Webbia Journal of Plant Taxonomy and Geography ISSN: 0083-7792 (Print) 2169-4060 (Online) Journal homepage: http://www.tandfonline.com/loi/tweb20 Scirpus L. and related genera (Cyperaceae) in Italy Lia Pignotti To cite this article: Lia Pignotti (2003) Scirpus L. and related genera (Cyperaceae) in Italy, Webbia, 58:2, 281-400, DOI: 10.1080/00837792.2003.10670754 To link to this article: http://dx.doi.org/10.1080/00837792.2003.10670754 Published online: 14 Apr 2013. Submit your article to this journal Article views: 44 View related articles Citing articles: 5 View citing articles Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tweb20 Download by: [Università di Pisa], [L. Peruzzi] Date: 19 February 2016, At: 01:17 Webbia 58(2): 281-400.2003 Scirpus L. and related genera (Cyperaceae) in Italy LIA PIGNOTTI Dipartimento di Biologia Vegetale dell'Universita Via G. La Pira 4, 1·50121 Firenze Ricevuto i/5 Luglio 2003 Accettato I'll Novembre 2003 Scirpus L. e i generi affini (Cyperaceae) in Italia.- Nella presente revisione viene compiuta una valutazione a livello generico, infragenerico e specifico delle entita italiane dd gruppo Scirpus s.l., sulla base di un'analisi morfologica comparata svolta su popolazioni naturali e sugli exsiccata conservati nei principali erbari italiani non­ che alla luce delle pili recenti acquisizioni sulla sistematica di questo complesso gruppo, diffuso su scala mondiale e ricchissimo di specie. Le 22 specie qui ricono­ sciute per il territorio italiano vengono attribuite a 7 generi: Blysmus Panz., Bolbo­ schoenus (Rchb.) Palla, Scirpoides Seg., Scirpus L., Schoenoplectus (Rchb.) Palla, Iso­ lepis R Br., Trichophorum Pers. Le specie italiane di Schoenoplectus vengono attri­ buite ai sottogeneri Schoenoplectus e Actaeogeton (Rchb.) Oteng-Y eb. Le specie ita­ liane di Isolepis vengono attribuite al solo sottogenere Isolepis e aile sezioni Isolepis ed Eleogiton (Link) Pax. Key words: Cyperaceae, Italy, nomenclature, Scirpus s.l., taxonomy. The genus Scirpus L. has been interpreted differently by various authors, ei­ ther in a wide sense as a single, heterogeneous genus (KoYAMA, 1958; ScHULTZE-MoTEL, 1967, 1971; DE FILIPPS, 1980), or in a strict sense, as a small genus segregated from related genera, viz. Bolboschoenus (Rchb.) Palla} Scir­ poides Seg.J Schoenoplectus (Rchb.) Palla, Isolepis R.Br.J Trichophorum Pers. (see e.g., RAYNAL, 1973; WILSON, 1981; HAINES & LYE, 1983; KUKKoNEN, 1996, 1998; GoETGHEBEUR, 1998). Blysmus (L.) Panz. was also included in Scirpus by KoYAMA (1958) and ScHULTZE-MoTEL (1967). KoYAMA (1958) included even Eriophorum L. and Fuirena Rottb. in Scirpus, but his interpretation has never been widely ac­ Downloaded by [Università di Pisa], [L. Peruzzi] at 01:17 19 February 2016 cepted. Eleocharid species, originally included in Scirpus by Linnaeus and seg­ regated under Eleocharis R.Br. (BRoWN, 1810), have been since then appraised as a separate genus by every author. The present work deals with the segregates of Scirpus sensu ScHULTZE-Mo­ TEL (1967), viz. Scirpus s.str., Bolboschoenus, Scirpoides, Schoenoplectus, Iso­ lepis} Trichophorum and Blysmus 1• A segregation of genus-level groups within Scirpus s.l. is here accepted, in accordance with most recent authors' interpretation. 1 This group is referred to as Scirpus sensu lato (s.l.) in the paper. 282 L. PIGNOTTI Such interpretation relies on a large amount of studies which, since nine­ teenth century, have been bringing more and more evidence that Scirpus s.l. is an unnatural genus (WILSON, 1981), "un genre residu" after RAYNAL (1973). Heterogeneity was at first pointed out on a morphological ground, e.g. by BROWN (1810) who segregated Isolepis from Scirpus on the basis of the absence of perianth bristles. Most of Scirpus segregates, born either as genera or as in­ frageneric taxa, date back to the first half of the nineteenth century or little later and are morphologically founded. At the end of the nineteenth century, anatomical studies on culms of Scirpus s.l. were carried out, which confirmed its heterogeneity and exhibited good ac­ cordance with the previous, morphologically settled groups (PALLA, 1888, 1889, 1907). In 1965, VAN DER VEKEN made embryological studies in Cyperoideae. He distinguished six embryo-types, all of which were represented in Scirpus s.l. The occurrence of the embryo-types and relative variants in the segregates of Scirpus s.l. for Italian species is recorded in table 1. A close correspondence be­ tween embryo-types on one hand, and morphological segregates of Scirpus s.l. was pointed out by Vander Veken's studies. An important result was the con­ firmation of the correct positioning of the critical species T richophorum alpi­ num (L.) Pers. in Trichophorum