Taxonomy and New Taxa in Eurasian Carex (Section Phaestoglochin, Cyperaceae

Taxonomy and New Taxa in Eurasian Carex (Section Phaestoglochin Cyperaceae)

Ana Molina1 Carmen Acedo2 and Feacutelix Llamas3

Department of Plant Biology University of Leoacuten E-24071 Leoacuten SpainAuthor for correspondence (1amolgunileones 2cacedounileones 3fllamasunileones)

Communicating Editor Thomas G Lammers

AbstractmdashSupported by detailed phenetic macro- and micromorphological characters ecology chorological studies numerical and phy-logenetic analysis and all known data about Eurasian species of Carex muricata group (sect Phaestoglochin) together with their biogeographywe have redefined this group and its subordinate aggregates Four new taxa are described illustrated and compared with putative relatedspecies Carex spicata subsp andresii endemic to the inner part of the Iberian Peninsula has the perigynum beak smooth and slightly corkyat the base Carex omeyica from the southern mountains of Spain and North Africa has erect-spreading oval perigynia and oval brownglumes without scarious margins Two new subspecies of C muricata are described C muricata subsp cesanensis from European moun-tains has a big trullate perigynium 45ndash525 mm long with a short beak and C muricata subsp ashokae from the Eastern EuropeanMountains to the Himalaya has an oval perigynium 5ndash6 mm long with a longer beak On the other hand C pairae is not a taxon subordinateto C muricata A key to the new and related taxa belonging to the aggregate is included

Keywordsmdashcomparative morphology new species taxonomy

Carex L section Phaestoglochin Dumort is a poorly definedpolyphyletic group of species occurring in North Americarepresented by some 25 species (Ball 2002) and in Eurasia bythe Carex muricata sl group The common traits that distin-guish this section are rhizomes short or absent inflorescencesimple racemose or with 1ndash2 proximal branches lowestbracts setaceous spikes androgynous and perigynia plano-convex in cross section The boundary with several othersections in subgenus Vignea [sects Vulpinae (J Carey) ChristHeleoglochin Dumort or Multiflorae Kunth] is not clear

The Carex muricata group sl has scarcely been investi-gated Most studies are partial and made in limited areas(Vollmann 1903 De Langhe 1944 Nelmes 1947 Hartvig 1986Stoeva and Popova 1997 Repka 2003) Only Kuumlkenthal (1909)made a worldwide study of this group The nomenclature ofthe group has a history of confusion and misunderstanding(David 1976 Loos 1996 Repka and Danihelka 2005)

Chater (1980) based on papers by David and Kelcey(1975) David (1976) and David and Chater (1977) recog-nized five European taxa belonging to three species in theCarex muricata group Carex spicata Huds C muricata Lsubsp muricata C muricata L subsp lamprocarpa Celak Cdivulsa Stokes subsp divulsa and C divulsa Stokes subspleersii (Kneuck) W Koch and mentioned C chabertii F WSchultz described from Germany This classification is ac-cepted by most European monographs and floras (Jermy etal 1982 David and Kelcey 1985 Nilsson 1985 Lucentildeo 1994Kukkonen 1998) On the other hand eastern European bota-nists (Hadac 1961 Malyschev and Peschkova 1990 Egorova1999 2000) recognize some of these taxa at the species levelChater (1980) indicated the need for further research in thisgroup to reach a deeper understanding of its taxonomy

In a preliminary study Molina et al (2006a) studied 67morphological characters in one hundred specimens of theCarex muricata group from Europe The results of this taxo-nomic investigation suggested that previous works over-looked several cryptic but distinct taxa as recently observedin other sections of Carex (Naczi and Ford 2001 Saarela andFord 2001)

Within the Carex muricata group we have found threemain clusters corresponding to the C divulsa aggregate (in-cluding at least C divulsa Stokes and C leersii F W Schultz)the C spicata aggregate and the C muricata aggregate Carex

coriogyne Nelmes is isolated without a clear position The lasttwo aggregates are treated in this paper in which we hy-pothesize four species C spicata (with two subspecies) Cmuricata (with three subspecies) C pairae F W Schultz anda new species C omeyica

MATERIALS AND METHODS

Taxa Included Within the Scope of this StudymdashAll taxa closely relatedto the Carex spicata and Carex muricata aggregates treated by Kuumlkenthal(1909) Kreczetovicz (1935) De Langhe (1944) Nelmes (1947) David(1976) David and Chater (1977) Chater (1980) David and Kelcey (1985)Hooper (1985) Nilsson (1985) Malyschev and Peschkova (1990) Kuk-konen (1998) and Egorova (1999 2000) were studied C foliosa D Don Cmuricata L subsp lamprocarpa Celak C muricata L subsp muricata Cpolyphylla Kar amp Kir C spicata Huds and C wendelboi Nelmes

Taxon SamplingmdashField work was carried out during 2001ndash2005 mainlyin the Alps Cantabrian and Pyrenean Mountains Morphological inves-tigations and range maps were made based on collected material andsheets from B CAG FCO TFC GDA-GDAC H JACA K LEB LISILISU MA P SANT SEV W and WU More than 1000 specimens wereexamined in this study (list available on request)

MorphologymdashAll characters used to distinguish taxa in former studiesof Carex sect Phaestoglochin (Webber and Ball 1984 Stoeva and Popova1997 Ball 2002 Repka 2003 Molina et al 2006a) were checked with her-barium material and with our own specimens to assess their taxonomicvalue Additional characters were included according to our observa-tions For this study 61 characters were selected that together with someratios gave a total of 71 variables (Table 1) Ninety specimens represent-ing a wide range of morphological and geographical variation were cho-sen for the numerical analysis Achene color a character with low taxo-nomic value was impossible to observe in many specimens and thus wasonly used in the phylogenetic analysis The explanations for some char-acters and methods of measurement can be found in Molina et al (2006a)Three characters related to the perigynium beak margin were surveyedwith a Jeol 6100 Scanning Electron Microscope (SEM)

Numerical and Statistical AnalysesmdashEvery pair of characters of thedataset was subjected to a Pearsonrsquos or Kendall Tau correlation coefficientdepending on their quantitative or qualitative status Variables with highcorrelation coefficient ( 075) were eliminated to avoid redundant in-formation To detect groups among the specimens we performed a prin-cipal coordinate analysis (PCoA) because it allows us to include missingdata and both quantitative and qualitative characters We carried out ahierarchical clustering with 101 specimens to confirm our hypothesisusing Gowerrsquos coefficient and UPGMA algorithms of the SYN-TAX 2000(Podani 2001) We also analyzed some specimens of Carex foliosa and Cpoyphylla because they are sometimes included in this group and Cotrubae Podp (sect Vulpinae) because it is a species closely related to sectPhaestoglochin The analysis was performed in two parts the first partincluding all species treated and the second part with only the specimensbelonging to C muricata One-way analysis of variance (ANOVA) and thepost hoc Tukey test were conducted on quantitative characters to assess

Systematic Botany (2008) 33(2) pp 237ndash250copy Copyright 2008 by the American Society of Plant Taxonomists

237

divergence among species among subspecies within a species and toidentify the best diagnostic characters for them Because the variance ofseveral variables was heterogeneous as the Levenersquos test indicated thesedata were logarithmically transformed for ANOVA Qualitative charac-ters were studied with analogous Kruskal-Wallis analysis Statisticalanalyses were performed with SPSS 130

Phylogenetic AnalysismdashPhylogenetic analysis was performed using 44characters Only the statistically significant variables among those listedin Table 1 were included in the phylogenetic analyses and all variableswere equally weighted As we have few qualitative characters availablefor this study we used quantitative data because one can recover phy-logenetic information from these data (Thiele 1993) Qualitative charac-ters were coded as in former analyses To avoid arbitrarity the quanti-tative data were coded according to the groups in ANOVA post-hoc testAlthough the results of phylogenetic analyses are better if all data aretreated as unordered we think that the quantitative characters evolve

progressively Values of quantitative characters were coded in a range of0ndash3 and treated as ordered multistate characters (Table 2) Carex otrubaeC foliosa and C polyphylla were used as outgroups Heuristic search wasused and the robustness of clades was assessed by means of bootstrapanalysis Maximum parsimony analysis was performed with PAUP40b10 (Swofford 2002) Data are deposited in TreeBASE (study numberS2026)

BiogeographymdashSpecimen label data were used to document or delimitthe geographic distribution of each taxon Information from Clarke(1894) Maire (1957) Hylander (1966) Mouterde (1966) Chater (1980)Jermy et al (1982) Pignatti (1982) Hooper (1985) Nilsson (1985) Hartvig(1986) Hulteacuten and Fries (1986) Malyschev and Peschkova (1990) Lucentildeo(1994) Kukkonen (1998) Egorova (1999 2000) and Aeschimann et al(2004) was compared with our own data to infer the real distribution ofthe taxa The distribution maps are based on herbarium specimens Thedata and distribution maps were plotted using ArcGIS 9

RESULTS

In the preliminary revision of herbarium material wefound little morphological variation among taxa The char-acters that allow us to include a specimen in these aggregatesare leaves 25ndash45 mm wide auricles absent culm diameter075ndash20 mm inflorescences 18ndash45 mm long with spikes 5ndash11close together and sessile (except sometimes the lowest)bract short-setaceous glumes acuminate or acute brownwithout arista perigynum plano-convex 33ndash60 times 18ndash275mm margin winged or not beak up to 15 (175) mm

According to previously published data Carex foliosa Cpolyphylla and C wendelboi do not belong in Carex sect Phaes-toglochin Carex wendelboi has a tight inflorescence that iselongate ellipsoidal or ovoid with compound spikes pistil-late scales acute or aristate and perigynia 27ndash3 times 1ndash14 mmCarex foliosa has leaves up to 7 mm wide with auricles in-florescences 30ndash100 mm long with compound spikes havingmany flowers and perigynia 27ndash33 times 12ndash15 mm subtendedby an awned glume Neither the protologue of Carex poly-phylla nor the type conserved at Kew have the diagnosticcharacters of this section mainly because it is a very robustplant and its inflorescence has compound spikes Thereforethese taxa were not included in the PCoA and ANOVA tests

A high Pearsons correlation was found in the distance be-tween the lowest spikes number of flowers nerves in theperigynium body and width of the perigynium marginsConsequently only 58 characters (marked with rsquorsquo in Table 1)were used for further analysis four additional characterswere excluded in the PCoA because they did not show varia-tion within the Carex muricata aggregates

The scatter plot of the samples in the first two axes of thePCoA shows two main groups (Fig 1) Axis 1 provides sepa-ration of Carex spicata from the rest by its corky perigyniumbase distance from base to widest point of perigynium andligule apex while C muricata and C pairae are separated inaxis 2 mainly by perigynium length perigynium width andthe ratio perigynium length perigynium width Specimensof C omeyica appear quite close to C muricata The first threecomponents explain 5301 of variation (axis 1 = 3028 axis2 = 1601 and axis 3 = 672) PCoA shows distinct cluster-ing of the species but there is a considerable overlap betweensubspecies and no clear separation

The hierarchical clustering (Fig 2) shows that Carex otru-bae C foliosa and C polyphylla are separated from the aggre-gates and the four species of section Phaesthoglochin appeargrouped in a different cluster with two parts The first is theCarex spicata aggregate with two subunits representing Cspicata subsp andresii and C spicata subsp spicata and the

TABLE 1 Morphological characters and character states studied L =length W = width + = yes - = no r 075

