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A taxonomic survey based on revision of herbarium material and critical study of bibliography is conducted to elucidate the taxonomic and nomenclatural status of four problematic Carex names: C. eremitica, C. mediterranea, C. orbicularis subsp. kotschyana var. caucasica, and C. transcaucasica, all described from SW Asia. We find that C. eremitica should be considered a synonym of C. stenophylla (section Boernerae). We perform the lectotypification of C. mediterranea, a synonym of C. hispida (section Thuringiacae). We discuss the taxonomic affinities of C. transcaucasica (section Phacocystis) and combine this name under C. nigra (C. nigra subsp. transcaucasica). We synonymize C. orbicularis subsp. kotschyana var. caucasica to C. nigra subsp. transcaucasica. Eventually, we indicate that populations previously believed to be C. orbicularis actually belong to C. nigra subsp. transcaucasica, confirming its presence in Iran, and expanding its range to Iraq.
Phytotaxa 219 (2): 183–189
Copyright © 2015 Magnolia Press PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Muthama Muasya: 27 Jun. 2015; published: 8 Jul. 2015
Taxonomic notes on some problematic Carex (Cyperaceae) names from SW Asia
1 School of Biological Sciences, Washington State University, 99164, Pullman, WA, USA.
2 Department of Botany, Iranian Research Institute of Plant Protection, P.O. Box 19395-1454, Tehran, Iran
3 Botany Area, Department of Molecular Biology and Biochemical Engineering, Pablo de Olavide University, ctra. de Utrera km. 1,
41013, Sevilla, Spain
* Corresponding author:,
A taxonomic survey based on revision of herbarium material and critical study of bibliography is conducted to elucidate the
taxonomic and nomenclatural status of four problematic Carex names: C. eremitica, C. mediterranea, C. orbicularis subsp.
kotschyana var. caucasica, and C. transcaucasica, all described from SW Asia. We find that C. eremitica should be consid-
ered a synonym of C. stenophylla (section Boernerae). We perform the lectotypification of C. mediterranea, a synonym of C.
hispida (section Thuringiacae). We discuss the taxonomic affinities of C. transcaucasica (section Phacocystis) and combine
this name under C. nigra (C. nigra subsp. transcaucasica). We synonymize C. orbicularis subsp. kotschyana var. caucasica
to C. nigra subsp. transcaucasica. Eventually, we indicate that populations previously believed to be C. orbicularis actually
belong to C. nigra subsp. transcaucasica, confirming its presence in Iran, and expanding its range to Iraq.
Key words: Caucasus, Eastern Mediterranean, Iran, Iraq, Middle East, nomenclature, Turkey, typification
The flora of southwestern Asia is relatively well-known from a taxonomic point of view. Several regional floras give
a relatively good and updated coverage for most of the families in the territory (e.g. Flora Iranica (Rechinger 1963-in
progress; Assadi et al. 1988-in progress), Flora of Turkey (Davies 1965–1985), Flora Palaestina (Zohary & Feinbrun-
Dothan 1966–1986), Nouvelle Flora du Liban et de la Syrie (Mouterde 1966–1970), Flora of Iraq (Townsend & Guest
1966–1985), Flora of the Arabian Peninsula and Socotra (Miller & Cope 1996)). However, taxonomic treatments of
controversial groups in this area are being constantly revised (Amini-Rad et al. 2010, Eker et al. 2014, Jiménez-Mejías
et al. 2014, among others), and new species continue to be frequently described (Vural et al. 2012, Martín-Bravo &
Jiménez-Mejías 2013, Ranjbar 2010, among others).
Carex Linnaeus (1753: 972) is the largest genus in the temperate zone and one of the largest in the world (Global
Carex Group in press). In Europe and the Mediterranean Basin it is represented by around 220 species (Jiménez-Mejías
& Luceño 2011a). In this paper we address some taxonomic problems on four names in Carex. We also reassess the
taxonomic and geographical delimitation between C. orbicularis (Boott 1845: 254) and C. nigra (L.) Reichard (1778:
96) in the Middle East.
