Botanical Journal of the Linnean Society, 2008, 156, 385–409. With 10 figures
Taxonomy and new taxa of the Carex divulsa aggregate
in Eurasia (section Phaestoglochin, Cyperaceae)
ANA MOLINA*, CARMEN ACEDO and FÉLIX LLAMAS
Department of Biodiversity and Environment Management, University of León, E-24071, León, Spain
Received 6 March 2006; accepted for publication 27 September 2007
The Carex divulsa aggregate, belonging to the Carex muricata group in section Phaestoglochin (Cyperaceae), was
studied using 60 macro- and micromorphological characters over a wider geographical range than has been attempted
previously, and the status or identity of several taxa commonly associated with this aggregate was examined. The
results of numerical analysis support the recognition of six new species within the Carex divulsa aggregate, which
are also in accordance with their geographical circumscription. Carex cyprica is endemic to Cyprus, C. enokii is
from the Mediterranean Basin, C. magacis from the mountains of Spain and France, C. nordica from Northern
Europe, and C. egorovae and C. otomana from Eastern Europe and Central Asia. The morphological data were
coded and used in a phylogenetic analysis to discover the relationships between taxa. A key to all taxa belonging
to the aggregate is included. Diagnostic characters, illustrations, and detailed descriptions of all species are given.
Notes on the habitat, distribution, phenology, etymology, and conservation status of each taxon are also presented.
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409.
ADDITIONAL KEYWORDS: biogeography – Carex muricata group – habitat – morphological characters –
numerical taxonomy – phylogenetic analyses – sedges.
INTRODUCTION
Carex divulsa and related taxa were placed in Carex
section Phaestoglochin by Dumortier (1827), which,
later, was named section Muehlenbergiae by Tuckerman (1843). The section is represented in Eurasia by
the C. muricata group, a problematic and scarcely
researched group, which can be distinguished by its
simple inflorescence, rarely with one to two short
branches at the base, lowest bracts setaceous, and
androgynous sessile spikes with perigynia planoconvex in cross-section (Clapham, Tutin & Moore,
1987). As in similar sections (Starr & Ford, 2001), the
morphological characters that differentiate species
are mostly quantitative and continuous. In studying
the C. muricata group in Eurasia (Molina, Acedo &
Llamas, 2008a), we have found three main cores
corresponding to the C. divulsa, C. spicata, and
C. muricata aggregates, and also an isolated species,
*Corresponding author. E-mail: [email protected],
[email protected]
C. coriogyne Nelmes, without a clear position. The
C. divulsa aggregate can be separated from the others
by its longer inflorescence with lower spikes not overlapping and/or hyaline female glumes.
Carex divulsa was described by Stokes (1787) and
C. leersii by Schultz (1870). Chater (1980) admitted
only one species, with two subspecies: C. divulsa ssp.
divulsa and C. divulsa ssp. leersii (Kneuck.) W. Koch.
Nilsson (1985) accepted this taxonomic treatment, but
both authors indicated that it must be regarded as
provisional because the C. muricata group is in need of
thorough study. Egorova (1999) considered these taxa
at the species level. Further studies of C. muricata
group based on morphological characters (Stoeva &
Popova, 1997; Repka, 2003) resulted in different opinions related to the taxonomic level of these taxa. One
problem in most studies and taxonomic classifications
is the restricted area covered, because most investigations are of local floras. Therefore, to understand and
explain this conflicting group, we need to determine
the variation over the entire distribution area.
Carex divulsa is easy to recognize (David, 1976),
and can be distinguished without problems. However,
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
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A. MOLINA ET AL.
C. leersii has always been a problematic taxon,
having a perigynium similar in size to that of C. spicata Huds. (Schultz, 1871: 23) and an inflorescence
similar in length to that of C. divulsa (Nelmes, 1947:
101). The identification of this taxon using the available literature is difficult, because there are discrepancies between authors about the range of character
variation; it seems that each author is talking about
a different plant. Historically, there has been much
confusion about the nomenclature of this taxon. Later,
it was renamed C. leerseana by Rouy (1912) and
C. leersiana by Rauschert (1973). At other times, it
has been treated as a synonym of C. polyphylla Kar.
& Kir. (Nelmes, 1947; Karjagin, 1952; Kern & Reichgelt, 1954; Garcke, 1972; David & Chater, 1977; Pignatti, 1982; etc.), C. guestphalica (Boenn. ex Rchb.)
O. Lang (Loos, 1996; Aeschimann et al., 2004), or
C. chabertii F.W. Schultz (Loos, 1996; Lambinon,
2004). In addition to the name, the rank is a source
of disagreement amongst taxonomists, who have
considered it to be subordinated to Carex muricata
L. (Ascherson & Graebner, 1902; Marshall, 1907;
Guinochet, 1978; Stoeva & Popova, 1997), C. pairae F.
W. Schultz (Vollmann, 1914; Fournier, 1961; Nyárády,
1966; Schultze-Motel, 1968; Garcke, 1972; Soó, 1973),
or C. divulsa (Hylander, 1966; David & Chater, 1977;
Chater, 1980; Jermy, Chater & David, 1982; Hooper,
1985; Nilsson, 1985; Feinbrun-Dothan, 1986; Clapham
et al., 1987; Aeschimann & Burdet, 1994; Duhamel,
1994; Luceño, 1994; Sell & Murrell, 1996; Kukkonen,
1998, 2002; Lambinon, 2004).
The main objective of this study was to clarify the
taxonomic situation. Consequently, new taxa have
been revealed with an interesting geographical distribution. As defined here, the C. divulsa aggregate
includes eight taxa – C. divulsa, C. leersii, and
six new species: C. egorovae, C. enokii, C. magacis,
C. cyprica, C. nordica, and C. otomana.
MATERIAL AND METHODS
TAXA
INCLUDED WITHIN THE SCOPE OF THIS STUDY
All taxa related to C. divulsa and C. leersii mentioned
by Vollmann (1903), Kükenthal (1909), Nelmes (1939,
1947), De Langhe (1944), Maire (1957), Hadac (1961),
Schultze-Motel (1968), David (1976), David & Chater
(1977), Chater (1980), David & Kelcey (1985), Hooper
(1985), Nilsson (1985), Malyschev & Peschkova
(1990), Strid & Tan (1991), Loos (1996), Kukkonen
(1998), Egorova (1999, 2000), and Repka (2003) were
studied here.
TAXON
SAMPLING
Some 1000 specimens from B, CAG, FCO, FI, GDAGDAC, H, JACA, JBVN, K, LEB, LISI, LISU, MA, P,
PAL, RNG, SANT, SEV, TFC, W, and WU were examined. The herbaria abbreviations are those of
Holmgren, Holmgren & Barnett (1990). Fieldwork
was performed during 2001–05, mainly in the Alps,
and Cantabrian and Pyrenean Mountains, to complete the morphological information.
MORPHOLOGY
For the morphological study, 113 specimens representing the entire geographical distribution and morphological variation of the aggregate were selected for
the numerical and statistical analyses. The characters used to distinguish the species in other Carex
section Phaestoglochin studies (Webber & Ball, 1984;
Stoeva & Popova, 1997; Ball, 2002; Repka, 2003;
Molina, Acedo & Llamas, 2006) were checked in a
group of selected specimens in order to assess their
taxonomic value. Six additional characters (7, 8, 10,
26, 43, 48) were included according to our observations. Finally, 61 variables were selected from the
53 studied characters (Table 1). Characters 1 and 2,
which have no variation inside the C. divulsa aggregate, were only used in the phylogenetic analysis,
and variable 7 (colour of ripe achene), which was
not available in all the studied specimens, was not
used in stepwise discriminant analysis (SDA). The
explanations of some characters and the method of
measurement are given in Molina et al. (2006). Characters 15 and 23, related to beak margin, were surveyed with a Jeol 6100 scanning electron microscope.
NUMERICAL
AND STATISTICAL ANALYSES
Characters were tested for significance using a pairwise t-test (P < 0.01) between 29 specimens of C. divulsa and 22 specimens of C. leersii. In order to group
more than 60 specimens which were difficult to identify, hierarchical clustering with all 113 specimens
was carried out to obtain clusters using Gower’s coefficient and unweighted pair group method with arithmetic averaging (UPGMA) algorithms of SYN-TAX
2000 (Podani, 2001).
The data set of all specimens was subjected to a
stepwise discriminant analysis to identify the best
characters to separate the clusters. Finally, one-way
analysis of variance (ANOVA) and the post hoc Tukey
test were conducted on the quantitative characters to
assess the divergence between them, and to identify
the best diagnostic characters for the clusters. Because
Levene’s test indicated that several variables showed
heterogeneous variance, they were logarithmically
transformed for ANOVA. Qualitative characters were
studied by analogous Kruskal–Wallis analysis. These
statistical analyses were performed with SPSS 13.0.
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
THE CAREX DIVULSA AGGREGATE IN EURASIA
Table 1. Studied morphological characters and character states (L, length; W, width; +, yes; -, no)
(1)
(2)
(3)
(4)
(5)
(6)
(7)
(8)
(9)
(10)
(11)
(12)
(13)
(14)
(15)
(16)
(17)
(18)
(19)
(20)
(21)
(22)
(23)
(24)
(25)
(26)
(27)
(28)
(29)
(30)
(31)
(32)
(33)
(34)
(35)
(36)
(37)
(38)
(39)
(40)
(41)
(42)
(43)
(44)
(45)
(46)
(47)
(48)
(49)
(50)
(51)
(52)
(53)
(54)
(55)
(56)
Awn 씸 glume: 0 -, 1 +
Auricle leaf blade: 0 -, 1 +
Spikes contiguous: 0 +, 1 Perigynium beak colour: 0 green, 1 brown
Lowest spike branched: 0 -, 1+
Peduncle lowest spike: 0 -, 1 +
Ripe achene brown: 0 -, 1 +
Beginning of anthesis: 0 spring, 1 summer
Scarious margin of female glume: 0 -, 1 + laterally, 2 complete
Shape spike: 0 globular, 1 globular-elliptic, 2 elliptic, 3 oblong
Apex of female glume: 0 rounded-obtuse, 1 acute, 2 apiculate, 3 acuminate
Colour 씸 glume: 0 hyaline, 1 gold-beige, 2 light brown, 3 dark brown, 4 purplish brown
Shape perigynium: 0 rounded, 1 elliptic, 2 ovate, 3 trullate, 4 oblong, 5 obovate
Apex of beak: 0 bidentate, 1 bifid, 2 variable
Beak margins: 0 smooth, 1 serrulate, 2 irregular serrulate, 3 scarcely serrulate
Shape achene: 0 rounded, 1 elliptic, 2 ovate, 3 pentagonal, 4 obovate
Shape female glume: 0 elliptic, 1 ovate, 2 obovate
Perigynium base corky: 0 -, 1 little, 2 +.
L leafs/L culm: 1 smaller, 2 similar, 3 larger
Veins perigynium body (abaxial): 0 -, 1 up to middle, 2 up to the top
Veins perigynium body (adaxial): 0 -, 1 up to middle, 2 up to the top
Attenuation perigynium beak: 0 gradually, 1 gradually to abrupt, 2 abrupt
Distribution denticles beak: 0 -, 1 only beak, 2 up to summit, 3 over 1/3 body, 4 only summit
Position of perigynium: 0 erect, 1 erect-patent, 2 patent, 3 very patent
Winged margin of perigynium body: 0 -, 1 from summit to 1/2 body, 2 up to the top, 3 wing variable
Orientation of spike: 0 less than 40°, 1 40°-50°, 2 more that 50°
L inflorescence: 0 up to 30, 1 30–50, 2 50–70, 3 more than 70
W of widest leaf
Diameter of culm (at the middle)
L lowest bract
L longest inflorescence
Distance between lowermost spikes
Maximum distance between second and third spikes
No. branches first order
No. perigynia in I2
No. male flowers I2
L perigynium
W perigynium
L perigynium beak
Maximum no. abaxial veins
Maximum no. adaxial veins
L apical tooth (abaxial)
No. denticles beak/mm
L female glume
W female glume
L male glume
Distance from base to widest point of perigynium
Difference (W perigynium - W achene)/2
L achene
W achene
L longest leaf
L apex of female glume
W inflorescence
L perigynium/L 씸 glume
L perigynium/W perigynium
L achene/W achene
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
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A. MOLINA ET AL.