(1) Awn glume 0 - 1 + (2) Auricle leaf blade 0 - 1 + (3)Lower spikes branched 0 - 1 + (4) Spikes contiguous 0 + 1 -(5) Basal sheaths purplish tinged 0 - 1 + (6) Perigynium beakcolor 0 green 1 brown (7) Lowest spike branched 0 - 1 + (8)Peduncle lowest spike 0 - 1 + (9) Margins of perigynium beak 0smooth 1 scabrid (10) Ripe achene brown 0 - 1 + (11) Scariousmargin of pistillate scale 0 - 1 + laterally 2 complete (12) Shapespike 0 globular 1 globular-elliptic 2 elliptic 3 oblong (13) Apexof pistillate scale 0 rounded-obtuse 1 acute 2 apiculate 3acuminate (14) Color glume 0 hyaline 1 gold-beige 2 lightbrown 3 dark brown 4 purplish brown (15) Perigynium shape 0rounded 1 oval 2 ovate 3 trullate 4 oblong 5 obovate (16) Apexof beak 0 bidentate 1 bifid 2 variable (17) Beak margins 0smooth 1 serrulate 2 irregular serrulate 3 scarcely serrulate (18)Shape achene 0 rounded 1 oval 2 ovate 3 pentagonal 4 obovate(19) Culm outline 0 rounded 1 obtusely trigonous 2 sharplytrigonous 3 winged (20) Inflorescence shape 0 lobulate 1 oblong2 elliptic (21) Pistillate scale shape 0 oval 1 ovate 2 obovate(22) Perigynium base corky 0 - 1 little 2 + (23) Ligule apex 0rounded 1 subacute 2 acute (24) L leafsL culm 1 lesser 2similar 3 bigger (25) Veins perigynium body (abaxial) 0 - 1 upto middle 2 all body (26)Perigynia with veins (abaxial side) 0none 1 several 2 all perigynia (27) Strength veins (abaxial side) 0veinless 1 faint 2 well marked 3 strongly (28) Veins perigyniumbody (adaxial) 0 - 1 up to middle 2 all body (29) Perigynia withveins (adaxial side) 0 none 1 several 2 all perigynia (30) Strengthof veins (adaxial side) 0 veinless 1 faint 2 well marked 3strongly (31)Attenuation perigynium beak 0 gradually 1gradually to abrupt 2 abrupt (32) Distribution denticles beak0 - 1 only beak 2 up to summit 3 over 13 body 4 only summit(33) Position of perigynium 0 erect 1 erectndashspreading 2spreading 3 very spreading (34) Winged margin of perigyniumbody 0 - 1 from summit to 1frasl2 body 2 all over body 3 wingvariable (35) Orientation of spike 0 lt 40deg 1 40deg-50deg 2 gt 50deg (36)L inflorescence 0 up to 30 1 30ndash50 2 50ndash70 3 more than 70 (37)W of widest leaf (38) Diameter of culm (at the middle) (39) Llongest ligule (40) L lowest bract (41) L longestinflorescence (42) Distance between lowermost spikes (43)Maximum distance between 2sd 3th spikes (44) No branchesfirst order (45) Number of perigynia in I2 (46) No male flowersI2 (47) L perigynium (48) W perigynium (49) L perigyniumbeak (50) Maximum no abaxial veins (51) Minimum noabaxial veins (52) Maximum no adaxial veins (53) Minimumno adaxial veins (54) L apical tooth (abaxial) (55) L apicaltooth (adaxial) (56) No denticles beakmm (57) L pistillatescale (58) Distance from base to widest point of perigynium(59) Difference (W perigynium ndash W achene)2 (60) Lachene (61) W achene (62) L perigyniumL glume (63) LperigyniumW perigynium (64) L acheneW achene (65) LperigyniumL beak (66) L perigyniumL achene (67) L longestinflorescenceDistance between lowermost spikes (68) Distancebetween lowermost spikesMaximum distance between secondand third spikes (69) No perigyniaNo flowers I2 (70) LperigyniumDistance from base to widest point ofperigynium (71) W wingW achene

238 SYSTEMATIC BOTANY [Volume 33

second includes the species belonging to the C muricata ag-gregate C omeyica C pairae and C muricata These speciesare separated by a dissimilarity value of 024 Only if weanalyze the Carex muricata cluster alone (Fig 3) can we sepa-rate the three subspecies C muricata subsp ashokae is clearlyseparated and the other two taxa C muricata subsp muricataand C muricata subsp cesanenesis are more similar to oneanother

ANOVA and Kruskal Wallis corroborate that this is a veryhomogeneous group The results for 12 variables are not sig-nificant From the remaining 59 17 have overlapping rangesbut each of 42 variables allows us to identify a taxon Thevariables that best separate the taxa are the perigynium shapeand length Each of the main clusters separated with PCoAand hierarchical clustering can be related to a set of characterstates (Table 3) Carex spicata is the best differentiated specieswith 16 variables C omeyica and C muricata appear togetherin most analyses and C spicata and C muricata seem to be the

FIG 2 Hierarchical cluster of 101 specimens of the Carex muricata andC spicata aggregates and C otrubae C polyphylla C foliosa and 58 vari-ables from UPGMA using Gowerrsquos coefficient

TA

BL

E2

Dat

am

atri

xus

edin

the

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ogen

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ysis

Pol

ymor

phic

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acte

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aby

C

hara

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refe

rto

Tab

le1

Ch

arac

ter

nu

mb

er1

23

45

67

910

11

12

13

14

15

16

18

19

20

21

22

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25

28

32

33

34

35

39

41

43

44

46

47

48

49

56

57

58

60

62

63

65

66

70

Cf

olio

sa1

11

10

01

10

23

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51

41

00

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AA

30

10

03

21

20

00

20

20

01

01

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pol

yphy

lla0

01

00

11

1

22

32

2A

12

10

10

21

22

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03

21

22

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21

32

11

01

0C

otr

ubae

11

10

01

11

00

20

22

02

31

12

22

12

20

01

22

12

21

21

20

10

10

22

Cs

pica

tasu

bsp

spic

ata

00

00

11

01

11

13

42

A3

21

12

22

12

10

11

10

00

21

22

22

20

20

11

Cs

pica

tasu

bsp

andr

esii

00

00

11

00

11

13

42

02

22

11

22

00

10

11

00

00

21

20

22

20

10

10

Co

mey

ica

00

00

00

01

00

12

31

02

11

00

01

12

10

10

10

00

21

02

23

20

01

00

Cp

aira

e0

00

00

1A

10

A0

32

20

11

12

00

10

22

01

00

0A

A1

00

21

21

00

10

0C

mur

icat

asu

bsp

mur

icat

a0

00

00

10

10

00

23

00

01

10

00

11

33

22

00

00

A1

10

20

11

10

10

0C

mur

icat

asu

bsp

cesa

nens

is0

00

00

10

10

00

23

31

11

10

00

21

33

22

00

00

02

11

21

32

10

10

0C

mur

icat

asu

bsp

asho

kae

00

00

01

01

10

02

31

11

11

00

00

03

32

20

10

00

21

22

13

22

20

10

FIG 1 Scatter plot of scores on axis 1 and 2 from principal coordinateanalysis of 90 members of the Carex muricata aggregates and 54 variables = C muricata = C omeyica = C pairae = C spicata

2008] MOLINA ET AL NEW TAXA IN CAREX 239

most different species Subspecies are differentiated mainlyby quantitative rather than qualitative characters

The phylogenetic analysis resulted in two most parsimo-nious trees with a tree length of 116 (Fig 4) Forty-one char-acters are informative with a consistency index (CI) of 067excluding uninformative characters and a retention index of060 Monophyly of the group is supported by four synapo-morphies spikes not branched inflorescence length up to 30mm spikes contiguous and a low number of first orderbranches

Biogeographymdash(Figs 5ndash7) Carex spicata and C muricata aresympatric and the two most widespread taxa in the groupCarex spicata is also sympatric with C pairae hybridizingmany times Carex spicata subsp spicata has the widest dis-tribution It occurs from Spain (2degW) to Altai (USSR 85degE)and from southern Scandinavia (60degN) to northern Iran(35degN) It is absent in southernmost Europe we did not findany specimen from Italy south of the Alps and only a fewfrom montane localities in Greece (Pindus and Rodopi) andthe mountains north of Ankara and the Kars in Turkey Thistaxon also occurs in a few localities in the Caucasus

Carex pairae also has a wide distribution (though not aswide as C spicata) It is also a more thermophilous plantmore frequent in Southern Europe (Greece and the AegeanIslands and southern Turkey but absent from other Medi-terranean islands) and reaching the Atlas Mountains in Mo-

rocco and Madeira and it is scarcely represented in the north(Denmark and southern Sweden) It is found in neither Nor-way nor Finland The eastern limit of its distribution area isnot clear It is partially allopatric with C muricata but Cpairae grows at lower altitudes

Carex muricata occurs over a larger area than C pairae butit is less frequent and associated with colder areas in the highmountains of Southern Europe and at lower altitudes in theNorth Our results confirm that C muricata subsp muricatalives only on limestone soils

DISCUSSION

Carex wendelboi was included in sect Phaestoglochin byEgorova (1999) who considered it is related to C spicata butstudying only a single specimen in LE she probably had anerroneous concept of this species After a meticulous study ofthe type material (K) we consider in agreement with otherauthors (Kukkonen 1998) that it must be treated as a syn-onym of C vulpinaris Nees (sect Vulpinae)

Another species sometimes included (Clarke 1894 Kuumlken-thal 1909 Kukkonen 1998) in the section is Carex foliosa butalthough some specimens are small and seem to be similar toC muricata it belongs to sect Multiflorae because its culm isunwinged and less than 1 mm in diameter its inflorescencevery branched with separate spikes and its perigynum smalland not corky at the base

Against common opinion (Kreczetovicz 1935 Kern amp Reich-gelt 1954 David and Chater 1977 Chater 1980 Jermy et al1982 Nilsson 1985 Malyschev and Peschkova 1990 Loos1996 Kukkonen 1998 Egorova 1999 2000) we agree withHadac (1961) and Hooper (1985) that C polyphylla is not re-lated to the Carex muricata aggregate and we consider it tobelong to sect Vulpinae Both C foliosa and C polyphylla ap-pear separated from section Phaestoglochin and next to Cotrubae in the hierarchical clustering

PCoA and hierarchical clustering support the recognitionof four species in two aggregates and clearly separated thetaxa at the species level The dissimilarity index at this levelhas a value up to 025 because the taxa are closely relatedBelow this value of dissimilarity we treated the different taxaas subspecies because of their morphological similarity andthe existence of specimens that are morphologically interme-diate

Hybridization is common in Carex and more frequent atlower taxonomic ranks We agree with Schmid (1983) thathybridization plays an important role in Carex evolution bygenerating new genetic combinations Hybrids are welldocumented in taxa of sect Phaestoglochin (Wallace 1975Jermy et al 1982 OrsquoMahony 1989 Stoeva and Popova 1997)We also found several specimens with hybrid characters inthe contact areas and included some of them in our numeri-cal analyses

Both qualitative and quantitative characters are needed todefine taxa in this section as other authors have indicated(Stoeva and Popova 1997 Repka 2003) In summary allanalyses support four species in the Carex spicata and C mu-ricata aggregates that have a set of very homogeneous mor-phological features Variables without significant differences(leaf width diameter of culm length of lowest bract inflo-rescence length etc) are used to characterize the aggregatesOnly 13 quantitative variables according to post-hoc testhave non-overlapping values in at least one group Perhaps

FIG 3 Hierarchical cluster from UPGMA using Gowerrsquos coefficient of34 specimens of Carex muricata and 54 variables = C muricata subspmuricata = C muricata subsp cesanenesis = C muricata subsp ashokae


frequent hybridization that we have observed contributes tothis homogeneity In the diagrams (Figs 3 4) subspecies areonly clearly identified by the hierarchical clustering Accord-ing to van de Wouw et al (2003) the characters that contrib-ute the most to the clustering at the subspecies level arequantitative

Former studies (Stoeva and Popova 1997 Repka 2003 Mo-lina et al 2006a) and all analyses in this paper confirm thatCarex spicata is the most strongly distinguished species insection Phaestoglochin in Eurasia Carex omeyica C pairae andC muricata are closely related to one another and differ infew characters The sharpest diagnostic features include theperigynium (size shape wing length and denticles of beak)color of pistillate scale shape and length of achene andspikes (shape and position in relation to inflorescence axis)The most important characters for distinguishing subspeciesare perigynium outline and size and presence of denticles onthe beak Perigynium size (length width lengthwidth and

perigynium lengthpistillate scale length) were also consid-ered important by other researchers who studied the Carexmuricata group (Stoeva and Popova 1997 Repka 2003) butStoeva and Popova (1997) observed important variation inthe the length of stem leaves the obvious number of bisexualspikes and male glume width This last character is verydifficult to measure and does not appear to vary betweentaxa in this group

The phylogenetic tree seems to be similar to the hierarchi-cal clustering diagram placing C muricata subsp cesanensiscloser to C muricata subsp ashokae than to subsp muricataThe synapomorphies for C muricata are those quoted abovespikes not branched inflorescence length up to 30 mmspikes contiguous and fewer than 10 first order branches

The differences between the Carex spicata aggregate andthe C muricata aggregate are the purplish-tinged basalsheaths globular-elliptic spike purplish-brown femaleglume culm sharply trigonous in outline perigynium withcorky base ligule with acute apex ligule up to 35 mmperigynium beak up to 125 mm and female glume more

FIG 4 Phylogram based on 67 characters of relationship among taxaCarex muricata aggregates using heuristic search with C otrubae C foliosaand C polyphylla as the outgroup Bootstrap support indicated onbranches The tree is 116 steps long with a consistency index of 067 anda retention index of 060 excluding uninformative characters

FIG 5 Geographic distribution of Carex spicata C spicata subsp an-dresii (left diagonal lines) and C spicata subsp spicata (right diagonallines)

TABLE 3 Distinctive characters for taxa Means plusmn SD for quantitative characters in mm Within a row means with different superscripts differsignificantly (ANOVA p lt 005) L = length DBWPP = Distance from base to widest point of perigynium