Carex eremitica Paine
Carex eremitica Paine (1875: 126) is currently an accepted species presumably distributed in the southeastern
Mediterranean basin (Israel-Palestine and Jordan; Govaerts 2014, Jiménez-Mejías & Luceño 2011a). Its taxonomic
circumscription and sectional placement have been obscure, and the actual taxonomic status of this species has not
been revised to date. It was described in the Natural History section of the Identification of Mount Pisgah (Paine 1875),
based on materials probably collected by Paine himself in Balqa (“Belqa”) plain, today northwestern Jordan.
Paine provided a detailed description of the species, remarking its androgynous spikes congested in a broadly
capitate inflorescence (“spicis in capitulum lato-ovatum spice masculum confertis”), which allows identifying it as
184 Phytotaxa 219 (2) © 2015 Magnolia Press
a member of the subgenus Vignea (P. Beauv. ex Lestib) Petermann (1849: 602). Among other critical characters,
linear leaves, dark (“dusky”) purple glumes with hyaline margins, and utricles “almost smooth”, “rhomboidal”, and
“cuspidate beaked” were specified. The species was later recorded by Post (1986) in his Flora of Syria, Palestine and
Sinai, including a vague description that probably just reproduced Paine’s record, as it was reported only from Balqa.
Paine considered this plant to be allied to C. praecox Schreber (1771: 63), a species included in section Ammoglochin
Dumortier (1827: 146). However, no species from this group is currently known from Jordan. According to checklists
(Govaerts 2014, Jiménez-Mejías & Luceño 2011a) and floristic treatments of neighboring regions (Egorova 1999,
Kukkonen 1998, Nilsson 1986), there are only two Carex species that could potentially match Paine’s description:
C. stenophylla Wahlenberg (1803: 24) (section Boernerae V. Krecz. ex Egorova (1965: 70)) or C. pachystylis J. Gay
(1838: 101) (section Physodeae Meinshausen (1901: 280, 312)). This possibility was already considered by Feinbrun-
Dothan (1986) in Flora Palaestina, who considered C. stenophylla to be conspecific with C. pachystylis, and listed
“?C. eremitica Paine” under the synonyms list.
We have requested or in situ studied materials assignable to C. eremitica Paine (including materials classified as
C. stenophylla and C. pachystylis) from all herbaria where Paine’s collections have been reported to be deposited (G,
GH, K, MICH, NYS, PH; cf. Stafleu & Cowan 1976–1988). However, all searches of C. eremitica type material were
Carex pachystylis and C. stenophylla can be difficult to distinguish from each other when ripe fruits are not
available, as the main feature that allow discriminating between them are the coriaceous utricles, not easily detachable
from the spike in C. stenophylla, while papyraceous and deciduous in C. pachystylis (Egorova 1999, Kukkonen 1998).
Despite Paine (1875) did not give any information about this character, he provided a key clue that allows us to figure
out the identity of his species: “almost smooth appressed perigynia”, a character that contrasts with the regularly
serrulate beak of C. praecox Schreb. (cf. Luceño et al. 2008), to which Paine compared its C. eremitica. It implies
that the utricle beak of C. eremitica was not totally smooth but, at least, very sparsely scabrid. We have studied the
scarce available materials of both C. stenophylla and C. pachystylis from Israel-Palestine and Jordan from K and E
herbaria (Appendix I). In all the studied samples, C. pachystylis displayed completely smooth utricles, whereas C.
stenophylla sometimes had utricles somewhat scabrid at apex. Both characters have been previously recorded for each
species in other treatments (Kukkonen 1998, Nilsson 1986). Thus, we conclude that Paine’s C. eremitica is probably a
heterotypic synonym of C. stenophylla, so it should be removed from checklists as an accepted species name.
Carex mediterranea C.B. Clarke ex Post
The name Carex mediterranea C.B. Clarke ex Post (1896: 837) displays a somewhat similar problematic situation to
that of C. eremitica. The species, distributed in Syria and Jordan, is currently considered as accepted in regional and
global checklists (Govaerts 2014, Jiménez-Mejías & Luceño 2011a), although it has previously been synonymized to
C. hispida Willd. (in Schkurh 1801: 63) (section Thuringiacae G. Don (in Loudon 1830: 376)) by Mouterde (1966).
Carex mediterranea was described on the first edition of Post’s Flora of Syria, Palestine and Sinai (Post 1896). The
indicatio locotypica states “between Burmah and Gerash (Gilead); Lattakia”, today in Jordan and Syria, respectively.
