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Annales Botanici Fennici 33: 129-136 Ann. Bot. Fennici 33: 129–136 ISSN 0003-3847 Helsinki 18 June 1996 © Finnish Zoological and Botanical Publishing Board 1996 Bolboschoenus yagara (Cyperaceae) newly reported for Europe Jane Browning, Kathleen D. Gordon-Gray, S. Galen Smith & Johannes van Staden Browning, J. & van Staden, J., Natal University Research Unit for Plant Growth and Development, Botany Department, University of Natal Pietermaritzburg, Private Bag X01, Scottsville 3209, Republic of South Africa Gordon-Gray, K. D., Botany Department, University of Natal, Private Bag X01, Scottsville 3209, Republic of South Africa Smith, S. G., Department of Biology, University of Wisconsin-Whitewater, Whitewater, Wisconsin 53190-1790, USA Received 8 January 1996, accepted 26 February 1996 We report the Asian species Bolboschoenus yagara (Ohwi) A. E. Kozhevnikov (Cyperaceae) from Europe, where only B. maritimus (L.) Palla has previously been recorded. Critical differences between these species lie mainly in achene structure, particularly anatomy of the pericarp, and in the persistence of the perianth bristles on the achenes, which we illustrate with SEM micrographs. We also report putative hy- brids (B. maritimus x yagara) on grounds of the intermediate structure of the achenes. The presence of B. yagara and hybrids in Europe bears on the typification of Scirpus maritimus L., which is problematic. We recognize B. affinis (Roth) Drobov pending further research. Key words: Bolboschoenus affinis, B. maritimus, B. yagara, Cyperaceae, Europe, Scirpus, taxonomy INTRODUCTION 1991), in part because of the paucity of reliable macromorphological features available for infrage- neric classification. In particular, B. maritimus (L.) Bolboschoenus (Ascherson) Palla is herein accepted Palla is differently constituted by different workers as a distinct genus following Goetghebeur and in different parts of its range on several continents Simpson (1991). Bolboschoenus is taxonomically (e.g., Koyama 1958, 1980, Browning et al. 1995). difficult (Wilson 1981, Goetghebeur & Simpson Norlindh’s (1972) valuable account for Eurasia drew 130 Browning et al. • ANN. BOT. FENNICI 33 (1996) attention to the variability of the inflorescence, the E Europe and Asia a greater number of species are number of style branches, and the shape of the generally recognised, but there are differences of achenes, all of which have been used frequently in opinion (e.g. Koyama 1958, 1980, Ohwi 1965, the delimitation of B. maritimus or its infraspecific Egorova 1976a, 1976b, Hultén & Fries 1986, taxa. Because of the variability he found, Norlindh Kozhevnikov 1988); nor is the taxonomy of the ge- doubted the value of these characters in the defini- nus stabilized in Africa (Browning & Gordon-Gray tion of species. Robertus-Koster (1969) illustrated 1992, 1993); in North America (e.g. Browning et al. the variability in achene shape for populations in 1995); and in the S Pacific (e.g. Wilson 1981). Holland. Browning and Gordon-Gray (1993), fol- Bolboschoenus yagara (Ohwi) A. E. Kozhev- lowing initial work by Oteng-Yeboah (1974), nikov (basionym Scirpus yagara Ohwi: type from showed that S African plants, previously collectively Japan, Kyoto, Ohwi 9238; holotype KYO, isotype identified as Bolboschoenus maritimus (Browning TNS) has previously been reported only from Asia & Gordon-Gray 1992), were divisible into two enti- (Koyama 1958, 1980, Hultén & Fries 1986, ties on the basis of gross morphology of the achenes, Koshevnikov 1988). Koyama (1958, 1980) reduced pericarp anatomy, and the persistence of the peri- B. yagara to Scirpus (Bolboschoenus) fluviatilis var./ anth bristles on the shed achenes. Subsequently subsp. yagara and Ohwi (1965) placed it in syn- Browning et al. (1995) used these criteria to help onymy under Scirpus fluviatilis. Other authors (e.g. clarify the species limits within Bolboschoenus, in- Hultén & Fries 1986), however, treat B. yagara as a cluding B. maritimus, in the area of the Flora of North distinct species restricted to Asia and B. fluviatilis America (FNA). They showed that B. maritimus and restricted to North America. In this paper we treat B. robustus(Pursh) Soják may be clearly distin- B. yagara as a species distinct from B. fluviatilis guished from other North American species on the pending further research. basis of achene structure and the non-persistence of Bolboschoenus yagara, as characterised by perianth bristles on the achenes. Koyama (1958, 1980) and Ohwi (1965), differs from In W Europe, most taxonomists in recent years B. maritimus and B. affinis. Distinguishing features have recognized only one species in Bolboschoenus, are outlined in the summary that follows. The dif- namely, Scirpus (Bolboschoenus) maritimus (e.g. ferences distinguishing B. yagara and B. fluviatilis Ascherson & Graebner 1904, Suessenguth 1939, from B. maritimus and B. affinis appear to be more Hermann 1956, Schultze-Motel 1967, Robertus- strongly marked in our opinion than are the differ- Koster 