Taxonomy, Distribution and Ecology of Bolboschoenus in Europe
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Ann. Bot. Fennici 44: 81–102 ISSN 0003-3847 Helsinki 26 April 2007 © Finnish Zoological and Botanical Publishing Board 2007 Taxonomy, distribution and ecology of Bolboschoenus in Europe Zdenka Hroudová1, Petr Zákravský1, Michal Ducháček2 & Karol Marhold3,4 1) Institute of Botany, Academy of Sciences of the Czech Republic, CZ-252 43 Průhonice, Czech Republic (e-mail: [email protected], [email protected]) 2) National Museum, Department of Botany, CZ-252 43 Průhonice, Czech Republic (e-mail: [email protected]) 3) Institute of Botany, Slovak Academy of Sciences, Dúbravská cesta 14, SK-845 23 Bratislava, Slovak Republic (e-mail: [email protected]) 4) Department of Botany, Charles University, Benátská 2, CZ-128 01 Praha, Czech Republic Received 15 July 2005, revised version received 24 May 2006, accepted 29 Aug. 2006 Hroudová, Z., Zákravský, P., Ducháček, M. & Marhold, K. 2007: Taxonomy, distribution and ecol- ogy of Bolboschoenus in Europe. — Ann. Bot. Fennici 44: 81–102. The five species of Bolboschoenus — B. glaucus, B. laticarpus, B. maritimus, B. planiculmis and B. yagara — occurring in Europe were studied. A detailed taxonomic and nomenclatural account is provided for these taxa, together with an identification key, distribution data and maps based on a revision of herbarium specimens from 36 herbaria and on field observations. Because ecological differentiation influences the distribution of the taxa concerned, detailed information on their ecology is also pro- vided. Key words: Bolboschoenus, Cyperaceae, distribution, Europe, nomenclature, tax- onomy Introduction ican (Browning et al. 1995) and Australasian (Browning et al. 1997) taxa. They reported B. The taxonomy of Bolboschoeneus (Cyperaceae; yagara to be a new taxon for Europe, as well as formerly included in Scirpus) has advanced con- its putative hybrid with B. maritimus (Browning siderably over the last decades. DeFilipps (1980) et al. 1996). treated the taxa now placed in Bolboschoenus Browning and Gordon-Gray (2000) provided within the broadly conceived genus Scirpus. an overview of the Bolboschoenus fruit morphol- Since the treatment by DeFilipps (1980), who ogy. Hroudová and her co-authors dealt with the recognized in Europe only Scirpus maritimus. inflorescence structure (Hroudová et al. 1998a), with two subspecies (subsp. maritimus and fruit anatomy (Hroudová et al. 1997, 1998b), subsp. affinis), the taxonomy has been stud- growth response to different trophic conditions ied extensively in different parts of the world. (Zákravský & Hroudová 1996), chromosome Browning and her co-authors dealt in detail with numbers (Jarolímová & Hroudová 1998) and the South African (Browning & Gordon-Gray ecological differentiation related to plant com- 1993, 1999, Browning et al. 1998), North Amer- munities (Hroudová et al. 1999a) in the central 82 Hroudová et al. • ANN. BOT. FENNICI Vol. 44 European taxa. They reported B. glaucus as a B, BIL, BP, BRA, BRNM, BRNU, GJO, GLM, new species for the Czech Republic (Hroudová GZU, JE, KL, KRA, KRAM, LBL, LD, LE, LI, et al. 1999b) and described a new species, B. LISI, LOD, M, MSB, P, PR, PRC, SAV, SLO, laticarpus (Marhold et al. 2004), corresponding SO, SOM, SZCZ, TRN, UGDA, W, WA, WRSL, to the previously reported alleged hybrid of B. WU, and ZV. Specimens from the private col- yagara and B. maritimus. Hroudová et al. (2001) lections of K. F. Günther, H. Manitz, H. Melzer, and Hroudová (2002) reported for the first time H.-J. Zündorf, specimens collected by us in for central Europe B. koshewnikowii (currently the Czech Republic, Slovakia, Hungary, Austria, synonymous with B. planiculmis; see Egorova & Poland, Germany and Portugal, and specimens Tatanov 2003). collected by our colleagues, namely O. Clever- For the area of the former Soviet Union ing, E. Collias, R. Hrivnák, M. Janišová, V. Egorova and Tatanov (2003) provided detailed Jarolímová, Z. Kaplan, P. Koutecký, F. Krahulec, data on B. planiculmis, and Tatanov (2003a) on W. Lazowski, L. Moravcová, J. Sádlo and J. B. yagara. They also solved some nomenclatural Štěpánková were also studied. The localities problems concerning these two taxa. Tatanov from the Czech Republic included in the maps (2003b) mapped the distribution of B. glaucus were found during a detailed study of the dis- in parts of eastern Europe that belonged to tribution of Bolboschoenus species in the coun- the former Soviet Union. Additionally, Tatanov try (Ducháček 2002, Ducháček et al. 2006). (2003c) provided a short account of Bolbosch- Selected localities representing the distribution oenus in the former Soviet Union, evaluated the of Bolboschoenus species in European countries taxonomic importance of the carpological char- are given in the