Table 1. Continued
(57)
(58)
(59)
(60)
(61)
L perigynium/L beak
L perigynium/L achene
L longest inflorescence/distance between lowermost spikes
No. perigynia/no. 씹 flowers I2
L perigynium/distance from base to widest point of perigynium
PHYLOGENETIC
ANALYSIS
Forty statistically significant variables amongst the
taxa were chosen to conduct the phylogenetic analysis
(Table 2). As few qualitative characters were available
for this study, quantitative data were also used,
because they have given good results in other taxonomic studies (Van de Wouw, Maxted & Ford-Lloyd,
2003). Qualitative characters were coded as in the
numerical analysis. Values of the quantitative characters were transformed according to the groups
resulting from the post hoc Tukey test (ANOVA), and
coded in the range of 0–4. Both types of character
were treated as unordered multistate, but the four
ratios were coded in the range of 0–9 and treated as
ordered multistate. Carex coriogyne was included
because it is closely related to the C. divulsa
aggregate, and C. otrubae Podp. [section Vulpinae
(J. Carey) Christ.] was used as outgroup because it is
closely related to section Phaestoglochin. All variables
were equally weighted. An heuristic search was
carried out using stepwise addition, and the robustness of the clades was assessed by means of bootstrap
analysis. Maximum parsimony analysis was performed for the data set with PAUP* 4.0b10 (Swofford,
2002). The data have been deposited in TreeBASE.
DISTRIBUTION
Specimen label data were used to estimate the geographical distribution of each taxon. Information from
Kreczetovicz (1935), Karjagin (1952), Maire (1957),
Hylander (1966), Mouterde (1966), Nyárády (1966),
Schultze-Motel (1968), Soó (1973) Chater (1980),
Pignatti (1982), David & Kelcey (1985), Greuter,
Matthäs & Risse (1985), Hooper (1985), Meikle
(1985), Nilsson (1985), Feinbrun-Dothan (1986),
Hultén & Fries (1986), Strid & Tan (1991), Luceño
(1994), Viney (1996), Kukkonen (1998, 2002), Egorova
(1999), Aeschimann et al. (2004), and Lambinon
(2004) were crossed with our own data to infer the
real distribution of the taxa. The distribution maps
were plotted using ArcGis 9.
RESULTS AND DISCUSSION
Preliminary revision of the herbarium material
showed the following: (1) often, herbarium specimens
(putatively employed in some previous studies) are
misidentified; (2) mistakes are frequent in every
taxon belonging to the C. muricata group; (3) sheets
showing sterile and depauperate flowers are difficult
to assign to any taxon; and (4) the taxa are morphologically similar, and seem to form a coherent group.
Keeping all this in mind, the main taxonomic characters for the C. divulsa aggregate are as follows:
narrow leaves 2.5–4.5 mm without auricles, culm
diameter 0.75–2.0 mm, inflorescences 50–155 mm
long, with 4–13 more or less separated sessile spikes,
except for the lowest which sometimes is pedunculate,
lowest bracts setaceous sometimes longer than their
spikes, acuminate hyaline to brown glumes without arista, perigynium plano-convex, 3.3–6.0 ¥ 1.8–
2.75 mm, sometimes a little spongy tissue at the
base, with or without a winged margin, and beak up
to 1.5(1.75) mm.
According to these data, C. canariensis Kük., historically linked with this aggregate, does not belong
to the C. divulsa aggregate (section Phaestoglochin).
Kükenthal (1900) described it from the Canary
Islands, and considered that it was related to C. divulsa. Both were included in Carex section Muehlenbergianae (Kükenthal, 1909) a synonym of section
Phaestoglochin. Carex canariensis has leaves more
than 4 mm wide, inflorescence long and paniculate,
usually much branched. Perigynia 2.7–3 ¥ 1–1.4 mm,
strongly veined and convex on both sides, long and
serrulate beak, female glumes reddish. Therefore, it
could be included in Carex section Heleoglochin
Dumort., although some specimens which have an
inflorescence with few branches may look like
C. divulsa.
In the literature, there is confusion between C. polyphylla Kar. & Kir. and C. leersii. Thus, Kükenthal
(1909: 156) included C. polyphylla Kar. & Kir. as a
possible synonym under C. contigua Hoppe. Nelmes
(1947), Karjagin (1952), and others have considered
C. polyphylla as a synonym of C. leersii. Moreover,
Kreczetovicz (1935) mentioned the ferruginose glumes
of C. polyphylla and its perigynium ovate with long
beak, and quoted C. leersii as a synonym; however,
according to the different characters listed in the
protologue by Schultz (1871: 25), it is another plant.
Hooper (1985) treated C. leersii and C. polyphylla
as two different species. David & Kelcey (1985)
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
1
2
1
6
3
3
3
4
4
9
9
5
7
5
4
5
4
3
1
0
2
4
5
4
7
7
6
6
6
7
1
4
3
3
5
6
4
8
7
6
1
0
1
1
0
1
2
2
2
3
1
2
3
1
2
2
1
3
3
4
3
1
3
0
2
2
2
3
1
1
3
0
2
0
2
1
4
1
1
4
4
1
4
0
2
2
2
3
1
0
3
0
3
0
1
1
2
3
1
2
2
1
3
0
1
1
3
2
1
3
3
1
3
0
1
2
1
3
1
3
2
0
1
0
2
2
2
2
1
1
0
1
1
0
2
1
2
3
3
2
0
1
1
0
2
2
2
3
3
3
4
1
4
0
2
2
4
3
1
1
3
0
3
0
1
2
2
4
2
3
0
1
2
3
1
1
1
2
1
3
0
3
3
1
2
2
2
1
1
1
0
3
3
1
3
2
2
1
1
0
1
4
4
1
1
2
2
2
2
3
0
3
3
1
1
1
2
2
1
1
1
0
A
0
2
0
2
0
0
0
3
2
1
1
3
A
3
2
1
2
2
A
2
1
1
0
1
0
1
0
0
0
0
0
0
A
0
1
1
2
1
0
1
0
0
A
0
1
1
1
1
3
3
1
1
1
1
1
3
2
1
0
0
1
1
A
1
1
1
A
4
1
0
1
1
1
3
2
2
2
0
0
0
0
2
1
1
2
0
3
1
3
3
2
3
3
2
3
3
2
0
1
1
2
0
1
0
1
1
2
0
0
0
0
1
1
2
2
0
A
0
A
0
0
1
1
1
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
1
0
0
1
0
1
1
0
0
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
C. coriogyne
C. divulsa
C. enokii
C. cyprica
C. nordica
C. leersii
C. magacis
C. otomana
C. egorovae
C. otrubae
1 2 3 4 7 8 9 10 11 12 13 14 15 17 18 22 24 25 27 30 32 33 34 37 38 39 42 43 44 45 46 47 48 49 52 53 55 56 57 61
Character number
Table 2. Data matrix used in the phylogenetic analysis. Polymorphic characters are indicated by A. Character numbers are the same as in Table 1
THE CAREX DIVULSA AGGREGATE IN EURASIA
389
recognized that plants of C. polyphylla, grown from
seed supplied by the Botanic Gardens in Tashkent
(Uzbekistan), showed different features from plants of
C. leersii. Kükhonen (1998: 184) indicated that materials from Khorassan (Iran) had morphological features different from the variation range of European
C. leersii, but was reluctant to separate a different
taxon because of the small number of specimens
available. Studying chromosome numbers, Toderash
(1979) reported 2n = 48 for C. polyphylla, and other
authors have counted 2n = 58 (Luceño, 1994; Stoeva
& Popova, 1997) or 2n = 57–59 (Hartvig, 1986) in
C. leersii. Thus, there are sufficient data to differentiate between C. leersii and C. polyphylla. Furthermore, we have already concluded (Molina et al.,
2008a) that C. polyphylla does not belong to the
C. muricata group s.l.
A different case is represented by C. coriogyne
Nelmes, which Nilsson (1985) combined as a subspecies under C. divulsa, emphasizing its inflorescence
3–18 cm long. After a meticulous examination of the
type specimen conserved at Kew, and other available
specimens from Turkey, we can conclude that the type
is the only known specimen of C. coriogyne, because
the remainder were misidentified. Therefore, C. coriogyne does not belong to the C. divulsa aggregate
because of its small inflorescence, shorter than
2.5 cm. It cannot be included in any aggregate of the
C. muricata group, and is an isolated taxon.
Nelmes (1939) described C. persica Nelmes from
Guilan (Iran), a taxon closely related to C. foliosa D.
Don. The only specimen found according to the diagnosis was the typus conserved at Kew. After a careful
study, we verified that C. persica is the same taxon as
C. divulsa Stokes, agreeing with Nilsson (1985) and
Kükkonen (1998).
Schultz (1871) described C. chabertii F. W. Schultz.
The original material supporting its description was
collected by Chabert in the Drôme and belonged to
the Schultz Normale Herbarium, now dispersed in
several herbaria. From a study of this material, no
difference from C. leersii could be found. We also
identified specimens belonging to C. spicata Huds.
mixed with this material.
All material collected by Porta in the Alps (Val de
Vestino), labelled as C. chabertii, also belongs to
C. leersii; moreover, after several collections, we did
not find any different specimens from the Pyrenean
range, as indicated by Schultz (1871) and Kükenthal
(1909).
After Schultz (1871), Repka (2003: 245) was the
only author who considered that C. chabertii was a
different taxon, although he was not sure of its
adequate status. In our numerical analysis, specimens labelled C. chabertii were in the same cluster as
C. leersii. Our results are in agreement with those
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
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A. MOLINA ET AL.
of Loos (1996), who considered C. chabertii as a
synonym of C. leersii. The problem is that the name
C. chabertii is the legitimate name for C. leersii F. W.
Schultz, but that it is invalid because Willdenow
(1787) used it previously for the species currently
named C. echinata Murray. Taking into account that
C. leersii is the name in use, we are preparing a
proposal for its conservation (Molina et al., 2008b).
The taxonomic rank of C. divulsa and C. leersii has
been discussed several times (David, 1976; Chater,
1980; Nilsson, 1985; Stoeva & Popova, 1997; Molina
et al., 2006). However, certain differences between
the two taxa, such as perigynium colour (Chater,
1980; Jermy et al., 1982; Clapham et al., 1987; Sell &
Murell, 1996), leaf width (De Langhe, 1944; Nelmes,
1947; Chater, 1980; Clapham et al., 1987; Sell &
Murrell, 1996), ligule (De Langhe, 1944; Nelmes,
1947; Chater, 1980; David & Kelcey, 1985; Clapham
et al., 1987; Duhamel, 1994; Sell & Murell, 1996),
position of the perigynium (De Langhe, 1944; Chater,
1980; Jermy et al., 1982; David & Kelcey, 1985; Sell
& Murell, 1996; Lambinon, 2004), and leaf sheath
(David & Kelcey, 1985), have been reported, but we
did not find them to be statistically significant.
However, the results of a pairwise t-test indicated
that C. divulsa must be considered as separate from
C. leersii at the species level. Eight quantitative characters and four ratios showed statistically significant
differences between C. divulsa and C. leersii, and four
qualitative variables were different. The main differences are that C. divulsa has a smaller perigynium
(3.25–4 mm) than C. leersii (4.5–5.5 mm), and a
longer (5–13 cm) and more slender (6–9 mm) inflorescence than C. leersii (3.5–7 cm ¥ 8–10 mm). The
achene size is 2.0–2.5 mm in C. divulsa and 2.2–
3.0 mm in C. leersii. Qualitative characters, such as
scarcely serrulate beak margins (Fig. 1), quoted
already by Nelmes (1947), are unique to C. divulsa.