Mouterde (1966) studied two type collections from Post’s herbarium (“Burma-Jérash, avril 1886” and “Lattaquié,
iun 1884”), probably housed at BEI (cf. Stafleu & Cowan 1976–1988). He identified them as C. hispida, although he
did not perform any formal typification. In despite of this synonymization was already taken up by Flora Palaestina
(Feinbrun-Dothan, 1986), Danin (2000) again considered C. mediterranea as a distinct species, and stated that a “sedge
expert” determination would be needed to verify a previous record (El-Oqlah & Lahham, 1985).
We have found a voucher at K herbarium confirmed by C.B. Clarke as C. mediterranea, whose label (“Burma—
Gerash; May 4, 1886”) matches part of the indicatio locotypica in the protologue. This specimen, collected by J.E.
Dinsmore, unequivocally belongs to C. hispida Willd., which corroborates Mouterde’s (1966) synonymization. We
hereby lectotypify the name C. mediterranea C.B. Clarke ex Post on this K material, and reassert that it is a heterotypic
synonym of C. hispida Willd.
Carex mediterranea C.B. Clarke ex Post in Flora of Syria, Palestine and Sinai: 837 (1896).
Lectotypus (designated here): “Burma—Gerash, 4 May 1886, J.E. Dinsmore” (K!).
Carex nigra in SW Asia east of Anatolia
Carex nigra belongs to the taxonomically problematic section Phacocystis Dumortier (1827: 146). The taxonomy of
C. nigra and its allies has been a complicate issue, mainly due to its wide morphological variability, that has led to
NOTES ON SOME CAREX NAMES FROM SW ASIA Phytotaxa 219 (2) © 2015 Magnolia Press 185
the description of many taxa below species level but also at species rank (see Chater (1980) for a detailed commented
list). One of the most outstanding examples has been the recognition of the tussock-forming plants as a separate
species (C. juncella (Fries) Th. Fries (1857: 207)), by caricologists as prominent as Egorova (1999). It is a growth-
form that strikingly contrasts with the typical rhizome-creeping plants. However, recent studies have shown that these
plants are genetically indistinguishable from C. nigra s.s. and that tussock-forming growth forms may be induced by
environmental conditions (Jiménez-Mejías et al., 2012; Košnar et al., 2013).
In the Middle East, C. nigra has been confused with C. orbicularis, and two additional taxa (C. transcaucasica
Egor. and C. orbicularis subsp. kotschyana var. caucasica Ö. Nilsson) were described to accommodate plants considered
to be problematic by regional authors.
1) Carex transcaucasica Egor.
Carex transcaucasica was described by Egorova (1989: 11) to accommodate the Caucasian populations of what she
considered to be distinct C. nigra-like plants. She stressed as differences between C. transcaucasica and C. nigra
the relative length of the lower bract (shorter than the inflorescence vs. longer or about as long as the inflorescence,
respectively), the utricle indumentum (without papillae vs. papillose), and the basal sheaths color (generally black-
purple vs. brown, reddish or yellowish-brown). Those populations are relatively geographically isolated from the
typical C. nigra populations from the neighboring non-Caucasian Russia and Anatolia (cf. Hultén 1950, Nilsson 1986,
Jiménez-Mejías et al. 2013).
Samples referable to C. transcaucasica were included in a previous genetic study (Jiménez-Mejías et al., 2013)
and found to be embedded within the genetic variation of the typical European forms of C. nigra. Therefore, these
Caucasian populations should be better considered as conspecific with C. nigra. Nonetheless, we studied eastern
Turkish and Caucasian materials of C. nigra-like plants (Appendix I) and observed that the distinct morphological
reproductive characteristics pointed out by Egorova (1999) are rather constant. Differences in the basal sheaths
(Egorova 1999) were found to be inconsistent, probably due to the different conservation of the sheaths in wet and
drier soils, as observed in other Carex sect. Phacocystis species (pers. obs.). In any case, the morpho-geographic
compartmentalization of the Caucasian populations stresses it as a taxonomic unit within C. nigra (cf. Stuessy 1990).