Pers. (sensu SALMENKALLIO & KuKKONEN, 1989) rather than in Eriophorum L. Also embryological differences within Sch­ oenoplectus [between species of sect. Actaeogeton Rchb. and species of sect. Schoenoplectus] and within Isolepis [between sect. Isolepis and sect. Eleogiton (Link) Rchb.] were pointed out by Van der Veken. The author did not make any nomenclatural change, but he gave his opinion on the opportunity that each genus should be represented by not more than one embryo-type, even though different genera could share the same embryo-type. The aim was achievable by simply reappraising several ancient genera, wrongly included in Scirpus L. (VANDER VEKEN, 1965). SEM (scanning electron microscope) studies conducted by ScHUYLER (1971) on the micromorphology of fruit (in particular on inner periclinal walls Downloaded by [Università di Pisa], [L. Peruzzi] at 01:17 19 February 2016 of epidermal cells) also pointed out the heterogeneity of Scirpus s.l. The writer and M. Mariotti Lippi (PIGNOTTI & MARIOTTI LIPPI, 2004 in press) carried out SEM studies on glumes, fruit surface, perianth bristles and pollen morphology of species of Scirpus s.l. from Southwestern Europe. The comparative analysis of these characters, besides giving value to major segregates, pointed out a sharp distinction between Schoenoplectus s.str. and Actaeogeton within Sch­ oenoplectus. Meaningful information came from studies performed by SA VILE (1979) on fungal parasites of Cyperaceae. Some species of rusts of Puccinia and Uromyces SCIRPUS L. AND RELATED GENERA IN ITALY 283 exhibited a genus-specificity within Scirpus, Bolboschoenus, Schoenoplectus and Trichophorum. The occurrence of fungal parasites is to be considered as a pseudocharacter (see e.g., BRUHL, 1995) and thus not more than an aid in plant classification (SAVILE, 1979). Furthermore, Savile's survey was not complete on Scirpus s.l. Nevertheless, a general accordance between these data and the current taxonomy of the group is apparent (WILSON, 1981). Later, classifications of Cyperaceae relying on a large dataset of characters from morphology, anatomy, embryology, phytochemistry, physiology, geo­ graphical distribution and ecology were proposed by BRUHL (1995) and GoET­ GHEBEUR (1998). These studies evidenced heterogeneity even at tribal level within the genera in question. BRUHL (1995) subdivided Cyperaceae in 2 sub­ families and 12 tribes. After his classification, all of the genera in question be­ long to the tribe Scirpeae (subfamily Cyperoideae), but the tribe is defined by the author as polyphyletic and the one "in greatest need of taxonomic effort in Cyperaceae". GoETGHEBEUR (1998) singled out 4 subfamilies and 17 tribes, giv­ ing a narrow circumscription to the tribe Scirpeae. After his classification, the genera at issue belong to 4 different tribes of subfamily Cyperoideae: Scirpeae (Scirpus, Trichophorum), Fuireneae (Bolboschoenus, Schoenoplectus), Cypereae (Isolepis, Scirpoides), Dulichieae (Blysmus). Recently, phylogenetic analysis of DNA sequence data in Cyperaceae was carried out at suprageneric level (MuAsYA et al., 1998), as well as within Scirpus s.l. (MuASYA et al., 2000b), Isolepis (MuAsYA et al., 2001) and Schoenoplectus (YouNG et al., 2002). Nor the tribe Scirpeae, neither Scirpus s.l. are supported as monophyletic groups, and DNA data are in close accordance with embryologi­ cal data. The splitting of Scirpus s.l. in several genera is thus supported by em­ bryological and DNA data. Even Isolepis turned out as not monophyletic, al­ though just minor difference was pointed out between Isolepis and Eleogiton, as it had happened (see table 1) as regarded embryo-types (VAN DER VEKEN, 1965). Isolepis and Scirpoides resulted closer to Cypereae than to Scirpeae (MuASYA et al., 2001), as it had resulted from their Cyperus-type embryo and in accordance with GoETGHEBEUR (1998). Schoenoplectus turned out to be Downloaded by [Università di Pisa], [L. Peruzzi] at 01:17 19 February 2016 polyphyletic too, represented by "two well supported monophiletic clades", one including representatives of section Schoenoplectus, the other one includ­ ing representatives of sections Actaeogeton (Rchb.) Raynal and Supini (Cherm.) Raynal (YOUNG et al., 2002). A combined DNA and morphological analysis performed in Cyperaceae by MuASYA et al. (2000a), which resolved the heterogeneous tribe Scirpeae s.l. in five clades differing from both the current tribal classifications of BRUHL's (1995) and GoETGHEBEuR's (1998), confirmed the extant uncertainty of tribal limits in Scirpeae s.l. 284 L. PIGNOTTI TABLE 1- Embryo-types in Italian species of Scirpus and related genera (after VANDER VEKEN, (1965). Embryo-types Variants of Scirpus s.l. segregates Italian species