Usually, C. divulsa has the lowest spike pedunculate,
and this feature is not present in C. leersii, which
rarely has a shorter peduncle. Studying the morphological characters, Stoeva & Popova (1997) and
Molina et al. (2006) found that C. leersii is the closest
taxon to C. pairae. Hendrichs et al. (2004), in a phylogenetic analysis, found that C. leersii was closer to
C. muricata than to C. divulsa. We consider that
C. divulsa and C. leersii are very similar taxa, but
there are sufficient characters to separate them as
species, although some intermediate specimens may
be found.
Hierarchical clustering (Fig. 2) shows three main
units separated by the inflorescence length and distance between spikes. The clusters can be differentiated by the perigynium size and shape, and glume
colour. The first two units join taxa with separate
spikes. The specimens grouped first belong to C. di-
Figure 1. Beak of perigynium: A, Carex divulsa (LEB
32466); B, C. leersii (LEB 82622).
vulsa and C. enokii with longer inflorescences. The
second group includes four species, C. magacis, C. nordica, C. leersii, and C. otomana, all with shorter inflorescences. Finally, the third group includes two taxa
with contiguous spikes: C. cyprica and C. egorovae.
The value of dissimilarity separating species is 0.27.
Carex magacis appears to be the closest taxon to
C. leersii. We can separate the two species by differences in the perigynium position and shape, but, as
they have a lower dissimilarity value, we prefer to keep
all specimens of C. divulsa in the same rank.
A scatter plot of the scores in the discriminant
analysis (Fig. 3) shows eight taxa within the C. divulsa
aggregate. The first three components explained 75.4%
of the variation, and were correctly identified in 100%
of cases. The variable with the highest loading on the
first component was the presence of contiguous spikes,
which separated C. egorovae and C. cyprica from all
the remaining taxa. Variables with the highest loading
on the second component were scarious margins of the
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
THE CAREX DIVULSA AGGREGATE IN EURASIA
391
Figure 2. Hierarchical cluster from unweighted pair group method with arithmetic averaging (UPGMA) using Gower’s
coefficient of 113 specimens of the Carex divulsa aggregate and 59 variables.
Figure 3. Scatter plot of scores on axes 1 and 2 of discriminant function analysis of 113 specimens of the Carex divulsa
aggregate.
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
392
A. MOLINA ET AL.
female glume, shape of the perigynium, and width of
the inflorescence, which separated C. magacis and
C. otomana. Carex divulsa, C. enokii, C. nordica, and
C. leersii were separated by the third component of
length and width of the perigynium. Other important
characters by load were perigynium beak and female
glume colour in the fourth component, and margin
winged in the perigynium body and length of the
inflorescence in the fifth component.
Obviously, there is no variable that is statistically
significant for every taxon simultaneously in ANOVA,
but there is one – maximum number of adaxial veins
– which is not significant. The width of the culm,
length of the longest inflorescence, number of male
flowers, length and width of the perigynium, and
length of the achene were the best variables, because
they clearly separated two or more groups and
allowed the identification of a single taxon or a pair
of taxa. Qualitative variables, such as contiguous
spikes, perigynium beak colour, beginning of anthesis,
scarious margins and shape of the female glume,
perigynium base corky, position of the perigynium,
and winged margin of the perigynium, clearly separate taxa without overlap. Each of the taxa separated
in the hierarchical cluster and SDA, and identified by
ANOVA and the Kruskall–Wallis test, can be related
to a set of character states (Table 3). The best diagnostic characters for the differentiation of species are
the perigynium (shape, size, and wing), beak perigynium (colour, length, and denticles), female glume
(colour, shape, and size), and spikes (shape, position
in relation to inflorescence axis, and distances). Most
are also diagnostic in other aggregates of the
C. muricata group (Molina et al., 2008a).
Although hierarchical clustering, SDA, and ANOVA
support the recognition of six new species in the
aggregate, the limits of C. leersii do not seem clear, and
several specimens identified under this name could
be hybrids with other taxa. Hybrids are common in
Carex (Cayoutte & Catling, 1992), and several authors
(Wallace, 1975; Jermy et al., 1982; O’Mahony, 1989;
Luceño, 1994; Stoeva & Popova, 1997) have documented them in section Phaestoglochin.
Carex cyprica is the most differentiated species in
this aggregate, because of its smaller perigynium
(3–3.5 mm), contiguous spikes, four lowest branches,
and large number of flowers per spike. Materials from
Cyprus included some specimens that had separate
spikes and sterile perigynia, but none was C. divulsa.
The main differences of C. enokii are the largest
erect-patent perigynium (4.5–5.5 mm), ovate to oblong,
sometimes slightly winged, apex tapering abruptly
into a beak, and beak bidentate with a scariose border.
Carex otomana and C. egorovae have the perigynium ovate, a little corky at the base, with a beak
longer than the remaining taxa within the aggregate.
Carex egorovae has contiguous spikes and the shortest inflorescence, and C. otomana has the perigynium
beak, female glume, and ripe achene brown and the
longest perigynium (4.75–6 mm) of the aggregate.
Carex magacis and C. nordica, the most closely
related taxa to C. leersii, can be more easily separated
by qualitative rather than quantitative characters
(Table 3). They are both unique in having globular
spikes tilted more than 50° from the main axis of
inflorescence, and a partial or total perigynium
winged body. The perigynium body is trullate in
C. magacis, and ovate to elliptic in C. nordica. Carex
magacis usually has a light perigynium beak, female
glume, and ripe achene, whereas in C. nordica they
are brown.
PHYLOGENETIC
ANALYSIS
The phylogenetic analysis resulted in two single
most-parsimonious trees, with a tree length of 179
(Fig. 4). Only 38 characters were informative, with a
consistency index (CI) of 0.65, excluding uninformative characters, and retention index of 0.51. Carex
coriogyne was included in the C. divulsa aggregate,
forming a well-supported clade with C. enokii (bootstrap support, 71%). Carex leersii and its closely
related taxa were also well supported (bootstrap
support, 86%). Carex egorovae and C. otomana do not
appear to be directly related to any other taxon. The
CI value was similar to that of another morphological
study in Carex (Naczi & Ford, 2001). Starr & Ford
(2001) obtained a better value including internal
transcribed spacer (ITS) sequences. The homoplasy
index was quite high (0.35). Other studies with
molecular data must be made to confirm the results.
HABITAT
AND DISTRIBUTION
Taking into account all the available data and ecological behaviour of the species belonging to the Eurasian Phaestoglochin section, it can be concluded that
they are frequent in hemerophilic habitats, such as
road sides, pastures, or waste areas (David & Kelcey,
1985; Aeschimann et al., 2004). Carex divulsa and
C. leersii were reported as pioneers by David &
Kelcey (1985), which is supported by our observations, with a generalist strategy and limited power of
persistence in dense vegetation.
The entire aggregate occurs in Eurasia, from the
Iberian Peninsula to Kazakhstan, and the morphological variation probably represents adaptation
to different climates. Carex divulsa and the closely
related taxa, C. cyprica and C. enokii (Fig. 5), are
adapted to warm climates. Carex divulsa lives mainly
in a Mediterranean climate and is widely distributed
from Madeira and the Canary Islands (30°W) to
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
No
White
Spring
Globular
Elliptic
Bidentate
Ovate
Gradual
Patent
No
50–150
79.8b ± 25.2
12–45
25.6d ± 8.2
3.5–4.25
3.8b ± 0.3
1.75–2.3
2.0a,b ± 0.2
0.5–1.0
0.80a,b ± 0.1
2–14
8.5a ± 3.8
3.0–4.0
3.4a,b ± 0.4
1.75–2.3
2.0b,c ± 0.1
1.25–1.75
1.5b ± 0.1
2.0–2.5
2.2b ± 0.2
Spikes contiguous
Beak colour
Beginning of anthesis
Shape spike
Shape perigynium
Apex beak
Shape female glume
Attenuation beak
Perigynium in ripe
Winged body
L inflorescence
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
L achene
Distance from base
W female glume
L female glume
No. denticles beak/mm
L beak
W perigynium
L perigynium
Distance lowermost spikes
C. divulsa
Character
Yes
White
Spring
Elliptic
Elliptic-ovate
Bifid
Elliptic
Gradual
Erect-patent
No
40–75
48.7a ± 13.8
6–12
8.9a ± 2.3
3.0–3.5
3.3a ± 0.1
1.6–2.0
1.9a ± 0.1
0.4–0.9
0.7a ± 0.1
8–14
10.8a,b ± 1.9
2.75–3.25
3.0a ± 0.2
1.2–2.0
1.7a ± 0.2
0.75–1.25
1.0a ± 0.2
1.8–2.2
2.0a ± 0.1
C. cyprica
No
White
Spring
Intermediate
Ovate-oblong
Bidentate
Ovate
Abrupt
Erect-patent
No
40–145
79.4b ± 30.2
12–55
24.1c,d ± 13.1
4.5–5.5
4.8d ± 0.3
2.5–2.75
2.6d ± 0.1
0.75–1.0
0.9a,b ± 0.1
8–18
11.5a–c ± 3.6
3.5–5
4.0c,d ± 0.5
2.0–2.5
2.2c ± 0.2
1.75–2.12
1.9e ± 0.1
2.5–3.0
2.8e ± 0.2
C. enokii
No
White to brown
Summer
Intermediate
Elliptic
Variable
Variable
Gradual
Erect-patent
No
35–60 (75)
54.7a ± 13.7
8–27
16.7a–c ± 4.8
4.5–5.5
4.7c,d ± 0.3
1.75–2.25
2.1b,c ± 0.2
0.67–1.25
1.0b ± 0.2
10–18
15.8c,d ± 2.1
3.0–4.5
3.6b,c ± 0.4
1.6–2.3
1.9b,c ± 0.1
1.5–1.87
1.65b–d ± 0.1
2.2–3.0
2.6d ± 0.2
C. leersii
No
Brown
Summer
Globular
Ovate
Bifid
Elliptic
Gradual
Patent
Yes
35–55 (75)
48.2a ± 10.4
13–30
19.3b–d ± 6.0
4.0–4.75
4.4b ± 0.2
1.75–2.5
2.2b,c ± 0.1
0.75–1.35
1.0b ± 0.2
12–20
15.1b–d ± 2.7
2.75–3.75
3.3a,b ± 0.3
1.5–2.2
1.8b ± 0.3
1.37–1.75
1.6b,c ± 0.1
2.0–2.75
2.5c,d ± 0.1
C. nordica
No
White
Summer
Globular
Trullate
Bifid
Elliptic
Gradual
Very patent
Yes
40–60
51.8a ± 6.4
14–21
17.2a–d ± 2.6
4.5–5.5
4.8c,d ± 0.1
2.25–2.75
2.5d ± 0.1
0.75–1.25
1.0b ± 0.1
14–18
17.0e ± 1.4
3.0–3.6
3.3a,b ± 0.2
2.0–2.5
2.2c ± 0.03
1.5–2.12
1.8c–e ± 0.2
2.5–3.0
2.7d,e ± 0.1
C. magacis
No
Brown
Spring
Intermediate
Ovate
Bifid
Ovate
Gradual
Erect-patent
No
40–65 (85)
59.2a,b ± 13.2
10–17
13.6a,b ± 2.2
5.0–6.0
5.5e ± 0.4
2.0–2.5
2.3c ± 0.2
1.0–1.6
1.3c ± 0.2
14–18
15.5c,d ± 1.2
3.25–5.25
4.2d ± 0.6
1.75–2.3
2.1c ± 0.3
1.5–2.25
1.8d,e ± 0.2
2.5–3.0
2.8e ± 0.2
C. otomana
Frequent
White
Spring
Intermediate
Ovate
Bifid
Ovate
Gradual to abrupt
Erect-patent
No
30–50 (68)
44.8a ± 9.2
6–13 (20)
12.7a,b ± 3.9
4.25–5.5
4.7c,d ± 0.4
1.80–2.5
2.0b,c ± 0.2
1.0–1.75
1.3c ± 0.2
4–16
12.2a–c ± 5.9
3.0–4.0
3.45a,b ± 0.3
1.6–2.25
1.8b ± 0.1
1.12–1.75
1.5b ± 0.1
2.0–2.7
2.3b,c ± 0.2
C. egorovae
Table 3. Quantitative and qualitative distinctive characters for the Carex divulsa aggregate. Ranges and means ± standard deviation for quantitative characters
in millimetres. Within a row, means with different superscripts differ significantly (ANOVA, P ⱕ 0.05) (L, length; W, width)
THE CAREX DIVULSA AGGREGATE IN EURASIA
393
394
A. MOLINA ET AL.
Figure 4. Phylogram based on 40 characters of the relationship between taxa of the Carex divulsa aggregate using an
heuristic search with C. otrubae as outgroup. Bootstrap support is indicated on the branches. The tree is 179 steps long
with a consistency index of 0.65 and a retention index of 0.51, excluding uninformative characters.