C spicata

C pairei C omeyica

C muricata

ssp spicata ssp andresii ssp muricata ssp cesanensis ssp ashokae

Basal sheaths purplish purplish no purplish no purplish no purplish no purplish no purplishBeak brown brown brown green brown brown brownMargins beak scabrid smooth scabrid scabrid scabrid scabrid scabridShape spike elliptic elliptic globular-elliptic globular-elliptic globular globular globularColor glume purplish purplish light brown dark brown dark brown dark brown dark brownShape perigynium ovate ovate oval oval rounded trullate ovalCulm outline sharp sharp obtuse obtuse obtuse obtuse obtuseInflorescence oblong elliptic oblong oblong oblong oblong oblongPistillate scale ovate ovate obovate oval oval oval ovalPerigynium corky present little absent absent absent absent absentApex of ligule acute acute no no no no noAttenuation beak gradual gradual intermediate intermediate abrupt intermediate intermediatePosition perigynium spreading erect spreading spreading erect spreading very spreading very spreading very spreadingWinged perigynium absent absent absent absent completely completely completelyL inflorescence 23ndash42 15ndash35 21ndash30 25ndash34 17ndash35 22ndash30 25ndash45 (-50)Longest ligule 590 plusmn 295b 465 plusmn 102b 183 plusmn 063a 240 plusmn 055a 186 plusmn 075a 189 plusmn 085a 142 plusmn 085a

L perigynium 517 plusmn 029d 477 plusmn 015c 371 plusmn 022a 465 plusmn 016c 406 plusmn 018b 458 plusmn 026c 536 plusmn 041d

Width perigynium 223 plusmn 018bc 217 plusmn 010ab 203 plusmn 014a 242 plusmn 016cd 234 plusmn 018bcd 251 plusmn 017d 233 plusmn 014bcd

L beak 143 plusmn 014bc 128 plusmn 013b 075 plusmn 014a 083 plusmn 005a 088 plusmn 015a 092 plusmn 015a 150 plusmn 010c

L pistillate scale 419 plusmn 033d 417 plusmn 026d 333 plusmn 028ab 392 plusmn 018cd 299 plusmn 033a 335 plusmn 024ab 354 plusmn 038bc

L perigynium glume 124 plusmn 012ab 115 plusmn 010a 112 plusmn 010a 118 plusmn 007a 137 plusmn 015b 137 plusmn 008b 153 plusmn 018c

Lwidth perigynium 233 plusmn 024b 220 plusmn 014b 184 plusmn 016a 193 plusmn 013a 175 plusmn 014a 183 plusmn 013a 230 plusmn 011b

L perigyniumbeak 363 plusmn 025a 375 plusmn 034a 506 plusmn 084bc 561 plusmn 048c 471 plusmn 067b 510 plusmn 078bc 359 plusmn 034a

L perigyniumachene 211 plusmn 010c 200 plusmn 015bc 168 plusmn 011a 173 plusmn 004a 184 plusmn 017ab 181 plusmn 011a 199 plusmn 013bc

L perigyniumDBWPP 366 plusmn 031c 298 plusmn 037b 256 plusmn 024a 249 plusmn 010a 293 plusmn 025b 282 plusmn 032ab 304 plusmn 023b


than 4 mm The bootstrap values support the C spicata ag-gregate by 98 but the C muricata aggregate by only 82The consistency index (CI) is not very low (067) if we com-pare it with other similar morphological studies in Carex(Naczi amp Ford 2001 Starr amp Ford 2001) and the homoplasyindex is quite high (033) In conclusion as only a few mor-phological characters are available for phylogenetic analyseswe intend to pursue molecular studies to confirm these re-sults

The uncommon red color of the basal sheaths in subgenusVignea (Naczi and Ford 2001 Ball 2002) the long and acuteligule and the corky base of the perigynium allow recogni-tion of Carex spicata and have been commonly considered asdiagnostic of this species (eg Chater 1980 Jermy et al 1982)These features separate it from the remaining species of theC muricata aggregate Similar results were obtained in a phy-logenetic study by Hendrichs et al (2004) Within Carex spi-cata micromorphological characters and chorology separateC spicata subsp andresii The smooth beak margin (Fig 8)small corky base few veins smaller perigynium and shorterinflorescence differentiate this subspecies well While speci-mens of C spicata subsp spicata are variable C spicata subspandresii is more homogeneous and occurs in a restricted dis-tribution area

Carex omeyica shows an exclusive set of characters notshared with the other two species in the C muricata aggre-gate It differs from C muricata because it has a perigynumwith unwinged margin and semierect spikes and from Cpairae by its bigger perigynium and darker pistillate scalewith a shorter apex Although the number of veins in theperigynium body was significantly different from that ofother taxa we chose to ignore this character since it proved tobe highly variable in every taxon studied

The taxonomic range of Carex pairae and C muricata has

been discussed several times (David 1976 Chater 1980 Nils-son 1985 Stoeva and Popova 1997 Molina et al 2006a) Allanalysis indicated that C pairae and C muricata are very simi-lar taxa sometimes considered conspecific because one canfind intermediate specimens in the mountains The resultsindicate that C pairae must be considered a species distinctfrom C muricata Nine statistically significant characters canbe used to define C pairae It is unique in having a small(325ndash40 times 175ndash225) perigynium and unique qualitative fea-tures such as the obovate acuminate light brown pistillatescale and spreading oval to ovate perigynium We find nosignificant differences in either spike shape (Chater 1980) orflowering time (Jermy et al 1982 David and Kelcey 1985)The difference in the number of female and male flowers isstatistically significant but we studied only a few specimensand we prefer not to consider this character Thus there areseveral exclusive quantitative and qualitative characters tosupport the segregation of C pairae as a distinct species Thechromosomal results obtained by Hartvig (1986) also supportthis treatment

Diagnostic characters for Carex muricata are those used byChater (1980) and Jermy et al (1982) mentioned under Cmuricata subsp muricata short glume globose spikes andwinged perigynium that is strongly spreading when ripe Cmuricata subsp ashokae differs from the other two subspecies

FIG 6 Geographic distribution of Carex pairae (right diagonal lines)and C omeyica ()

FIG 7 Geographic distribution of C muricata C muricata subsp mu-ricata () C muricata subsp cesanensis (right diagonal lines isolated lo-calities ) and C muricata subsp ashokae (left diagonal lines)

FIG 8 Perigynium beak of Carex spicata a Carex spicata subsp spicata(LEB 82671) b Carex spicata subsp andresii (LEB 49969)


by its large inflorescence (up to 45 cm) and a perigyniumwith a long beak and from C muricata subsp muricata by itsbigger perigynia Malyschev and Peschkova (1990) only rec-ognize a single taxon C muricata L

The hierarchical clustering clearly differentiates Carex mu-ricata subsp muricata and C muricata subsp cesanensis andthe low distance can be explained by the morphological simi-larity C muricata subsp cesanensis has bigger trullate perigy-nia while C muricata subsp muricata has a small roundedperigynium Only Carex muricata subsp muricata is found oncalcareous soils so the separation of C muricata subsp ce-sanensis is clear

Taking into account all available data we suggest all spe-cies have a similar ecology living frequently in man-madehabitats such as roadsides pastures or waste places (Davidand Kelcey 1985 Aeschimann et al 2004) on soils having amoderate to high content of organic matter Carex pairae Cmuricata subsp muricata and C spicata are pioneers (Davidand Kelcy 1985 pers obs) with a generalist strategy Wethink this is typical of section Phaestoglochin in Eurasia

Carex sect Phaestoglochin is widespread in Eurasia fromthe Iberian Peninsula to Kashmir (India) and Altai (USSR)but all of the taxa have different distributions and some areseparated altitudinally Carex spicata and C muricata are themost widespread taxa Carex spicata is the most widespreadspecies It occurs between 500ndash1000 m needs higher moisturein the soil than the other taxa but does not live near the coastCarex omeyica has a restricted distribution and more data areneeded to confirm its full distribution Carex pairae is alsowidespread Although Egorova (1999) supposes it occurs inWest Russia and Hartvig (1986) quoted it from SouthernNorway we did not find any specimens from those places It

grows between 0ndash1500 m but it is not tolerant of very coldenvironments It is more abundant than C muricata whichhas a narrower habitat and prefers high mountains The onlytaxon with a restricted habitat is C muricata subsp muricatabecause it only grows on calcareous soils (Nelmes 1947Chater 1980 Jermy et al 1982 David and Kelcey 1985)

Carex muricata subsp muricata is considered NT (nearthreatened) according to UICN categories because of its re-stricted distribution in Great Britain (Cheffings and Farrell2005) In Spain (Villar 2003) where its populations are con-sidered in regression it was cataloged as DD (deficient data)but after studying all populations we consider it VU (vul-nerable) In other countries because of the uncertain tax-onomy of the C muricata aggregate there is not a correctassignment of the threatened category for this taxon Finallythe limited range of C omeyica indicates it merits protection

From the phylogenetic analysis we can see that Carex spi-cata the most differentiated taxon was probably the firsttaxon to diverge within the C muricata group The apomor-phies allow us to distinguish it Within the Carex muricataaggregate C omeyica differentiated when ice withdrawal iso-lated the Sierra Nevada and the Atlas from the remainingNorthern mountains Carex muricata and C pairae are veryclosely related species probably still in the speciation processjudging by the frequent hybrids and some authors have jus-tified considering only one species On the other hand as aconsequence of the montane habitat the populations of Cmuricata were segregated in three subspecies subsp muricatawhich is adapted to limestone soils subsp cesanenesis occur-ring in western areas and subsp ashokae (several times mis-identified as C polyphylla) in the East

TAXONOMIC TREATMENT

KEY TO THE EURASIAN TAXA OF CAREX SECT PHAESTOGLOCHIN

1 Sheaths basal leaves base of culm and sometimes pistillate scales purplish tinged Ligule acute longer than wide C spicata 22 Perigynium swollen corky at base beak serrulate subsp spicata2 Perigynium slightly corky at base beak smooth subsp andresii

1 Sheaths basal leaves base of culm and pistillate scales not tinged Ligule as wide as or wider than long 33 Perigynium erect-spreading with a green beak C omeyica3 Perigynium spreading or widely spreading with a brown beak 4

4 Perigynium spreading not winged 325ndash40 mm Pistillate scale almost as long as the perigynium light brown ovate toobovate C pairae

4 Perigynium strongly spreading winged 35ndash6 mm Pistillate scale shorter than the perigynium dark brown ovate C muricata 55 Perigynium body rounded 35ndash45 mm beak abrupt subsp muricata5 Perigynium body oval to trullate 425ndash6 mm beak not abrupt 6

6 Perigynium trullate 425ndash5 times 225ndash275 mm beak 075ndash110 mm inflorescence up to 30 mm subsp cesanensis6 Perigynium oval 5ndash6 times 20ndash25 mm beak 125ndash175 mm inflorescence up to 45 mm subsp ashokae

Sect PHAESTHOGLOCHIN Dumort Fl Belg 146 1827 (lectotypedesignated by Jones (1994) Carex muricata L)

CAREX SPICATA AGGREGATE

CAREX SPICATA Huds Fl Angl 349 1762mdashTYPE UNITEDKINGDOM Bedfordshire Eaton Socon on wasteground in gravel field 20 Jul 1946 E Milne-Redhead 5579(Neotype designated by Molina et al 2006b K)

Plants without or with short rhizomes Culms 20ndash80 cmtimes 125ndash200 mm at midheight sharply trigonous slightly sca-brid above Leaves basal sheaths and sheaths red or purpletinged blades about 35 as long as culms widest leaf blades25ndash4 mm plicate to flat ligule 35ndash13 mm longer than wideacute at apex Inflorescence elliptic to oblong 15ndash42 times 8ndash14

mm with 7ndash10(ndash12) elliptic spikes single at nodes sessileusually spikes overlapping the lowest spikes 2ndash8 mm dis-tant bracts glume like the proximal setaceous shorter thaninflorescence 4ndash16(ndash27) mm long Pistillate scale purplish-brown with a narrow green to brown midrib and with orwithout laterals narrow scarious margins 40ndash50 times 20ndash25mm ovate apex apiculate to acuminate 01ndash04 mm Stami-nate glume 35ndash45 times 15ndash20 mm narrower than female An-thers 3 25ndash30 mm long Perigynium erect-spreading tospreading greenish to pale yellowish-brown body ovate toovate-elliptic body nerveless to well marked veins at base40ndash55(ndash6) times 20ndash25 mm base rounded and corky graduallytapered into a beak Beak 115ndash160 mm long brown to red-dish-brown distally serrulate or smooth apex bidentate api-


cal teeth 025ndash05 mm Achenes pentagonal to oval-ovatedark brown when ripe 215ndash275 times 175ndash220 mm Stigmas 220ndash25 mm long

CAREX SPICATA Huds subsp SPICATA (Fig 9AndashG)Carex contigua Hoppe in J Sturm Deutschl Fl 61 1833Carex muricata genuina Gren amp Godr Fl France 3 394

