Botanical Journal of the Linnean Society, 2008, 156, 385–409. With 10 figures Taxonomy and new taxa of the Carex divulsa aggregate in Eurasia (section Phaestoglochin, Cyperaceae) ANA MOLINA*, CARMEN ACEDO and FÉLIX LLAMAS Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 Department of Biodiversity and Environment Management, University of León, E-24071, León, Spain Received 6 March 2006; accepted for publication 27 September 2007 The Carex divulsa aggregate, belonging to the Carex muricata group in section Phaestoglochin (Cyperaceae), was studied using 60 macro- and micromorphological characters over a wider geographical range than has been attempted previously, and the status or identity of several taxa commonly associated with this aggregate was examined. The results of numerical analysis support the recognition of six new species within the Carex divulsa aggregate, which are also in accordance with their geographical circumscription. Carex cyprica is endemic to Cyprus, C. enokii is from the Mediterranean Basin, C. magacis from the mountains of Spain and France, C. nordica from Northern Europe, and C. egorovae and C. otomana from Eastern Europe and Central Asia. The morphological data were coded and used in a phylogenetic analysis to discover the relationships between taxa. A key to all taxa belonging to the aggregate is included. Diagnostic characters, illustrations, and detailed descriptions of all species are given. Notes on the habitat, distribution, phenology, etymology, and conservation status of each taxon are also presented. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409. ADDITIONAL KEYWORDS: biogeography – Carex muricata group – habitat – morphological characters – numerical taxonomy – phylogenetic analyses – sedges. INTRODUCTION C. coriogyne Nelmes, without a clear position. The C. divulsa aggregate can be separated from the others Carex divulsa and related taxa were placed in Carex by its longer inflorescence with lower spikes not over- section Phaestoglochin by Dumortier (1827), which, lapping and/or hyaline female glumes. later, was named section Muehlenbergiae by Tucker- Carex divulsa was described by Stokes (1787) and man (1843). The section is represented in Eurasia by C. leersii by Schultz (1870). Chater (1980) admitted the C. muricata group, a problematic and scarcely only one species, with two subspecies: C. divulsa ssp. researched group, which can be distinguished by its divulsa and C. divulsa ssp. leersii (Kneuck.) W. Koch. simple inflorescence, rarely with one to two short Nilsson (1985) accepted this taxonomic treatment, but branches at the base, lowest bracts setaceous, and both authors indicated that it must be regarded as androgynous sessile spikes with perigynia plano- provisional because the C. muricata group is in need of convex in cross-section (Clapham, Tutin & Moore, thorough study. Egorova (1999) considered these taxa 1987). As in similar sections (Starr & Ford, 2001), the at the species level. Further studies of C. muricata morphological characters that differentiate species group based on morphological characters (Stoeva & are mostly quantitative and continuous. In studying Popova, 1997; Repka, 2003) resulted in different opin- the C. muricata group in Eurasia (Molina, Acedo & ions related to the taxonomic level of these taxa. One Llamas, 2008a), we have found three main cores problem in most studies and taxonomic classifications corresponding to the C. divulsa, C. spicata, and is the restricted area covered, because most investiga- C. muricata aggregates, and also an isolated species, tions are of local floras. Therefore, to understand and explain this conflicting group, we need to determine the variation over the entire distribution area. *Corresponding author. E-mail: [email protected], Carex divulsa is easy to recognize (David, 1976), [email protected] and can be distinguished without problems. However, © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409 385 386 A. MOLINA ET AL. C. leersii has always been a problematic taxon, PAL, RNG, SANT, SEV, TFC, W, and WU were exam- having a perigynium similar in size to that of C. spi- ined. The herbaria abbreviations are those of cata Huds. (Schultz, 1871: 23) and an inflorescence Holmgren, Holmgren & Barnett (1990). Fieldwork similar in length to that of C. divulsa (Nelmes, 1947: was performed during 2001–05, mainly in the Alps, 101). The identification of this taxon using the avail- and Cantabrian and Pyrenean Mountains, to com- able literature is difficult, because there are discrep- plete the morphological information. ancies between authors about the range of character variation; it seems that