Figure 5. Geographical distribution of Carex divulsa ( ), C. cyprica (䊉), and C. enokii (夹).
south-east of the Caspian Sea (55°E), and from the
coastal strip of northern Africa (Algeria and Morocco,
30°N) to Northern Europe (45°-52°N). Carex enokii
inhabits areas throughout the southern and eastern
Mediterranean Basin, from Mauritania to Turkey.
Carex cyprica is endemic to Cyprus. Carex leersii and
its closely related taxa, C. magacis and C. nordica
(Fig. 6), are distributed in the Atlantic temperate
regions of Europe, but are not common in Southern
Europe. They are adapted to the cold conditions of
Northern and Middle Europe. Carex leersii has a
narrower distribution than C. divulsa, from northern
Spain to the south of the Scandinavian Peninsula and
to eastern Turkey. Carex nordica is limited to a strip
zone in Europe between 50° and 60°N from Great
Britain to Sweden, and scattered in Central Europe.
Carex magacis occurs in a small area in Pyrenean and
other Iberian Mountains (Spain, Andorra, and France).
Carex otomana and C. egorovae occur from Eastern
Europe to Middle Asia (Fig. 6). Carex otomana occurs
from the east of Bulgaria and Greece to Tyan Shan
(Kazakhstan), and C. egorovae in three disjointed
areas from Hungary to Iran.
ORIGIN
No data are available on the origin and evolution of
this aggregate. The eastern part of the Mediterranean
Basin (Greece, Turkey), where most taxa of the group
live, could be a hot spot of diversity (Médail & Quézel,
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
THE CAREX DIVULSA AGGREGATE IN EURASIA
395
Figure 6. Geographical distribution of Carex leersii ( ), C. magacis ( ), C. nordica (夹), C. otomana ( ), and
C. egorovae (䊉).
1997) and the speciation zone of the aggregate.
According to Nelmes (1947), C. leersii could be the
oldest taxon, from which C. muricata and C. divulsa
have diversified. In the phylogenetic tree, C. leersii is
also separated before C. divulsa and its relatives.
Carex cyprica was putatively segregated from C. divulsa. A reduction in the internode length and an
increase in the number of flowers per spike might
have led to the evolution of C. cyprica. Its isolated
distribution has avoided introgressions with C. divulsa. According to the phylogenetic analysis, C. coriogyne is the closest species to C. enokii, and they both
have similar perigynia; however, their relation to
C. divulsa is not clear. Carex nordica and C. magacis
could be of hybrid origin between C. leersii and other
taxa of the C. muricata group.
In conclusion, if Carex originated in the early Tertiary and the subgenus Vignea evolved in cooler climates, probably in the mid-Tertiary (Ball, 1990), it is
possible that the C. muricata group may have shown
rapid diversification, and may have adapted to different climatic conditions, helped by its opportunist
strategy. The recognition of six new species in the
C. divulsa aggregate is the result of an in-depth study
of the entire C. muricata group in a broad geographical context. Other studies may discover more diversity, mainly in the eastern distribution area because
of the lack of sampling. Hybrids and morphological
intermediates often occur, and isolated reproduction
is not effective, but our results show that there
are several species supported by morphological
characters.
SECTION PHAESTOGLOCHIN DUMORT. FL. BELG. 146.
1827. (LECTOTYPE DESIGNATED BY JONES (1994):
CAREX MURICATA L.)
1. Carex divulsa Stokes in With., Bot. Arr. Brit. Pl.
ed. 2, 2: 1035. 1787. (Fig. 7A–F)
= Carex canescens sensu Huds., Fl. Angl.: 405. 1778,
non L. 1753.
= Carex virens Lam., Encycl., 3: 384. 1792.
= Carex divulsa Gooden., Trans. Linn. Soc. 2: 160.
1794.
= Carex guestphalica (Boenn. ex Rchb.) O. Lang, Flora
26: 147. 1843. Neotype (designated by Loos): Feddes
Repert. 107(1–2): 74. 1996 (MSTR!).
= Carex persica Nelmes, Bull. Misc. Inform. Kew
1939: 185. Holotype (designated by Nelmes): Bull.
Misc. Inform. Kew 1939: 185. 1939. (K!).
Lectotype (designated here): [icon] ‘Carex nemorosa,
fibrosa radice, caule exquisite triangulari, spica longa
divulsa seu interrupta, capitulis solitariis praeterquam ultimo’ in Micheli, Nov Pl. Gen.: t. 33, f. 10.
1729. – Epitype (designated here): ‘Gram. Cyp. spicatum minus spicâ longâ/divulsâ seu interrupta Ray
1279. 2 Petiver’ (No. 17274, FI-M!).
Description: Caespitose perennial tufted plant with or
without short rhizomes c. 6 mm in diameter. CULMS
15–70 cm ¥ 0.8–1.25 mm at mid-height, obtusely trigonous, slightly scabrid above; sheaths pale to dark
brown. LIGULE 1.25–4.0 mm long, wider than long,
usually rounded at apex. LEAF blades about three-
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
396
A. MOLINA ET AL.
TAXONOMIC TREATMENT
KEY
TO THE
EURASIAN
TAXA OF THE
CAREX
DIVULSA AGGREGATE
1. Inflorescence with spikes contiguous, only the lowest separated; glumes pale...............................................2
1. Inflorescence with more spikes separated...............................................................................................3
2. Inflorescence with three to four basal spikes compound, perigynia 3–3.5 mm long, short beak (0.7–1 mm), and apical
teeth 0.2–0.5 mm long....................................................................................................2. Carex cyprica
2. Inflorescence with zero to two basal spikes compound, perigynia more than 4 mm long, long beak (1–1.75 mm), and
apical teeth 0.6–1.0 mm long ......................................................................................... 8. Carex egorovae
3. Perigynium completely or partially winged ............................................................................................ 4
3. Perigynium not winged.......................................................................................................................5
4. Perigynium elliptic, 4–4.75 mm long, female glume light to dark brown ................................. 5. Carex nordica
4. Perigynium trullate, 4.25–5.5 mm long, female glume hyaline to light brown ......................... 6. Carex magacis
5. Perigynium 3.5–4.25 mm long ......................................................................................... 1. Carex divulsa
5. Perigynium 4.25–6 mm long ................................................................................................................ 6
6. Perigynia 4.75–6 mm long, slightly corky at base, and brown glumes ................................... 7. Carex otomana
6. Perigynia 4.5–5.5 mm long, not corky at base, and pale glumes.................................................................7
7. Perigynium ovate to oblong, abruptly contracted in a beak.....................................................3. Carex enokii
7. Perigynium elliptic to ovate, gradually tapering in a beak ..................................................... 4. Carex leersii
Figure 7. Carex divulsa (A, from GDA 9116; B–F, from GDA 38629): A, inflorescence; B, male glume with anther; C,
female glume; D, perigynium, abaxial side; E, achene; F, perigynium with abaxial side removed to show achene. Carex
cyprica (from holotypus MA 495407): G, inflorescence; H, male glume with anther; I, female glume; J, perigynium,
abaxial side; K, achene; L, perigynium with abaxial side removed to show achene. All figures drawn by X. Magaz. Scale
bar, 1 mm.
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
THE CAREX DIVULSA AGGREGATE IN EURASIA
quarters or as long as culms, dark green to grey green,
widest leaf blades 2–3 mm, plicate to flat. INFLORESCENCE oblong, 50–135(150) ¥ 6–9 mm, with 7–12
globular sessile and unbranched spikes, except the
lowest that is usually branched and pedunculate.
Lowest internode 12–45 mm long, the second one
6–28 mm. BRACTS glume-like, the proximal setaceous,
shorter than inflorescence, 6–50(-63) mm long.
FEMALE GLUME hyaline to golden-yellow, with a midrib
green, 3.0–4.0 ¥ 1.75–2.30 mm, ovate, apex acuminate
0.15–0.3 mm long. STAMINATE GLUME 2.75–3.5 ¥ 1.4–
2.0 mm, similar to the female. ANTHERS 3, 1–2.5 mm
long. PERIGYNIUM patent to erect-patent, greenish,
body elliptic to rounded, nerveless or faintly veined,
3.5–4.0(4.25) ¥ 1.75–2.30 mm, acute at base, gradually
tapered into a beak. BEAK 0.5–1.0 mm long, white
distally, with scarcely serrulate margins, apex bidentate, apical teeth 0.3–0.5 mm. ACHENE elliptic to
rounded, shining brown when ripe, 2.0–2.5 ¥ 1.5–
2.25 mm. STIGMA 2, 2.0–3.0 mm long.
Phenology: Flowering begins in April, fruiting from
May to September, wind pollinated. All of our
data indicate that C. divulsa flowers 1 month before
C. leersii, disagreeing with Jermy et al. (1982) and
David & Kelcey (1985), who reported it 1 month later.
Habitat: Carex divulsa seems to need some moisture
in the soil. It grows in shady habitats, such as forest
and disturbed places (Guinochet, 1978; Chater, 1980;
Clapham et al., 1987), but also in sunny open habitats (David & Kelcey, 1985). In the latter case, specimens are harder. Carex divulsa occurs at low
altitudes (0–700 m), although in the warmest, southern part of the distribution area it can be found at
1000–1200 m. Greuter et al. (1985) quoted 1800 m.
According to David & Kelcey (1985) and our own
observations, it tolerates a wide range of soil acidity
(pH 5.3–7.4).
Distribution: Carex divulsa is the most common
species of the group, widely distributed (Fig. 5) from
Madeira to Iran (55°E). It is frequently found in
Mediterranean areas in Spain, France, Italy, Greece,
Turkey and the islands (Balearic, Corsica, Sicily, and
Crete), northern Africa (from Morocco to Algeria), and
in the Canary and Madeira Islands. It is scarce in
temperate zones, and very scarce towards the northeast of Europe, for example it is rare in Belgium
(Lambinon, 2004). In Russia, it appears only in the
south-west and Caucasus region, and it extends from
Iran to south-east of the Caspian Sea. No specimens
have been found in Iraq.
397
Observations: Carex divulsa presents some variation
throughout its distribution area. In Europe, the perigynium beak is short, bidentate, and very scarcely
serrulate. It is longer towards the east, almost
bifid and more regularly serrulate. Inflorescences are
usually long, but near the Mediterranean coast are
sometimes shorter (even less than 4 cm), probably
caused by introgression with C. pairae. Specimens
with a rounded perigynium body are sometimes found
in Southern Europe. Intermediates between C. divulsa and C. leersii have been named C. divulsa var.
polycarpa Vollmann (Lambinon, 2004). Carex divulsa
is also sympatric with C. magacis, C. egorovae, and
C. enokii. In mixed populations, it maintains its distinctiveness, with the perigynia of C. divulsa being
slightly larger or having a longer beak. It is allopatric
with C. cyprica, C. leersii, and C. otomana. According
to Nelmes (1947), when this species is located on
calcareous soils, the perigynia are slightly larger. We
studied the variation of the perigynium length in
calcareous soils and found no clear relationship:
longer and shorter perigynia were found on both
calcareous and noncalcareous soils.
