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Sect. Actaeogeton, Cyperaceae) and Comparison with Eastern Asian Allies

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Sect. Actaeogeton, Cyperaceae) and Comparison with Eastern Asian Allies New Combinations within North American Schoenoplectus smithii and S. purshianus (sect. Actaeogeton, Cyperaceae) and Comparison with Eastern Asian Allies S. Galen Smith Department of Biological Sciences, University of Wisconsin-Whitewater, Whitewater, Wisconsin 53190, U.S.A. [email protected] Eisuke Hayasaka Botanical Garden, Graduate School of Science, Tohoku University, Sendai, 980-0862 Japan. [email protected] ABSTRACT. The new combinations Schoenoplectus phorum, and Websteria. The largest of these in purshianus (Fernald) M. T. Strong var. williamsii North America is Schoenoplectus with about 70 spe- (Fernald) S. G. Smith, S. smithii (A. Gray) SojaÂk cies worldwide and 17 in North America. var. setosus (Fernald) S. G. Smith, and S. smithii Schoenoplectus is diverse morphologically and var. levisetus (Fassett) S. G. Smith are made. Di- has been divided into several infrageneric taxa agnostic features, mainly from spikelet dimensions, (Oteng-Yeboah, 1974; Raynal, 1976; Smith & Hay- perianth bristles, and achenes, are provided that asaka, 2001). Smith and Hayasaka (2001) recog- distinguish the eastern North American S. smithii nized and provided a key to four sections within and S. purshianus, and the eastern Asian and Pa- Schoenoplectus: sect. Schoenoplectus, sect. Actaeo- ci®c island S. hotarui (Ohwi) Holub, S. juncoides geton (Reichenbach) J. Raynal, sect. Malacogeton (Roxburgh) Palla, and S. komarovii (Roshevitz) So- (Ohwi) S. G. Smith & Hayasaka, and sect. Supini jaÂk, all ®ve of which have been confused with each (Chermezon) J. Raynal. other. A description of Schoenoplectus sect. Actaeo- Upon their transfer from Scirpus to Schoenoplec- geton (Reichenbach) J. Raynal is provided based tus, the infraspeci®c taxa of the two native North mostly on North American and eastern Asian spe- American species of Schoenoplectus sect. Actaeo- cies. geton, S. purshianus (Fernald) M. T. Strong and S. Key words: Asia, Cyperaceae, North America, smithii (A. Gray) SojaÂk, require new combinations. Schoenoplectus, Schoenoplectus sect. Actaeogeton. These species also have been confused with each other as well as with three species of eastern Asia This paper derives from the senior author's work and adjacent western Paci®c islands (Koyama, on his treatment of Schoenoplectus (Reichenbach) 1958, 1962). In addition, the section Actaeogeton Palla for the upcoming volume 23, Cyperaceae, of has never been clearly delineated. This paper ad- the Flora of North America (FNA), during which he dresses these nomenclatorial and taxonomic prob- studied numerous herbarium specimens from about lems. 52 herbaria (Smith, 1995), as well as from the ju- 1. Schoenoplectus purshianus (Fernald) M. T. nior author's ongoing work toward a monograph of Strong, Novon 3: 202. 1993. Scirpus purshi- Schoenoplectus, during which he studied numerous anus Fernald, Rhodora 44: 479. 1942. Scirpus herbarium specimens from the herbaria listed in debilis Pursh, Fl. Amer. Sept. 1: 55. 1814, non the acknowledgments. Scirpus debilis Lamarck, Illustr. 1: 141. 1791. Scirpus L. s.l. has been segregated worldwide in Scirpus erectus Poiret var. debilis Camus, Fl. recent years based on embryo as well as vegetative Gen. Indo-Chine 7: 136. 