1856Carex contigua varremota F W Schultz Flora 53 459 1870Carex muricata L var genuina subvar incrassata Creacutep

Notes Pl Rar Belgique 126 1859Carex muricata L var virens subvar incrassata Creacutep Notes

Pl Rar Belgique 126 1859Carex muricata L var contigua (Hoppe) Kneuck in Seubert amp

Klein Excurs-Fl Baden 5 52 1891Carex muricata subsp macrocarpa typica Neuman Sver Fl

716 1901Carex muricata L var typica Asch amp Graebn Syn Mitteleur

Fl 2(2) 39 1902Carex muricata L f submonostachya Asch amp Graebn Syn

Mitteleur Fl 2(2) 39 1902

Carex muricata L f mediterranea Asch amp Graebn Syn Mitte-leur Fl 2(2) 39 1902

Carex muricata f pseudoguestephalica Asch amp Graebn SynMitteleur Fl 2(2) 39 1902

Carex muricata L race II- C lumnitzeri Rouy in G Rouy amp JFoucaud Fl France 13 412 1912

Carex lumnitzeri (Rouy) V I Krecz in V L Komarov (ed) FlURSS 3 154 1935

Ligule 45ndash13 mm Inflorescence oblong 23ndash42 times 8ndash10 mmwith 8ndash10 spikes the lowest spikes 3ndash8 mm distant the proxi-mal bracts setaceous shorter than inflorescence 4ndash16(ndash27)mm long Pistillate scale 40ndash50 times 20ndash23 mm Staminateglume 40ndash45 times 15ndash20 mm Perigynium body ovate almostall body with faint or well marked veins at base (45)50ndash55(ndash6) times 20ndash25 mm base rounded swollen and corky Beak115ndash160 mm long irregular serrulate apex usually biden-tate rarely bifid Achene usually pentagonal 225ndash275 times180ndash220 mm Stigmas 2 25 mm long

Representative Specimens ExaminedmdashAUSTRIA Otztaler Alpen amWeg von den Stables-Wiessen Nordtirol nach Nauders silikat an ein-erlessteinmauer 1650 m 9 Aug 1979 A Polatschek (W 1979ndash15840)

BELGIUM Pont-aacute-Celles bord du route 5 Jul 1967 coll P Sotiaux (MA627306)

BULGARIA Bei Tirnoivo 20 May 1898 Prof Urumoff (WU)FINLAND Aboumlensis Korppoo Lohm bordure prairie alluviale anci-

ennement pacirctureacutee 8 Jul 1961 Kukkonnen 678 (MA 274612)FRANCE Vosges Ventron lieux humides 30 Jun 1962 G Gavelle (MA

194487) Miracle 1 Jun 1907 Marcet (MA 016930) Hautes Pyreacuteneacutees Asteacuteafueras del pueblo prado siega borde huacutemedo 600 m 10 Jul 2004 AMolina (LEB 82671) Marecayas pregraves Gles 18 Jun 1861 Meanceau (sn) (P)

GEORGIA Kartli Bakuriani dry places 23 Jul 1918 I Kemularia (MA575399)

GERMANY Bayern Untere Hochebene Verswaschener 1 Jun 1952 WFreiberg (MA 321060)

GREECE Ep Metsovou Katara Pass Ioanninon 9ndash10 km from thesummit summit along main road to Ioannina damp to wet meadows inopening Fagus forest Serpentine 1350 m EK2 9 Jul 1985 M Salmenkallioamp V Salo 1073 (H 1557072)

IRAN East Azerbaijan East side of Hasi Amir Pass on Russian border29 km NE f Ardebil shale cliff 1600 m 21 Jul 1964 M Grant 16396 (W1965ndash17302)

IRELAND Roscommon 1 mile east of Roscommon roadside verge 28Jun 1962 M Mc Callum Wekter 7603 (K)

ITALY Trieste Carey (K)MACEDONIA Gostivar SW of Gostivar near Recane Fagus zone

1300 m 9 Jul 1968 H Den Held (MA 195630)NETHERLANDS Nimwegen Ooy near Nimwegen roadside 28 Jun

1925 herb J Kern and B Reichgelt (K)NORWAY Oslo Grefsen Glads veg 1 Jul 1952 Johanneslid (K)POLAND Breslau Oderdaumlmme bei Carlowitz 15 Jul 1895 Callier (K)ROMANIA Craiova Oltet Luca Mofleni Dolj in silvis sparsis Q ro-

bori-Carpinetum rivulo Jiu 95 m 1 Jun 1965 D Cicircrtu amp M Cicircrtu (MA237002)

SPAIN Huesca Tramacastilla de Tena camino del Iboacuten 1410 m30TYN1832 19 Jul 1986 L Villar amp al (JACA 219783)

SWEDEN Uppland Hamman SE of Svaumlrlinge ca 5 km NW of RimboFasterna Parish Norrtaumllje kommun 27 Jul 1996 S Ortiz A Anderberg ampM X Martiacutenez (SANT 33621)

SWITZERLAND Romanshorn Thurgau 9 Jun 1968 Schatz amp SulgerBuumlel (MA 321059)

TURKEY Ankara A4 31 km N Kicilcahamam Pinus sylvestris wald-drand 1500 m 16 Jul 1977 Dr F Sorger 77ndash51ndash5 (W 1992ndash01219)

UNITED KINGDOM Northern Ireland 39 Co Antrim near BelfastMacedon Point abundant in marshy ground 21 Aug 1945 D Meikle 10(K)

RUSSIA Siberia Altai dist Maima oppidi Gornoaltaisk (Ulala) invicinitate oppidi Gornoalta loco ldquoMotkin pikhtachrdquo dicto 400ndash800 m 10Jul 1972 V Vasaacutek (K)

Carex spicata Huds subsp andresii A Mol Acedo amp Lla-mas subsp novmdashTYPE SPAIN Leoacuten Codornillos30TUM39 850 m 17 Jun 1984 J Andreacutes (holotype LEB

FIG 9 AndashG Carex spicata subsp spicata A Inflorescence (SANT033621) B Pistillate scale (SANT 033621) C Male glume with anther(LEB SANT 033621) D Ligule (LEB 82671) E Perigynium side abaxial(JACA 523286) F Achene (JACA 523286) G Perigynium with side re-moved to show achene (JACA 523286) HndashN Carex spicata subsp andresiiH Inflorescence (JACA 88164) I Pistillate scale (LEB 49969) J Maleglume with anther (LEB 49969) K Ligule (LEB 49969) L Perigyniumside abaxial (LEB 83953) M Achene (LEB 49969) N Perigynium withside removed to show achene (LEB 83953) Bar = 1mm


49969 isotypes ABH BCN HVR JACA LOU MAMACB MAF SANT VAL VIT) Figure 9HndashM

Carice spicatae subsp spicatae similis sed ab ea differt in-florescentia compactiore perigyniis brevioribus 45ndash50 mmlongitudinis habet basim paulum spongiosam in rostroutriculorum denticulis caret

Ligule 35ndash70 mm Inflorescences elliptic to oblongdensely capitate 15ndash35 times 10ndash14 mm with 7ndash10 (-12) spikesspikes usually overlapping difficult to distinguish rarely thelowest spikes 2ndash6 mm distant the proximal bracts setaceous6ndash15 mm long Pistillate scale 39ndash45 times 20ndash25 mm Staminateglume 35ndash425 times 20 mm Perigynium body ovate to ovate-elliptic nerveless or with faint veins at base (4)45ndash50(55) times20ndash25 mm base rounded slightly swollen and corky Beak125ndash160 mm long smooth apex bidentate Achene usuallyoval to ovate 225ndash275 times 16ndash20 mm Stigmas 2 2ndash25 mmlong

Phenology and Floral BiologymdashFruiting from June to JulyWind pollinated

HabitatmdashCarex spicata subsp andresii has a wide ecologicalrange in very different dry communities borders or paths inQuercus pyrenaica Willd forest borders of cultured fieldsmeadows wasteland and moisture zones in the border oflakes

DistributionmdashThis subspecies is endemic from the innerpart of the Iberian Peninsula lives between 500ndash1000 m andis missing in the coast and the Southern areas (Algarve andmost of Andalusia) and disjunct from C spicata subsp spicata(Fig 5)

EtymologymdashThe new subspecies is named in honor of theProf Jaime Andreacutes who began the Carex research in the Uni-versity of Leoacuten

Representative Specimens ExaminedmdashPORTUGAL Tras os Montes eAlto Douro Arredores de Tondella Lobao 1 May 1892 A Moller (MA016936) Braganccedila 17 May 1942 P Barros Carneiro (MA 194485) Miran-dela Lameirado pastagem permanente 4 May 1951 Malato Beliz e Ruivo(MA 268784) Serra do Soajo Senhora da Peneda 1 Jul 1890 A Moller(WU ACQ-J-Nr1188- nordm828)

SPAIN Avila Ramacastantildeas vaguada en dehesa 30TUK2050 350 m10 May 1987 Lucentildeo amp Vargas (MA 349775) Pinar Hoyocasero bosque dePinus sylvestris 1350 m 30TUK3273 9 Jul 1984 M Lucentildeo (MA 293734)Caacuteceres Bantildeos Herbario Antiguo sd Simoacuten de Rojas Clemente (MA143572) Valle Jerte Plasencia bois des chataigniers 1 May 1863 EBourgeau (MA 016922) Ciudad Real Fuentecaliente Sierra Madronavalle del arroyo del Herradero 30SUH9059 740 m 31 May 1997 R Garciacutea(MA 596360) Coacuterdoba Sierra Morena Los Patalos prado sobre suelopizarroso fresco 600 m UH8823 30 Abr 1992 M Melendo (GDA-GDAC42316) Guadalajara La Fuensavintildean 29 Jun 1983 Carrasco Monge Rom-ero amp Velayos (MA 477226) Puebla de Belentildea navajo de Puebla de Belentildea956 m 30TVL8125 1 Jun 1996 E Alvaro amp L Medina LMP 195 (MA649874) Leoacuten Ardoacuten 30TTN80 6 Jun 1978 A Penas (LEB 11870) Avia-dos prado de siega huacutemedo 1000 m 30TN9851 26 Jun 2004 A Molina(LEB 82668) Aviados terreno removido 950 m 30TTN9851 12 Jun 2005A Molina (LEB 83953) Celadilla del Paacuteramo 30TTN61 8 May 1988 JAndreacutes (LEB 19463) Fasgar 29TQH24 18 Jun 1988 J Andreacutes (LEB 39410)Gradefes 30TUN12 19 May 1985 J Andreacutes (LEB 19448) Montejos delCamino Km 14 camino 12 Jun 2002 A Molina (LEB 78166) Murias deRechivaldo 29TQH30 6 Jun 1978 J Andreacutes amp R Carboacute (LEB 27879) No-cedo de Gordoacuten 30TTN84 12 Jun 1983 C Peacuterez Morales (LEB 24911)Ponferrada San Lorenzo finca de las Concabias prado de siega 550 m1 Jun 2001 A Molina (LEB 83920) Ibidem 8 May 2002 A Molina (LEB79022) Ibidem finca los Perros prado de siega 8 May 2002 A Molina(LEB 79021) Riantildeo 30TUN36 12 Jun 1969 J Andreacutes amp R Carboacute (LEB27881) Sahechores 30TUN12 19 May 1985 J Andreacutes (LEB 49909) Sahe-lices del Payuelo 30TUN30 3 Jul 1988 J Andreacutes (LEB 48413) Torenoborde de rebolar (Q pyrenaica) 750 m 29TQH039295 27 May 2005 CAcedo C Lence amp A Molina (LEB 83696) Valle de San Pelayo Buroacuten30TUN36 7 Jul 1988 J Andreacutes (LEB 19467 SANT 28793) Velilla de laReina 22 Jun 1989 J Andreacutes (LEB 50254) Villadangos del Paacuteramo

30TTN71 8 May 1988 J Andreacutes (LEB 39435) Logrontildeo Logrontildeo Jun 1903I Zubia (MA 16862) El Rasillo de Cameros 28 Jun 1875 I Zubia (MA16912) Lugo Monforte de Lemos en prado 29TPH2309 5 Jun 1989 JAmigo amp M I Romero (SANT 19928) Madrid Buitrago in humidis aculiginosis 29 May 1918 C Vicioso (MA 16901) Cadalso de los Vidrios 10Jul 1973 Izco amp Costa (MA 314689) Canal del Manzanares sd Pereda(MA 573185) El Escorial May 1897 C Pau (MA 16904) Quijorna regatoa 1 Km de la localidad 550 m 30TVK0875 14 May 1982 P Monserrat ampD Goacutemez (JACA 017182) Somosierra 9 Jul 1963 M Mayor (FCO 07429)Ourense Rubiaacute Vilardesilva praderas permanentes de Arrhenatheretaliadebajo del pueblo 25 May 1988 J Amigo amp J Jimeacutenez (SANT 27342)Salamanca Valdelosa 19 May 1978 J Saacutenchez (MA 516721) SegoviaFresno de la Fuente borde de una charca 30TVL4485 1100 m 20 Jun1985 A Izuzquiza (MA 506903) Ibidem 20 Jun 1985 A Izuzquiza amp A RBurgaz (MA 314788) Soria Pinar Grande 24 Jun 1959 A Segura Zubi-zarreta (MA 374403) Teruel Villar del Salz pr El Collado 1350 mXL2302 10 Jun 1989 G Mateo (MA 475213) Toledo Talavera carretera aAlcaudete a 10 Km 30SUK4018 17 May 1967 P Monserrat (JACA025967) Zamora Muga de Sayago 760 m 29TQF3486 21 May 1964 PMonserrat (JACA 88164) Tabara alrededores prado de siega 750 m30TTM5335 3 May 1996 B Hernaacutendez (MA 651945)