Representative specimens examined: FRANCE. Corsica,
Evisa, kastanienhain oberhalb des Ortes, 900 m,
23.vii.1956, W. Rössler (MA 274203). Haute-Corse,
Ghisonaccia,
bord
N-198,
bosquet
rudéral
d’Eucalyptus, 1.vi.1997, J. Lambinon (MA 595765).
Haute Garonne, Rieux Volvestre camping, sombra
sotobosque de Quercus pubescens, 600 m, 14.vii.2002,
A. Molina (LEB 78511). CROATIA. Dalmacia, Lapad
peninsula, edge of path round end of Peninsula,
19.iv.1935, A. K. Jackson & W. B. Turrill 132 (K).
GREECE. Crete, Dist. Chania, in saxosis schist,
humidis inter Skines & Nea Rumata, 300 m, 29.v.1942,
K. H. Rechinger fill. 13384 (K). IRAN. 5′S of Chalus,
Caspian rain forest, 500 ft, 27.vi.1962, P. Furse 2888
(K). E. Mazandaran, Centre, 5 km W of Tangegol,
c. 1 km NE Cheshme-e-, NE Cheshme-e-AdamChaqran (spring), in Crataegus forest (Golestan
National Park), 980 m, 37°24′N, 55°54′E, 3.viii.1994,
H. Akhani 9799 (H 1695179). Gongân, Jangal-eGolestân, Tang-e-Râh, 580 m, 17.v.1976, Termeh &
Matin 34797 (H 1585245). North Persia, ad radices m.
Elbrus, pr. pagum Darake, umbrosis. 25.iv.1843, C.G.
T. Kotschy 76 (P 00281847). IRELAND. Limerick, Sroolane near Foynes, hedgebank, 22.vi.1952, R. B. Drummond 736 (K). ITALY. Toscana, Firenze, waste ground,
24.v.1964, R. K. Brummitt 4554 (K). MOROCCO. pr. El
Araix, el Guedira dicto, in locis humidis, 20.v.1930,
Font Quer (GDA-GDAC 02646). Vertiente mediterranea del Gurugu, sobre la segunda caseta, 1.vi.1915,
A. Caballero (MA 017149). Zaian, maison forestière
d’Assoual, ravine humide au pied de la maison,
26.iv.1952, Ch. Sauvage (SANT 48676). NETHER-
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A. MOLINA ET AL.
LANDS. Limburgh, Plasmolen at Mook, roadside,
1.v.1926, Herb. Kern & Reichgelt (J. K & B. R) (K).
ROMANIA. Dolj, Oltenia, rivuli Jiu, pagum Bratovoesti,
silvis humidis Q. robori–Carpinatum, 65 m, 6.vi.1971,
D. & Mariana Cirtu (MA 236897). SPAIN. Albacete,
Villapaladios, 14.vi.1993, G. Aragón et al. (LEB
52179). Almería, Sierra de Gador, Fondón, praderas,
herbazales, acequias y arroyos, ambiente muy
húmedo, 900 m, 14.vi.1992, A. Hervás (GDA 38269).
Barcelona, Massif du Tibidabo à S. Genis, 30.v.1912,
F. Senen (MA 016815). Cáceres, El Portanchito,
29SQD27, 7.v.1983, M. Ladero & M. Santos (LEB
24812). Granada, Orgiva, salida del pueblo, borde de
acequia, 8.iv.1980, M. Ladero & J. Molero (GDA 9116).
León, Lago Sumido, borde lago, 700 m, 29TPH826037,
1.v.2002, A. Molina (LEB 79002). Pombriego,
29TPG89, 7.v.1985, I. Jiménez (LEB 32466). Mallorca,
Pont d’Inca, Torrent, herbazal ruderalizado, 6.v.1987,
L. Villar & C. Chocarr (JACA 028187). TURKEY. Zonguldak, Kel tepe above Yenice, Abies–Fagus forest,
1200 m, 20.viii.1962, Davis, Coode & Yaltarik 37793
(K). UNITED KINGDOM. 41 Glamorgan, between
Melling Griffith and Tongwylais, dry bank in canal,
22.vi.1949, E. Nelmes 960 (K).
2. Carex cyprica A. Mol., Acedo & Llamas,
sp. nov. (Fig. 7G–L)
Carice divulsa similes, sed ab ea differt inflorescentia
brevioribus, 4–5 cm longitudinis, spicae elipsoideus,
multiflorae; spicae basarum proximus, perigyniis
brevioribus, 3–3.25 mm longitudinis.
Holotype: Cyprus, Distr. Paphos, station forestière
Paphos, Ayia, milieu anthropisé humide, sous-bois de
Platane, 650 m, 23.iv.1991, Iter Mediterraneum IV
1284 (MA 495407!) [sub. Carex muricata L. ssp. lamprocarpa]. Isotypes: B!, FI!, JBVN!, PAL!, RNG?.
Description: Caespitose perennial tufted plant with
short rhizomes c. 4 mm in diameter. CULMS 28–
75 cm ¥ 1.0–1.5 mm at mid-height, obtusely trigonous, slightly scabrid above; sheaths pale to dark
brown. LIGULE 0.5–1.25 mm long, wider than long,
rounded at apex. LEAF blades pale green about
three-quarters to as long as culms, widest leaf blades
3–3.5 mm, plicate to flat. INFLORESCENCE oblong,
40–75 ¥ 8–9 mm, with 10–14 elliptic, sessile spikes,
the first four branched. Lowest internode 6–12 mm
long, the second one 6–8 mm, other spikes more
crowded. BRACTS glume-like, the proximal setaceous,
shorter than inflorescence, 6–18 mm long. FEMALE
GLUME hyaline to yellow-golden, with a midrib
green, 2.75–3.25 ¥ 1.2–2.0 mm, elliptic, apex apiculate 0.2 mm long. STAMINATE GLUME 2.5–3.0 ¥ 1.5–
2.0 mm, similar to the female. ANTHERS 3, 1.5–2 mm
long. PERIGYNIUM erect-patent to erect, pale green-
ish, body elliptic to ovate, nerveless to well-marked
veins, 3.0–3.5 ¥ 1.6–2.0 mm, rounded base, gradually
to abrupt tapered into a beak. BEAK 0.4–0.5(0.9) mm
long, white distally, with serrulate winged margins
reaching the summit, apex bifid, apical teeth
0.20–0.50 mm. ACHENE elliptic to ovate, shining,
light greenish to brown, 1.8–2.25 ¥ 1.5–1.75 mm.
STIGMATA 2, 1.5–2.5 mm long.
Etymology: The new species is named from Cyprus,
where this plant is endemic.
Phenology: Flowering begins in April, fruiting from
May to August, wind pollinated.
Habitat: Carex cyprica occurs in warm and humid
places, near brooks, springs, or streams between 200
and 1200 m, and is indifferent to soil composition.
Distribution: Endemic to Cyprus (Fig. 5).
Observations: Specimens from Cyprus identified as
C. divulsa by Meikle (1985) do not agree with the
diagnostic characters of C. divulsa. Meikle recognized
problems in the taxonomy of the complex, but stated,
‘The temptation to describe another closely allied
taxon is resisted because much remains to be discovered of the citology and crossing behaviour of this
group’.
Representative specimens examined: CYPRUS. Near
Inia, Akamas, quartzite, growing in the crevices,
8.iv.1957, L. F. H. Merton 3021 (K). Distr. Kyrenia,
Laphitos, loco umbroso juncto rivulum, 13.vi.1939, H.
Lindberg iter 1939 (K) (H 1302858) and (H 1302856).
Distr. Limassol, foot of Aphamia, on calcareous rock,
2900 ft, 20.v.1939, E. W. Kennedy 1739 (K). Between
Platres & Aphamis, by a spring in garigue, 946 m,
22.iv.1941, P. H. Davis 3192 (H 1205849). Distr.
Prodhromos, in nemorosis, 4500 ped, 1862, Ms. Merii
& T. Kotschy 855 (K). Ytronas river, by stream,
4000 ft, 3.v.1962, R. D. Meikle 2828 (K). Ayia valley,
roadside by a stream in the forest, 2000 ft, 28.ix.1936,
L. F. H. Merton 2831 (K). Mandria, on igneous rock,
3000 ft, 21.iv.1939, E. W. Kennedy 1438 (K).
3. Carex enokii A. Mol., Acedo & Llamas,
sp. nov. (Fig. 8A–F)
Carice divulsa similes, sed ab ea differt perigynia
maioribus est, 4.5–5 mm longitudinis, oblonguis
formam habens atque in rostrum abruptim contractas.
Holotype: Planta Lybicae ex Herbarium Universitatis
Florentinae n°1156 (sub. Carex divulsa Gooden.)
Libia Cirenaica. Cirene- U. Belgadir, 26.iv.1934, R.
Pampanini & R. Pichi-Sermolli 1156 (K!).
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THE CAREX DIVULSA AGGREGATE IN EURASIA
399
Figure 8. Carex enokii (from holotypus R. Pampanini & R. Pichi-Sermoli 1156, K): A, inflorescence; B, male glume with
anther; C, female glume; D, perigynium, abaxial side; E, achene; F, perigynium with abaxial side removed to show achene.
Carex leersii (G, from Hennipman et al. 1756, K; H–L, from LEB 82662): G, inflorescence; H, male glume with anther;
I, female glume; J, perigynium, abaxial side; K, achene; L, perigynium with abaxial side removed to show achene. All
figures drawn by X. Magaz. Scale bar, 1 mm.
Description: Caespitose perennial plant with short
rhizomes c. 3–4 mm in diameter. CULMS 25–
70 cm ¥ 1.0–2 mm at mid-height, obtusely trigonous,
slightly scabrid above; sheaths brown. LIGULE 1.0–
3.0 mm long, subacute to acute at apex. LEAF blades
grey green, about three-quarters as long as culms,
widest leaf blades 3–4 mm, plicate to flat. INFLORESCENCE oblong, 40–90(145) ¥ 10–12 mm, with 7–12
globular-elliptic spikes. Lowest spikes usually compound and shortly pedunculate, upper spikes single
at nodes, sessile, and contiguous. The lowest internode 12–55 mm long, the second one 7–17 mm. BRACTS
glume-like, the proximal setaceous, shorter than
the inflorescence, 6–55 mm long. FEMALE GLUMES
hyaline, with a midrib green, 3.5–5.0 ¥ 2.0–2.5 mm,
ovate, apex acuminate 0.3–0.5 mm long. STAMINATE
GLUMES 3.2–4.25 ¥ 1.75–2.5 mm, similar to the
female. ANTHERS 3, 1.75–3.0 mm long. PERIGYNIUM
erect-patent to patent, pale greenish, body ovate
to oblong, veinless or faintly veined, 4.5–5.5 ¥ 2.5–
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A. MOLINA ET AL.
2.75 mm, rounded base, sometimes slightly winged,
and abruptly tapered into a beak. BEAK 0.75–1.0 mm
long, white-scarious distally, with scarcely serrulate
margins reaching the summit, apex bidentate,
apical teeth 0.3–0.5 mm. ACHENE elliptic, shining,
light brown, 2.5–3.0 ¥ 2.0–2.5 mm. STIGMATA 2, 2.0–
3.0 mm long.
Phenology: Flowering begins in April, fruiting extends
from May to July, wind pollinated.
Habitat: There is little information about the habitat
of C. enokii. The plants inhabit waste lands and forest
borders, such as savin woods. They also grow in shady
and wet places by streams. The substrates mentioned
for C. enokii include marl and volcanic rocks, and so
it appears to need rich nutrient soils.