1912. TYPE: U.S.A. and reproductive characters (Bruhl, 1995; Goetgh- Pennsylvania: Muhlenberg Herbarium, ``No. ebeur & Simpson, 1991; Hayasaka & Ohashi, 65'' and ``m. 33'' (lectotype, designated by 2000; Smith & Hayasaka, 2001; Smith & Yatskiev- Strong (1993: 203), PH, Herb. Muhl. sheet ych, 1996; Wilson, 1981). Nine segregate genera 345±74, photograph seen; isolectotype, PH, will be recognized for the upcoming Flora of North Herb. Muhl. sheet 345±75, photograph seen). America: Amphiscirpus, Blysmus, Bolboschoenus, Is- olepis, Oxycaryum, Schoenoplectus, Scirpus, Tricho- Muhlenberg ®rst published the name Scirpus de- NOVON 12: 106±111. 2002. Volume 12, Number 1 Smith & Hayasaka 107 2002 North American Schoenoplectus bilis without a description in his 1813 catalog, and The names Scirpus smithii A. Gray var. williamsii Pursh apparently appropriated Muhlenberg's earlier (Fernald) Beetle and Scirpus juncoides Roxburgh name. The lectotype and duplicate sheet may have var. williamsii (Fernald) T. Koyama were misap- been seen by Pursh (Merrill & Hu, 1949), or the plied by their authors to the entire species S. pur- specimens may even have been collected by Pursh shianus. Koyama (1963: 457) stated that his new (C. E. Smith, 1962: 458). A. E. Schuyler and J. combination Scirpus juncoides Roxburgh var. digyn- Macklin at PH have kindly provided us with pho- us (BoÈckler) T. Koyama has priority over Scirpus tographs of the types and closely examined the debilis var. williamsii. As Koyama referred to his specimens. On the lectotype sheet, ``Scirpus debilis 1962 paper, he apparently intended his new vari- lupulinus Sm.'' is written on the label; ``345±74'' is etal combination to apply to the whole species S. at the bottom of the sheet. The annotation directly purshianus. BoÈckler's type of the basionym Scirpus on the sheet reads ``Scirpus purshianus Fern. Type supinus L. [var.] digynus BoÈckler needs to be ex- of Scirpus debilis Pursh, AE Schuyler 1976''; this amined to determine the speci®c and varietal iden- sheet bears 15 in¯orescence-bearing culms, 3 of tity of Scirpus juncoides var. digynus (BoÈckler) T. which are attached together at their bases, and 12 Koyama. are without bases. A second sheet bears labels and Perianth bristles absent. Reported habitats: annotations similar to those on the lectotype with shores of lakes, reportedly with substantial water- ``345±75'' written at the bottom of the sheet, and level ¯uctuations. This variety is known only from the annotation on the sheet reads ``Scirpus purshi- the type locality in eastern Massachusetts, several anus Fern. TYPE MATERIAL of Scirpus debilis counties in southwestern Michigan, one county in Pursh, Fl. Am. Sept. 1: 55. 1813[1814]. AES adjacent Indiana, and two counties in Wisconsin. 1976.'' It bears 14 culms with in¯orescences, and is evidently part of the same collection as the lec- 2. Schoenoplectus smithii (A. Gray) SojaÂk, Cas. totype. All culms on both sheets are very similar, NaÂr. Mus. Odd. Prir. 141: 62. 1972, not and several bear achenes with attached bristles, Schoenoplectus smithii (A. Gray) J. Raynal, which are typical of Schoenoplectus purshianus (A. Adansonia, n.s., 16(4): 530. 1977. Scirpus smi- E. Schuyler & J. Macklin, pers. comm. 2001). thii A. Gray, Manual ed. 5: 563. 