Therefore, we propose combining C. transcaucasica as a C. nigra subspecies (see below).
2) Carex orbicularis subsp. kotschyana var. caucasica Ö. Nilsson
Nilsson’s (1986) account of Carex sect. Phacocystis Dumort. for Flora of Turkey was one of the most important
contributions towards an accurate treatment of this taxonomically complex group in the Middle East. However, the
faint morphological boundaries between species, as well as the hybridization between them, have highly obscured the
different taxonomic units in this area. This, together with the probably limited availability of reference materials to
compare at the moment, misled Nilsson to a few wrong conclusions. For example, he conceived C. acuta Linnaeus
(1753: 978) and C. kurdica Handel-Mazzetti (1914: 23) as taxa linked to different phytogeographical regions, Euro-
Siberian and Irano-Turanian, respectively. However, some of the Turkish populations that he considered to be C.
acuta match the morphological variation of C. kurdica (Jiménez-Mejías et al. 2014), which increases the area of this
taxon from the Kurdistan westwards to the Mediterranean. Similarly, he overlooked the presence of C. buekii Wimmer
(1852: 83) in Turkey, due to its confusion with C. elata subsp. omskiana (Meinsh.) Jalas (in Jalas & Hirvelä 1964: 49)
(see Jiménez-Mejías & Luceño 2011b; Jiménez-Mejías & Rodríguez-Palacios 2014).
The case of C. orbicularis subsp. kotschyana var. caucasica Nilsson (1985: 156) (from herein C. orbicularis
var. caucasica) is somewhat similar to what happened with C. kurdica. Carex orbicularis is a southern Asian species
morphologically resembling both C. nigra and C. bigelowii Torr. ex Schweinitz (1824: 166) s.l. Its taxonomical
identity has already been supported by molecular studies (Schönswetter et al. 2008; Jiménez-Mejías 2011). Two
widely distributed geographical races have been recognized: the eastern subsp. orbicularis (ranging roughly from
Afghanistan to the Himalayas), and the western subsp. kotschyana (Boiss. & Hohen.) Kukkonen (1984: 389), from
the Caucasus to the Iranian mountain ranges called the Alborz and Zagros mountains (Kukkonen 1998; Egorova 1999;
Amini-Rad 2011). Nilsson (1986) described C. orbicularis var. caucasica to accommodate the northeastern Turkish
populations (Caucasian-bordering) of what he thought to be deviant forms of C. orbicularis subsp. kotschyana. On
the contrary, he considered the populations from the Armenian Highlands of southeastern Turkey to be typical C.
orbicularis subsp. kotschyana var. kotschyana (from herein var. kotschyana). Remarkably, despite he reported C.
nigra from the Turkish Caucasus, he did not notice the affinities of his C. orbicularis var. caucasica with C. nigra. It
186 Phytotaxa 219 (2) © 2015 Magnolia Press
is interesting to note that he distinguished C. orbicularis from C. nigra using characters that broadly overlapped with
those that he also used to distinguish var. kotschyana from var. caucasica: “utricles broadly elliptic to almost orbicular
[…] basal sheaths dark reddish brown” in C. orbicularis, vs. “utricles ovate to obovate-elliptic […] basal sheaths
pale brown” in C. nigra; “basal sheaths dark reddish-brown to dark brown […] utricles […] rounded at apex […]”
in C. orbicularis var. kotschyana, vs. “basal sheaths pale brown to dark-greyish brown […] utricles […] cuneate at
apex […]” in var. caucasica. Egorova (1999) already noticed the remarkable differences between the var. caucausica
and the var. kotschyana, and considered the former to “take an intermediate position between var. kotschyana and C.