Distribution: Carex enokii is found (Fig. 5) throughout
the eastern and southern Mediterranean Basin from
Turkey, through Palestine and Libya, to Mauritania,
and also in Corsica, Sicily, and the Canary Islands.
Observations: Mouterde (1966) and Erteeb & Sherif
(1985) do not mention any Carex with these characteristics. Carex muricata ssp. divulsa var. divulsa f.
guestphalica sensu Maire (1957) could be a similar
plant. Feinbrun-Dothan (1986) included under C. divulsa ssp. leersii plants from Palestine that have
the same characters as C. enokii (perigynium size,
broadly ovate, abruptly tapering beak). Its scarce
presence in herbaria informs us about its rarity.
There are robust specimens with longer inflorescences
from Sardinia and Tenerife that may be hybrids with
C. divulsa.
Representative specimens examined: ITALY. Sardinia,
Rio Baccu Mannu, Domus de Maria (CA), 11.v.2004, R.
Angius, R. Mandis & M. Orrú (CAG). LIBYA. Cirenaica,
Cirene, U. Belgadir, 26.iv.1934, R. Pampanini & R.
Pichi-Sermoli 1156 (K). MAURITANIA. Fainyer?,
1.iv.1921, C. Pau (MA 016856). PALESTINE. Above Acie
Fit, woods, 9.v.1911, F. S. Meyers & J.E. Dinsmore
91781 (K). SPAIN. Canarias: Tenerife, Anaga, Sabinar
de Afur, 28.i.2004, J. A. Reyes Betancort
(TFC 45312). Tenerife, El Moquinal-Pedro Alvarez,
borde carretera, 15.v.2004, M. A. Padrón Mederos (TFC
45313). Tenerife, La Orotava, 30.vii.1983, E. Hernández (LEB 62448). Tenerife, La Orotava, Teide, bei El
Bebedero, c. 740 m, 28RCS 521 395, 18.vi.1985, E.
Vitek (W 2001-1476i). TURKEY. Antalya, Antalya, E. of
the town, waste places, along road, partly shaded
marl-soil, wet roadside, 35 m, 14.iv.1959, Hennipman
et al. 366 (Iter leydense) (K). Puertes Celiciennes,
11.vii.1855, B. Balanse (K). Saziantifs, Gazianlifs,
Kara Tash, nonline, rock volcanic, 3200 ft, 3.v.1935,
E. K. Balls B2218 (K).
4. Carex leersii F. W. Schultz, Flora 53: 459. 1870,
non Willd. 1787 (Fig. 8G–L)
= Carex canescens sensu Leers, Fl. Herborn. 712. t. 14.
f. 3. 1789, non L. 1753.
= Carex virens Lam. & DC, Fl. France, vol. 3: 106.
1805, non Lam. 1792.
= Carex divulsa Gooden b virens Durieu, Bull. Soc.
Bot. France 6: 633. 1859.
= Carex duriaei F. W. Schultz, Flora 53: 548. 1870,
non Steudel, 1841.
= Carex chabertii F. W. Schultz, Flora 54: 21. 1871.
Lectotype (designated by Loos): Feddes Repert.
107(1–2): 67. 1996 (B!). Isolectotype e.g.: BR, C, GB,
K!, L, M, MA!, MANCH, P!, W!.
= Carex divulsa Gooden. var. chabertii (F. W. Schultz)
Kneuck. in Seub. & Klein, Exk.-Fl. Baden: 52. 1891.
= Carex muricata var. leersii Kneuck. in Seub. &
Klein, Exk.-Fl. Baden: 52. 1891.
= Carex divulsa Gooden. ssp. chabertii (F. W. Schultz)
Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 42. 1902.
= Carex muricata race C. leersiana Rouy(‘leerseana’),
Fl. France 13: 413. 1912.
= Carex divulsa ssp. leersii (Kneuck.) W. Koch, Mitt.
Bad. Landesvereins Naturk. Naturschutz Freiburg
11: 259. 1923.
= Carex leersiana Rauschert, Feddes Repert. 83(9–10):
660. 1973. Lectotype (designated by Loos sub. C. leersii F. W. Schultz): Feddes Repert. 107(1–2): 67. 1996
(B!). Isolectotype e.g.: K!, L, M, MANCH, P!, W!.
Description: Caespitose perennial tufted plant with
or without short rhizomes. CULMS 39–90 cm ¥ 0.8–
2.0 mm at mid-height, obtusely trigonous, slightly
scabrid above; sheaths pale to dark brown. LIGULE
0.75–3.5 mm long, wider than long, rounded or subacute at apex. LEAF blades green to grey green, about
three-quarters to as long as culms, widest leaf blades
(2)2.25–3.75(4) mm, plicate to flat. INFLORESCENCE
oblong, 35–60(75) ¥ 8–10 mm, with 6–11(13) globularelliptic spikes. The lowest spike sometimes branched and shortly pedunculate, upper spikes single
at nodes, sessile, and contiguous, first internode
8–27 mm long, the second one 5–15 mm. BRACTS
glume-like, the proximal setaceous, shorter than the
inflorescence, 6–30 mm long. FEMALE GLUMES hyaline
to golden-yellow, with a midrib green, 3.0–4.5 ¥
1.6–2.30 mm, ovate or elliptic, apex acuminate
0.15–0.3 mm long. STAMINATE GLUMES 2.75–3.5 ¥
1.5–2.5 mm, similar to the female. ANTHERS 3, 1.8–
3.0 mm long. PERIGYNIUM erect-patent to patent,
greenish, body elliptic to ovate, nerveless or faintly
veined, 4.5–5.5 ¥ 1.75–2.25(2.5) mm, acute at base,
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THE CAREX DIVULSA AGGREGATE IN EURASIA
sometimes slightly corky at the base, gradually tapering into a beak. BEAK 0.67–1.25 mm long, white to
brown distally, with serrulate winged margins reaching the summit, apex bifid or bidentate, apical teeth
0.3–0.6 mm. ACHENE elliptic to ovate, shining brown,
2.25–3.0 ¥ 1.5–2.1 mm. STIGMAS 2, 2.0–3.0 mm long.
Phenology: Flowering begins in May, fruiting from
June to August, wind pollinated.
Habitat: Carex leersii occurs in areas with a temperate oceanic climate. Although most authors (De
Langhe, 1944; Nelmes, 1947; Chater, 1980; Jermy
et al., 1982; David & Kelcey, 1985; Clapham et al.,
1987; Sell & Murell, 1996) and most of the studied
materials indicate that it lives on calcareous or rich
soils, other authors, such as Luceño (1994), do not
agree. Chabert stated that he sampled his specimens
of C. chabertii from ‘coteaux granitiques de Laveyron.
Drôme’. Therefore, this taxon is not exclusively
calcicolous. We found C. leersii on well-drained, drysummer soils, in sunny open places, and often in
disturbed habitats (ravines or roadsides).
Distribution: Carex leersii occurs (Fig. 6) from northern Spain and Italy (41°N) to Great Britain and the
Scandinavian Peninsula (60°N), and Turkey in the
east, between 300 and 1000 m. It is more common in
France, Austria, and Germany, but is rare in Eastern
Europe.
Observations: Schultz (1871: 25) gave a very detailed
description of C. leersii based on his own material.
The best diagnostic characters of C. leersii are
perigynium elliptic to ovate c. 5 mm, and hyaline
elliptic female glume. Table 3 demonstrates that
C. leersii shows significant morphological variation.
Frequently, specimens labelled as C. leersii are young,
sterile, or damaged, and, according to Jermy et al.
(1982) and David & Kelcey (1985), when animals
have cropped them, they have depauperate flowering
stems and are difficult to identify. Specimens from
Greece and Turkey frequently show intermediate
characters between C. leersii and C. otomana. Moreover, in the lowest part of the Pyrenees, specimens
with intermediate characteristics between C. leersii
and C. magacis were found.
Representative specimens examined: BELGIUM.
Buissons, 1.vi.1907, Izn (MA 016826). CROATIA. Ins.
Korcula (of pines in maquis), under light shade, on
limestone hills, 1.v.1976, C. C. Townsend 76/36 (K).
FRANCE. Haute-Saône, Port-sur-Saône, au pied d’un
mur, 30.v.1930. V. Madiot (P). Isère, Rivoire de Allemont, carretera local, talud carretea, a pleno sol, 1000 m,
30.vii.2004, A. Molina (LEB 82667). GERMANY. Baden,
401
Buntsandstein, Weinberg, bei Ettlingen, auf berasten,
with C. divulsa, C. praecrox, 180 m, 48°57′N. Br.,
26°6′E v. F., 16.vi.1895, A. Kneucker (K). GREECE.
Epirus, inter Joannina et Dodona, sust. Calc., 1000–
1200 m, 26.v.1964, K. H. Rechinger (MA 419448).
HUNGARY. Villany, Comut. Bourounya, in monte Harsanyhegy, 16.v.1907, Herb. Dr A. de Degen: Koces (K).
ITALY. Sudtirol, Val Vestino, bei Bolone, an Wiesenund Wegrändern, 700 m, 1.vi.1901, P. Porta (MA
236899). Tirol australis, Val Vestino, près Bollone,
1.vi.1879. P.Porta (P). SPAIN. León, Aviados, entre
talud de la carretera y bosquete de Quercus pyrenaica,
c. 1000 m, 30TN 9851, 26.vi.2004, A. Molina (LEB
82662). Lérida, Valle de Arán, Pontant, Quercion
robori-petraea, 660 m, 31TCH 1445, 8.vii.1992, P. Monserrat, J. L. Benito & J. A. Sese (JACA 223192).
TURKEY. Trabzon, near 20 km W of Trabzon, marshy
grasses along pool, 4.vi.1959, E. Hennipmann et &.
1756 (K). SWEDEN. Skárva, Blekinge, Stauraby,
14.vi.1926, B. Holmgren (K). SWITZERLAND. Branson,
Valais, 26.iv.1863, L. Favrat (K). UNITED KINGDOM.
Norfolk, Appleton, grassy roadside bank, on chalk,
28.vi.1942, C. E. Hubbard (K).
5. Carex nordica A. Mol., Acedo & Llamas,
sp. nov. (Fig. 9A–F)
Carice leersii similes, sed ab ea differt perigyniis
minoribus atque alatus, squamae feminea fusca,
spicae patens et separatus.
Holotype: Sweden: Skåne, Paroecia, Hyby Bökebergsslätt, in agare viae ferreae (sub. C. divulsa
Stokes subsp. leersii) 21.vi.1922, G. Samuelsson (K!).
Description: Caespitose perennial plant with or
without short rhizomes. CULMS 30–80 cm ¥ 0.8–
1.5 mm at mid-height, obtusely trigonous, slightly
scabrid above; sheaths light to dark brown. LIGULE
1–3 mm long, wider than long, subacute to rounded at
apex. LEAVES green to dark green, about threequarters as long as culms, widest leaf blades
2.0–3.0 mm, plicate to flat. INFLORESCENCE oblong,
35–55(70) ¥ 7–10 mm, with six to nine globular spikes,
sessile, single at nodes, first internode 13–30 mm
long, the second one 7–14 mm, the remaining spikes
contiguous. BRACTS glume-like, the proximal setaceous, shorter than the inflorescence, 10 mm long.
