Taxonomy and New Taxa of the Carex Divulsa Aggregate in Eurasia (Section Phaestoglochin, Cyperaceae)

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,

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 distribution. 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 identify, 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.

Table 1. Studied morphological characters and character states (L, length; W, width; +, yes; -, no)

(1) Awn Ǩ glume: 0 -,1+ (2) Auricle leaf blade: 0 -,1+ (3) Spikes contiguous: 0 +,1- (4) Perigynium beak colour: 0 green, 1 brown (5) Lowest spike branched: 0 -,1+ (6) Peduncle lowest spike: 0 -,1+ (7) Ripe achene brown: 0 -,1+ (8) Beginning of anthesis: 0 spring, 1 summer

(9) Scarious margin of female glume: 0 -,1+ laterally, 2 complete Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 (10) Shape spike: 0 globular, 1 globular-elliptic, 2 elliptic, 3 oblong (11) Apex of female glume: 0 rounded-obtuse, 1 acute, 2 apiculate, 3 acuminate (12) Colour Ǩ glume: 0 hyaline, 1 gold-beige, 2 light brown, 3 dark brown, 4 purplish brown (13) Shape perigynium: 0 rounded, 1 elliptic, 2 ovate, 3 trullate, 4 oblong, 5 obovate (14) Apex of beak: 0 bidentate, 1 bifid, 2 variable (15) Beak margins: 0 smooth, 1 serrulate, 2 irregular serrulate, 3 scarcely serrulate (16) Shape achene: 0 rounded, 1 elliptic, 2 ovate, 3 pentagonal, 4 obovate (17) Shape female glume: 0 elliptic, 1 ovate, 2 obovate (18) Perigynium base corky: 0 -, 1 little, 2 +. (19) L leafs/L culm: 1 smaller, 2 similar, 3 larger (20) Veins perigynium body (abaxial): 0 -, 1 up to middle, 2 up to the top (21) Veins perigynium body (adaxial): 0 -, 1 up to middle, 2 up to the top (22) Attenuation perigynium beak: 0 gradually, 1 gradually to abrupt, 2 abrupt (23) Distribution denticles beak: 0 -, 1 only beak, 2 up to summit, 3 over 1/3 body, 4 only summit (24) Position of perigynium: 0 erect, 1 erect-patent, 2 patent, 3 very patent (25) Winged margin of perigynium body: 0 -, 1 from summit to 1/2 body, 2 up to the top, 3 wing variable (26) Orientation of spike: 0 less than 40°, 1 40°-50°, 2 more that 50° (27) L inflorescence: 0 up to 30, 1 30–50, 2 50–70, 3 more than 70 (28) W of widest leaf (29) Diameter of culm (at the middle) (30) L lowest bract (31) L longest inflorescence (32) Distance between lowermost spikes (33) Maximum distance between second and third spikes (34) No. branches first order

(35) No. perigynia in I2

(36) No. male ﬂowers I2 (37) L perigynium (38) W perigynium (39) L perigynium beak (40) Maximum no. abaxial veins (41) Maximum no. adaxial veins (42) L apical tooth (abaxial) (43) No. denticles beak/mm (44) L female glume (45) W female glume (46) L male glume (47) Distance from base to widest point of perigynium (48) Difference (W perigynium - W achene)/2 (49) L achene (50) W achene (51) L longest leaf (52) L apex of female glume (53) W inﬂorescence (54) L perigynium/L Ǩ glume (55) L perigynium/W perigynium (56) L achene/W achene

Table 1. Continued

(57) L perigynium/L beak (58) L perigynium/L achene (59) L longest inﬂorescence/distance between lowermost spikes

(60) No. perigynia/no. ǩ ﬂowers I2 (61) L perigynium/distance from base to widest point of perigynium

PHYLOGENETIC ANALYSIS (putatively employed in some previous studies) are Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 Forty statistically significant variables amongst the misidentified; (2) mistakes are frequent in every taxa were chosen to conduct the phylogenetic analysis taxon belonging to the C. muricata group; (3) sheets (Table 2). As few qualitative characters were available showing sterile and depauperate flowers are difficult for this study, quantitative data were also used, to assign to any taxon; and (4) the taxa are morpho- because they have given good results in other taxo- logically similar, and seem to form a coherent group. nomic studies (Van de Wouw, Maxted & Ford-Lloyd, Keeping all this in mind, the main taxonomic char- 2003). Qualitative characters were coded as in the acters for the C. divulsa aggregate are as follows: numerical analysis. Values of the quantitative char- narrow leaves 2.5–4.5 mm without auricles, culm acters were transformed according to the groups diameter 0.75–2.0 mm, inflorescences 50–155 mm resulting from the post hoc Tukey test (ANOVA), and long, with 4–13 more or less separated sessile spikes, coded in the range of 0–4. Both types of character except for the lowest which sometimes is pedunculate, were treated as unordered multistate, but the four lowest bracts setaceous sometimes longer than their ratios were coded in the range of 0–9 and treated as spikes, acuminate hyaline to brown glumes with- ordered multistate. Carex coriogyne was included out arista, perigynium plano-convex, 3.3–6.0 ¥ 1.8– because it is closely related to the C. divulsa 2.75 mm, sometimes a little spongy tissue at the aggregate, and C. otrubae Podp. [section Vulpinae base, with or without a winged margin, and beak up (J. Carey) Christ.] was used as outgroup because it is to 1.5(1.75) mm. closely related to section Phaestoglochin. All variables According to these data, C. canariensis Kük., his- were equally weighted. An heuristic search was torically linked with this aggregate, does not belong carried out using stepwise addition, and the robust- to the C. divulsa aggregate (section Phaestoglochin). ness of the clades was assessed by means of bootstrap Kükenthal (1900) described it from the Canary analysis. Maximum parsimony analysis was per- Islands, and considered that it was related to C. di- formed for the data set with PAUP* 4.0b10 (Swofford, vulsa. Both were included in Carex section Muehlen- 2002). The data have been deposited in TreeBASE. bergianae (Kükenthal, 1909) a synonym of section Phaestoglochin. Carex canariensis has leaves more than 4 mm wide, inflorescence long and paniculate, DISTRIBUTION usually much branched. Perigynia 2.7–3 ¥ 1–1.4 mm, Specimen label data were used to estimate the geo- strongly veined and convex on both sides, long and graphical distribution of each taxon. Information from serrulate beak, female glumes reddish. Therefore, it Kreczetovicz (1935), Karjagin (1952), Maire (1957), could be included in Carex section Heleoglochin Hylander (1966), Mouterde (1966), Nyárády (1966), Dumort., although some specimens which have an Schultze-Motel (1968), Soó (1973) Chater (1980), inflorescence with few branches may look like Pignatti (1982), David & Kelcey (1985), Greuter, C. divulsa. Matthäs & Risse (1985), Hooper (1985), Meikle In the literature, there is confusion between C. poly- (1985), Nilsson (1985), Feinbrun-Dothan (1986), phylla Kar. & Kir. and C. leersii. Thus, Kükenthal Hultén & Fries (1986), Strid & Tan (1991), Luceño (1909: 156) included C. polyphylla Kar.&Kir.asa (1994), Viney (1996), Kukkonen (1998, 2002), Egorova possible synonym under C. contigua Hoppe. Nelmes (1999), Aeschimann et al. (2004), and Lambinon (1947), Karjagin (1952), and others have considered (2004) were crossed with our own data to infer the C. polyphylla as a synonym of C. leersii. Moreover, real distribution of the taxa. The distribution maps Kreczetovicz (1935) mentioned the ferruginose glumes were plotted using ArcGis 9. of C. polyphylla and its perigynium ovate with long beak, and quoted C. leersii as a synonym; however, according to the different characters listed in the RESULTS AND DISCUSSION protologue by Schultz (1871: 25), it is another plant. Preliminary revision of the herbarium material Hooper (1985) treated C. leersii and C. polyphylla showed the following: (1) often, herbarium specimens as two different species. David & Kelcey (1985)

recognized that plants of C. polyphylla, grown from seed supplied by the Botanic Gardens in Tashkent (Uzbekistan), showed different features from plants of C. leersii. Kükhonen (1998: 184) indicated that materials from Khorassan (Iran) had morphological features different from the variation range of European C. leersii, but was reluctant to separate a different taxon because of the small number of specimens available. Studying chromosome numbers, Toderash (1979) reported 2n = 48 for C. polyphylla, and other authors have counted 2n = 58 (Luceño, 1994; Stoeva Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 & Popova, 1997) or 2n = 57–59 (Hartvig, 1986) in C. leersii. Thus, there are sufficient data to differentiate between C. leersii and C. polyphylla. Further- more, we have already concluded (Molina et al., 2008a) that C. polyphylla does not belong to the C. muricata group s.l. A different case is represented by C. coriogyne Nelmes, which Nilsson (1985) combined as a subspecies under C. divulsa, emphasizing its inflorescence 3–18 cm long. After a meticulous examination of the type specimen conserved at Kew, and other available specimens from Turkey, we can conclude that the type is the only known specimen of C. coriogyne, because the remainder were misidentified. Therefore, C. coriogyne does not belong to the C. divulsa aggregate because of its small inflorescence, shorter than 2.5 cm. It cannot be included in any aggregate of the C. muricata group, and is an isolated taxon. Nelmes (1939) described C. persica Nelmes from

01222122212211212216753 Guilan (Iran), a taxon closely related to C. foliosa D. Don. The only specimen found according to the diag- nosis was the typus conserved at Kew. After a careful study, we verified that C. persica is the same taxon as C. divulsa Stokes, agreeing with Nilsson (1985) and Kükkonen (1998). Schultz (1871) described C. chabertii F. W. Schultz. 2120201301331321332041436709 The original material supporting its description was collected by Chabert in the Drôme and belonged to the Schultz Normale Herbarium, now dispersed in several herbaria. From a study of this material, no 2111001321132112222111222205743 0111001101111300000000000113456 00031021A3433234111333423313571 010300A203433101110110101204452 0411102310100034002323433111291 0213111101211121331111111327614 2211110202211243332323313328634 1311001322222124222132242124643 difference from C. leersii could be found. We also identified specimens belonging to C. spicata Huds. mixed with this material. All material collected by Porta in the Alps (Val de Vestino), labelled as C. chabertii, also belongs to C. leersii; moreover, after several collections, we did not find any different specimens from the Pyrenean range, as indicated by Schultz (1871) and Kükenthal (1909). 000001 1 1 3 1 1 A 1 A A 0 A 000111 1 0 3 001000 0 2 2 000000 0 1 3 00000A0 1 3 001000 0 0 1 Character number 123478 9 101112131415171822242527303233343738394243444546474849525355565761 111101 A2 2 3 2 A 001000 0 1 3 000110 2 1 3 000001 2 0 2 After Schultz (1871), Repka (2003: 245) was the only author who considered that C. chabertii was a Data matrix used in the phylogenetic analysis. Polymorphic characters are indicated by A. Character numbers are the same as in Table 1 different taxon, although he was not sure of its adequate status. In our numerical analysis, specimens labelled C. chabertii were in the same cluster as C. leersii C. nordica C. cyprica C. enokii C. divulsa C. coriogyne Table 2. C. otrubae C. egorovae C. otomana C. magacis C. leersii. Our results are in agreement with those