Despite the holotype specimen of C. orbicularis var. caucasica (see below) seems to be lost (L. Glancy, E
herbarium, pers. comm.), we were able to study three paratype vouchers (Appendix I). Their morphology does match
that of C. nigra (amphistomatic leaves; utricles nerved, cuneate at the top and attenuated into a beak), especially the
subsp. transcaucasica, rather than C. orbicularis (hypostomatic leaves; utricles nerveless or faintly nerved, abruptly
contracted into a short beak) (see Egorova (1999) and Jiménez-Mejías et al. (2014)). In addition, a sample matching
var. caucasica morphology was included in a molecular study (Jiménez-Mejías 2011), and found to be placed within
C. nigra genetic variability, with no apparent relation with C. orbicularis. It also helps to reject that C. orbicularis
var. caucasica has a hybrid origin between C. nigra and C. orbicularis. Thus, we conclude that C. orbicularis var.
caucasica should be considered a heterotypic synonym of C. nigra subsp. transcaucasica.
3) Carex nigra subsp. transcaucasica in Iran and Iraq
Following a phytogeographical criterion, most authors have regarded southwestern Asian populations of C. nigra-like
plants outside Turkey as belonging to C. orbicularis.
For Iran, Assadi (1988) reported the presence of C. nigra in Azerbaijan province (Arasbaran Protected Area).
However, Amini-Rad (2011) rejected this record and considered it to be C. orbicularis. Kukkonen (1998) neither
included Assadi’s record nor listed C. nigra in his Flora Iranica treatment. For Iraq, Hopper (1985) only listed two
species of the section Phacocystis: C. elata Allioni (1785: 272) and C. orbicularis. The records of C. elata from Iraq
probably correspond to C. kurdica (cf. Jiménez-Mejías et al., 2014). For C. orbicularis, Hopper (1985) noted that
previous authors reported some records to be C. orbicularis and others C. nigra (as C. dacica Heuffel (1835: 247), a
name formerly misapplied to C. nigra (cf. Egorova, 1999)). However, she preferred to be synthetic and considered all
them as C. orbicularis. After the molecular confirmation of Iranian specimens from Azerbaijan province as C. nigra
(Jiménez-Mejías et al., 2013), we carefully examined C. nigra / C. orbicularis-like materials from the Middle East
following Egorova (1999) and Jiménez-Mejías et al. (2014). The populations listed in Appendix I match the variability
of C. nigra, in particular subsp. transcaucasica. The following key, modified from both treatments, readily allows the
distinction of C. orbicularis and C. nigra. For the distinction between these two taxa and the other species from section
Phacocystis co-occurring in this area, the reader must be referred to Jiménez-Mejías et al. (2014).
1. Utricles faintly nerved to nerveless, very rarely nerved, orbicular to suborbicular in outline, rounded at the top and abruptly con-
tracted into a short beak, biconvex to inflated-biconvex, arranged from erect-ascendent to patent in the spike; leaves hypostomatic,
more rarely amphistomatic ............................................................................................................................................ C. orbicularis
- Utricles conspicuously nerved, rarely faintly nerved, ovate to elliptic-obovoid in outline, cuneate at the top and gradually attenu-
ated into a short beak, narrowly biconvex to plano-convex, arranged erect-ascendent in the spike; leaves epistomatic or amphisto-
matic, very rarely hypostomatic .............................................................................................................................................C. nigra
To sum up, the study of materials allows us to confirm the presence of C. nigra in Iran, and spread its range further south
to Iraq. Conversely, for C. orbicularis, the Iraqi records must be considered doubtful and needed of confirmation.
4) Nomenclature
Carex nigra subsp. transcaucasica (Egor.) Jim.-Mejías, G.E. Rodr., Amini-Rad & Martín-Bravo, comb. & stat. nov.
Basionym: Carex transcaucasica Egor. (1989: 11). Type:—(RUSSIA) Caucasus septentrionalis, reservatum Teberdense,
in prato ad declive austro-orientale vallis fl. Mukhu, ad rivulum, 2000 m, 19 July 1936, D. Wolgunov 206 (holotype
LE, photo!).
= C. orbicularis subsp. kotschyana var. caucasica Nilsson (1985: 156). Holotype: (TURKEY) Erzurum, Bingöl Dağ, 16 July 1980, A.
Engin 387 (E, probably lost (see above)). Paratypes in E and K (see Appendix I).