FEMALE GLUME brown, with a midrib green to brown,
with scarious margins, 2.75–3.75 ¥ 1.5–2.25 mm,
elliptic, apex acuminate 0.25–0.3 mm long. STAMINATE GLUME 2.75–3.5 ¥ 1.5–2.0 mm, similar to the
female. PERIGYNIUM patent to very patent, greenish
to brown, body ovate to elliptic, nerveless or faintly
veined, 4.0–4.75 ¥ 1.75–2.5 mm, base rounded, gradually tapered into a beak. BEAK 0.75–1.35 mm long,
brown distally, with serrulate winged margins reach-
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402
A. MOLINA ET AL.
Figure 9. Carex nordica (A, C–F, from LEB 48138; B, from MA 387223): A, inflorescence; B, male glume; C, female
glume; D, perigynium, abaxial side; E, achene; F, perigynium with abaxial side removed to show achene. Carex magacis
(from holotypus LEB 82649): G, inflorescence; H, male glume with anther; I, female glume; J, perigynium, abaxial side;
K, achene; L, perigynium with abaxial side removed to show achene. All figures drawn by X. Magaz. Scale bar, 1 mm.
ing the body, the winged margins going to the middle
or the full body, apex bifid, apical teeth 0.3–0.6 mm.
ACHENE elliptic, shining, dark brown, 2.0–2.75 ¥
1.25–2.0 mm. STIGMATA 2, 2.0–2.25 mm long.
Distribution: Carex nordica is scarcely distributed in
a narrow zone from Great Britain to Sweden, and in
some scattered localities in Central Europe (Fig. 6).
Etymology: The name comes from the European area
in which it lives.
Representative specimens examined: AUSTRIA. Osttirol, Deferegen Tal: zwischen Huben und Hopfgarten,
Quellfluren, Felsen, Schutthalden, 6.vi.1967, F. Krendl
(W 1985-10548). Vorarlberg, Tosters westlich Feldkirch am Weg zur Ruine Tosters, 1.viii.1968. A.
Polatschek. (W 1969-2947). BELGIUM. Laeken, parc
public, bord chemin, 27.vi.1924, N. Gunz (K). Liège,
Amay, talus s, exp S; sol silicieux, avec Arrhenatherum,
6.vii.1972, W. Bellotte (MA 387223) and (LEB 48138).
Amay, 6.vii.1972, Malato Baliz (MA 274200). Hainaut,
Jamioulx, talus herbeux à exposition S, rue Francois
Phenology: Flowering begins in June, fruiting from
July to August, wind pollinated.
Habitat: It occurs in disturbed habitats and the
borders of tracks. The scarce data available indicate
human altered habitats, such as car parks, railways,
parks, and roads. It lives in temperate hyperoceanic
to oceanic areas.
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THE CAREX DIVULSA AGGREGATE IN EURASIA
Vandanme, 29.vi.1972, Duvigneaud & Lebeau (MA
387222). Hainaut, Vaulx, limite d’Antoing, friche herbeuse sur calcaire, 18.vi.1972, J. Duvigneaud (MA
274405). Namur, Purnode, près de la gare, vallée du
Bocq, antique voie ferrée, avec C. spicata, friche herbeuse termophile, 2.vii.1979, Duvigneaud (MA 236900)
and (MA 387225). DENMARK. Dyrharnen, Klamperburg, Sjaelland, park, 15.viii.1947, W. R. Price 2035
(K). Fyen, Nyborg, 1.viii.1868, B. J. Cöster (K). Fyen,
Faaborg, D32: Dyreborg Skov, rides in plantation by
seashore, 24.vii.1952, D. P. Young 4479 (K). NETHERLANDS. South-Limburgh, between Gud-Valkenburg
and Schin op Geul, 23.vii.1932, A. de Trever in Kern
& Reichgelt herbarium (K). SWEDEN. Gotlandia,
Wisby, 1.vii.1868, F. Ahlberg (K). Skåne, Sk. Stehag,
1.viii.1878, N. H. Nilsson (K). Skåne, Ringsgen,
1.vii.1828, N. H. Nilsson (K). UNITED KINGDOM. Derby,
vc. 57: Millersdale. 27.vi.1943, N. F. D., Herb. G. V. C.
Last 753/60 (K). Wales: Pembrokeshire, Caldey Island,
between high cliff and park area, 1.vi.1952, F. N.
Hepper (K).
6. Carex magacis A. Mol., Acedo & Llamas,
sp. nov. (Fig. 9G–L)
Carice leersii similes, sed ab ea differt perigyniis
rhombi formam habens; habet basim truncate,
2.25 mm latioribus, atque alatus. Carice muricatae
similis, sed ab ea differt perigyniis longioribus,
squamae feminea hyalina, denticuli rostrum dilatati
in perigynia corpore.
Holotype: Spain, Navarra, Selva de Irati, Senda 69,
c. río Urtxuria, sobre grietas de caliza en borde de
camino umbroso, 800 m, 42°57′N, 1°03′W, 5.vii.2004,
A. Molina (LEB 82649).
Description: Caespitose perennial plant with short
rhizomes c. 6 mm in diameter. CULMS 26–86 cm ¥
1.0–2.0 mm at mid-height, obtusely trigonous, slightly
scabrid above; sheaths pale to dark brown. LIGULE
0.5–3 mm long, wider than long, rounded to subacute
at apex. LEAVES pale green, about as long as culms,
widest leaf blades 3.0–4.25 mm, plicate to flat.
INFLORESCENCE oblong, 40–60 ¥ 10–12 mm, with 7–12
globular spikes, usually the first one branched and
shortly pedunculate, upper spikes sessile, single at
nodes, and contiguous, first internode 14–21 mm long,
the second one 7–19 mm. BRACTS glume-like, the
proximal setaceous, shorter than the inflorescence,
6–18(48) mm long. FEMALE GLUME golden-yellow to
light brown, with a midrib green, with scarious
margins, 3.0–3.6 ¥ 2.0–2.5 mm, elliptic, apex apiculate
0.10–0.25 mm long. STAMINATE GLUME 3.0–4.0 ¥ 1.5–
2.0 mm, similar to the female. ANTHERS 3, 2.0–2.5 mm
long. PERIGYNIUM very patent to patent, greenish,
body trullate to ovate-trullate, with a broad winged
403
margin, nerveless or faintly veined, 4.5–5.5 ¥ 2.25–
2.75 mm, base cuneate, gradually tapered into a beak.
BEAK 0.75–1.25 mm long, white distally, with serrulate winged margins reaching the body, apex bifid,
apical teeth 0.3–0.7 mm. ACHENE elliptic, shining
brown, 2.5–3.0 ¥ 1.75–2.25 mm. STIGMATA 2, 2.5–
3.25 mm long.
Etymology: The new subspecies is named in honour of
the family Magaz.
Phenology: Flowering begins in June, fruiting from
July to August, wind pollinated.
Habitat: Carex magacis inhabits rocky places, such as
ravines, stony areas, and open spaces in forests
(scotch pine, beech, etc.) in a wide range of montane
level communities [(600)780–1200 m; mean temperature, 2.5–7.5 °C; high precipitation (700–1200 mm)]
in temperate oceanic sub-Mediterranean zones. It can
grow on rich calcareous soils, but its resistance to
acidic conditions is not known.
Distribution: Carex magacis is scarce (Fig. 6) and is
found in a small area in the Pyrenean Mountains and
Iberian Range (Spain, Andorra, and France).
Conservation status: Carex magacis is an endemic
plant. Its distribution indicates that it is a rare taxon
which, according to IUCN (2001), and because of the
few localities and small area in which it is found,
must be catalogued as vulnerable (VU).
Representative specimens examined: ANDORRA. Parroquia S. Julià de Lòria, Cova de l’Ossa, taludes,
roquedos, cultivos abandonados, esquistos, 1020 m,
42°27′41″N, 1°29′12″E, 30.v.2004, C. Aedo, Aizpuru,
Barriego, Pedrol & Molina (MA 714438). Sta Julia
de Loria, Pont de la Marginada, ambiente de
encinar, suelo pobre en bases, 980 m, 31T CH7604,
6.vii.1992, S. Castroviejo et al. (MA 512056).
FRANCE. Hautes Pyrénées, St Marie de Capman,
subida a Pont de la Moulette, bord chemin calizas,
1000–1200 m, 10.vii.2004, A. Molina (LEB 82663).
SPAIN. Castellón: Entre Mas de la Cabra y el río
Monleón
(Vistabella),
1100 m,
30TYK
2768,
14.vi.1987, C. Fabregat (MA 467749). Huesca: Aniés,
Senda a ermita Virgen de la Peña, calizas abrigadas, 1080–1100 m, 30TYM 0187, 3.vi.1992, L. Villar
J. A. Sesé et al. (JACA 79392). Casbas de Huesca,
Sierra Guara, collado, relleno y peñascos, 970 m,
30TYM 2981, 30.vi.1972, P. Monserrat & L. Villar
(JACA 379772). Castillonroy, Barranco de la acequia
de Salellas, Baldellou, 500–650 m, 31TBG 9545,
27.v.1987, G. Monserrat (JACA 434787). Fanlo,
Añisclo, bosque mixto, 950–1200 m, 31BTH 5817,
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
404
A. MOLINA ET AL.
2.vii.1973, P. Monserrat & L. Villar (JACA 263873).
Jusen, Graus, alrededor del pueblo, 780–820 m,
31TBG 8263, 1.vi.1986, G. Monserrat (JACA
658486). Nueno, Sierra Guara, Nocito, Bco. de
Lapillera, glera, solana cálida, hayedo en hoya,
1050 m, 30TYM 2987, 11.vii.1974, P. Monserrat
(JACA 368274). Oroel, Jaca, abetal con hayas
escasas, 1200 m, 30TYM 0410, 25.viii.1988, L.
Villar, D. Gómez et al. (JACA 271888). San Juan de
la Peña, camino del mirador, pinar de P. sylvestris,
1240 m, 30TXN 9109, 14.vi.1970, P. Monserrat & L.
Villar (JACA 223470). San Juan de Plan, pradería,
prado de siega, 1150–1200 m, 31TBH 8219,
19.vi.1981, P. & G. Monserrat, D. Gómez & F. Fillat
(JACA 066481). Navarra: Burgui-Salvatierra, Foz de
Forniellos, 1050 m, 30TXN 6830, 1.vii.1975, L. Villar
(JACA 10069075). Teruel: Rubrelos, 23.vi.1897,
Benedicto (MA 16803). Segura de Baños, cantiles y
rellenos, 1000 m, 30TXL 7235, 11.vii.1973, P. Monserrat, J. Erviti & L. Villar (JACA 318173). Zaragoza: Luesia, alto de la Sierra de Sto. Domingo,
herbazales, 1250–1300 m, 30TXN 6902, 21.vi.1987,
J. Pedrol (MA 437584). Salvatierra de Esca, Foz de
Forniellos, cantiles solanos y carrascal con quejigo,
750–800 m, 30TXN 6829, 22.vi.1973, L. Villar
(JACA 100118673).
7. Carex otomana A. Mol., Acedo & Llamas,
sp. nov. (Fig. 10A–F)
Carice leersii similes, sed ab ea differt perigyniis
ovate sunt, maioribus, 5.5 mm longitudines, corpore
glaucous, squamae feminea fusca, in rostrum dilatati
utriculorum.
Holotype: [Kazakhstan], Asia centralis: Alma-Ata,
Tyan-Shan montes ‘Zailijki Alatau’, in valle fluminis
‘Malaia Almatinka’, loco Medeo dito, 1400–1750 m,
30.v.1974 (sub Carex polyphylla Kar. & Kir.), V. Vasák,
Iter centrali-asiaticum 1974 (W 11223!).
Description: Caespitose perennial plant with or
without short rhizomes. CULMS 50–90 cm ¥ 1.5–
2.5 mm at mid-height, obtusely trigonous, scabrid
above; sheaths pale yellow to dark brown. LIGULE
1.5–4.0 mm long, wider than long, subacute to rounded
at apex. LEAVES grey green, about three-fifths as long
as culms, widest leaf blades 2.25–4 mm, plicate to flat.
INFLORESCENCE oblong, 40–65(85) ¥ 9–14 mm, with
8–13 globular-elliptic spikes, sometimes the first spike
branched and shortly pedunculate, the remaining
spikes single at node, sessile, and contiguous, first
internode 10–17 mm long, the second one 7–17 mm.