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409 390 A. MOLINA ET AL. of Loos (1996), who considered C. chabertii as a synonym of C. leersii. The problem is that the name C. chabertii is the legitimate name for C. leersii F. W. Schultz, but that it is invalid because Willdenow (1787) used it previously for the species currently named C. echinata Murray. Taking into account that C. leersii is the name in use, we are preparing a proposal for its conservation (Molina et al., 2008b). The taxonomic rank of C. divulsa and C. leersii has been discussed several times (David, 1976; Chater, 1980; Nilsson, 1985; Stoeva & Popova, 1997; Molina Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 et al., 2006). However, certain differences between the two taxa, such as perigynium colour (Chater, 1980; Jermy et al., 1982; Clapham et al., 1987; Sell & Murell, 1996), leaf width (De Langhe, 1944; Nelmes, 1947; Chater, 1980; Clapham et al., 1987; Sell & Murrell, 1996), ligule (De Langhe, 1944; Nelmes, 1947; Chater, 1980; David & Kelcey, 1985; Clapham et al., 1987; Duhamel, 1994; Sell & Murell, 1996), position of the perigynium (De Langhe, 1944; Chater, 1980; Jermy et al., 1982; David & Kelcey, 1985; Sell & Murell, 1996; Lambinon, 2004), and leaf sheath (David & Kelcey, 1985), have been reported, but we did not find them to be statistically significant. However, the results of a pairwise t-test indicated that C. divulsa must be considered as separate from C. leersii at the species level. Eight quantitative characters and four ratios showed statistically significant differences between C. divulsa and C. leersii, and four qualitative variables were different. The main differences are that C. divulsa has a smaller perigynium (3.25–4 mm) than C. leersii (4.5–5.5 mm), and a longer (5–13 cm) and more slender (6–9 mm) inflores- Figure 1. Beak of perigynium: A, Carex divulsa (LEB cence than C. leersii (3.5–7 cm ¥ 8–10 mm). The 32466); B, C. leersii (LEB 82622). achene size is 2.0–2.5 mm in C. divulsa and 2.2– 3.0 mm in C. leersii. Qualitative characters, such as scarcely serrulate beak margins (Fig. 1), quoted vulsa and C. enokii with longer inflorescences. The already by Nelmes (1947), are unique to C. divulsa. second group includes four species, C. magacis, C. nor- Usually, C. divulsa has the lowest spike pedunculate, dica, C. leersii, and C. otomana, all with shorter inflo- and this feature is not present in C. leersii, which rescences. Finally, the third group includes two taxa rarely has a shorter peduncle. Studying the morpho- with contiguous spikes: C. cyprica and C. egorovae. logical characters, Stoeva & Popova (1997) and The value of dissimilarity separating species is 0.27. Molina et al. (2006) found that C. leersii is the closest Carex magacis appears to be the closest taxon to taxon to C. pairae. Hendrichs et al. (2004), in a phy- C. leersii. We can separate the two species by differ- logenetic analysis, found that C. leersii was closer to ences in the perigynium position and shape, but, as C. muricata than to C. divulsa. We consider that they have a lower dissimilarity value, we prefer to keep C. divulsa and C. leersii are very similar taxa, but all specimens of C. divulsa in the same rank. there are sufficient characters to separate them as A scatter plot of the scores in the discriminant species, although some intermediate specimens may analysis (Fig. 3) shows eight taxa within the C. divulsa be found. aggregate. The first three components explained 75.4% Hierarchical clustering (Fig. 2) shows three main of the variation, and were correctly identified in 100% units separated by the inflorescence length and dis- of cases. The variable with the highest loading on the tance between spikes. The clusters can be differenti- first component was the presence of contiguous spikes, ated by the perigynium size and shape, and glume which separated C. egorovae and C. cyprica from all colour. The first two units join taxa with separate the remaining taxa. Variables with the highest loading spikes. The specimens grouped first belong to C. di- on the second component were scarious margins of the

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409 THE CAREX DIVULSA AGGREGATE IN EURASIA 391 Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021

Figure 2. Hierarchical cluster from unweighted pair group method with arithmetic averaging (UPGMA) using Gower’s coefficient of 113 specimens of the Carex divulsa aggregate and 59 variables.

Figure 3. Scatter plot of scores on axes 1 and 2 of discriminant function analysis of 113 specimens of the Carex divulsa aggregate.

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409 392 A. MOLINA ET AL. female glume, shape of the perigynium, and width of Carex egorovae has contiguous spikes and the short- the inflorescence, which separated C. magacis and est inflorescence, and C. otomana has the perigynium C. otomana. Carex divulsa, C. enokii, C. nordica, and beak, female glume, and ripe achene brown and the C. leersii were separated by the third component of longest perigynium (4.75–6 mm) of the aggregate. length and width of the perigynium. Other important Carex magacis and C. nordica, the most closely characters by load were perigynium beak and female related taxa to C. leersii, can be more easily separated glume colour in the fourth component, and margin by qualitative rather than quantitative characters winged in the perigynium body and length of the (Table 3). They are both unique in having globular inflorescence in the fifth component. spikes tilted more than 50° from the main axis of Obviously, there is no variable that is statistically inflorescence, and a partial or total perigynium significant for every taxon simultaneously in ANOVA, winged body. The perigynium body is trullate in Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 but there is one – maximum number of adaxial veins C. magacis, and ovate to elliptic in C. nordica. Carex – which is not significant. The width of the culm, magacis usually has a light perigynium beak, female length of the longest inflorescence, number of male glume, and ripe achene, whereas in C. nordica they flowers, length and width of the perigynium, and are brown. length of the achene were the best variables, because they clearly separated two or more groups and allowed the identification of a single taxon or a pair PHYLOGENETIC ANALYSIS of taxa. Qualitative variables, such as contiguous The phylogenetic analysis resulted in two single spikes, perigynium beak colour, beginning of anthesis, most-parsimonious trees, with a tree length of 179 scarious margins and shape of the female glume, (Fig. 4). Only 38 characters were informative, with a perigynium base corky, position of the perigynium, consistency index (CI) of 0.65, excluding uninforma- and winged margin of the perigynium, clearly sepa- tive characters, and retention index of 0.51. Carex rate taxa without overlap. Each of the taxa separated coriogyne was included in the C. divulsa aggregate, in the hierarchical cluster and SDA, and identified by forming a well-supported clade with C. enokii (boot- ANOVA and the Kruskall–Wallis test, can be related strap support, 71%). Carex leersii and its closely to a set of character states (Table 3). The best diag- related taxa were also well supported (bootstrap nostic characters for the differentiation of species are support, 86%). Carex egorovae and C. otomana do not the perigynium (shape, size, and wing), beak perigy- appear to be directly related to any other taxon. The nium (colour, length, and denticles), female glume CI value was similar to that of another morphological (colour, shape, and size), and spikes (shape, position study in Carex (Naczi & Ford, 2001). Starr & Ford in relation to inflorescence axis, and distances). Most (2001) obtained a better value including internal are also diagnostic in other aggregates of the transcribed spacer (ITS) sequences. The homoplasy C. muricata group (Molina et al., 2008a). index was quite high (0.35). Other studies with Although hierarchical clustering, SDA, and ANOVA molecular data must be made to confirm the results. support the recognition of six new species in the aggregate, the limits of C. leersii do not seem clear, and several specimens identified under this name could HABITAT AND DISTRIBUTION be hybrids with other taxa. Hybrids are common in Taking into account all the available data and eco- Carex (Cayoutte & Catling, 1992), and several authors logical behaviour of the species belonging to the Eur- (Wallace, 1975; Jermy et al., 1982; O’Mahony, 1989; asian Phaestoglochin section, it can be concluded that Luceño, 1994; Stoeva & Popova, 1997) have docu- they are frequent in hemerophilic habitats, such as mented them in section Phaestoglochin. road sides, pastures, or waste areas (David & Kelcey, Carex cyprica is the most differentiated species in 1985; Aeschimann et al., 2004). Carex divulsa and this aggregate, because of its smaller perigynium C. leersii were reported as pioneers by David & (3–3.5 mm), contiguous spikes, four lowest branches, Kelcey (1985), which is supported by our observa- and large number of flowers per spike. Materials from tions, with a generalist strategy and limited power of Cyprus included some specimens that had separate persistence in dense vegetation. spikes and sterile perigynia, but none was C. divulsa. The entire aggregate occurs in Eurasia, from the The main differences of C. enokii are the largest Iberian Peninsula to Kazakhstan, and the morpho- erect-patent perigynium (4.5–5.5 mm), ovate to oblong, logical variation probably represents adaptation sometimes slightly winged, apex tapering abruptly to different climates. Carex divulsa and the closely into a beak, and beak bidentate with a scariose border. related taxa, C. cyprica and C. enokii (Fig. 5), are Carex otomana and C. egorovae have the perigy- adapted to warm climates. Carex divulsa lives mainly nium ovate, a little corky at the base, with a beak in a Mediterranean climate and is widely distributed longer than the remaining taxa within the aggregate. from Madeira and the Canary Islands (30°W) to

Table 3. Quantitative and qualitative distinctive characters for the Carex divulsa aggregate. Ranges and means ± standard deviation for quantitative characters in millimetres. Within a row, means with different superscripts differ signiﬁcantly (ANOVA, P Յ 0.05) (L, length; W, width)

Character C. divulsa C. cyprica C. enokii C. leersii C. nordica C. magacis C. otomana C. egorovae