- C. orbicularis sensu Hopper (1985: 404) pro parte, non Boott (1845: 254).
NOTES ON SOME CAREX NAMES FROM SW ASIA Phytotaxa 219 (2) © 2015 Magnolia Press 187
We would like to acknowledge to M. Escudero and an anonymous reviewer for their comments that greatly improved
the quality of this manuscript; the curators and staff from the herbaria G, GH, MICH, NYS, and PH for checking the
existence of C. eremitica vouchers among their collections; the curators and staff from E, FI, G, K, IRAN, M, MSB,
NY, and TARI herbaria for support when in situ studying their collections and/or sending us materials on loan; P.
Volkova for providing the protologue and pictures of the holotype of C. transcaucasica; and E. von Raab-Straube
from the Berlin Botanical Garden for their critical indications about previous literature records for the studied names;
funding was provided by CGL 2012-38744 project from the Spanish Ministry of Economy and Competitiveness, and
by a US National Science Foundation (NSF Award-1256033) postdoctoral fellowship towards P.J.M.
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List of additional studied materials.
C. nigra subsp. transcaucasica
ARMENIA: Aragatsotn province, Mt. Aragats, route from lake at end of the road along eastern flank of Mt. Hamberd
to castle Hamberd, 12 July 2003, H. Ter-Voskanian & E. Vitek 03-1640 (FI).
GEORGIA: Bodganovka, Aspaza village, lake Taparavani, 13 July 1949, Zamtaradze (E). Desheti, Azagui river source,
Mleta pastures, 1 August 1971, Zamtaradze (E). Kazbegi, Truso gorge, 25 July 2005, Abdaladze 588 (NY).
IRAQ: NW corner of Erbil Liwa, Ser Kurawa, 11 August 1947, Gillett 9781 (K). Al-Gurd Dagh, 22 July 1932 Guest
28855 (K). Erbil Liwa, Al-Gurd Dagh, 5 August 1947, Gillett 9554 (K).
IRAN: Mazandaran province: 20 km S of Ramsar, between Tanoorekash and Janat-Rudbar, 14 July 1984, Assadi
& Maassoumi 51378 (TARI); S of Ramsar, E of Lapasar, 12 August 1976, Runemark & Maassoumi 21684 (TARI).
Azerbaijan province: Arasbaran, Doghroon mountain, 2600 m, 21 June 2004, Amini-Rad 37226 (IRAN); Meshkin
Shahr, Ghotour Soui, Shabil, 2650–2800 m, 21 July 2005, Amini-Rad 41625 (IRAN); Khoy, Avrin mountain, 2824
m, 19 July 2011, Amini-Rad & Torabi 57618 (IRAN); Orumieh, Dizaj, Gisian, Dalamper mt., 2773 m, 22 July 2011,
Amini-Rad & Torabi 57619 (IRAN); Orumieh, Silvana, Khalil Kuh, 2600 m, 3 July 2010, Amini-Rad & Torabi 54786
(IRAN); 14 km Piranshahr to Naqade, Silveh village, Sefid Kuh, 2500–3175 m, 1 July 2010, Amini-Rad 54783 (IRAN);
Ardabil, Mounta Sabalan, August 1961, G.N. Harrington 279 (K).
RUSSIA: Kabardino-Balkar Republic: Karasu, 17 August 1925, Y.N. Bush (G); Khyzny-su, 4 August 1927, Y.N. Bush (G).
TURKEY: Hakkari, Kara Dag, 15 August 1954, Davis & Polunin 24391 (E). Szandschak Gümüschkhane, 31 July
1894, C. Haussknecht 7394 (E). Szandschak Gümüschkhane, 31 July 1894, C. Haussknecht 7398 (E) [classified by
Ö. Nilsson as C. orbicularis var. caucasica]. Hakkari, Samdi Dag, July 1965, E.M. Rik 232 (K). Erzurum, Ispir to
Ikisdar, 14 July 1981, K. Tan 780 (E) [C. orbicularis var. caucasica paratype]. Rize, Çat, 30 July 1981, K. Tan 1199
(E) [C. orbicularis var. caucasica paratype]. Haldizan Kagh, 27 August 1934, E.K. Balls 1865B (K) [C. orbicularis
var. caucasica paratype].