each author is talking about a different plant. Historically, there has been much MORPHOLOGY confusion about the nomenclature of this taxon. Later, For the morphological study, 113 specimens repre- Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 it was renamed C. leerseana by Rouy (1912) and senting the entire geographical distribution and mor- C. leersiana by Rauschert (1973). At other times, it phological variation of the aggregate were selected for has been treated as a synonym of C. polyphylla Kar. the numerical and statistical analyses. The charac- & Kir. (Nelmes, 1947; Karjagin, 1952; Kern & Reich- ters used to distinguish the species in other Carex gelt, 1954; Garcke, 1972; David & Chater, 1977; Pig- section Phaestoglochin studies (Webber & Ball, 1984; natti, 1982; etc.), C. guestphalica (Boenn. ex Rchb.) Stoeva & Popova, 1997; Ball, 2002; Repka, 2003; O. Lang (Loos, 1996; Aeschimann et al., 2004), or Molina, Acedo & Llamas, 2006) were checked in a C. chabertii F.W. Schultz (Loos, 1996; Lambinon, group of selected specimens in order to assess their 2004). In addition to the name, the rank is a source taxonomic value. Six additional characters (7, 8, 10, of disagreement amongst taxonomists, who have 26, 43, 48) were included according to our observa- considered it to be subordinated to Carex muricata tions. Finally, 61 variables were selected from the L. (Ascherson & Graebner, 1902; Marshall, 1907; 53 studied characters (Table 1). Characters 1 and 2, Guinochet, 1978; Stoeva & Popova, 1997), C. pairae F. which have no variation inside the C. divulsa aggre- W. Schultz (Vollmann, 1914; Fournier, 1961; Nyárády, gate, were only used in the phylogenetic analysis, 1966; Schultze-Motel, 1968; Garcke, 1972; Soó, 1973), and variable 7 (colour of ripe achene), which was or C. divulsa (Hylander, 1966; David & Chater, 1977; not available in all the studied specimens, was not Chater, 1980; Jermy, Chater & David, 1982; Hooper, used in stepwise discriminant analysis (SDA). The 1985; Nilsson, 1985; Feinbrun-Dothan, 1986; Clapham explanations of some characters and the method of et al., 1987; Aeschimann & Burdet, 1994; Duhamel, measurement are given in Molina et al. (2006). Char- 1994; Luceño, 1994; Sell & Murrell, 1996; Kukkonen, acters 15 and 23, related to beak margin, were sur- 1998, 2002; Lambinon, 2004). veyed with a Jeol 6100 scanning electron microscope. The main objective of this study was to clarify the taxonomic situation. Consequently, new taxa have been revealed with an interesting geographical dis- tribution. As defined here, the C. divulsa aggregate NUMERICAL AND STATISTICAL ANALYSES includes eight taxa – C. divulsa, C. leersii, and Characters were tested for significance using a pair- six new species: C. egorovae, C. enokii, C. magacis, wise t-test (P < 0.01) between 29 specimens of C. di- C. cyprica, C. nordica, and C. otomana. vulsa and 22 specimens of C. leersii. In order to group more than 60 specimens which were difficult to iden- tify, hierarchical clustering with all 113 specimens MATERIAL AND METHODS was carried out to obtain clusters using Gower’s coef- TAXA INCLUDED WITHIN THE SCOPE OF THIS STUDY ficient and unweighted pair group method with arith- All taxa related to C. divulsa and C. leersii mentioned metic averaging (UPGMA) algorithms of SYN-TAX by Vollmann (1903), Kükenthal (1909), Nelmes (1939, 2000 (Podani, 2001). 1947), De Langhe (1944), Maire (1957), Hadac (1961), The data set of all specimens was subjected to a Schultze-Motel (1968), David (1976), David & Chater stepwise discriminant analysis to identify the best (1977), Chater (1980), David & Kelcey (1985), Hooper characters to separate the clusters. Finally, one-way (1985), Nilsson (1985), Malyschev & Peschkova analysis of variance (ANOVA) and the post hoc Tukey (1990), Strid & Tan (1991), Loos (1996), Kukkonen test were conducted on the quantitative characters to (1998), Egorova (1999, 2000), and Repka (2003) were assess the divergence between them, and to identify studied here. the best diagnostic characters for the clusters. Because Levene’s test indicated that several variables showed heterogeneous variance, they were logarithmically TAXON SAMPLING transformed for ANOVA. Qualitative characters were Some 1000 specimens from B, CAG, FCO, FI, GDA- studied by analogous Kruskal–Wallis analysis. These GDAC, H, JACA, JBVN, K, LEB, LISI, LISU, MA, P, statistical analyses were performed with SPSS 13.0. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409 THE CAREX DIVULSA AGGREGATE IN EURASIA 387 Table 1. Studied morphological characters and character states (L, length; W, width; +, yes; -, no) (1) Awn Ǩ glume: 0 -,1+ (2)