BRACTS glume-like, the proximal setaceous, shorter
than inflorescence, 6–20 mm long. FEMALE GLUME
light brown, with a midrib green, with scarious
margins, 3.25–5.25 ¥ 1.75–2.3 mm, ovate to elliptic,
apex acuminate 0.2–0.5 mm long. STAMINATE GLUME
3.75–4.75 ¥ 1.5–2.0 mm, similar to the female.
ANTHERS 3, 1.75–2.0 mm long. PERIGYNIUM erectpatent to patent, greenish, body ovate, nerveless or
faintly veined, 5.0–6.0 ¥ 2.0–2.5 mm, base rounded to
cuneate, slightly swollen and corky, gradually tapered
into a beak. BEAK 1.0–1.6 mm long, brown distally,
with serrulate winged margins reaching the body, apex
bifid, apical teeth 0.5–0.75 mm. ACHENE elliptic to
ovate, shining dark brown, 2.5–3.0 ¥ 1.75–2.25 mm.
STIGMAS 2, 2.0–2.25 mm long.
Etymology: Species dedicated to the Ottoman people,
who developed an empire in part of the area in which
the species lives.
Phenology: Flowering begins in May, fruiting from
June to July, wind pollinated.
Habitat: Carex otomana occurs between 700 and
1850 m in wet mountain areas (ravine forest,
meadows, and springs) in open forest or above the
timberline in all types of soil.
Distribution: Carex otomana occurs (Fig. 6) from east
of the Black Sea (Bulgaria) and Greece through
the Turkish mountains and the Caucasus to the
mountains on the west side of Tyan Shan in Central
Asia (Kazakhstan).
Observations: Greek specimens of C. otomana are
similar to C. leersii with regard to perigynium size
and shape, but can be easy differentiated by the
brown glumes. Towards the east of the area, the
specimens have larger perigynia that are clearly
ovate and slightly corky at the base. The characters
quoted by Nilsson (1985) for C. divulsa ssp. leersii can
also be assigned to C. leersii and C. otomana. Nevertheless, C. divulsa ssp. leersii sensu Kukkonen (1998)
has characteristics from C. otomana (female glume
triangular, utricule ovoid, base slightly suberous,
beak 1.4–2.3 mm, long teeth), although some specimens reported by him are outside the distribution
area of this species. Similar characters have also been
reported in C. polyphylla sensu Kreczetovicz (1935)
(scales ovate, ferrugineous, perigynium ovoid, long
beak).
Representative specimens examined: ARMENIA. TakAchac, north-east shore Seran Ist ravine, lower boundary of forest, 16.vi.1930, O. Polianskaya (leg por Krecz)
(K). BULGARIA. Varnenski Okrug, Varna, Slatni Pjasatsi, glade of a shrubby woodland slope, 50 m, NH3,
7.vi.1973, J. Suominen 3056 (H 1664673). GREECE.
Serron, Serron, Mt. Vrondous, summit area (meadows)
rocky slopes (and rock crevices) at and above timber-
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
THE CAREX DIVULSA AGGREGATE IN EURASIA
405
Figure 10. Carex otomana (from J. Suominen 3056, H 1664673): A, inflorescence; B, male glume with anther; C, female
glume; D, perigynium, abaxial side; E, achene; F, perigynium with abaxial side removed to show achene. Carex egorovae
(G–I, from O. Polunin 5110 K; J–L, from R.W. Haines W 676 K): G, inflorescence; H, male glume with anther; I, female
glume; J, perigynium, abaxial side; K, achene; L, perigynium with abaxial side removed to show achene. All figures drawn
by X. Magaz. Scale bar, 1 mm.
line (formed by Pinus sylvestris), granite, 1700–
1840 m, GL1, 6.vii.1985, M. Salmenkallio & V. Salo
895 (H 1557247). IRAN. E. Mazandaran, N. Semman,
southern slopes of Alu-Baq Mountain: around Cheshm,
Cheshmehe Alu-Baq (spring) (Golestan National
Park), 1600–1650 m, 37°19′N, 55°53′15″E, 7.viii.1995,
H. Akhani 11898 (H 1695176). Gorgan, Mohammed
Reza Shah National Park, forest S. of Tang-e Gol,
subst. calcareo, in quercetis (Q. castaneifolia) apertis
ad cascades, 750 m, Iter Iranicum IX (1975), 4.vi.1975,
K. H. Rechinger 52649 (WU 024145, WU 024144, and
W 05994). Gorgan, Shahpasand, 73 km. Shahpasand,
750 m, 5.vi.1975, Termeh, Herbarium Minesterii
Iranici Agriculturae EVIN n°36871 (H 1686153).
NW Khorasan, Centre, 6 km E of Tangegol, Golzar,
in Zelkova carpinifolia and Carpinus forest (Golestan
National Park), 800–950 m, 37°22′N, 56°00′E,
9.vii.1994, H. Akhani 9604 (H 1695181). Gorgan,
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
406
A. MOLINA ET AL.
Mohammed Reza Shah Wildlife Park, forest S of
Tang-e Gol, 700–1000 m, Ariamehr Bot. Garden,
21.vi.1974, Wendelbo & Foroughi 12795 (W 21498).
KIRGUISTAN. Songarica, 1.ix.1870, Schrenk (K).
MACEDONIA. NW Macedonia, Drnova, Angkatholow,
bank of ravine, 2800 ft, 13.vi.1932, A. H. G. Alston & N.
Y. Sandwith 296 (K). TURKEY. Bursa A2 (A), above
Bursa on the road to Uludag, roadside in Castaneo–
Fagetum, 730 m, 8.vii.1978, P. Uotila 27146 (H
1203994). Prov. Adana, Amanus, 1500 m, 26.viii.1949,
P. Davis 16442 (K). UZBEKISTAN. Asia Media, distr.
Andizham, Fergana, in silva prope Arslanbob,
25.v.1899, Litrovinov (K).
8. Carex egorovae A. Mol., Acedo & Llamas,
sp. nov. (Fig. 10G–L)
Carice leersii similes, sed ab ea differt inflorescentia
brevioribus, 4–5 mm longitudine; spicae erectopatens, pauciflorae; perigynia rostrum longiore et
bifidus.
Holotype: Iraq. MAM, Shariffa near Amadia, shady
meadow, 4000 ft, 22.iv.1958 (sub. Carex divulsa
Stokes ssp. leersii (Kneuck.) W. Koch), O. Polunin
5110 (K!). Isotype: O. Polunin 5110b (K!).
Description: Caespitose perennial plant with or
without short rhizomes c. 5 mm in diameter. CULMS
14–50 cm ¥ 0.8–1.25 mm at mid-height, obtusely trigonous, slightly scabrid above; sheaths pale to dark
brown. LIGULE 1.5–3.0 mm long, wider than long,
rounded or subacute at apex. LEAF blades grey green,
about three-quarters to longer than culms, widest leaf
blades 2–3 mm, plicate to flat. INFLORESCENCE oblong,
30–50(68) ¥ 9–13 mm, with 6–11 globular-elliptic,
sessile spikes, the lowest spike sometimes branched,
upper spikes single at nodes, first internode 6–
13(20) mm long, the second one 3–9(13) mm. BRACTS
glume-like, the proximal setaceous, shorter than the
inflorescence, 5–38 mm long. FEMALE GLUME hyaline,
with a midrib green, 3.0–4.0 ¥ 1.6–2.25 mm, ovate,
apex acuminate 0.3–0.5 mm long. STAMINATE GLUME
3.0–3.5 ¥ 1.4–2.0 mm, similar to the female. ANTHERS
3, 1.5–2 mm long. PERIGYNIUM erect-patent, pale
greenish, body ovate, nerveless or with well-marked
veins, 4.25–5.5 ¥ 1.8–2.5 mm, rounded base, some of
them slightly swollen and corky, gradually to abrupt
tapered into a beak. BEAK 1.0–1.75 mm long, white
distally, with serrulate margins reaching the summit,
apex bifid, apical teeth 0.6–1.25 mm. ACHENE elliptic,
shining, brown, 2.0–2.7 ¥ 1.6–1.8 mm. STIGMATA 2,
2.0–2.5 mm long.
Etymology: This new species is named in honour of
Dr Tatiana Egorova, a Russian specialist in Carex.
Phenology: Flowering begins in April, fruiting from
April to June, wind pollinated.
Habitat: Damp and shady places between 700 and
1360 m: stony hillsides, brook sides, meadows, and
forests.
Distribution: Carex egorovae occurs in three disjointed areas (Fig. 6): Eastern Europe (from Hungary
to north-west Turkey), Middle East (Iraq and Iran),
and Cyprus.
Observations: Specimens from Europe have slightly
different characters to those from the other two areas.
European plants have longer and more serrulate
beaks and are not corky at the perigynium base.
Given the scarce number of available sheets, they are
all given the same rank. The plants from Iraq that
were included in C. divulsa ssp. leersii by Hooper
(1985) seem to be the same plant.
Representative specimens examined: CROATIA, Montes
Velebit, pars austr., Nationali Park Paklenica, in valle
augusta rivi Velika Paklenica dispersim, Flora Balcanica: Croatica, 24.v.1996, Mil. Marek (WU). CYPRUS.
Kythrea, in irrigation channel, 500 ft, 13.iv.1962, R. D.
Meikle 2556 (K). HUNGARY. Nr Podhragy in Ungarn,
Holuby (K). Nr Podhragy in Cominztronam, 1876,
Holuby (K). Herkulesbad im Banat in Ungarn, auf der
Coroninihöle, con Genista tinctoria, Thymus montanus, etc., 500 m, 30.vi.1902, L. Ritcher (K). GREECE.
Thasos, S coast, wood by stream, 100 ft, 24.v.1972, W.
R. Price 1224 (K). IRAN. Asterabad, Bender Ges. in
dumetis, 30.iii.1901, P. E. Sintesis 1466. (P 00281845).
Trorth Gonbad, near village Golidagh, shady area,
under trees, 1080 m, 25.v.1976, T. F. Henr 3872 (K).
Gorgan, Persia borealis, Asterabad, Bender Ges,
30.iii.1901, J. Freyn (K). IRAQ. MSU district, Tawila,
stony hillside, 1360 m, 16.vi.1957, A. Rawi 21896 (K).
Kurdistan, Sulaimaniya, in ditione pagi Tawilla, Mte.
Avroman at confines Persiae, Sosakan, in platanetis,
1200 m, Itinera Orientalis 1956/57, 18.vi.1957, K. H.
Rechinger 12137a,b (W 01174, W 01146). MRO district,
Shaqlawa, irrigated orchard-pastures in rough grass,
2800 ft, 15.v.1956, Univ Herb Baghdad, R. W. Haines
(W 01133). Shaqlawa, shady bank on roadside, 2800 ft,
12.v.1956, R. W. Haines W676 (K). Saisang, N slope
Gara, mountain grassy, 1300 m, 20.vi.1968, Herb.
College of Agric. & Forestry, Univ Mossul, Anders 2359
(W 1945). TURKEY. Bilecik B2, c. 40 km from Inegöl
to Bozüyuk, Aksutekke, by the main road, brookside
meadow in the service station area, 860 m, 20.v.1983,
M. Salmenkallio 268 (H 1492360).
ACKNOWLEDGEMENTS
We are grateful to the staff of the consulted herbaria
for their help with the location of material. We also
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 385–409
THE CAREX DIVULSA AGGREGATE IN EURASIA
wish to express special gratitude to Professor David
Simpson for his kind help during our visits to the Kew
Herbarium. We especially thank Ms Maria Elizabeth
Bailey who corrected the final English version, Mr
Jorge Magaz for helping us with the drawings, and
Professor Benito Valdes and Professor Stephen Jury
for their advice and valuable comments. Finally, we
thank Junta de Castilla y León for granting a High
Studies Licence to the first author, and for grant LE
025A05 that partially supported our research.
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