Spikes contiguous No Yes No No No No No Frequent Beak colour White White White White to brown Brown White Brown White Beginning of anthesis Spring Spring Spring Summer Summer Summer Spring Spring Shape spike Globular Elliptic Intermediate Intermediate Globular Globular Intermediate Intermediate Shape perigynium Elliptic Elliptic-ovate Ovate-oblong Elliptic Ovate Trullate Ovate Ovate oaia ora fteLnenSociety Linnean the of Journal Botanical Apex beak Bidentate Bifid Bidentate Variable Bifid Bifid Bifid Bifid Shape female glume Ovate Elliptic Ovate Variable Elliptic Elliptic Ovate Ovate Attenuation beak Gradual Gradual Abrupt Gradual Gradual Gradual Gradual Gradual to abrupt Perigynium in ripe Patent Erect-patent Erect-patent Erect-patent Patent Very patent Erect-patent Erect-patent Winged body No No No No Yes Yes No No L inflorescence 50–150 40–75 40–145 35–60 (75) 35–55 (75) 40–60 40–65 (85) 30–50 (68) THE 79.8b ± 25.2 48.7a ± 13.8 79.4b ± 30.2 54.7a ± 13.7 48.2a ± 10.4 51.8a ± 6.4 59.2a,b ± 13.2 44.8a ± 9.2 Distance lowermost spikes 12–45 6–12 12–55 8–27 13–30 14–21 10–17 6–13 (20) AE DIVULSA CAREX 25.6d ± 8.2 8.9a ± 2.3 24.1c,d ± 13.1 16.7a–c ± 4.8 19.3b–d ± 6.0 17.2a–d ± 2.6 13.6a,b ± 2.2 12.7a,b ± 3.9 L perigynium 3.5–4.25 3.0–3.5 4.5–5.5 4.5–5.5 4.0–4.75 4.5–5.5 5.0–6.0 4.25–5.5 3.8b ± 0.3 3.3a ± 0.1 4.8d ± 0.3 4.7c,d ± 0.3 4.4b ± 0.2 4.8c,d ± 0.1 5.5e ± 0.4 4.7c,d ± 0.4 W perigynium 1.75–2.3 1.6–2.0 2.5–2.75 1.75–2.25 1.75–2.5 2.25–2.75 2.0–2.5 1.80–2.5 2.0a,b ± 0.2 1.9a ± 0.1 2.6d ± 0.1 2.1b,c ± 0.2 2.2b,c ± 0.1 2.5d ± 0.1 2.3c ± 0.2 2.0b,c ± 0.2 L beak 0.5–1.0 0.4–0.9 0.75–1.0 0.67–1.25 0.75–1.35 0.75–1.25 1.0–1.6 1.0–1.75 2008, , 0.80a,b ± 0.1 0.7a ± 0.1 0.9a,b ± 0.1 1.0b ± 0.2 1.0b ± 0.2 1.0b ± 0.1 1.3c ± 0.2 1.3c ± 0.2

No. denticles beak/mm 2–14 8–14 8–18 10–18 12–20 14–18 14–18 4–16 EURASIA IN AGGREGATE a a,b a–c c,d b–d e c,d a–c

156 8.5 ± 3.8 10.8 ± 1.9 11.5 ± 3.6 15.8 ± 2.1 15.1 ± 2.7 17.0 ± 1.4 15.5 ± 1.2 12.2 ± 5.9 L female glume 3.0–4.0 2.75–3.25 3.5–5 3.0–4.5 2.75–3.75 3.0–3.6 3.25–5.25 3.0–4.0 385–409 , 3.4a,b ± 0.4 3.0a ± 0.2 4.0c,d ± 0.5 3.6b,c ± 0.4 3.3a,b ± 0.3 3.3a,b ± 0.2 4.2d ± 0.6 3.45a,b ± 0.3 W female glume 1.75–2.3 1.2–2.0 2.0–2.5 1.6–2.3 1.5–2.2 2.0–2.5 1.75–2.3 1.6–2.25 2.0b,c ± 0.1 1.7a ± 0.2 2.2c ± 0.2 1.9b,c ± 0.1 1.8b ± 0.3 2.2c ± 0.03 2.1c ± 0.3 1.8b ± 0.1 Distance from base 1.25–1.75 0.75–1.25 1.75–2.12 1.5–1.87 1.37–1.75 1.5–2.12 1.5–2.25 1.12–1.75 1.5b ± 0.1 1.0a ± 0.2 1.9e ± 0.1 1.65b–d ± 0.1 1.6b,c ± 0.1 1.8c–e ± 0.2 1.8d,e ± 0.2 1.5b ± 0.1 L achene 2.0–2.5 1.8–2.2 2.5–3.0 2.2–3.0 2.0–2.75 2.5–3.0 2.5–3.0 2.0–2.7 2.2b ± 0.2 2.0a ± 0.1 2.8e ± 0.2 2.6d ± 0.2 2.5c,d ± 0.1 2.7d,e ± 0.1 2.8e ± 0.2 2.3b,c ± 0.2

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Figure 4. Phylogram based on 40 characters of the relationship between taxa of the Carex divulsa aggregate using an heuristic search with C. otrubae as outgroup. Bootstrap support is indicated on the branches. The tree is 179 steps long with a consistency index of 0.65 and a retention index of 0.51, excluding uninformative characters.

Figure 5. Geographical distribution of Carex divulsa ( ), C. cyprica (᭹), and C. enokii (ଙ). south-east of the Caspian Sea (55°E), and from the Britain to Sweden, and scattered in Central Europe. coastal strip of northern Africa (Algeria and Morocco, Carex magacis occurs in a small area in Pyrenean and 30°N) to Northern Europe (45°-52°N). Carex enokii other Iberian Mountains (Spain, Andorra, and France). inhabits areas throughout the southern and eastern Carex otomana and C. egorovae occur from Eastern Mediterranean Basin, from Mauritania to Turkey. Europe to Middle Asia (Fig. 6). Carex otomana occurs Carex cyprica is endemic to Cyprus. Carex leersii and from the east of Bulgaria and Greece to Tyan Shan its closely related taxa, C. magacis and C. nordica (Kazakhstan), and C. egorovae in three disjointed (Fig. 6), are distributed in the Atlantic temperate areas from Hungary to Iran. regions of Europe, but are not common in Southern Europe. They are adapted to the cold conditions of Northern and Middle Europe. Carex leersii has a ORIGIN narrower distribution than C. divulsa, from northern No data are available on the origin and evolution of Spain to the south of the Scandinavian Peninsula and this aggregate. The eastern part of the Mediterranean to eastern Turkey. Carex nordica is limited to a strip Basin (Greece, Turkey), where most taxa of the group zone in Europe between 50° and 60°N from Great live, could be a hot spot of diversity (Médail & Quézel,

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Figure 6. Geographical distribution of Carex leersii ( ), C. magacis ( ), C. nordica (ଙ), C. otomana ( ), and C. egorovae (᭹).

1997) and the speciation zone of the aggregate. SECTION PHAESTOGLOCHIN DUMORT. FL. BELG. 146. According to Nelmes (1947), C. leersii could be the 1827. (LECTOTYPE DESIGNATED BY JONES (1994): oldest taxon, from which C. muricata and C. divulsa CAREX MURICATA L.) have diversified. In the phylogenetic tree, C. leersii is 1. Carex divulsa Stokes in With., Bot. Arr. Brit. Pl. also separated before C. divulsa and its relatives. ed. 2, 2: 1035. 1787. (Fig. 7A–F) Carex cyprica was putatively segregated from C. di- = Carex canescens sensu Huds., Fl. Angl.: 405. 1778, vulsa. A reduction in the internode length and an non L. 1753. increase in the number of flowers per spike might = Carex virens Lam., Encycl., 3: 384. 1792. have led to the evolution of C. cyprica. Its isolated = Carex divulsa Gooden., Trans. Linn. Soc. 2: 160. distribution has avoided introgressions with C. di- 1794. vulsa. According to the phylogenetic analysis, C. cori- = Carex guestphalica (Boenn. ex Rchb.) O. Lang, Flora ogyne is the closest species to C. enokii, and they both 26: 147. 1843. Neotype (designated by Loos): Feddes have similar perigynia; however, their relation to Repert. 107(1–2): 74. 1996 (MSTR!). C. divulsa is not clear. Carex nordica and C. magacis = Carex persica Nelmes, Bull. Misc. Inform. Kew could be of hybrid origin between C. leersii and other 1939: 185. Holotype (designated by Nelmes): Bull. taxa of the C. muricata group. Misc. Inform. Kew 1939: 185. 1939. (K!). In conclusion, if Carex originated in the early Ter- tiary and the subgenus Vignea evolved in cooler climates, probably in the mid-Tertiary (Ball, 1990), it is Lectotype (designated here): [icon] ‘Carex nemorosa, possible that the C. muricata group may have shown fibrosa radice, caule exquisite triangulari, spica longa rapid diversification, and may have adapted to differ- divulsa seu interrupta, capitulis solitariis praeter- ent climatic conditions, helped by its opportunist quam ultimo’ in Micheli, Nov Pl. Gen.: t. 33, f. 10. strategy. The recognition of six new species in the 1729. – Epitype (designated here): ‘Gram. Cyp. spica- C. divulsa aggregate is the result of an in-depth study tum minus spicâ longâ/divulsâ seu interrupta Ray of the entire C. muricata group in a broad geographi- 1279. 2 Petiver’ (No. 17274, FI-M!). cal context. Other studies may discover more diversity, mainly in the eastern distribution area because Description: Caespitose perennial tufted plant with or of the lack of sampling. Hybrids and morphological without short rhizomes c. 6 mm in diameter. CULMS intermediates often occur, and isolated reproduction 15–70 cm ¥ 0.8–1.25 mm at mid-height, obtusely trigo- is not effective, but our results show that there nous, slightly scabrid above; sheaths pale to dark are several species supported by morphological brown. LIGULE 1.25–4.0 mm long, wider than long, characters. usually rounded at apex. LEAF blades about three-

TAXONOMIC TREATMENT KEY TO THE EURASIAN TAXA OF THE CAREX DIVULSA AGGREGATE 1. Inflorescence with spikes contiguous, only the lowest separated; glumes pale...... 2 1. Inflorescence with more spikes separated...... 3 2. Inflorescence with three to four basal spikes compound, perigynia 3–3.5 mm long, short beak (0.7–1 mm), and apical teeth 0.2–0.5 mm long...... 2. Carex cyprica 2. Inflorescence with zero to two basal spikes compound, perigynia more than 4 mm long, long beak (1–1.75 mm), and apical teeth 0.6–1.0 mm long...... 8. Carex egorovae 3. Perigynium completely or partially winged ...... 4 3. Perigynium not winged...... 5 Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 4. Perigynium elliptic, 4–4.75 mm long, female glume light to dark brown ...... 5. Carex nordica 4. Perigynium trullate, 4.25–5.5 mm long, female glume hyaline to light brown ...... 6. Carex magacis 5. Perigynium 3.5–4.25 mm long ...... 1. Carex divulsa 5. Perigynium 4.25–6 mm long...... 6 6. Perigynia 4.75–6 mm long, slightly corky at base, and brown glumes ...... 7. Carex otomana 6. Perigynia 4.5–5.5 mm long, not corky at base, and pale glumes...... 7 7. Perigynium ovate to oblong, abruptly contracted in a beak...... 3. Carex enokii 7. Perigynium elliptic to ovate, gradually tapering in a beak ...... 4. Carex leersii

Figure 7. Carex divulsa (A, from GDA 9116; B–F, from GDA 38629): A, inflorescence; B, male glume with anther; C, female glume; D, perigynium, abaxial side; E, achene; F, perigynium with abaxial side removed to show achene. Carex cyprica (from holotypus MA 495407): G, inflorescence; H, male glume with anther; I, female glume; J, perigynium, abaxial side; K, achene; L, perigynium with abaxial side removed to show achene. All figures drawn by X. Magaz. Scale bar, 1 mm.