CAREX MURICATA AGGREGATE

CAREX PAIRAE F W Schultz Flora 51 303 1868mdashTYPEFRANCE Bas-Rhin Dans les focircrets Brumath F WSchultz Herb Norm 1160 27 Jun et 9 Jul 1868 M Paira(lectotype designated by David 1976 K isolectotypesBRNM BRNU PR PRC) Figure 10AndashF

FIG 10 AndashF Carex pairae A Inflorescence (MA 257384) B Pistillatescale (LEB 19505) C Male glume with anther (LEB 82657) D Perigy-nium side abaxial (LEB 19505) E Achene (LEB 19505) F Perigyniumwith side removed to show achene (LEB 19505) GndashL C omeyica (MA410621) G Inflorescence H Pistillate scale I Male glume with anther JPerigynium side abaxial K Achene L Perigynium with side removed toshow achene Bar = 1mm


Carex loliacea Schkuhr Riedgr 22 tab Ee no 91 1801 non L1753

Carex cuprina auct non (I Saacutendor ex Heuff) Nendtv ex AKern 1863

Carex muricata L subsp pairae (F W Schultz) Celak KvetOkoliacute Praž 43 1870

Carex muricata L subsp lamprocarpa Celak Anal KvetCeskaacute 88 1879

Carex bullockiana Nelmes Bot Mater Gerb Bot Inst Ko-morova Akad Nauk SSS R 19 77 1959

Plants without or with short rhizomes Culms 20ndash70 cm times08ndash125 mm at midheight obtusely trigonous slightly sca-brid above sheaths brown Leaves blades about 12 or 13as long as culms widest leaf blades 25ndash3(ndash35) mm plicate toflat ligule 10ndash35 mm wider than long round or subacute atapex Inflorescence oblong 21ndash30 times 8ndash10 mm with 6ndash10(ndash11)globular spikes single at nodes except the lowest sessilelowest spikes usually separated 4ndash9 mm other crowdedbracts glume like the proximal setaceous shorter than inflo-rescence 6ndash15 mm long Pistillate scale light brown with anarrow midrib green to brown with or without lateral scari-ous margins 275ndash36 times 175ndash225 mm obovate to ovate apexacuminate 03ndash04 mm long Staminate glume 30ndash35 times 15ndash20 mm narrower than the female Anthers 3 2ndash25 mm longPerigynium spreading greenish to brownish body roundedto oval-rounded nerveless or faintly veined without wingedmargins 30ndash40 times 175ndash225 mm rounded base more or lesstapered into a beak Beak 05ndash09 mm long brown distallywith serrulate margins reaching the summit apex bifid api-cal teeth 025ndash05 mm Achene oval to ovate yellowishbrown 20ndash275 times 15ndash220 mm Stigmas 2 225ndash30 mm long

Representative Specimens ExaminedmdashBOSNIA-HERZEGOVINA Invalle Bioca prope Sarein 840 m 23 Jun 1932 Maly WU(sn)

CZECHIA Moravia Septentrional Olomouc Palazovreck in Hlubokyacutezleb prope Naacutemest in declivi fruticoso-silvatico 250 m 9 Jun 1932 JOtruba (K)

DENMARK Near Hammel Pot road side 56deg25N 9deg50E 18 Jun1979 I Nielsen et al (MA 314912)

FRANCE LrsquoAude Pla de Bac-Estable Le Caunil solana algo seca congayuba y oreacutegano 1300 m 31T DH4136 6 Jun 1983 P Monserrat y LVillar (MA 257383) Var Le Garde- Frenet 17 Jun 1911 A et E G Camus510 (P)

GERMANY Malente Schleswig-Holstein trockener Waldrandnemoral 14 Jun 1965 G Langer (MA 388490)

GREECE Mavrolithari vallis Arkoudi reuma in humdiiusculis sil-varum 1100 m 28 Jul 1906 coll Halacsy Herb Graecum R Maire et MPetitmengin (WU)

HUNGARY Insula Csepreg prope pagum Tokul en silvaticum um-brosis 25 May 1872 J A Tauschen (WU)

MOROCCO Tetouan Au N de la M F de Talassemtrane preacutes de lapiste 1600 m 35deg06N 5deg06W 21 Jun 1980 Jacquemond et Jeanmonod (MA257384)

NETHERLANDS Nimwegen Habert roadside 5 Jul 1925 J Kern andB Riechgelt (K)

PORTUGAL Castelo Bon berrocal graniacutetico con Quercus pyrenaica 700m 29TPE 7597 13 May 1973 P Monserrat (JACA 078273) Madeira EntreSanto da Serra et Lamareiros talus de terra C Menezes 4 (P) Choupanapar leacuteteacute 700 m C Menezes 7 (P) Prie Ruins ad rupes umbrosis 15 May1902 Gandoger (P) Tras os Montes Braganccedila 29TPG83 28 May 1991 CAguiar (LEB 61347)

ROMANIA Transilvania distr Turda prope oppid Tura solo calc-hum in fisura ldquoCheia Turziirdquo 400 m 22 May 1921 Al Borza and M Peacuteterfi(K)

SPAIN A Coruntildea Corcubioacuten plaial de Quenxe xunto da fonte prado10 m 6 Jun 2004 A Molina (LEB 82657) Burgos Pineda de la Sierra 26Jun 1914 sr (MA 016910) Caacuteceres Cantildeaveral Puerto de los Castantildeossuelo aacutecido alcornocal 450 m 29SQE2612 7 May 1983 E Bayoacuten et al (MA483174) Leoacuten Cadafresnas 29TPH71 7 Jul 1985 J Andreacutes (LEB 19458)Canseco 30TTN96 10 Jul 1987 J Andreacutes (LEB 54544) Morla 29TQG28 8

Jul 1993 M E Garciacutea (LEB 54932) Morredero 29TQG09 25 Jun 1989 FGoacutemiz (LEB 42477) Pentildea de Valdorria 30TUN05 5 Jul 1986 J Andreacutes(LEB 19505) Redilluera 30TTN96 29 Jun 1986 J Andreacutes (LEB 19449)Subida Morredero desde Villar de los Barrios cuneta sobre pizarra 1500m 29T QG09 19 Jun 1981 G Nieto Feliner et al (MA 317397) SalamancaPentildea de Francia La Alberca 4 Jul 1946 A Caballero (MA 016924) SoriaCovaleda riacuteo Quesos humedales siliacuteceos 10 Sep 1971 A Segura Zubi-zarreta (MA 321049)

SWEDEN Hammersta Soumldermanland Oumlsmo 13 Jul 1932 E Asplund(K)

TURKEY Adana Feke Damps roks Dodds cetik Sancan Dere be-tween Gurumze and Suphandere 1000ndash1200 m 1 Jul 1952 Davis 19608(K)

UNITED KINGDOM Surrey sandy bank lower Eashing 1 Jul 1951 NY Sandwith (MA 158554)

Carex omeyica A Mol Acedo amp Llamas sp novmdashTYPESPAIN Granada Lanjaroacuten Sierra Nevada barranco delriacuteo Lanjaroacuten Querceto pyrenaico solo siliceo 1600 m30SVF59 15 Jul 1975 Fernaacutendez Casas amp Saacutenchez Garciacutea(holotype MA 410621) Figure 10GndashL

A Carice spicatae differt perigyniis ovatis sine texto spon-gioso ad basim a Carice pairae propter inflorescentiam atqueperigynias longiores a Carice muricatae quia perigyniae eiuscarent alis et quod illa squamam feminam longiorem habet

Plants with short rhizomes Culms 30ndash70 cm times 08ndash125mm at midheight obtusely trigonous slightly scabrid abovesheaths brown to dark brown Leaves blades about 12ndash13as long as culms widest leaf blades 25ndash30 mm plicate toflat ligule 10ndash30mm wider than long round or subacute atapex Inflorescences oblong 25ndash34 mm times 8ndash11 mm with 6ndash9globular-elliptic spikes one per node sessile spikes usuallyoverlapping the lowest spikes 4ndash9 mm distant bracts glumelike the proximal setaceous shorter than inflorescence 5ndash12mm long Pistillate scales dark brown with a narrow green tobrown midrib and without scarious margins 375ndash425 times 20ndash23 mm oval apex apiculate 01ndash02 mm Staminate glume40ndash45 times 175ndash20 mm narrower than the female Anthers 3175ndash3 mm long Perigynium erect-spreading pale greenishto brown body oval nerveless or whit faint veins in thebody 44ndash50 times 225ndash26 mm base rounded not corky apexmore or less gradually tapered into a beak Beak 075ndash10 mmlong green to brown distally with serrulate margins reach-ing the summit apex bidentate apical teeth 025ndash06 mmAchene ovate to oval light brown 25ndash28 times 175ndash225 mmStigmas 2 20ndash250 mm long

Phenology and Floral BiologymdashFlowering begins in MayFruiting from June to July Wind pollinated

HabitatmdashCarex omeyica occurs in forests of middle-highmountain 1500ndash1700 m in gullies on cedar or oak forestsQuerceto pyrenaicum on acid soils

DistributionmdashEndemic to the Sierra Nevada in the South-ern Spain and Atlas Range in the Moroccan Rif (Fig 6)

EtymologymdashThe new species is named from the Omeyasa family of caliphs in Cordoba (Al-Andalus) between 929ndash1031 AD

Representative Specimens ExaminedmdashMOROCCO Tazzeka cidraiedu Jebel Tazzeka partie inferieur 1750 m 22 Jun 1952 Ch Sauvage (MA274379)

CAREX MURICATA L Sp Pl 974 1753mdashTYPE Habitat in Eu-ropae nemoribus humentibus (lectotype designated byMarshall 1907 LINN 110026)

Plants without or with short rhizomes Culms 20ndash100 cm times10ndash20 mm at midheight obtusely trigonous slightly scabridabove basal sheaths and sheaths brown to dark brown


Leaves blades half as long to as long as culms widest leafblades 225ndash45 mm plicate to flat ligule 05ndash35 mm widerthan long round or subacute at apex Inflorescence oblong17ndash45(ndash50) times 8ndash13 mm with 5ndash10 globular spikes single atnodes sessile usually lowest spikes separated 3ndash15 mmother crowded bracts glume like the proximal setaceousshorter than inflorescence 4ndash20 mm long Pistillate scale darkbrown some times reddish with a narrow green to brownmidrib and without scarious margins 225ndash375 times 175ndash225mm ovate apex acute to acuminate 0ndash04 mm long Stami-nate glume 275ndash425 times 15ndash2 mm narrower than female An-thers 3 175ndash25 mm long Perigynium strongly spreadingdull greenish to yellowishbrown body round trullate oroval nerveless or faint veins with a winged margin 30ndash60 times20ndash275 mm base rounded to cuneate abruptly or no ta-pered into a beak Beak 065ndash175 mm long brown distallywith serrulate margins reaching the summit or the body ofthe perigynium apex bidentate to bifid apical teeth 025ndash06mm Achene round to oval light brown to dark brown 19ndash35 times 160ndash225 mm Stigmas 2 15ndash25 mm long

CAREX MURICATA L subsp MURICATA (Fig 11AndashD)Carex echinata sensu Kuumlk in Engler Pflanzenr IV (20) 160

1909 non Murray 1770C pairae F W Schultz subsp borealis Hyl Nord Kaumlrlvaumlxtfl

2 386 1966

Culms 20ndash87 cm times 10ndash15 mm at midheight Leaves ligule125ndash25 mm blades about 1frasl2 or 13 as long as culms widestleaf blades 225ndash4 mm Inflorescence 17ndash35 times 8ndash12 mm with(5)7ndash10 spikes usually lowest spikes separated 3ndash10 mmproximal bracts 4ndash10 mm long Pistillate scale 225ndash35 times 18ndash20 mm oval apex acute to apiculate 0ndash02 mm long Stami-nate glume 275ndash325 times 15ndash2 mm Anthers 3 175ndash2 mm longPerigynium body round to oval-round with a broad wingedmargin 375ndash425 times 20ndash275 mm rounded base more or lessabruptly tapered into a beak Beak 065ndash115 mm long withserrulate margins sometimes reaching the summit apex bi-dentate to bifid apical teeth 025ndash06 mm Achene round toovate light brown 19ndash26 times 160ndash225 mm Stigmas 2 175ndash2mm long