C. pachystylis
ISRAEL-PALESTINE: Jordan, near Jisr el Majami bridge, 1 February 1942, P.H. Davis 3883 (E, K). Mountain slopes
east of Al Bireh, 21 January 1987, L.J. Musselman (E). Bersheba, 10 March 1922, F.S. Meyer & J.E. Dinsmore 8384
JORDAN: Edom, 19 April 1945, Davis 8876 (E). Khirbet Faria, 29 March 1995, F.N. Hepper 10048 (K). 15 km E of
Azraq lake, 19 April 1963, J.B. Gillet 15726 (K). Track to the Amman road 17 km NW of Azraq Shishan, 19 April
1965, C.C. Towsend 65/133 (K).
C. stenophylla
JORDAN: Mashita, 25 April 1911, F.S. Meyer & J.E. Dinsmore M384 (E). 99 km north of Aqaba, 12 March 1974, L.
Boulos et al. (E). Jarash, 8 January 1974, L. Boulos 5982 (E). Transjordan, 19 April 1945, P.H. Davir 8707 (K).
... The nomenclature used follows Egorova [19] and Koopman [20], except for C. curvata [21], C. hartmaniorum A.Cajander [22], and C. nigra subsp. transcaucasica (T.V.Egorova) Jim.Mejías, G.E.Rodr.-Pal., Amini Rad & Martín-Bravo [23]. The names of sections used follow Egorova [19], Reznicek [24], and Ball & Reznicek [1]. ...
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Counting chromosomes is the first step towards a better understanding of the karyotype evolution and the role of chromosome evolution in species diversification within Carex; however , the chromosome count is not known yet for numerous sedges. In this paper chromosome counts were performed for 23 Carex taxa from Armenia, Austria, the Czech Republic, and Poland. Chromosome numbers were determined for the first time in three species (Carex cilicica, 2n = 54; C. phyllostachys, 2n = 56; C. randalpina, 2n = 78), two subspecies (C. muricata subsp. ashokae, 2n = 58; C. nigra subsp. transcaucasica, 2n = 84) and two hybrids (C. ×decolorans, 2n = 74; C. ×walasii, 2n = 108). Among the taxa whose number of chromosomes had been known before, the largest difference was found in C. hartmaniorum (here 2n = 52) and C. aterrima subsp. medwedewii (here 2n = 52). A difference in the chromosome count was demonstrated for C. cilicica (2n = 54) versus the species of the section Aulocystis (2n = 30 to 40) and for C. tomentosa (2n = 48) versus the species of the section Acrocystis (2n = 18 to 38). The results of this study indicate that the position of C. cilicica in Aulocystis section may raise doubts. Attention was paid to the relationship between C. phyl-lostachys and taxa of the subgenus Carex section Gynobasidae.
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Carex section Phacocystis (Cyperaceae) is one of the most diverse and taxonomically complex groups of sedges (between 116‐147 spp.) with a worldwide distribution in a wide array of biomes. It has a very complicated taxonomic history, with numerous disagreements among different treatments. We studied the biogeography and niche evolution in a phylogenetic framework to unveil the relative contribution of geographical and ecological drivers to diversification of the group. We used a large species sampling of the section (82% of extant species) to build a phylogeny based on four DNA regions, constrained with a phylogenomic HybSeq tree and dated with six fossil calibrations. Our phylogenetic results recovered sect. Phacocystis s.s. (core Phacocystis) as sister to section Praelongae. Ancestral area reconstruction points to the N Pacific as the cradle for the crown diversification of section Phacocystis during the Middle Miocene. Wide distributions were recurrently inferred across deep nodes. Large Northern Hemisphere lineages with geographical congruence were retrieved, pointing to the importance of allopatric divergence at deep phylogenetic levels, whereas within‐area speciation emerges as the predominant pattern at shallow phylogenetic level. The Southern Hemisphere (Neotropics, SW Pacific) was colonized several times from the Northern Hemisphere. The global expansion of Carex section Phacocystis did not entail major ecological changes along the inner branches of the phylogeny. Nevertheless, ecological differentiation seems to gain importance towards recent times. This article is protected by copyright. All rights reserved.