1969, Casper & Krausch 1980). They included ences that distinguish B. fluviatilis from B. yagara within the limits of S. maritimus plants with capitate and B. maritimus from B. affinis. All these taxa re- to open inflorescences, bifid or trifid styles, and len- quire more extensive study. ticular to trigonous achenes with caducous to attached Bolboschoenus yagara: Inflorescence compound perianth bristles. Several authors have also pointed or rarely head-like, usually with 3–8 branches each out that B. maritimus occurs both in saline to brack- carrying 2 or 3 spikelets; glumes reddish to darker ish (maritime and inland) habitats and in fresh-water brown; perianth bristles rather strong, remaining at- inland habitats in Europe as described by Hejny tached to mature achenes, mostly ca. equalling (1960), Schultze-Motel (1967), Robertus-Koster achene; styles trifid; achenes nearly equilaterally (1969) and Casper and Krausch (1980). In Flora strongly trigonous, apex tapered to a stout beak Europaea, DeFilipps (1980) recognized only Scirpus 0.2–0.6 mm long. maritimus but with two subspecies, namely subsp. Bolboschoenus fluviatilis differs from B. yagara maritimus with “reddish or dark brown glumes [and] mainly in its larger overall plant size and in its larger nut plano-convex or trigonous”, and subsp. affinis achenes. Achene sizes as given by Koyama (1980) (Roth) T. Norlindh, with “glumes whitish-yellow, are 2.5–3.5 × 1.8–2.2 mm for B. fluviatilis subsp. stramineous or silvery membranous [and] nut len- yagara and 3.8–4.2 × 2.0–2.5 mm for subsp. ticular”. This interpretation clearly follows that of fluviatilis; Browning et al. (1995) give 3.8–5.5 × Norlindh (1972), who stated that he considered 2.0–2.9 mm for North American B. fluviatilis. Scirpus affinis Roth to be only a geographical race Bolboschoenus maritimus: Inflorescence head- of S. maritimus. We herein recognize Bolboschoenus like or with 1 or 2 short branches; glumes reddish or affinis (Roth) Drobov pending further research. For dark brown; perianth bristles weak, caducous, to ANN. BOT. FENNICI 33 (1996) • Bolboschoenus yagara newly reported for Europe 131 Fig. 1. Scanning electron micrographs of achenes. — A–D: Bolboschoenus maritimus (L.) Palla (from Nilsson 9515, H). — E–H: B. affinis (Roth) Drobov (from Kukkonen 12727, H). — I–L: B. yagara (Ohwi) A. E. Kozhevnikov (from Baenitz II Lf. 109, MO). — A, E, I: Abaxial views. B, F, J: Surfaces of exocarp. C, G, K: Trans- verse sections. D, H, L: Transverse sections of pericarp, exocarp at top. Scale bars: A, C, E, G, I, K = 500 µ m. B, D, F, H, J, L = 25 µm. about 2/3 of achene length; styles bifid, or trifid, or mens, Bolboschoenus affinis differs from B. mari- variable in a spikelet; achenes flattened, plano-con- timus mainly in its smaller overall plant size; its pale vex to biconvex to obscurely trigonous to trigonous, whitish-yellow, stramineous or silvery membranous apex abruptly contracted to a mucro 0.2–0.4 mm glumes (generally bright brown in B. maritimus); long. Almost all plants of B. maritimus in North and its consistently bifid styles and lenticular achenes. America have bifid styles and plano-convex to len- The purposes of this paper are to show 1) that ticular achenes (Browning et al. 1995), and all of Bolboschoenus yagara occurs in Europe; 2) that it the rather few specimens we have seen from Asia has previously been incorporated within B. maritimus and South America on which style number or there; 3) that B. maritimus and B. yagara may read- achenes can be observed are digynous. In contrast, ily be distinguished by characters of the achene sur- trifid styles and trigonous achenes are more com- face and pericarp anatomy in addition to the gross mon in Europe and in Africa (Browning et al. 1995). morphological characters given above; 4) and that According to Norlindh (1972), and our unpub- the problem of species differentiation in W Europe lished observations of a limited number of speci- is exacerbated by the presence of putative B. mari- 132 Browning et al. • ANN. BOT. FENNICI 33 (1996) Fig. 2. Scanning electron micro- graphs of achenes of putative Bolbo- schoenus mari- timus (L.) Palla x yagara (Ohwi) A. E. Kozhevnikov hybrids. — A–D: (from Firket s.n., WIS) — E – L: Trigonous and len- ticular achenes (from Schuhwerk 7039/2, NY). — A, E, I: Abaxial views. B, F, J: Surfaces of exocarp. C, G, K: Transverse sec- tions. D, H, L: Transverse sec- tions of pericarp, exocarp at top; endocarp not visib- le in H and L. Scale bars: A, C, E, G, I, K = 500 µ m. B, D, F, H, J, L = 25 µm. timus x yagara hybrids which are recognizable as dissecting microscopes and others using SEM at the Univer- such on the basis of their achene morphology. sity of Natal, Pietermaritzburg, according to the methods de- scribed by Browning and Gordon-Gray (1993). We identi- fied the specimens using the diagnostic characteristics of the MATERIALS AND METHODS achenes as described by taxonomists. Data on these collec- tions are given at the end of this paper.
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