ObservationsmdashDavid (1976) made a nomenclatural revi-sion of Carex muricata group Many authors have mistaken Cspicata C muricata and C pairae From Marshall (1907) Cspicata and C muricata are recognized as different taxaNelmes (1947) recognize C muricata and C pairae as differenttaxa Hylander (1966) and later David (1976) consider C mu-ricata and C pairae at subspecies rank This last treatment isfollowed by Chater (1980) and most of modern authors(Jermy et al 1982 David amp Kelcey 1985 Lucentildeo 1994 Sell ampMurrell 1996 Lambinon 2004)

Representative Specimens ExaminedmdashAUSTRIA NiederoumlsterreichWinzerdorf 8 Jun 1941 H Noumlthig (W 1967ndash10168)

BELGIUM Liegravege Neu-Moresnet entre la Gueule et lrsquoancien route Aatalus herbeux route a Aachen ancien halde calaminaire 28 May 1975 PAuquier W Bellotte et J Duvigneaud (MA 387226) Namur Dion (al S deWinenne) coupe forestiegravere sur le plateau sol riche 29 May 1971 J Du-vigneaud (MA 274352)

DENMARK Hjorto 1864 Leffley (W 1961ndash14286)FINLAND Liuhto Varsinais-Suomi Kisko ca 300 m from Suoranta

farmhouse steep slope with deciduous bushes basic rock 50 m 27degE669303099 FM 1 27 Jun 1985 I Kukkonen (MA 367574)

FRANCE Alpes de Savoie Brezon marais 1 Jul 1848 E Bourgeau (K)Saogravene et Loire Le Bourgneuf murgers des vignes auteur du chacircteau 4Jun 1888 Ch Ozanon (MA 016806)

GERMANY Bayern Ries beim Brennhof 1 Jun 1981 R Fischer (MA387224)

MACEDONIA Crnogora Montenegro Durmitor near Zabljak be-tween village and Crno Jezero 24 Jul 1973 Hooper 3473 (K)

NORWAY Akirshusamt Vold Cfoke Barum 26 Jun 1917 RS Fridlz(K)

POLAND Albertusoka Hill near R Vistula chalk cliff 1 Jul 1976 BeyerSchilling amp Keesing 18 (K)

RUSSIA Pskow Borissowiczi in decliviis 13 Jun 1900 W Andrejew(K)

SLOVAKIA Moravia Oriental Carpathi m Vsetin in monte Rybnickyin fageto 750 m 1 Jun 1930 G Piacutecan (K)

SPAIN Leoacuten Marantildea circo glaciar de Mampodre repisa sobre grietascalizas algo nitrofilo 1550 m 3 Jul 2004 C Lence amp A Molina (LEB82647) Teruel Collado de la Gitana Sierra de Guacutedar Valdelinares um-briacutea 1860 m 5 Jul 1957 P Monserrat (MA 169375)

SWEDEN Uplandia N J Andersson (MA 059967)UNITED KINGDOM Yorkshire near Gordale 64 mid west Yorkshire

limestones slopes 26 Jun 1934 E Milne Redhead amp N Y Sandwith 2016 (K)

Carex muricata L subsp cesanensis A Mol Acedo amp Lla-

FIG 11 AndashD Carex muricata subsp muricata A Pistillate scale (LEB070509) B Achene (JACA 543271) C Perigynium side abaxial (JACA10120975) D Inflorescence (JACA 88164) EndashH C muricata subsp ce-sanensis (LEB 80889) E Pistillate scale F Achene G Perigynium sideabaxial H Inflorescence IndashL C muricata subsp ashokae (H 1498323) IPistillate scale J Achene K Perigynium side abaxial L InflorescenceBar = 1 mm


mas subsp novmdashTYPE FRANCE Les Hautes AlpesLac Lauvitel (Parc National des Ecrins) borde caminoladera seca sobre roca aacutecida 1600 m 3 Aug 2004 AMolina (holotype LEB 82653) Figure 11EndashH

Carice muricatae subsp muricatae similis sed ab ea differtperigyniis maioribus 45ndash525 mm rhombi formam habensatque aquenios ovatos

Culms 20ndash85 cm times 10ndash15 mm at midheight Leaves wid-est leaf blades 3ndash4 mm ligule 10ndash35 mm blades about 12or 13 as long as culms Inflorescence oblong 22ndash30 times 9ndash13mm with 5ndash7(9) spikes usually lowest spikes separated 6ndash10mm proximal bracts setaceous 4ndash11 mm long Pistillatescales 30ndash35 times 175ndash21 mm apex acute to apiculate 0ndash02mm long Staminate glume 30ndash425 times 15ndash2 mm long An-thers 3 175ndash2 mm long Perigynium body trullate to ovate-trullate with a broad winged margin 45ndash525 times 225ndash275mm base cuneate more or less tapered into a beak Beak075ndash125 mm long with serrulate margins reaching the sum-mit apex bifid apical teeth 03ndash06 mm Achene oval yel-lowish brown 2ndash3 times 175ndash225 mm Stigmas 2 2ndash250 mmlong

Phenology and Floral BiologymdashThe flowering begins inMay Fruiting from June to July Wind pollinated

HabitatmdashCarex muricata subsp cesanensis is adapted tomountains between 850 and 1800 m It grows on neutral oracid materials of forest clearings (beech pine etc) pasturesmoist cliffs way sides and other disturbance habitats

DistributionmdashMountains of Southern Europe (PyreneesAlps Balcans and Carpates) to South Turkey (Toros Dagli-ari) and also from Northern Europe (Norway Estonia) out ofmountain areas (Fig 7) We think that in colder areas it canlive at lower altitude as it occurs with C muricata subspmuricata

ObservationsmdashThe limits between the two subspecies arenot clear Toward the East the peryginium size increasesbecoming similar to C muricata subsp ashokae and there is noimportant geographic barrier to isolate both subspecies Thespecimens from the Carpathans are bigger and have bigperigynia similar in size to C muricata subsp ashokae buttrullate in outline Currently and as we have not seen a lot ofmaterial from this area we consider them to belong to Cmuricata subsp cesanensis In Turkey the southern specimenshave a trullate outline and short beaks and are closer to Cmuricata subsp cesanenesis but those from the Northeast(Kars) have oval perigynia and longer beaks and we considerthem subsp ashokae although the achenes are less than 5 mmlong

Conservation StatusmdashLocally a frequent plant but inSpain with only three populations it should be consideredEN (endangered)

EtymologymdashThe new species is named after Cesana theplace where the first specimen was found

Representative Specimens ExaminedmdashAUSTRIA Stubaier Alpen Un-teres Gurgler Tal rechte Talseite silicatat an einem Wiesenzaun 1650 m23 Aug 1987 A Polatschek (W 1987ndash7007) Flora Vindobonienssis Vienna2 Jun 1880 Carolus Aust (W 1978ndash01008) Niederosterreich GagravenserudorfMarderfeld 1 May 1966 G H Gute (W 1968ndash20267) Nordtirol ZillertalesAlpen Zillergrund silikat 1100 m (W 1976ndash14541) Von KaumlrntenKarawanken Freibachtal 13 Jul 1970 M Pull (W 1972ndash17810)

BOSNIA-HEZERGOVINA Near Sarajevo Trebevia 1500 m 8 Jul1960 S S Hooper 760 (K)

BULGARIA Rila prope monasterium Rila in margine siroce 6 Aug1939 H Lindberg (H 1301868)

FRANCE Isereacute Rivoire de Allemont claro bosque 1100 m 30 Jul 2004A Molina (LEB 82650)

ESTONIA Laane Wesenberg (Rakvere) Peithof in prato sicco litto-rale 25 Jun 1914 E E Ditmer (K)

ITALY Piamonte Cesane Turinese talud boscoso 44deg57N 6deg48E 1750m 3 Aug 2003 A Molina (LEB 80889)

NORWAY Satersdalen Aardal Fonekleiven 24 Jul 1903 Arkell Roske-land (K)

RUSSIA Saratov district (Area 1) 4 Jul 1968 Lovelius (K)SPAIN Huesca Turbon al pie del cantil herboso y huacutemedo 1800 m

9 Jul 1952 P Monserrat (MA 168226) Seira SW de Sierra de la ChiaMonte de la Carlania 1740ndash1820 m 31TBH 8810 10 Jul 1985 G Mon-serrat (JACA 842185) Collado de Bonanza hayedo pinar y pastos sobreel Collado 1360ndash1480 m 31TCG 0699 3 Jul 1987 J A Seseacute amp J MMonserrat (JACA 644687) Leacuterida Val de Tredos Araacuten prados secosumbrosos 1350 m 1 Jul 1995 A Pallareacutes (MA 561289) Eriste subidarefugio de Forcau Pirineo Central 1600ndash2000 m 31T BH92 19 Jul 1987G Nieto Feliner amp al (MA 480032 and MA 374477)

TURKEY Tauria Distr Alushta inter pylas Angara et clivum orien-talem montis Chatyr-dag 800ndash1200 m 1 Aug 1977 V Vasaacutek (W 1986ndash01638)

Carex muricata L subsp ashokae A Mol Acedo amp Llamassubsp novmdashTYPE India Kashmir Alibad 9000 feet 9July 1876 C B Clarke 28644 (holotype K) Figure 11IndashL)

Carice muricatae subsp muricatae similis sed ab ea differtquia planta robusta est perigynias maiores habens 5ndash6 mmin rostrum longiorem gradatim contractas

Culms 35ndash100 cm times 15ndash2 mm at midheight Leaf blades aslong or 34 as long as culms widest leaf blades 30ndash45 mmligule 05ndash20 mm Inflorescence 25ndash45 (-50) times 10ndash12 mm with5ndash9 spikes lowest spikes usually separated 7ndash15 mm proxi-mal bracts setaceous 5ndash20 mm long Pistillate scales 30ndash375times 175ndash225 mm long apex apiculate to acuminate 02ndash04 mmlong Staminate glume 30ndash40 times 15ndash2 mm long Anthers 320ndash225 mm long Perigynium body oval to ovate with awinged margin 5ndash6 times 225ndash275 mm base rounded to cune-ate gradually tapered into a beak Beak 125ndash175 mm longwith serrulate margins reaching the body apex bifid apicalteeth 04ndash075 mm Achene oval light brown to dark brown25ndash35 times 175ndash225 mm Stigmas 2 15ndash2 mm long

Phenology and Floral BiologymdashThe flowering begins inMay Fruiting between June to July and August Wind pol-linated

HabitatmdashCarex muricata subsp ashokae occurs in cold andrainy places in high mountains at (700ndash)1300ndash2800 m in al-pine meadows rocky ravines rocky streams and gorges andalso in open woods pastures and subalpine steppe It ap-pears relatively indifferent as to soil

DistributionmdashMountains of Eastern Europe and theMiddle East from the Caucasus (Armenia and Georgia) andthe Kars (NE Turkey) towards Central Asia through theZagros Mountains (Iran) to the Pamirs (Kashmir India) andTargabatay (Tadzhikistan Fig 7)

ObservationsmdashSome reports of Carex polyphylla by Egor-ova (2000) from several localities in Central Asia must bereferred to Carex muricata subsp ashokae but we have notfound any materials from Tien Shan (Minusink East Siberia)and West Siberia (Altai) Since Karelin and Kirilow (1841)described C polyphylla from the Altai perhaps materials thatEgorova quoted (lc) were hybrids between these taxa

Conservation StatusmdashBecause C polyphylla is widespreadin high mountains there is no need of protective conserva-tion status for this species

EtymologymdashThe new subspecies is named from Ashokawhich means in Sanskrit without sorrow who was a famousemperor (-232 BC) of the Mauryan dynasty


Representative Specimens ExaminedmdashARMENIA Caucasus distrRazdan divi montis Ketandag in vecinatati pagi Charencavan 1700ndash2100 m 7 Jul 1975 V Vasaacutek (W 1983ndash05736)

GEORGIA Caucasus Aragac montis apud ruinas Amberd 2100ndash2300 m 23 Jul 1975 V Vasaacutek (W 12975) and (W 1993ndash00618) Araratmontes ldquoGegamski khrebetrdquo in vecinitate ruinarum pagi Akhkeng1800ndash2100 m 10 Jul 1975 V Vasaacutek (H 1455112) and (W 12973) Cartha-linia Abastuman 9 Jun 1881 AH amp VF Brotherus 872 (H 1301888)Tbilisi Dabahane gorge above Botanical institute 600ndash700 m 30 Jun1959 Davis 33887 (K) Kartli Westteil des Trialetischen Gebirges talein-schmitt am fuss des berges Kokhta gora ca 2 km E Bakuriani gegenMitarbi strassenrad waldrand 1650 m 41deg45N 43deg33E 15 Jul 1997 PSchoumlnswetter amp A Tribsch (WU)