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On the basis of previously published molecular data, and morphological observations, Carex nigra subsp. drukyulensis from the eastern Himalayas (Bhutan) is raised to specific rank as Carex drukyulensis .
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We present the first large-scale phylogenetic hypothesis for the genus Carex based on 996 of the 1983 accepted species (50.23%). We used a supermatrix approach using three DNA regions: ETS, ITS and matK. Every concatenated sequence was derived from a single specimen. The topology of our phylogenetic reconstruction largely agreed with previous studies. We also gained new insights into the early divergence structure of the two largest clades, core Carex and Vignea clades, challenging some previous evolutionary hypotheses about inflorescence structure. Most sections were recovered as non-monophyletic. Homoplasy of characters traditionally selected as relevant for classification, historical misunderstanding of how morphology varies across Carex, and regional rather than global views of Carex diversity seem to be the main reasons for the high levels of polyphyly and paraphyly in the current infrageneric classification.
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Carex (Cyperaceae), with an estimated 2000 species, nearly cosmopolitan distribution and broad range of habitats, is one of the largest angiosperm genera and the largest in the temperate zone. In this article, we provide argument and evidence for a broader circumscription of Carex to add all species currently classified in Cymophyllus (monotypic), Kobresia (c. 60 species), Schoenoxiphium (c. 15 species) and Uncinia (c. 70 species) to those currently classified as Carex. Carex and these genera comprise tribe Cariceae (subfamily Cyperoideae, Cyperaceae) and form a well supported monophyletic group in all molecular phylogenetic studies to date. Carex as defined here in the broad sensecurrently comprises at least four clades. Three are strongly supported (Siderostictae, core Vignea and core Carex), whereas the caricoid clade, which includes all the segregate genera, receives only weak to moderate support. The caricoid clade is most commonly split into two clades, one including a monophyletic Schoenoxiphium and two small clades of species of Carex s.s., and the other comprising Kobresia, Uncinia and mostly unispicate species of Carex s.s. Morphological variation is high in all but the Vignea clade, making it extremely difficult to define consistent synapomorphies for most clades. However, Carex s.l. as newly circumscribed here is clearly differentiated from the sister groups in tribe Scirpeae by the transition from bisexual flowers with a bristle perianth in the sister group to unisexual flowers without a perianth in Carex. The naked female flowers of Carex s.l. are at least partially enclosed in a flask-shaped prophyll, termed a perigynium. Carex s.s. is not only by far the largest genus in the group, but also the earliest published name. As a result, only 72 new combinations and 58 replacement names are required to treat all of tribe Cariceae as a single genus Carex. We present the required transfers here, with synonymy, and we argue that this broader monophyletic circumscription of Carex reflects the close evolutionary relationships in the group and serves the goal of nomenclatural stability better than other possible treatments.
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Reseda minoica Martín-Bravo & Jim. Mejías (Resedaceae), a new species from the eastern Mediterranean region, is described and illustrated. It is distributed in Crete (Gavdos Island), Cyprus and S Anatolia (Mersin), where it grows mostly on basic, occasionally schistose, substrates near the coast. It is included in Reseda sect. Phy-teuma, a taxonomically complex group mostly containing narrow endemics from the western or eastern Mediterranean region. Reseda minoica has been confused with R. odorata, R. orientalis and R. balansae in Crete, Cyprus and Turkey. It can be distin-guished from those by the lower number of stamens, seed size, colour of petals and indumentum. An identification key to the eastern Mediterranean taxa of Reseda sect. Phyteuma is provided.
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A new genus (Pseudodelphinium H.Duman, Vural, Aytaç & Adıgüzel) and 3 new species (Pseudodelphinium turcicum H.Duman, Vural, Aytaç & Adıgüzel; Iberis halophila Vural & H.Duman; and Frankenia salsuginea Adıgüzel & Aytaç) from the Tuz Gölü (Salt Lake) basin (Central Anatolia) are described. Illustrations, ecological habit, and a distribution map are given. Th e relationships of the new genus and species are discussed in terms of related taxa.