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409 THE CAREX DIVULSA AGGREGATE IN EURASIA 397 quarters or as long as culms, dark green to grey green, Observations: Carex divulsa presents some variation widest leaf blades 2–3 mm, plicate to flat. INFLORES- throughout its distribution area. In Europe, the peri- CENCE oblong, 50–135(150) ¥ 6–9 mm, with 7–12 gynium beak is short, bidentate, and very scarcely globular sessile and unbranched spikes, except the serrulate. It is longer towards the east, almost lowest that is usually branched and pedunculate. bifid and more regularly serrulate. Inflorescences are Lowest internode 12–45 mm long, the second one usually long, but near the Mediterranean coast are 6–28 mm. BRACTS glume-like, the proximal setaceous, sometimes shorter (even less than 4 cm), probably shorter than inflorescence, 6–50(-63) mm long. caused by introgression with C. pairae. Specimens FEMALE GLUME hyaline to golden-yellow, with a midrib with a rounded perigynium body are sometimes found green, 3.0–4.0 ¥ 1.75–2.30 mm, ovate, apex acuminate in Southern Europe. Intermediates between C. di- Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 0.15–0.3 mm long. STAMINATE GLUME 2.75–3.5 ¥ 1.4– vulsa and C. leersii have been named C. divulsa var. 2.0 mm, similar to the female. ANTHERS 3, 1–2.5 mm polycarpa Vollmann (Lambinon, 2004). Carex divulsa long. PERIGYNIUM patent to erect-patent, greenish, is also sympatric with C. magacis, C. egorovae,and body elliptic to rounded, nerveless or faintly veined, C. enokii. In mixed populations, it maintains its dis- 3.5–4.0(4.25) ¥ 1.75–2.30 mm, acute at base, gradually tinctiveness, with the perigynia of C. divulsa being tapered into a beak. BEAK 0.5–1.0 mm long, white slightly larger or having a longer beak. It is allopatric distally, with scarcely serrulate margins, apex biden- with C. cyprica, C. leersii,andC. otomana. According tate, apical teeth 0.3–0.5 mm. ACHENE elliptic to to Nelmes (1947), when this species is located on rounded, shining brown when ripe, 2.0–2.5 ¥ 1.5– calcareous soils, the perigynia are slightly larger. We 2.25 mm. STIGMA 2, 2.0–3.0 mm long. studied the variation of the perigynium length in calcareous soils and found no clear relationship: longer and shorter perigynia were found on both Phenology: Flowering begins in April, fruiting from calcareous and noncalcareous soils. May to September, wind pollinated. All of our data indicate that C. divulsa flowers 1 month before Representative specimens examined: FRANCE. Corsica, C. leersii, disagreeing with Jermy et al. (1982) and Evisa, kastanienhain oberhalb des Ortes, 900 m, David & Kelcey (1985), who reported it 1 month later. 23.vii.1956, W. Rössler (MA 274203). Haute-Corse, Ghisonaccia, bord N-198, bosquet rudéral d’Eucalyptus, 1.vi.1997, J. Lambinon (MA 595765). Habitat: Carex divulsa seems to need some moisture Haute Garonne, Rieux Volvestre camping, sombra in the soil. It grows in shady habitats, such as forest sotobosque de Quercus pubescens, 600 m, 14.vii.2002, and disturbed places (Guinochet, 1978; Chater, 1980; A. Molina (LEB 78511). CROATIA. Dalmacia, Lapad Clapham et al., 1987), but also in sunny open habi- peninsula, edge of path round end of Peninsula, tats (David & Kelcey, 1985). In the latter case, speci- 19.iv.1935, A. K. Jackson & W. B. Turrill 132 (K). mens are harder. Carex divulsa occurs at low GREECE. Crete, Dist. Chania, in saxosis schist, altitudes (0–700 m), although in the warmest, south- humidis inter Skines & Nea Rumata, 300 m, 29.v.1942, ern part of the distribution area it can be found at K. H. Rechinger fill. 13384 (K). IRAN.5′S of Chalus, 1000–1200 m. Greuter et al. (1985) quoted 1800 m. Caspian rain forest, 500 ft, 27.vi.1962, P. Furse 2888 According to David & Kelcey (1985) and our own (K). E. Mazandaran, Centre, 5 km W of Tangegol, observations, it tolerates a wide range of soil acidity c. 1 km NE Cheshme-e-, NE Cheshme-e-Adam- (pH 5.3–7.4). Chaqran (spring), in Crataegus forest (Golestan National Park), 980 m, 37°24′N, 55°54′E, 3.viii.1994, H. Akhani 9799 (H 1695179). Gongân, Jangal-e- Distribution: Carex divulsa is the most common Golestân, Tang-e-Râh, 580 m, 17.v.1976, Termeh & species of the group, widely distributed (Fig. 5) from Matin 34797 (H 1585245). North Persia, ad radices m. Madeira to Iran (55°E). It is frequently found in Elbrus, pr. pagum Darake, umbrosis. 25.iv.1843, C.G. Mediterranean areas in Spain, France, Italy, Greece, T. Kotschy 76 (P 00281847). IRELAND. Limerick, Srool- Turkey and the islands (Balearic, Corsica, Sicily, and ane near Foynes, hedgebank, 22.vi.1952, R. B. Drum- Crete), northern Africa (from Morocco to Algeria), and mond 736 (K). ITALY. Toscana, Firenze, waste ground, in the Canary and Madeira Islands. It is scarce in 24.v.1964, R. K. Brummitt 4554 (K). MOROCCO.pr.El temperate zones, and very scarce towards the north- Araix, el Guedira dicto, in locis humidis, 20.v.1930, east of Europe, for example it is rare in Belgium Font Quer (GDA-GDAC 02646). Vertiente mediterra- (Lambinon, 2004). In Russia, it appears only in the nea del Gurugu, sobre la segunda caseta, 1.vi.1915, south-west and Caucasus region, and it extends from A. Caballero (MA 017149). Zaian, maison forestière Iran to south-east of the Caspian Sea. No specimens d’Assoual, ravine humide au pied de la maison, have been found in Iraq. 26.iv.1952, Ch. Sauvage (SANT 48676). NETHER-

