, KUN, KYO, L, MBK, MHA, MO, NOU, Reip. Pop. Sin. 11: 34, 224 (1961). NU, NY, P, PNH, SAPS, SING, SKK, SNU, TI, Schoenoplectiella ×juncohotarui (Yashiro) TUS, TUSG, UPS, URO, YAMA. Hayasaka, comb. nov. References Schoenoplectus ×juncohotarui Yashiro in J. Bruhl J. J. 1995. Sedge genera of the world: relationships Jpn. Bot. 79: 97 (2004). and a new classification of the Cyperaceae. Aust. Syst. Schoenoplectiella ×oguraensis (T. Koyama) Bot. 8: 125–305. Hayasaka, comb. nov. Egorova T. V. 2005. Synopsis taxonomica generis Scirpus ×oguraensis T. Koyama in J. Fac. Schoenoplectus (Reichenb.) Palla (Cyperaceae) florae Eurasiae borealis. Novosti Sist. Vyssh. Rast. 37: 49–79. Sci. Univ. Tokyo, sect. 3, 7: 362 (1958). Gordon-Gray K. D. 1995. Cyperaceae in Natal. Strelitzia 2: Schoenoplectus ×oguraensis (T. Koyama) 1–218. Hayasaka & H. Ohashi in J. Jpn. Bot. 75: 224 Govaerts R. and Simpson D. A. 2007. World checklist of (2000). Cyperaceae. 765 pp. Kew Publishing, Royal Botanic Gardens, Kew. Schoenoplectiella ×osoreyamensis (M. Kikuchi) Haines R. W. and Lye K. A. 1983. The sedges and rushes Hayasaka, comb. nov. of East Africa. 404 pp. East African Natural History Scirpus ×osoreyamensis M. Kikuchi in Ann. Society, Nairobi. Rep. Gakugei Fac. Iwate Univ. 18: 131 (1961). Harris J. G. and Harris M. W. 2000. Plant identification Schoenoplectus ×osoreyamensis (M. terminology. An illustrated glossary, 2nd ed. 206pp. Spring Lake Publishing, Spring Lake. Kikuchi) Hayasaka in J. Jpn. Bot. 80: 308 Hayasaka E. 2002. Taxonomic revision of the genus (2005). Schoenoplectus (Cyperaceae), with special reference Schoenoplectiella ×trapezoidea (Koidz.) J. Jung to nutlet micromorphology. Doctoral Thesis. 309 pp., & H. K. Choi in Korean J. Pl. Taxon. 41: 27 9 tabs., 53 figs. Graduate School of Science, Tohoku University, Sendai. (2011). Hayasaka E. 2003. A new species of Schoenoplectus Schoenoplectiella ×uzenensis (T. Koyama) (Cyperaceae) from southern Africa. J. Jpn. Bot. 78: Hayasaka, comb. nov. 65–70. Scirpus ×uzenensis T. Koyama in J. Fac. Sci. Hayasaka E. 2009. A new species of Schoenoplectus Univ. Tokyo, sect. 3, 7: 363 (1958). (Cyperaceae) from Australia. J. Jpn. Bot. 84: 13–18. Hayasaka E. and Ohashi H. 2000. Prolification Schoenoplectus ×uzenensis (T. Koyama) in Schoenoplectus hondoensis (Ohwi) Soják Hayasaka & H. Ohashi in J. Jpn. Bot. 75: 224 (Cyperaceae). J. Jpn. Bot. 75: 376–377. (2000). Hayasaka E. and Sato C. 2004. A new species of Schoenoplectus (Cyperaceae) from Japan. J. Jpn. Bot. 79: 322–325. This paper derives from my dissertation Iokawa Y., Kohno K. and Daigobo S. 2004. A new variety submitted to Tohoku University in 2002, with of Schoenoplectus mucronatus (L.) Palla (Cyperaceae) additional data obtained afterward. I thank from Japan. J. Jpn. Bot. 79: 1–3. Jung J. and Choi H.-K. 2010. Systematic rearrangement of Mitsuo Suzuki and Hiroyoshi Ohashi at Tohoku Korean Scirpus L. s. l. (Cyperaceae) as inferred from University for their support and encouragement nuclear ITS and chloroplast rbcL sequences. J. Plant in the graduate school. I am grateful to S. Galen Biol. 53: 222–232. June 2012 Journal of Japanese Botany Vol. 87 No.3 185

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早坂英介：フトイ属から新たに分離されたホソガタホタ ルイ属（カヤツリグサ科）の分類概要 フトイ属 Schoenoplectus (Rchb.) Palla から新たに分 種，雑種の学名の 34 新組合せを発表した．ホソガタホ 離されたホソガタホタルイ属 Schoenoplectiella Lye を タルイ属をホソガタホタルイ節 Schoenoplectiella sect. 独立の属として受け入れ，Lye (2003) の定めたホソガタ Schoenoplectiella と カ ン ガ レ イ 節 Schoenoplectiella ホタルイ属の範囲を拡げてこれまでフトイ属カンガレ sect. Actaeogeton (Rchb.) Hayasaka の 2 節に分類して イ節 Schoenoplectus sect. Actaeogeton (Rchb.) J. Rayn. それぞれに 25 種を含め，節への検索表と節の記載文を およびフトイ属ホソガタホタルイ節 Schoenoplectus 示した．ホソガタホタルイ属を近縁属から区別する形態 sect. Supini (Cherm.) J. Rayn. に分類されていたすべて 形質，およびホソガタホタルイ属とそれに含まれる節を の種を新しい属に含めた．ホソガタホタルイ属の記載文 定義する形態形質について論じた． （福井総合植物園） を改訂してこの属に含まれる全 50 種を一覧し，種，変