Appendix. Centres of distribu- acters (Tatanov 2004a), and proposed an infra- tion as well as individual isolated localities are generic systematic arrangement of the genus included. A full list of herbarium specimens (Tatanov 2004b). Nevertheless, a comprehensive studied is available on request from the first treatment of Bolboschoenus for Europe (or Eura- author. In all lists only the localities, from which sia) is still lacking and relevant data are scattered the herbarium specimens with fruiting plants throughout numerous separate papers. were available are included (determination of The present paper aims to provide a syn- specimens without fruits is not reliable). Addi- thesis of the taxonomy and nomenclature of tionally, more detailed surveys of the occurrence Bolboschoenus in Europe (excluding B. affinis of Bolboschoenus in some European countries (= B. popowii), occurring in Europe only mar- were published with full lists of known locali- ginally, in Russia; see Egorova 1976), together ties; such surveys were made for Austria (Hrou- et al. et al. with the distribution data based on revision of dová 2006) and Poland (Hroudová 2005), and are in preparation for Germany, Slo- herbarium specimens and studies of plants in the vakia and Hungary. field. Because ecological differentiation plays Morphological and anatomical characters of a major role in the expansion of plants in new the studied species, and their quantitative ranges, localities and in their disappearance from some are based on measurements of plants from natu- areas, detailed data on the ecology of the taxa are ral populations from the Czech Republic, Slo- also provided. Our study concentrates on central vakia and Hungary, and of cultivated plants Europe, thus the distribution data for the other (results published by Hroudová et al. 1997, parts of Europe are not presented in detail; nev- 1998a, 1998b, 1999a, 1999b) and also on the ertheless, even the scattered data on their occur- studied herbarium material. rence provide important information reflecting their general European distribution pattern. Identification key to European species of Bolboschoenus Material and methods 1. Inflorescence branched, consisting of a central group of The distribution maps are based on the exam- sessile spikelets and of (1–)2–7(–12) rays bearing single ined specimens kept in the following herbaria: spikelets or fascicles of spikelets; rays mostly more ANN. BOT. FENNICI Vol. 44 • Taxonomy, distribution and ecology of Bolboschoenus in Europe 83 than twice longer than sessile spikelets; total number of spikelets on rays higher than or same as number of sessile spikelets; perianth bristles mostly persistent on ripe fruits; fruits (achenes) trigonous or convex with slightly developed edge on abaxial side; trigonous in cross section (rarely nearly flattened or slightly convex to subtrigonous on abaxial side); outer layer of pericarp (exocarp) thinner than mesocarp, formed by isodiametric or slightly elongated cells not always air-filled; achene surface smooth or with fine network of cell outlines (at 20¥ magnification) ....................................................... 2 1. Inflorescence head-like or branched, consisting of a central group of sessile spikelets and of 1–2(–4) rays bearing single spikelets or fascicles of spikelets; rays usually less than twice as long as sessile spikelets; total number of spikelets on rays lower than number of sessile spikelets; perianth bristles caducous; fruits (achenes) not trigonous, but concave, flattened, convex to subtrigonous on abaxial side; biconcave, lenticular, plano-convex to subtrigonous in cross section; outer layer of pericarp (exocarp) thicker than or as thick as mesocarp, formed by elongated cylindrical cells always air-filled; achene surface with a highly visible polygonal network of cell outlines (at 20¥ magnification) .................................... 4 2. Achenes mostly with a clear edge on abaxial side, trigonous in cross section; exocarp layer thin but visibly developed; floral scales never brownish-red or deep pur- plish-red ....................................................................... 3 Fig. 1. Bolboschoenus glaucus (from Košíře, Prague, 2. Achenes with only slightly developed edge on abaxial Czech Republic, (collected in 1983 and cultivated in side, plano-convex to subtrigonous in cross section; Průhonice), 2003 Z. Hroudová, PRA). — a: Inflores- exocarp slightly visible; floral scales frequently brown- cence. — b: Achene (abaxial view). — c: Achene ish-red or deep purplish-red .......................... B. glaucus cross-section. — d: Pericarp layers (ex = exocarp, m = 3. Achenes narrow (1.6–1.8 mm wide), equilaterally trigo- mesocarp, en = endocarp). nous in cross-section; exocarp very thin, formed of more-or-less isodiametric cells .....................