LANDS. Limburgh, Plasmolen at Mook, roadside, ish, body elliptic to ovate, nerveless to well-marked 1.v.1926, Herb. Kern & Reichgelt (J.K&B.R)(K). veins, 3.0–3.5 ¥ 1.6–2.0 mm, rounded base, gradually ROMANIA. Dolj, Oltenia, rivuli Jiu, pagum Bratovoesti, to abrupt tapered into a beak. BEAK 0.4–0.5(0.9) mm silvis humidis Q. robori–Carpinatum, 65 m, 6.vi.1971, long, white distally, with serrulate winged margins D.&Mariana Cirtu (MA 236897). SPAIN. Albacete, reaching the summit, apex bifid, apical teeth Villapaladios, 14.vi.1993, G. Aragón et al. (LEB 0.20–0.50 mm. ACHENE elliptic to ovate, shining, 52179). Almería, Sierra de Gador, Fondón, praderas, light greenish to brown, 1.8–2.25 ¥ 1.5–1.75 mm. herbazales, acequias y arroyos, ambiente muy STIGMATA 2, 1.5–2.5 mm long. húmedo, 900 m, 14.vi.1992, A. Hervás (GDA 38269). Barcelona, Massif du Tibidabo à S. Genis, 30.v.1912, Etymology: The new species is named from Cyprus, F. Senen (MA 016815). Cáceres, El Portanchito, where this plant is endemic. Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 29SQD27, 7.v.1983, M. Ladero & M. Santos (LEB 24812). Granada, Orgiva, salida del pueblo, borde de Phenology: Flowering begins in April, fruiting from acequia, 8.iv.1980, M. Ladero & J. Molero (GDA 9116). May to August, wind pollinated. León, Lago Sumido, borde lago, 700 m, 29TPH826037, 1.v.2002, A. Molina (LEB 79002). Pombriego, Habitat: Carex cyprica occurs in warm and humid 29TPG89, 7.v.1985, I. Jiménez (LEB 32466). Mallorca, places, near brooks, springs, or streams between 200 Pont d’Inca, Torrent, herbazal ruderalizado, 6.v.1987, and 1200 m, and is indifferent to soil composition. L. Villar & C. Chocarr (JACA 028187). TURKEY. Zon- guldak, Kel tepe above Yenice, Abies–Fagus forest, Distribution: Endemic to Cyprus (Fig. 5). 1200 m, 20.viii.1962, Davis, Coode & Yaltarik 37793 (K). UNITED KINGDOM. 41 Glamorgan, between Observations: Specimens from Cyprus identified as Melling Griffith and Tongwylais, dry bank in canal, C. divulsa by Meikle (1985) do not agree with the 22.vi.1949, E. Nelmes 960 (K). diagnostic characters of C. divulsa. Meikle recognized problems in the taxonomy of the complex, but stated, 2. Carex cyprica A. Mol., Acedo & Llamas, ‘The temptation to describe another closely allied sp. nov. (Fig. 7G–L) taxon is resisted because much remains to be discov- Carice divulsa similes, sed ab ea differt inflorescentia ered of the citology and crossing behaviour of this brevioribus, 4–5 cm longitudinis, spicae elipsoideus, group’. multiflorae; spicae basarum proximus, perigyniis brevioribus, 3–3.25 mm longitudinis. Representative specimens examined: CYPRUS. Near Inia, Akamas, quartzite, growing in the crevices, Holotype: Cyprus, Distr. Paphos, station forestière 8.iv.1957, L. F. H. Merton 3021 (K). Distr. Kyrenia, Paphos, Ayia, milieu anthropisé humide, sous-bois de Laphitos, loco umbroso juncto rivulum, 13.vi.1939, H. Platane, 650 m, 23.iv.1991, Iter Mediterraneum IV Lindberg iter 1939 (K) (H 1302858) and (H 1302856). 1284 (MA 495407!) [sub. Carex muricata L. ssp. lam- Distr. Limassol, foot of Aphamia, on calcareous rock, procarpa]. Isotypes: B!, FI!, JBVN!, PAL!, RNG?. 2900 ft, 20.v.1939, E. W. Kennedy 1739 (K). Between Platres & Aphamis, by a spring in garigue, 946 m, Description: Caespitose perennial tufted plant with 22.iv.1941, P. H. Davis 3192 (H 1205849). Distr. short rhizomes c. 4 mm in diameter. CULMS 28– Prodhromos, in nemorosis, 4500 ped, 1862, Ms. Merii 75 cm ¥ 1.0–1.5 mm at mid-height, obtusely trigo- & T. Kotschy 855 (K). Ytronas river, by stream, nous, slightly scabrid above; sheaths pale to dark 4000 ft, 3.v.1962, R. D. Meikle 2828 (K). Ayia valley, brown. LIGULE 0.5–1.25 mm long, wider than long, roadside by a stream in the forest, 2000 ft, 28.ix.1936, rounded at apex. LEAF blades pale green about L. F. H. Merton 2831 (K). Mandria, on igneous rock, three-quarters to as long as culms, widest leaf blades 3000 ft, 21.iv.1939, E. W. Kennedy 1438 (K). 3–3.5 mm, plicate to flat. INFLORESCENCE oblong, 40–75 ¥ 8–9 mm, with 10–14 elliptic, sessile spikes, 3. Carex enokii A. Mol., Acedo & Llamas, the first four branched. Lowest internode 6–12 mm sp. nov. (Fig. 8A–F) long, the second one 6–8 mm, other spikes more Carice divulsa similes, sed ab ea differt perigynia crowded. BRACTS glume-like, the proximal setaceous, maioribus est, 4.5–5 mm longitudinis, oblonguis shorter than inflorescence, 6–18 mm long. FEMALE formam habens atque in rostrum abruptim contractas. GLUME hyaline to yellow-golden, with a midrib green, 2.75–3.25 ¥ 1.2–2.0 mm, elliptic, apex apicu- Holotype: Planta Lybicae ex Herbarium Universitatis late 0.2 mm long. STAMINATE GLUME 2.5–3.0 ¥ 1.5– Florentinae n°1156 (sub. Carex divulsa Gooden.) 2.0 mm, similar to the female. ANTHERS 3, 1.5–2 mm Libia Cirenaica. Cirene- U. Belgadir, 26.iv.1934, R. long. PERIGYNIUM erect-patent to erect, pale green- Pampanini & R. Pichi-Sermolli 1156 (K!).

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409 THE CAREX DIVULSA AGGREGATE IN EURASIA 399 Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021

Figure 8. Carex enokii (from holotypus R. Pampanini & R. Pichi-Sermoli 1156, K): A, inflorescence; B, male glume with anther; C, female glume; D, perigynium, abaxial side; E, achene; F, perigynium with abaxial side removed to show achene. Carex leersii (G, from Hennipman et al. 1756, K; H–L, from LEB 82662): G, inflorescence; H, male glume with anther; I, female glume; J, perigynium, abaxial side; K, achene; L, perigynium with abaxial side removed to show achene. All figures drawn by X. Magaz. Scale bar, 1 mm.

Description: Caespitose perennial plant with short at nodes, sessile, and contiguous. The lowest intern- rhizomes c. 3–4 mm in diameter. CULMS 25– ode 12–55 mm long, the second one 7–17 mm. BRACTS 70 cm ¥ 1.0–2 mm at mid-height, obtusely trigonous, glume-like, the proximal setaceous, shorter than slightly scabrid above; sheaths brown. LIGULE 1.0– the inﬂorescence, 6–55 mm long. FEMALE GLUMES 3.0 mm long, subacute to acute at apex. LEAF blades hyaline, with a midrib green, 3.5–5.0 ¥ 2.0–2.5 mm, grey green, about three-quarters as long as culms, ovate, apex acuminate 0.3–0.5 mm long. STAMINATE widest leaf blades 3–4 mm, plicate to ﬂat. INFLORES- GLUMES 3.2–4.25 ¥ 1.75–2.5 mm, similar to the CENCE oblong, 40–90(145) ¥ 10–12 mm, with 7–12 female. ANTHERS 3, 1.75–3.0 mm long. PERIGYNIUM globular-elliptic spikes. Lowest spikes usually com- erect-patent to patent, pale greenish, body ovate pound and shortly pedunculate, upper spikes single to oblong, veinless or faintly veined, 4.5–5.5 ¥ 2.5–

2.75 mm, rounded base, sometimes slightly winged, 11.vii.1855, B. Balanse (K). Saziantifs, Gazianlifs, and abruptly tapered into a beak. BEAK 0.75–1.0 mm Kara Tash, nonline, rock volcanic, 3200 ft, 3.v.1935, long, white-scarious distally, with scarcely serrulate E. K. Balls B2218 (K). margins reaching the summit, apex bidentate, apical teeth 0.3–0.5 mm. ACHENE elliptic, shining, 4. Carex leersii F. W. Schultz, Flora 53: 459. 1870, light brown, 2.5–3.0 ¥ 2.0–2.5 mm. STIGMATA 2, 2.0– non Willd. 1787 (Fig. 8G–L) 3.0 mm long. = Carex canescens sensu Leers, Fl. Herborn. 712. t. 14. f. 3. 1789, non L. 1753. Phenology: Flowering begins in April, fruiting extends = Carex virens Lam. & DC, Fl. France, vol. 3: 106. 1805, non Lam. 1792. from May to July, wind pollinated. Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 = Carex divulsa Gooden b virens Durieu, Bull. Soc. Bot. France 6: 633. 1859. Habitat: There is little information about the habitat = Carex duriaei F. W. Schultz, Flora 53: 548. 1870, of C. enokii. The plants inhabit waste lands and forest non Steudel, 1841. borders, such as savin woods. They also grow in shady = Carex chabertii F. W. Schultz, Flora 54: 21. 1871. and wet places by streams. The substrates mentioned Lectotype (designated by Loos): Feddes Repert. for C. enokii include marl and volcanic rocks, and so 107(1–2): 67. 1996 (B!). Isolectotype e.g.: BR, C, GB, it appears to need rich nutrient soils. K!, L, M, MA!, MANCH, P!, W!. = Carex divulsa Gooden. var. chabertii (F. W. Schultz) Distribution: Carex enokii is found (Fig. 5) throughout Kneuck. in Seub. & Klein, Exk.-Fl. Baden: 52. 1891. the eastern and southern Mediterranean Basin from = Carex muricata var. leersii Kneuck. in Seub. & Turkey, through Palestine and Libya, to Mauritania, Klein, Exk.-Fl. Baden: 52. 1891. and also in Corsica, Sicily, and the Canary Islands. = Carex divulsa Gooden. ssp. chabertii (F. W. Schultz) Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 42. 1902. = Carex muricata race C. leersiana Rouy(‘leerseana’), Observations: Mouterde (1966) and Erteeb & Sherif Fl. France 13: 413. 1912. (1985) do not mention any Carex with these charac- = Carex divulsa ssp. leersii (Kneuck.) W. Koch, Mitt. teristics. Carex muricata ssp. divulsa var. divulsa f. Bad. Landesvereins Naturk. Naturschutz Freiburg guestphalica sensu Maire (1957) could be a similar 11: 259. 1923. plant. Feinbrun-Dothan (1986) included under C. di- = Carex leersiana Rauschert, Feddes Repert. 83(9–10): vulsa ssp. leersii plants from Palestine that have 660. 1973. Lectotype (designated by Loos sub. C. leer- the same characters as C. enokii (perigynium size, sii F. W. Schultz): Feddes Repert. 107(1–2): 67. 1996 broadly ovate, abruptly tapering beak). Its scarce (B!). Isolectotype e.g.: K!, L, M, MANCH, P!, W!. presence in herbaria informs us about its rarity. There are robust specimens with longer inflorescences Description: Caespitose perennial tufted plant with from Sardinia and Tenerife that may be hybrids with or without short rhizomes. CULMS 39–90 cm ¥ 0.8– C. divulsa. 2.0 mm at mid-height, obtusely trigonous, slightly scabrid above; sheaths pale to dark brown. LIGULE Representative specimens examined: ITALY. Sardinia, 0.75–3.5 mm long, wider than long, rounded or sub- Rio Baccu Mannu, Domus de Maria (CA), 11.v.2004, R. acute at apex. LEAF blades green to grey green, about Angius, R. Mandis & M. Orrú (CAG). LIBYA. Cirenaica, three-quarters to as long as culms, widest leaf blades Cirene, U. Belgadir, 26.iv.1934, R. Pampanini & R. (2)2.25–3.75(4) mm, plicate to flat. INFLORESCENCE Pichi-Sermoli 1156 (K). MAURITANIA. Fainyer?, oblong, 35–60(75) ¥ 8–10 mm, with 6–11(13) globular- 1.iv.1921, C. Pau (MA 016856). PALESTINE. Above Acie elliptic spikes. The lowest spike sometimes bran- Fit, woods, 9.v.1911, F. S. Meyers & J.E. Dinsmore ched and shortly pedunculate, upper spikes single 91781 (K). SPAIN. Canarias: Tenerife, Anaga, Sabinar at nodes, sessile, and contiguous, first internode de Afur, 28.i.2004, J. A. Reyes Betancort 8–27 mm long, the second one 5–15 mm. BRACTS (TFC 45312). Tenerife, El Moquinal-Pedro Alvarez, glume-like, the proximal setaceous, shorter than the borde carretera, 15.v.2004, M. A. Padrón Mederos (TFC inflorescence, 6–30 mm long. FEMALE GLUMES hyaline 45313). Tenerife, La Orotava, 30.vii.1983, E. Hernán- to golden-yellow, with a midrib green, 3.0–4.5 ¥ dez (LEB 62448). Tenerife, La Orotava, Teide, bei El 1.6–2.30 mm, ovate or elliptic, apex acuminate Bebedero, c. 740 m, 28RCS 521 395, 18.vi.1985, E. 0.15–0.3 mm long. STAMINATE GLUMES 2.75–3.5 ¥ Vitek (W 2001-1476i). TURKEY. Antalya, Antalya, E. of 1.5–2.5 mm, similar to the female. ANTHERS 3, 1.8– the town, waste places, along road, partly shaded 3.0 mm long. PERIGYNIUM erect-patent to patent, marl-soil, wet roadside, 35 m, 14.iv.1959, Hennipman greenish, body elliptic to ovate, nerveless or faintly et al. 366 (Iter leydense) (K). Puertes Celiciennes, veined, 4.5–5.5 ¥ 1.75–2.25(2.5) mm, acute at base,