1867. TYPE: 1a. Schoenoplectus purshianus var. purshi- U.S.A. New Jersey: in tidal mud on the N.J. anus shore of the Delaware below Red Bank oppo- site the River Schuylkill, July 1865, C. E. Perianth bristles 6, stout, tapered distally, about Smith s.n. (holotype, GH; isotype, PH). equaling achene, densely spinulose. Reported hab- itats: shores of inland ponds and ditches. Distrib- 2a. Schoenoplectus smithii var. smithii uted throughout the range of the species in north- eastern North America from Quebec to Georgia Perianth bristles absent or rudimentary. Report- westward to Minnesota, eastern Missouri, and ed habitats: fresh-water tidal shores of rivers and northern Louisiana. inland shores with large ¯uctuations in water level. This variety is known from Quebec (St. Laurence 1b. Schoenoplectus purshianus var. williamsii River near Quebec City) to New Jersey westward (Fernald) S. G. Smith, comb. nov. Basionym: to Minnesota. Scirpus debilis Pursh var. williamsii Fernald, Rhodora 3: 252. 1901. Scirpus smithii A. Gray 2b. Schoenoplectus smithii var. setosus (Fer- var. williamsii (Fernald) Beetle, Amer. J. Bot. nald) S. G. Smith, comb. nov. Basionym: Scir- 29: 655. Oct. 1942. Scirpus purshianus Fer- pus smithii A. Gray var. setosus Fernald, Rho- nald f. williamsii (Fernald) Fernald, Rhodora dora 3: 252. 1901. Scirpus smithii A. Gray f. 44: 479. Dec. 1942. Scirpus juncoides Rox- setosus (Fernald) Fernald, Rhodora 44: 479. burgh var. williamsii (Fernald) T. Koyama, 1942. TYPE: U.S.A. Illinois: Augusta, 1845, Canad. J. Bot. 40: 914. 1962. Schoenoplectus S. B. Mead s.n. (holotype, GH; isotypes, GH, smithii (A. Gray) SojaÂk subsp. williamsii (Fer- NY, PH). nald) SojaÂk, Cas. NaÂr. Mus. Odd. Prir. 148: 194. 1980. TYPE: U.S.A. Massachusetts: Perianth bristles 4 to 6, equaling or slightly ex- sandy shore of Massapoag Lake, Sharon, 7 ceeding achene, densely spinulose. Chromosome Sep. 1901, Plantae Exsiccatae Grayanae 26, number n 5 19 (Schuyler, 1972). Reported habi- E. F. Williams & M. L. Fernald s.n. (holotype, tats: wet places with little ¯uctuation in water level GH; isotypes, BRIT, CAS, GH, ILL, MO, such as mires and shores of small lakes or ponds. NEBC, SAPS, UC, WTU). This variety is known from Quebec (near Quebec 108 Novon City) to Delaware and western North Carolina west- the morphological differences are completely trivi- ward to Wisconsin and Minnesota. In Michigan, al. Wisconsin, and Minnesota it is much more common Schoenoplectus smithii var. setosus, S. smithii var. than S. smithii var. smithii. levisetus, and S. purshianus var. williamsii were ®rst described with no mention of ecological differences 2c. Schoenoplectus smithii var. levisetus (Fas- except for S. smithii var. levisetus, for which Fassett sett) S. G. Smith, comb. nov. Basionym: Scirpus (1921) found the fresh-water tidal habitat remark- smithii A. Gray var. levisetus Fassett, Rhodora able. When Fernald (1942: 483) reduced the vari- 19: 42. 1921. Scirpus smithii A. Gray f. levi- eties to forms he merely stated: ``. their differ- setus (Fassett) Fernald, Rhodora 44: 479. ences are in the presence or absence of 1942. TYPE: U.S.A. Maine: tidal ¯ats of the perianth-bristles, or in the case of S. smithii, forma Cathance River at Bowdoinham and at its levisetus, the lack of retrorse barbs on the bristles.'' mouth in Merrymeeting Bay, 25 Aug.±2 Sep. However, Schuyler (1972: 398±399) stated: ``In 1920, N. C. Fassett s.n. (holotype, GH; isotype, both S. purshianus and S. smithii the presence or NY). absence of barbed bristles appears to be correlated with the stability of the habitat. For example in Perianth bristles 1 to 3 (4), mostly much shorter eastern New York, New Jersey, eastern Pennsylva- than achene, mostly without spinules.
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