INDIA Kashmir Alibad 9000 feet 9 Jul 1892 C B Clarke 28644 (K)Gulmarg 8000 feet 31 Jul 1926 R R Steward (K) Pahlgam 8000 feet 31Jul 1945 R R Steward 21528 (K) Ramoo 6000 feet 10 Jul 1876 C BClarke 28533 (K) Above Utrot (Swat State N W F P) 8ndash9000 feet 21 Jul1953 R R Stewart amp A Rahman 25223 (K)

IRAN Kalardasht Flush alpine meadow limestone 3000 m 9 Aug1960 Spooner X5 (K) E Mazandaran NW Khorasan Center South side ofDivar Kaji Mountain summit (Golestan National Park) open Quercusmacranthera forest and subalpine steppe 2200ndash2300 m 37deg24N 56deg02E12 Jul 1995 H Akhani 11778 (H 1695169) Gilan Mountain above Damesheast of Rudbar(Herb Ariamehr Bot Garden) 1900 m low regenaratingFagus forest 21 Jun 1975 Wandelbo amp Ann Ala 18175 (W 03321)

PAKISTAN Chitral 9000 feet 25 May 1895 Sarg Lt Harris amp J Wlls16739 (K)

TURKEY Armenia turcica Guumlmuumlschkhane Szandschack 6 Aug 1894P Sintesis 7404 (K) and (WU) Kars Yalnizcam Daglari sleep meadowtutfed 2100ndash2300 m 19 Aug 1957 Davis amp Hedge D 32498 (H 1205643)Artvin Coruh mountain above Artvin igneus pasture at edge of Piceaforest tufted 1700 m 19 Jun 1957 Davis amp Hedge 29712 (K) N of Kas-tamonu side of Ilgaz Daglari igneus knoll 1950 m 28 Jul 1962 DavisCoode amp Yaltirik 38354 (K) Lazistan Djimil sous alpine 16 Jul 1866 BBalansa (P 00281835)

TADJIKISTAN Songaria Targabatai 1841 A G Schrenk (K) Kondaravalley Hissar Mts c 30 Km N of Dushanbe in Varsob valle just abovethe Acadm Sciences field station shady brooklet ravine in rather drymarginal area good stand 1300 m 22 Jun 1983 I Kukkonen (H 1498323)

ACKNOWLEDGMENTS We are grateful to the staff of the consulted her-baria for their help locating material We also want also to express specialgratitude to David Simpson for his kind help during our visits to KewHerbarium We specially thank Miguel Rubio and Manuel MarcosCasquero for the Latin diagnoses and Xurxo Magaz for helping us withthe drawings Finally we thank the Junta de Castilla y Leoacuten that granteda High Studies Licence to the first author and the grant LE025A05 thatpartially supported our research

LITERATURE CITED

Aeschimann D K Lauber D M Moser and J P Theurillat 2004 FloraAlpina vol 2 Paris Beliacuten

Ball P W 2002 Carex L section Phaestoglochin Dumortier Pp 285ndash297 inFlora of North America north of Mexico vol 23 eds Flora of NorthAmerica Editorial Committee New York Oxford University Press

Cheffings C F and L Farrell (eds) 2005 The vascular plant Red DataList for Great Britain Species Status Assessment Project 7 1ndash116

Chater A O 1980 Carex L Pp 290ndash323 in Flora Europaea vol 5 eds T GTutin V H Heywood N A Burges D M Moore S M Waters andD A Webb Cambridge Cambridge University Press

Clarke C B 1894 Carex L Pp 699ndash748 in Flora of British India vol 6 edJ D Hooker London L Reeve and Co

David R 1976 Nomenclature of the British taxa of the Carex muricata Laggregate Watsonia 11 59ndash65

David R and J G Kelcey 1975 Carex muricata L sensu Nelmes and Carexbullockiana Nelmes Watsonia 10(4) 412ndash414

David R and A O Chater 1977 Carex polyphylla Kar amp Kir and Carexleersiana Rauschert Watsonia 11 253ndash254

David R and J G Kelcey 1985 Carex muricata L aggregate (Biologicalflora of the British Isles) Journal of Ecology 73 1021ndash1039

De Langhe J E 1944 Sur le groupe du Carex muricata L en BelgiqueBulletin de la Socieacuteteacute Royal de Botanique de Belgique 76 39ndash50

Egorova T V 1999 The sedges (Carex L) of Russia and adjacent states(within the limits of the former USSR) ed A L Takhtajan St Peters-burg State Chemical-Pharmaceutical Academy

Egorova T V 2000 Plants of central Asia Plant collections from China andMongolia vol 3 Sedges and rushes ed VI Grubov Enfield NHPlymouth UK Science Publishers

Hadac E 1961 The family Cyperaceae in Iraq Bulletin of the College ofScience 6 1ndash27

Hartvig P 1986 Chromosome numbers in Nordic populations of theCarex muricata group (Cyperaceae) Acta Universitatis UpsaliensisSymbolae Botanicae Upsaliensis 27(2) 127ndash138

Hendrichs M S Michalski D Begerow F Oberwinkler and F H Hell-wig 2004 Phylogenetic relationship in Carex subgenus Vignea(Cyperaceae) based on ITS sequences Plant Systematics and Evolution246 109ndash125

Hooper S S 1985 Carex L Pp 386ndash406 in Flora of Iraq vol 8 eds C CTownsend and E Guest Baghdad Ministry of Agriculture andAgrarian Reform

Hulteacuten E and M Fries 1986 Atlas of North European vascular plants (northof the Tropic of Cancer) Koumlniegstein Koeltz Scientific Books

Hylander N 1966 Carex L Pp 84ndash91 in Nordisk Kaumlrlvaumlxtflora vol 2Stockholm Almqvist and Wiksell

Jermy A C A O Charter and R W David 1982 Sedges of the BritishIsles (a new edition of British sedges) Ed 2 Botanical Society of theBritish Isles 1 1ndash268

Jones S D 1994 A new species of Carex (Cyperaceae Phaestoglochin)from Oklahoma and Texas typification of section Phaestoglochin andnotes on sections Bracteosae and Phaestoglochin Sida 16 341ndash353

Karelin G and J Kirilow 1841 Enumeratio plantarum anno 1840 inregionibus altaicis et confinibus collectarum Bulletin de la SocieacuteteacuteImpeacuteriale des Naturalistes de Moscou 1841(3) 857ndash861

Kern J H and T J Reichgelt 1954 Carex L Pp 1ndash133 in Flora neerlandicavol 1(3) eds Van Th Weevers B H Danser and J Heimans Am-sterdam Koninklijke Nederlandsche Botanische Vereeniging

Kreczetovicz V L 1935 Carex L Pp 86ndash369 in Flora of the USSR vol 3 edV L Komorov Moscow Botanicheskii Institut Akadamiya NaukUSSR

Kuumlkenthal G 1909 Cyperaceae-Caricoidae Pp 1ndash824 in Das Pflanzen-reich IV 20 (Heft 38) ed A Engler Leipzig W Englemann

Kukkonen I 1998 Cyperaceae in Flora Iranica vol 173 ed K H Rech-inger Graz-Austria Akademische Druck-u Verlagsanstalt

Lambinon J 2004 Carex L Pp 829ndash860 in Nouvelle flore de la Belgique duGrand-Ducheacute de Luxembourg du Nord de la France et des Reacutegionsvoisines 5ordf edition eds J Lambinon L Delvosalle and J Du-vigneaud Meise Patrimoine du Jardin Botanique National de Bel-gique

Loos G H 1996 Zur identitaumlt von Carex leersiana Rauschert C chaberti FW Schultz C polyphylla Kar and Kir und C guestphalica (Boenn exRchb) Boenn ex O F Lang Feddes Repertorium 107(1ndash2) 61ndash74

Lucentildeo M 1994 Monografiacutea del geacutenero Carex L en la Peniacutensula Ibeacutericae Islas Baleares Ruizia 14 1ndash139

Maire R 1957 Caricoideae Pax Pp 97ndash180 in Flore de lAfrique du Nordvol 4 Paris Paul Lechevalier

Malyschev L I and G A Peschkova 1990 Flora Sibiri vol 3 Novosi-birsk Siberia Nauka Sibirskoe Otdelenie

Marshall E S 1907 Carex and Epilobium in the Linnean herbarium Jour-nal of Botany British and Foreign 45 363ndash368

Molina A C Acedo and F Llamas 2006a Delimitacioacuten taxonoacutemica deCarex grupo muricata (Cyperaceae) en Europa Resultados prelimin-ares Bulletin de la Socieacuteteacute de Histoire Naturelle Toulouse 141 57-61

Molina A C Acedo and F Llamas 2006b Typification of some Hudsonplant names in Carex Taxon 55 1009-1013

Mouterde P S J 1966 Nouvelle Flore du Liban et de la Syrie vol 1 BeirutImprimerie Catholique

Naczi R F C and B A Ford 2001 Systematics of the Carex jamesiicomplex (Cyperaceae sect Phyllostachyae) Sida 19(4) 853ndash884

Nelmes E 1947 Two critical groups of British sedges Reports of theBotanical Exchange Club British Isles 13 99ndash105

Nilsson Ouml 1985 Carex L Pp 73ndash158 in Flora of Turkey and East AegeanIslands vol 9 ed P H Davis Edinburgh Edinburgh UniversityPress

OrsquoMahony T 1989 Carex divulsa Stokes times C muricata L ocurring as aspontaneous garden hybrid and wild plant in Cork new to IrelandThe Irish Naturalistsrsquo Journal 23(4) 137ndash141

Pignatti S 1982 Carex L Pp 636ndash676 in Flora dItalia vol 3 BolognaEdagricole

Podani J 2001 SYN-TAX 2000 Computer programs for data analysis inEcology and Systematic Budapest Scientia Publishing

Repka R 2003 The Carex muricata aggregate in the Czech Republic mul-


tivariate analysis of quantitative morphological characters PresliaPraha 75 233ndash248

Repka R and J Danihelka 2005 Typification of the name Carex muricatavar lamprocarpa Wallr and its nomenclatural consequences PresliaPraha 77 129ndash136

Saarela J M and B A Ford 2001 Taxonomy of the Carex backii complex(Section Phyllostachyae Cyperaceae) Systematic Botany 26(4) 704ndash721

Schmid B 1983 Notes on the nomenclature and taxonomy of the Carexflava group in Europe Watsonia 14 309ndash319

Sell P and G Murrell 1996 Carex Pp 82ndash120 in Flora of Great Britain andIreland vol 5 Cambridge Cambridge University Press

Stoeva M and E Popova 1997 A taxonomic study of Carex sect Phaes-toglochin and sect Stellulatae (Cyperaceae) in Bulgaria Bocconea 5787ndash796

Starr J R and B A Ford 2001 The taxonomic and phylogenetic utilityof vegetative anatomy and fruit epidermal silica bodies in Carexsection Phyllostachys (Cyperaceae) Canadian Journal of Botany 79 362ndash379

Swofford D L 2002 PAUP phylogenetic analysis using parsimony ( andother methods) Version 40 beta 10 Sunderland Sinauer Associates

Thiele K 1993 The holy grail of the perfect character the cladistic treat-ment of morphometric data Cladistics 9 275ndash304

van de Wouw M N Maxted and B V Ford-Lloyd 2003 A multivariateand cladistic study of Vicia L ser Vicia (Fabaceae) based on analysisof morphological characters Plant Systematics and Evolution 237 19ndash39

Villar L 2003 Carex muricata L subsp muricata P 922 in Atlas y Libro Rojode la Flora Vascular Amenazada de Espantildea eds A Bantildeares G BlancaJ Guumlemes J M Moreno and S Ortiz Madrid D G C N

Vollmann F 1903 Der formenkreis der Carex muricata und seine Verbrei-tung in Bayern Denkschriften Koumlniglich Bayerische Botanischen Gesell-schaft in Regensburg 2(8) 55ndash90

Wallace E C 1975 Carex L Pp 513ndash540 in Hybridization and the flora of theBritish Isles ed C A Stace London Academic Press

Webber J M and P W Ball 1984 The taxonomy of the Carex rosea group(section Phaestoglochin) in Canada Canadian Journal of Botany 622058ndash2073


divergence among species among subspecies within a species and toidentify the best diagnostic characters for them Because the variance ofseveral variables was heterogeneous as the Levenersquos test indicated thesedata were logarithmically transformed for ANOVA Qualitative charac-ters were studied with analogous Kruskal-Wallis analysis Statisticalanalyses were performed with SPSS 130