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409 THE CAREX DIVULSA AGGREGATE IN EURASIA 401 sometimes slightly corky at the base, gradually taper- Buntsandstein, Weinberg, bei Ettlingen, auf berasten, ing into a beak. BEAK 0.67–1.25 mm long, white to with C. divulsa, C. praecrox, 180 m, 48°57′N. Br., brown distally, with serrulate winged margins reach- 26°6′E v. F., 16.vi.1895, A. Kneucker (K). GREECE. ing the summit, apex biﬁd or bidentate, apical teeth Epirus, inter Joannina et Dodona, sust. Calc., 1000– 0.3–0.6 mm. ACHENE elliptic to ovate, shining brown, 1200 m, 26.v.1964, K. H. Rechinger (MA 419448). 2.25–3.0 ¥ 1.5–2.1 mm. STIGMAS 2, 2.0–3.0 mm long. HUNGARY. Villany, Comut. Bourounya, in monte Har- sanyhegy, 16.v.1907, Herb. Dr A. de Degen: Koces (K). Phenology: Flowering begins in May, fruiting from ITALY. Sudtirol, Val Vestino, bei Bolone, an Wiesen- June to August, wind pollinated. und Wegrändern, 700 m, 1.vi.1901, P. Porta (MA 236899). Tirol australis, Val Vestino, près Bollone, Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 Habitat: Carex leersii occurs in areas with a tem- 1.vi.1879. P.Porta (P). SPAIN. León, Aviados, entre perate oceanic climate. Although most authors (De talud de la carretera y bosquete de Quercus pyrenaica, Langhe, 1944; Nelmes, 1947; Chater, 1980; Jermy c. 1000 m, 30TN 9851, 26.vi.2004, A. Molina (LEB et al., 1982; David & Kelcey, 1985; Clapham et al., 82662). Lérida, Valle de Arán, Pontant, Quercion 1987; Sell & Murell, 1996) and most of the studied robori-petraea, 660 m, 31TCH 1445, 8.vii.1992, P. Mon- materials indicate that it lives on calcareous or rich serrat, J. L. Benito & J. A. Sese (JACA 223192). soils, other authors, such as Luceño (1994), do not TURKEY. Trabzon, near 20 km W of Trabzon, marshy agree. Chabert stated that he sampled his specimens grasses along pool, 4.vi.1959, E. Hennipmann et &. of C. chabertii from ‘coteaux granitiques de Laveyron. 1756 (K). SWEDEN. Skárva, Blekinge, Stauraby, Drôme’. Therefore, this taxon is not exclusively 14.vi.1926, B. Holmgren (K). SWITZERLAND. Branson, calcicolous. We found C. leersii on well-drained, dry- Valais, 26.iv.1863, L. Favrat (K). UNITED KINGDOM. summer soils, in sunny open places, and often in Norfolk, Appleton, grassy roadside bank, on chalk, disturbed habitats (ravines or roadsides). 28.vi.1942, C. E. Hubbard (K).

Distribution: Carex leersii occurs (Fig. 6) from north- 5. Carex nordica A. Mol., Acedo & Llamas, ern Spain and Italy (41°N) to Great Britain and the sp. nov. (Fig. 9A–F) Scandinavian Peninsula (60°N), and Turkey in the Carice leersii similes, sed ab ea differt perigyniis east, between 300 and 1000 m. It is more common in minoribus atque alatus, squamae feminea fusca, France, Austria, and Germany, but is rare in Eastern spicae patens et separatus. Europe. Holotype: Sweden: Skåne, Paroecia, Hyby Böke- Observations: Schultz (1871: 25) gave a very detailed bergsslätt, in agare viae ferreae (sub. C. divulsa description of C. leersii based on his own material. Stokes subsp. leersii) 21.vi.1922, G. Samuelsson (K!). The best diagnostic characters of C. leersii are perigynium elliptic to ovate c. 5 mm, and hyaline Description: Caespitose perennial plant with or elliptic female glume. Table 3 demonstrates that without short rhizomes. CULMS 30–80 cm ¥ 0.8– C. leersii shows significant morphological variation. 1.5 mm at mid-height, obtusely trigonous, slightly Frequently, specimens labelled as C. leersii are young, scabrid above; sheaths light to dark brown. LIGULE sterile, or damaged, and, according to Jermy et al. 1–3 mm long, wider than long, subacute to rounded at (1982) and David & Kelcey (1985), when animals apex. LEAVES green to dark green, about three- have cropped them, they have depauperate flowering quarters as long as culms, widest leaf blades stems and are difficult to identify. Specimens from 2.0–3.0 mm, plicate to flat. INFLORESCENCE oblong, Greece and Turkey frequently show intermediate 35–55(70) ¥ 7–10 mm, with six to nine globular spikes, characters between C. leersii and C. otomana. More- sessile, single at nodes, first internode 13–30 mm over, in the lowest part of the Pyrenees, specimens long, the second one 7–14 mm, the remaining spikes with intermediate characteristics between C. leersii contiguous. BRACTS glume-like, the proximal seta- and C. magacis were found. ceous, shorter than the inflorescence, 10 mm long. FEMALE GLUME brown, with a midrib green to brown, Representative specimens examined: BELGIUM. with scarious margins, 2.75–3.75 ¥ 1.5–2.25 mm, Buissons, 1.vi.1907, Izn (MA 016826). CROATIA. Ins. elliptic, apex acuminate 0.25–0.3 mm long. STAMI- Korcula (of pines in maquis), under light shade, on NATE GLUME 2.75–3.5 ¥ 1.5–2.0 mm, similar to the limestone hills, 1.v.1976, C. C. Townsend 76/36 (K). female. PERIGYNIUM patent to very patent, greenish FRANCE. Haute-Saône, Port-sur-Saône, au pied d’un to brown, body ovate to elliptic, nerveless or faintly mur, 30.v.1930. V. Madiot (P). Isère, Rivoire de Allem- veined, 4.0–4.75 ¥ 1.75–2.5 mm, base rounded, gradu- ont, carretera local, talud carretea, a pleno sol, 1000 m, ally tapered into a beak. BEAK 0.75–1.35 mm long, 30.vii.2004, A. Molina (LEB 82667). GERMANY. Baden, brown distally, with serrulate winged margins reach-

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Figure 9. Carex nordica (A, C–F, from LEB 48138; B, from MA 387223): A, inflorescence; B, male glume; C, female glume; D, perigynium, abaxial side; E, achene; F, perigynium with abaxial side removed to show achene. Carex magacis (from holotypus LEB 82649): G, inflorescence; H, male glume with anther; I, female glume; J, perigynium, abaxial side; K, achene; L, perigynium with abaxial side removed to show achene. All figures drawn by X. Magaz. Scale bar, 1 mm. ing the body, the winged margins going to the middle Distribution: Carex nordica is scarcely distributed in or the full body, apex bifid, apical teeth 0.3–0.6 mm. a narrow zone from Great Britain to Sweden, and in ACHENE elliptic, shining, dark brown, 2.0–2.75 ¥ some scattered localities in Central Europe (Fig. 6). 1.25–2.0 mm. STIGMATA 2, 2.0–2.25 mm long.

Etymology: The name comes from the European area Representative specimens examined: AUSTRIA. Ost- in which it lives. tirol, Deferegen Tal: zwischen Huben und Hopfgarten, Quellﬂuren, Felsen, Schutthalden, 6.vi.1967, F. Krendl Phenology: Flowering begins in June, fruiting from (W 1985-10548). Vorarlberg, Tosters westlich Feld- July to August, wind pollinated. kirch am Weg zur Ruine Tosters, 1.viii.1968. A. Polatschek. (W 1969-2947). BELGIUM. Laeken, parc Habitat: It occurs in disturbed habitats and the public, bord chemin, 27.vi.1924, N. Gunz (K). Liège, borders of tracks. The scarce data available indicate Amay, talus s, exp S; sol silicieux, avec Arrhenatherum, human altered habitats, such as car parks, railways, 6.vii.1972, W. Bellotte (MA 387223) and (LEB 48138). parks, and roads. It lives in temperate hyperoceanic Amay, 6.vii.1972, Malato Baliz (MA 274200). Hainaut, to oceanic areas. Jamioulx, talus herbeux à exposition S, rue Francois

Vandanme, 29.vi.1972, Duvigneaud & Lebeau (MA margin, nerveless or faintly veined, 4.5–5.5 ¥ 2.25– 387222). Hainaut, Vaulx, limite d’Antoing, friche her- 2.75 mm, base cuneate, gradually tapered into a beak. beuse sur calcaire, 18.vi.1972, J. Duvigneaud (MA BEAK 0.75–1.25 mm long, white distally, with serru- 274405). Namur, Purnode, près de la gare, vallée du late winged margins reaching the body, apex bifid, Bocq, antique voie ferrée, avec C. spicata, friche her- apical teeth 0.3–0.7 mm. ACHENE elliptic, shining beuse termophile, 2.vii.1979, Duvigneaud (MA 236900) brown, 2.5–3.0 ¥ 1.75–2.25 mm. STIGMATA 2, 2.5– and (MA 387225). DENMARK. Dyrharnen, Klamper- 3.25 mm long. burg, Sjaelland, park, 15.viii.1947, W. R. Price 2035 (K). Fyen, Nyborg, 1.viii.1868, B. J. Cöster (K). Fyen, Etymology: The new subspecies is named in honour of Faaborg, D32: Dyreborg Skov, rides in plantation by the family Magaz. Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 seashore, 24.vii.1952, D. P. Young 4479 (K). NETHER- LANDS. South-Limburgh, between Gud-Valkenburg Phenology: Flowering begins in June, fruiting from and Schin op Geul, 23.vii.1932, A. de Trever in Kern July to August, wind pollinated. & Reichgelt herbarium (K). SWEDEN. Gotlandia, Wisby, 1.vii.1868, F. Ahlberg (K). Skåne, Sk. Stehag, Habitat: Carex magacis inhabits rocky places, such as 1.viii.1878, N. H. Nilsson (K). Skåne, Ringsgen, ravines, stony areas, and open spaces in forests 1.vii.1828, N. H. Nilsson (K). UNITED KINGDOM. Derby, (scotch pine, beech, etc.) in a wide range of montane vc. 57: Millersdale. 27.vi.1943, N. F. D., Herb. G. V. C. level communities [(600)780–1200 m; mean tempera- Last 753/60 (K). Wales: Pembrokeshire, Caldey Island, ture, 2.5–7.5 °C; high precipitation (700–1200 mm)] between high cliff and park area, 1.vi.1952, F. N. in temperate oceanic sub-Mediterranean zones. It can Hepper (K). grow on rich calcareous soils, but its resistance to acidic conditions is not known. 6. Carex magacis A. Mol., Acedo & Llamas, sp. nov. (Fig. 9G–L) Distribution: Carex magacis is scarce (Fig. 6) and is Carice leersii similes, sed ab ea differt perigyniis found in a small area in the Pyrenean Mountains and rhombi formam habens; habet basim truncate, Iberian Range (Spain, Andorra, and France). 2.25 mm latioribus, atque alatus. Carice muricatae similis, sed ab ea differt perigyniis longioribus, Conservation status: Carex magacis is an endemic squamae feminea hyalina, denticuli rostrum dilatati plant. Its distribution indicates that it is a rare taxon in perigynia corpore. which, according to IUCN (2001), and because of the few localities and small area in which it is found, Holotype: Spain, Navarra, Selva de Irati, Senda 69, must be catalogued as vulnerable (VU). c. río Urtxuria, sobre grietas de caliza en borde de camino umbroso, 800 m, 42°57′N, 1°03′W, 5.vii.2004, Representative specimens examined: ANDORRA. Parro- A. Molina (LEB 82649). quia S. Julià de Lòria, Cova de l’Ossa, taludes, roquedos, cultivos abandonados, esquistos, 1020 m, Description: Caespitose perennial plant with short 42°27′41″N, 1°29′12″E, 30.v.2004, C. Aedo, Aizpuru, rhizomes c. 6 mm in diameter. CULMS 26–86 cm ¥ Barriego, Pedrol & Molina (MA 714438). Sta Julia 1.0–2.0 mm at mid-height, obtusely trigonous, slightly de Loria, Pont de la Marginada, ambiente de scabrid above; sheaths pale to dark brown. LIGULE encinar, suelo pobre en bases, 980 m, 31T CH7604, 0.5–3 mm long, wider than long, rounded to subacute 6.vii.1992, S. Castroviejo et al. (MA 512056). at apex. LEAVES pale green, about as long as culms, FRANCE. Hautes Pyrénées, St Marie de Capman, widest leaf blades 3.0–4.25 mm, plicate to flat. subida a Pont de la Moulette, bord chemin calizas, INFLORESCENCE oblong, 40–60 ¥ 10–12 mm, with 7–12 1000–1200 m, 10.vii.2004, A. Molina (LEB 82663). globular spikes, usually the first one branched and SPAIN. Castellón: Entre Mas de la Cabra y el río shortly pedunculate, upper spikes sessile, single at Monleón (Vistabella), 1100 m, 30TYK 2768, nodes, and contiguous, first internode 14–21 mm long, 14.vi.1987, C. Fabregat (MA 467749). Huesca: Aniés, the second one 7–19 mm. BRACTS glume-like, the Senda a ermita Virgen de la Peña, calizas abriga- proximal setaceous, shorter than the inflorescence, das, 1080–1100 m, 30TYM 0187, 3.vi.1992, L. Villar 6–18(48) mm long. FEMALE GLUME golden-yellow to J. A. Sesé et al. (JACA 79392). Casbas de Huesca, light brown, with a midrib green, with scarious Sierra Guara, collado, relleno y peñascos, 970 m, margins, 3.0–3.6 ¥ 2.0–2.5 mm, elliptic, apex apiculate 30TYM 2981, 30.vi.1972, P. Monserrat & L. Villar 0.10–0.25 mm long. STAMINATE GLUME 3.0–4.0 ¥ 1.5– (JACA 379772). Castillonroy, Barranco de la acequia 2.0 mm, similar to the female. ANTHERS 3, 2.0–2.5 mm de Salellas, Baldellou, 500–650 m, 31TBG 9545, long. PERIGYNIUM very patent to patent, greenish, 27.v.1987, G. Monserrat (JACA 434787). Fanlo, body trullate to ovate-trullate, with a broad winged Añisclo, bosque mixto, 950–1200 m, 31BTH 5817,

2.vii.1973, P. Monserrat & L. Villar (JACA 263873). apex acuminate 0.2–0.5 mm long. STAMINATE GLUME Jusen, Graus, alrededor del pueblo, 780–820 m, 3.75–4.75 ¥ 1.5–2.0 mm, similar to the female. 31TBG 8263, 1.vi.1986, G. Monserrat (JACA ANTHERS 3, 1.75–2.0 mm long. PERIGYNIUM erect- 658486). Nueno, Sierra Guara, Nocito, Bco. de patent to patent, greenish, body ovate, nerveless or Lapillera, glera, solana cálida, hayedo en hoya, faintly veined, 5.0–6.0 ¥ 2.0–2.5 mm, base rounded to 1050 m, 30TYM 2987, 11.vii.1974, P. Monserrat cuneate, slightly swollen and corky, gradually tapered (JACA 368274). Oroel, Jaca, abetal con hayas into a beak. BEAK 1.0–1.6 mm long, brown distally, escasas, 1200 m, 30TYM 0410, 25.viii.1988, L. with serrulate winged margins reaching the body, apex Villar, D. Gómez et al. (JACA 271888). San Juan de bifid, apical teeth 0.5–0.75 mm. ACHENE elliptic to la Peña, camino del mirador, pinar de P. sylvestris, ovate, shining dark brown, 2.5–3.0 ¥ 1.75–2.25 mm. Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 1240 m, 30TXN 9109, 14.vi.1970, P. Monserrat & L. STIGMAS 2, 2.0–2.25 mm long. Villar (JACA 223470). San Juan de Plan, pradería, prado de siega, 1150–1200 m, 31TBH 8219, Etymology: Species dedicated to the Ottoman people, 19.vi.1981, P.&G. Monserrat, D. Gómez & F. Fillat who developed an empire in part of the area in which (JACA 066481). Navarra: Burgui-Salvatierra, Foz de the species lives. Forniellos, 1050 m, 30TXN 6830, 1.vii.1975, L. Villar (JACA 10069075). Teruel: Rubrelos, 23.vi.1897, Phenology: Flowering begins in May, fruiting from Benedicto (MA 16803). Segura de Baños, cantiles y June to July, wind pollinated. rellenos, 1000 m, 30TXL 7235, 11.vii.1973, P. Mon- serrat, J. Erviti & L. Villar (JACA 318173). Zara- Habitat: Carex otomana occurs between 700 and goza: Luesia, alto de la Sierra de Sto. Domingo, 1850 m in wet mountain areas (ravine forest, herbazales, 1250–1300 m, 30TXN 6902, 21.vi.1987, meadows, and springs) in open forest or above the J. Pedrol (MA 437584). Salvatierra de Esca, Foz de timberline in all types of soil. Forniellos, cantiles solanos y carrascal con quejigo, 750–800 m, 30TXN 6829, 22.vi.1973, L. Villar Distribution: Carex otomana occurs (Fig. 6) from east (JACA 100118673). of the Black Sea (Bulgaria) and Greece through the Turkish mountains and the Caucasus to the 7. Carex otomana A. Mol., Acedo & Llamas, mountains on the west side of Tyan Shan in Central sp. nov. (Fig. 10A–F) Asia (Kazakhstan). Carice leersii similes, sed ab ea differt perigyniis ovate sunt, maioribus, 5.5 mm longitudines, corpore Observations: Greek specimens of C. otomana are glaucous, squamae feminea fusca, in rostrum dilatati similar to C. leersii with regard to perigynium size utriculorum. and shape, but can be easy differentiated by the brown glumes. Towards the east of the area, the Holotype: [Kazakhstan], Asia centralis: Alma-Ata, specimens have larger perigynia that are clearly Tyan-Shan montes ‘Zailijki Alatau’, in valle fluminis ovate and slightly corky at the base. The characters ‘Malaia Almatinka’, loco Medeo dito, 1400–1750 m, quoted by Nilsson (1985) for C. divulsa ssp. leersii can 30.v.1974 (sub Carex polyphylla Kar. & Kir.), V. Vasák, also be assigned to C. leersii and C. otomana. Never- Iter centrali-asiaticum 1974 (W 11223!). theless, C. divulsa ssp. leersii sensu Kukkonen (1998) has characteristics from C. otomana (female glume Description: Caespitose perennial plant with or triangular, utricule ovoid, base slightly suberous, without short rhizomes. CULMS 50–90 cm ¥ 1.5– beak 1.4–2.3 mm, long teeth), although some speci- 2.5 mm at mid-height, obtusely trigonous, scabrid mens reported by him are outside the distribution above; sheaths pale yellow to dark brown. LIGULE area of this species. Similar characters have also been 1.5–4.0 mm long, wider than long, subacute to rounded reported in C. polyphylla sensu Kreczetovicz (1935) at apex. LEAVES grey green, about three-fifths as long (scales ovate, ferrugineous, perigynium ovoid, long as culms, widest leaf blades 2.25–4 mm, plicate to flat. beak). INFLORESCENCE oblong, 40–65(85) ¥ 9–14 mm, with 8–13 globular-elliptic spikes, sometimes the first spike Representative specimens examined: ARMENIA. Tak- branched and shortly pedunculate, the remaining Achac, north-east shore Seran Ist ravine, lower bound- spikes single at node, sessile, and contiguous, first ary of forest, 16.vi.1930, O. Polianskaya (leg por Krecz) internode 10–17 mm long, the second one 7–17 mm. (K). BULGARIA. Varnenski Okrug, Varna, Slatni Pjas- BRACTS glume-like, the proximal setaceous, shorter atsi, glade of a shrubby woodland slope, 50 m, NH3, than inflorescence, 6–20 mm long. FEMALE GLUME 7.vi.1973, J. Suominen 3056 (H 1664673). GREECE. light brown, with a midrib green, with scarious Serron, Serron, Mt. Vrondous, summit area (meadows) margins, 3.25–5.25 ¥ 1.75–2.3 mm, ovate to elliptic, rocky slopes (and rock crevices) at and above timber-