Phylogenetic AnalysismdashPhylogenetic analysis was performed using 44characters Only the statistically significant variables among those listedin Table 1 were included in the phylogenetic analyses and all variableswere equally weighted As we have few qualitative characters availablefor this study we used quantitative data because one can recover phy-logenetic information from these data (Thiele 1993) Qualitative charac-ters were coded as in former analyses To avoid arbitrarity the quanti-tative data were coded according to the groups in ANOVA post-hoc testAlthough the results of phylogenetic analyses are better if all data aretreated as unordered we think that the quantitative characters evolve

progressively Values of quantitative characters were coded in a range of0ndash3 and treated as ordered multistate characters (Table 2) Carex otrubaeC foliosa and C polyphylla were used as outgroups Heuristic search wasused and the robustness of clades was assessed by means of bootstrapanalysis Maximum parsimony analysis was performed with PAUP40b10 (Swofford 2002) Data are deposited in TreeBASE (study numberS2026)

BiogeographymdashSpecimen label data were used to document or delimitthe geographic distribution of each taxon Information from Clarke(1894) Maire (1957) Hylander (1966) Mouterde (1966) Chater (1980)Jermy et al (1982) Pignatti (1982) Hooper (1985) Nilsson (1985) Hartvig(1986) Hulteacuten and Fries (1986) Malyschev and Peschkova (1990) Lucentildeo(1994) Kukkonen (1998) Egorova (1999 2000) and Aeschimann et al(2004) was compared with our own data to infer the real distribution ofthe taxa The distribution maps are based on herbarium specimens Thedata and distribution maps were plotted using ArcGIS 9

RESULTS

In the preliminary revision of herbarium material wefound little morphological variation among taxa The char-acters that allow us to include a specimen in these aggregatesare leaves 25ndash45 mm wide auricles absent culm diameter075ndash20 mm inflorescences 18ndash45 mm long with spikes 5ndash11close together and sessile (except sometimes the lowest)bract short-setaceous glumes acuminate or acute brownwithout arista perigynum plano-convex 33ndash60 times 18ndash275mm margin winged or not beak up to 15 (175) mm

According to previously published data Carex foliosa Cpolyphylla and C wendelboi do not belong in Carex sect Phaes-toglochin Carex wendelboi has a tight inflorescence that iselongate ellipsoidal or ovoid with compound spikes pistil-late scales acute or aristate and perigynia 27ndash3 times 1ndash14 mmCarex foliosa has leaves up to 7 mm wide with auricles in-florescences 30ndash100 mm long with compound spikes havingmany flowers and perigynia 27ndash33 times 12ndash15 mm subtendedby an awned glume Neither the protologue of Carex poly-phylla nor the type conserved at Kew have the diagnosticcharacters of this section mainly because it is a very robustplant and its inflorescence has compound spikes Thereforethese taxa were not included in the PCoA and ANOVA tests

A high Pearsons correlation was found in the distance be-tween the lowest spikes number of flowers nerves in theperigynium body and width of the perigynium marginsConsequently only 58 characters (marked with rsquorsquo in Table 1)were used for further analysis four additional characterswere excluded in the PCoA because they did not show varia-tion within the Carex muricata aggregates

The scatter plot of the samples in the first two axes of thePCoA shows two main groups (Fig 1) Axis 1 provides sepa-ration of Carex spicata from the rest by its corky perigyniumbase distance from base to widest point of perigynium andligule apex while C muricata and C pairae are separated inaxis 2 mainly by perigynium length perigynium width andthe ratio perigynium length perigynium width Specimensof C omeyica appear quite close to C muricata The first threecomponents explain 5301 of variation (axis 1 = 3028 axis2 = 1601 and axis 3 = 672) PCoA shows distinct cluster-ing of the species but there is a considerable overlap betweensubspecies and no clear separation

The hierarchical clustering (Fig 2) shows that Carex otru-bae C foliosa and C polyphylla are separated from the aggre-gates and the four species of section Phaesthoglochin appeargrouped in a different cluster with two parts The first is theCarex spicata aggregate with two subunits representing Cspicata subsp andresii and C spicata subsp spicata and the

TABLE 1 Morphological characters and character states studied L =length W = width + = yes - = no r 075

(1) Awn glume 0 - 1 + (2) Auricle leaf blade 0 - 1 + (3)Lower spikes branched 0 - 1 + (4) Spikes contiguous 0 + 1 -(5) Basal sheaths purplish tinged 0 - 1 + (6) Perigynium beakcolor 0 green 1 brown (7) Lowest spike branched 0 - 1 + (8)Peduncle lowest spike 0 - 1 + (9) Margins of perigynium beak 0smooth 1 scabrid (10) Ripe achene brown 0 - 1 + (11) Scariousmargin of pistillate scale 0 - 1 + laterally 2 complete (12) Shapespike 0 globular 1 globular-elliptic 2 elliptic 3 oblong (13) Apexof pistillate scale 0 rounded-obtuse 1 acute 2 apiculate 3acuminate (14) Color glume 0 hyaline 1 gold-beige 2 lightbrown 3 dark brown 4 purplish brown (15) Perigynium shape 0rounded 1 oval 2 ovate 3 trullate 4 oblong 5 obovate (16) Apexof beak 0 bidentate 1 bifid 2 variable (17) Beak margins 0smooth 1 serrulate 2 irregular serrulate 3 scarcely serrulate (18)Shape achene 0 rounded 1 oval 2 ovate 3 pentagonal 4 obovate(19) Culm outline 0 rounded 1 obtusely trigonous 2 sharplytrigonous 3 winged (20) Inflorescence shape 0 lobulate 1 oblong2 elliptic (21) Pistillate scale shape 0 oval 1 ovate 2 obovate(22) Perigynium base corky 0 - 1 little 2 + (23) Ligule apex 0rounded 1 subacute 2 acute (24) L leafsL culm 1 lesser 2similar 3 bigger (25) Veins perigynium body (abaxial) 0 - 1 upto middle 2 all body (26)Perigynia with veins (abaxial side) 0none 1 several 2 all perigynia (27) Strength veins (abaxial side) 0veinless 1 faint 2 well marked 3 strongly (28) Veins perigyniumbody (adaxial) 0 - 1 up to middle 2 all body (29) Perigynia withveins (adaxial side) 0 none 1 several 2 all perigynia (30) Strengthof veins (adaxial side) 0 veinless 1 faint 2 well marked 3strongly (31)Attenuation perigynium beak 0 gradually 1gradually to abrupt 2 abrupt (32) Distribution denticles beak0 - 1 only beak 2 up to summit 3 over 13 body 4 only summit(33) Position of perigynium 0 erect 1 erectndashspreading 2spreading 3 very spreading (34) Winged margin of perigyniumbody 0 - 1 from summit to 1frasl2 body 2 all over body 3 wingvariable (35) Orientation of spike 0 lt 40deg 1 40deg-50deg 2 gt 50deg (36)L inflorescence 0 up to 30 1 30ndash50 2 50ndash70 3 more than 70 (37)W of widest leaf (38) Diameter of culm (at the middle) (39) Llongest ligule (40) L lowest bract (41) L longestinflorescence (42) Distance between lowermost spikes (43)Maximum distance between 2sd 3th spikes (44) No branchesfirst order (45) Number of perigynia in I2 (46) No male flowersI2 (47) L perigynium (48) W perigynium (49) L perigyniumbeak (50) Maximum no abaxial veins (51) Minimum noabaxial veins (52) Maximum no adaxial veins (53) Minimumno adaxial veins (54) L apical tooth (abaxial) (55) L apicaltooth (adaxial) (56) No denticles beakmm (57) L pistillatescale (58) Distance from base to widest point of perigynium(59) Difference (W perigynium ndash W achene)2 (60) Lachene (61) W achene (62) L perigyniumL glume (63) LperigyniumW perigynium (64) L acheneW achene (65) LperigyniumL beak (66) L perigyniumL achene (67) L longestinflorescenceDistance between lowermost spikes (68) Distancebetween lowermost spikesMaximum distance between secondand third spikes (69) No perigyniaNo flowers I2 (70) LperigyniumDistance from base to widest point ofperigynium (71) W wingW achene





TA

BL

E2

Dat

am

atri

xus

edin

the

phyl

ogen

etic

anal

ysis

Pol

ymor

phic

char

acte

rsar

ein

dic

ated

byA

mis

sing

dat

aby

C

hara

cter

num

bers

refe

rto

Tab

le1

Ch

arac

ter

nu

mb

er1

23

45

67

910

11

12

13

14

15

16

18

19

20

21

22

23

25

28

32

33

34

35

39

41

43

44

46

47

48

49

56

57

58

60

62

63

65

66

70

Cf

olio

sa1

11

10

01

10

23

11

51

41

00

02

AA

30

10

03

21

20

00

20

20

01

01

0C

pol

yphy

lla0

01

00

11

1

22

32

2A

12

10

10

21

22

01

03

21

22

11

21

32

11

01

0C

otr

ubae

11

10

01

11

00

20

22

02

31

12

22

12

20

01

22

12

21

21

20

10

10

22

Cs

pica

tasu

bsp

spic

ata

00

00

11

01

11

13

42

A3

21

12

22

12

10

11

10

00

21

22

22

20

20

11

Cs

pica

tasu

bsp

andr

esii

00

00

11

00

11

13

42

02

22

11

22

00

10

11

00

00

21

20

22

20

10

10

Co

mey

ica

00

00

00

01

00

12

31

02

11

00

01

12

10

10

10

00

21

02

23

20

01

00

Cp

aira

e0

00

00

1A

10

A0

32

20

11

12

00

10

22

01

00

0A

A1

00

21

21

00

10

0C

mur

icat

asu

bsp

mur

icat

a0

00

00

10

10

00

23

00

01

10

00

11

33

22

00

00

A1

10

20

11

10

10

0C

mur

icat

asu

bsp

cesa

nens

is0

00

00

10

10

00

23

31

11

10

00

21

33

22

00

00

02

11

21

32

10

10

0C

mur

icat

asu

bsp

asho

kae

00

00

01

01

10

02

31

11

11

00

00

03

32

20

10

00

21

22

13

22

20

10









DISCUSSION

















C spicata

C pairei C omeyica

C muricata






























TAXONOMIC TREATMENT




































































































2 386 1966

























































LITERATURE CITED





























































TA

BL

E2

Dat

am

atri

xus

edin

the

phyl

ogen

etic

anal

ysis

Pol

ymor

phic

char

acte

rsar

ein

dic

ated

byA

mis

sing

dat

aby

C

hara

cter

num

bers

refe

rto

Tab

le1

Ch

arac

ter

nu

mb

er1

23

45

67

910

11

12

13

14

15

16

18

19

20

21

22

23

25

28

32

33

34

35

39

41

43

44

46

47

48

49

56

57

58

60

62

63

65

66

70

Cf

olio

sa1

11

10

01

10

23

11

51

41

00

02

AA

30

10

03

21

20

00

20

20

01

01

0C

pol

yphy

lla0

01

00

11

1

22

32

2A

12

10

10

21

22

01

03

21

22

11

21

32

11

01

0C

otr

ubae

11

10

01

11

00

20

22

02

31

12

22

12

20

01

22

12

21

21

20

10

10

22

Cs

pica

tasu

bsp

spic

ata

00

00

11

01

11

13

42

A3

21

12

22

12

10

11

10

00

21

22

22

20

20

11

Cs

pica

tasu

bsp

andr

esii

00

00

11

00

11

13

42

02

22

11

22

00

10

11

00

00

21

20

22

20

10

10

Co

mey

ica

00

00

00

01

00

12

31

02

11

00

01

12

10

10

10

00

21

02

23

20

01

00

Cp

aira

e0

00

00

1A

10

A0

32

20

11

12

00

10

22

01

00

0A

A1

00

21

21

00

10

0C

mur

icat

asu

bsp

mur

icat

a0

00

00

10

10

00

23

00

01

10

00

11

33

22

00

00

A1

10

20

11

10

10

0C

mur

icat

asu

bsp

cesa

nens

is0

00

00

10

10

00

23

31

11

10

00

21

33

22

00

00

02

11

21

32

10

10

0C

mur

icat

asu

bsp

asho

kae

00

00

01

01

10

02

31

11

11

00

00

03

32

20

10

00

21

22

13

22

20

10









DISCUSSION

















C spicata

C pairei C omeyica

C muricata






























TAXONOMIC TREATMENT




































































































2 386 1966

























































LITERATURE CITED
































































DISCUSSION

















C spicata

C pairei C omeyica

C muricata






























TAXONOMIC TREATMENT




































































































2 386 1966

























































LITERATURE CITED


































































C spicata

C pairei C omeyica

C muricata






























TAXONOMIC TREATMENT




































































































2 386 1966

























































LITERATURE CITED











































































TAXONOMIC TREATMENT




































































































2 386 1966

























































LITERATURE CITED

















































































































































2 386 1966

























































LITERATURE CITED































































































LITERATURE CITED


























































Taxonomy and New Taxa in Eurasian Carex (Section Phaestoglochin, Cyperaceae

Documents

Transcript of Taxonomy and New Taxa in Eurasian Carex (Section Phaestoglochin, Cyperaceae