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409 THE CAREX DIVULSA AGGREGATE IN EURASIA 405 Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021

Figure 10. Carex otomana (from J. Suominen 3056, H 1664673): A, inflorescence; B, male glume with anther; C, female glume; D, perigynium, abaxial side; E, achene; F, perigynium with abaxial side removed to show achene. Carex egorovae (G–I, from O. Polunin 5110 K; J–L, from R.W. Haines W 676 K): G, inflorescence; H, male glume with anther; I, female glume; J, perigynium, abaxial side; K, achene; L, perigynium with abaxial side removed to show achene. All figures drawn by X. Magaz. Scale bar, 1 mm. line (formed by Pinus sylvestris), granite, 1700– ad cascades, 750 m, Iter Iranicum IX (1975), 4.vi.1975, 1840 m, GL1, 6.vii.1985, M. Salmenkallio & V. Salo K. H. Rechinger 52649 (WU 024145, WU 024144, and 895 (H 1557247). IRAN. E. Mazandaran, N. Semman, W 05994). Gorgan, Shahpasand, 73 km. Shahpasand, southern slopes of Alu-Baq Mountain: around Cheshm, 750 m, 5.vi.1975, Termeh, Herbarium Minesterii Cheshmehe Alu-Baq (spring) (Golestan National Iranici Agriculturae EVIN n°36871 (H 1686153). Park), 1600–1650 m, 37°19′N, 55°53′15″E, 7.viii.1995, NW Khorasan, Centre, 6 km E of Tangegol, Golzar, H. Akhani 11898 (H 1695176). Gorgan, Mohammed in Zelkova carpinifolia and Carpinus forest (Golestan Reza Shah National Park, forest S. of Tang-e Gol, National Park), 800–950 m, 37°22′N, 56°00′E, subst. calcareo, in quercetis (Q. castaneifolia) apertis 9.vii.1994, H. Akhani 9604 (H 1695181). Gorgan,

Mohammed Reza Shah Wildlife Park, forest S of Phenology: Flowering begins in April, fruiting from Tang-e Gol, 700–1000 m, Ariamehr Bot. Garden, April to June, wind pollinated. 21.vi.1974, Wendelbo & Foroughi 12795 (W 21498). Habitat: Damp and shady places between 700 and KIRGUISTAN. Songarica, 1.ix.1870, Schrenk (K). 1360 m: stony hillsides, brook sides, meadows, and MACEDONIA. NW Macedonia, Drnova, Angkatholow, bank of ravine, 2800 ft, 13.vi.1932, A. H. G. Alston & N. forests. Y. Sandwith 296 (K). TURKEY. Bursa A2 (A), above Distribution: Carex egorovae occurs in three dis- Bursa on the road to Uludag, roadside in Castaneo– jointed areas (Fig. 6): Eastern Europe (from Hungary Fagetum, 730 m, 8.vii.1978, P. Uotila 27146 (H to north-west Turkey), Middle East (Iraq and Iran), 1203994). Prov. Adana, Amanus, 1500 m, 26.viii.1949,

and Cyprus. Downloaded from https://academic.oup.com/botlinnean/article/156/3/385/2418228 by guest on 28 September 2021 P. Davis 16442 (K). UZBEKISTAN. Asia Media, distr. Andizham, Fergana, in silva prope Arslanbob, Observations: Specimens from Europe have slightly 25.v.1899, Litrovinov (K). different characters to those from the other two areas. European plants have longer and more serrulate 8. Carex egorovae A. Mol., Acedo & Llamas, beaks and are not corky at the perigynium base. sp. nov. (Fig. 10G–L) Given the scarce number of available sheets, they are Carice leersii similes, sed ab ea differt inflorescentia all given the same rank. The plants from Iraq that brevioribus, 4–5 mm longitudine; spicae erecto- were included in C. divulsa ssp. leersii by Hooper patens, pauciflorae; perigynia rostrum longiore et (1985) seem to be the same plant. bifidus. Representative specimens examined: CROATIA, Montes Velebit, pars austr., Nationali Park Paklenica, in valle Holotype: Iraq. MAM, Shariffa near Amadia, shady augusta rivi Velika Paklenica dispersim, Flora Balca- meadow, 4000 ft, 22.iv.1958 (sub. Carex divulsa nica: Croatica, 24.v.1996, Mil. Marek (WU). CYPRUS. Stokes ssp. leersii (Kneuck.) W. Koch), O. Polunin Kythrea, in irrigation channel, 500 ft, 13.iv.1962, R. D. 5110 (K!). Isotype: O. Polunin 5110b (K!). Meikle 2556 (K). HUNGARY. Nr Podhragy in Ungarn, Holuby (K). Nr Podhragy in Cominztronam, 1876, Description: Caespitose perennial plant with or Holuby (K). Herkulesbad im Banat in Ungarn, auf der without short rhizomes c. 5 mm in diameter. CULMS Coroninihöle, con Genista tinctoria, Thymus monta- 14–50 cm ¥ 0.8–1.25 mm at mid-height, obtusely trigo- nus, etc., 500 m, 30.vi.1902, L. Ritcher (K). GREECE. nous, slightly scabrid above; sheaths pale to dark Thasos, S coast, wood by stream, 100 ft, 24.v.1972, W. brown. LIGULE 1.5–3.0 mm long, wider than long, R. Price 1224 (K). IRAN. Asterabad, Bender Ges. in rounded or subacute at apex. LEAF blades grey green, dumetis, 30.iii.1901, P. E. Sintesis 1466. (P 00281845). about three-quarters to longer than culms, widest leaf Trorth Gonbad, near village Golidagh, shady area, blades 2–3 mm, plicate to flat. INFLORESCENCE oblong, under trees, 1080 m, 25.v.1976, T. F. Henr 3872 (K). 30–50(68) ¥ 9–13 mm, with 6–11 globular-elliptic, Gorgan, Persia borealis, Asterabad, Bender Ges, sessile spikes, the lowest spike sometimes branched, 30.iii.1901, J. Freyn (K). IRAQ. MSU district, Tawila, upper spikes single at nodes, first internode 6– stony hillside, 1360 m, 16.vi.1957, A. Rawi 21896 (K). 13(20) mm long, the second one 3–9(13) mm. BRACTS Kurdistan, Sulaimaniya, in ditione pagi Tawilla, Mte. glume-like, the proximal setaceous, shorter than the Avroman at confines Persiae, Sosakan, in platanetis, inflorescence, 5–38 mm long. FEMALE GLUME hyaline, 1200 m, Itinera Orientalis 1956/57, 18.vi.1957, K. H. with a midrib green, 3.0–4.0 ¥ 1.6–2.25 mm, ovate, Rechinger 12137a,b (W 01174, W 01146). MRO district, apex acuminate 0.3–0.5 mm long. STAMINATE GLUME Shaqlawa, irrigated orchard-pastures in rough grass, 3.0–3.5 ¥ 1.4–2.0 mm, similar to the female. ANTHERS 2800 ft, 15.v.1956, Univ Herb Baghdad, R. W. Haines 3, 1.5–2 mm long. PERIGYNIUM erect-patent, pale (W 01133). Shaqlawa, shady bank on roadside, 2800 ft, greenish, body ovate, nerveless or with well-marked 12.v.1956, R. W. Haines W676 (K). Saisang, N slope veins, 4.25–5.5 ¥ 1.8–2.5 mm, rounded base, some of Gara, mountain grassy, 1300 m, 20.vi.1968, Herb. them slightly swollen and corky, gradually to abrupt College of Agric. & Forestry, Univ Mossul, Anders 2359 tapered into a beak. BEAK 1.0–1.75 mm long, white (W 1945). TURKEY. Bilecik B2, c. 40 km from Inegöl distally, with serrulate margins reaching the summit, to Bozüyuk, Aksutekke, by the main road, brookside apex bifid, apical teeth 0.6–1.25 mm. ACHENE elliptic, meadow in the service station area, 860 m, 20.v.1983, shining, brown, 2.0–2.7 ¥ 1.6–1.8 mm. STIGMATA 2, M. Salmenkallio 268 (H 1492360). 2.0–2.5 mm long. ACKNOWLEDGEMENTS Etymology: This new species is named in honour of We are grateful to the staff of the consulted herbaria Dr Tatiana Egorova, a Russian specialist in Carex. for their help with the location of material. We also

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409 THE CAREX DIVULSA AGGREGATE IN EURASIA 407 wish to express special gratitude to Professor David Mongolia, 3. Enfield, NH/Plymouth: Science Publishers, Simpson for his kind help during our visits to the Kew 38–97. Herbarium. We especially thank Ms Maria Elizabeth Erteeb FB, Sherif AS. 1985. Cyperaceae. In: Jafri SMH, Bailey who corrected the final English version, Mr El-Gadi A, eds. Flora of Libya, Vol. 119-120. Tripoli: Faculty Jorge Magaz for helping us with the drawings, and of Sciences, 1–46. Professor Benito Valdes and Professor Stephen Jury Feinbrun-Dothan N. 1986. Carex L. In: Zohary M, for their advice and valuable comments. Finally, we Feinbrun-Dothan N, eds. Flora Palaestina 4. Jerusalem: thank Junta de Castilla y León for granting a High The Israel Academy of Sciences and Humanities, 369– 375. Studies Licence to the first author, and for grant LE Fournier P. 1961. Les quatre flores de la France. Corse 025A05 that partially supported our research.

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