Pl Syst Evol 267: 13–23 (2007)
DOI 10.1007/s00606-007-0576-4
Printed in The Netherlands
The genus Rhaponticoides Vaill. (Asteraceae) in Turkey:
a new species and first key
Ö. Eren
Department of Biology, Adnan Menderes University Aydın, Aydın, Turkey
Received 17 February 2006; Accepted 9 May 2007; Published online 8 August 2007
 Springer-Verlag 2007
Abstract. Turkish representatives of the genus
Rhaponticoides Vaill. are examined. A key and
distribution map of Rhaponticoides species occur-
ring in Turkey are provided for the first time. R.
hierroi Ö. Eren sp. nov., from Bakacak pass in the
Western Taurus range, SW Anatolia, allied to R.
mykalea (Hub.-Mor.) M. V. Agab. & Greuter is
described as a species new to science and illus-
trated. Its habitat conditions and conservation
status are considered. Pollen micrographs of R.
hierroi and R. mykalea are presented using scan-
ning electron microscopy.
Keywords: Asteraceae; Centaureinae; Rhaponticoi-
des; Rhaponticoides hierroi; Centaurea; Turkey; phy-
togeography; flora; SEM
During the first revision of Centaurea L. made
by Wagenitz for the Flora of Turkey and East
Aegean Island, C. amasiensis Bornm. was
classified under the sect. Centaurea L. as single
Turkish representative of a well defined group
based on the following morphological features:
mostly pinnatipartite leaves with serrate seg-
ments, nearly glabrous stem, branched above
Correspondence: Özkan Eren, Department of Biology, Faculty of Sciences and Arts, Adnan Menderes University, 09010,
Aydın, Turkey
e-mail: ozkaneren@adu.edu.tr
Plant Systematics
and Evolution
with a few large capitula, ovoid involucre,
coriaceous, nearly exappendiculate and gla-
brous involucral bracts with several dark
longitudinal nerves near an obtuse apex (Wage-
nitz 1975). The pollen of the ‘‘Centaurium’’ type
(Wagenitz 1955), a unique basic base chromo-
some number of x ¼ 15 (Agababian and Gouk-
asian 1994) and a double pappus, but no
bolster cells at the apex of achene (Wagenitz
and Hellwig 1996) are the other characteristics
of sect. Centaurea. Later, C. mykalea Hub.-
Mor. and C. iconiensis Hub.-Mor. were de-
scribed, and C. amplifolia Boiss. & Heldr. was
also added to the Flora of Turkey, accounting
as additional three members of the type section
(Huber-Morath 1979, 1980, 1981; Davis et al.
1988; Nydegger-Hügli 1994; Kaynak and
Tarımcılar 2001). C. pythiae Azn. & Bornm.
(Bornmüller 1927) was included as a synonymy
of C. amasiensis by Wagenitz (1975), in the
Flora of Turkey. This statement was not
supported by subsequent authors who traced
C. pythiae as a distinct species (Agababian
1997, Greuter 2003).
During the last years, many comparative
studies have been performed on the Asteraceae
14 Ö. Eren: The genus Rhaponticoides Vaill. (Asteraceae) in Turkey
systematic using morphological, anatomical
and molecular characters (for details, see the
survey papers by Dittrich 1966, 1968a, b, 1970;
Susanna et al. 1995, 1999, 2006; Wagenitz and
Hellwig 1996; Petit 1997; Vilatersana et al.
2000; Petit et al. 2001; Greuter et al. 2001;
Häffner 2000; Garcia-Jacas et al. 2000, 2001,
2002, 2006; Martins and Hellwig 2005a, 2005b;
Greuter et al. 2005; Hidalgo et al. 2006;
Wagenitz et al. 2006 and literature cited there).
Delimitation of taxa within the family has
always been controversial; consequently, cir-
cumscription of genera and tribes has repeat-
edly changed over time.
The distribution of Centaurea sect. Centau-
rea (including C. hajastana Tzvel., C. tamani-
anae Agababian, C. ruthenica Lam. and C.
rasdorsky Karyagin) has been studied in the
Caucasus by Agababian and Fajvush in 1991.
In particular, Agababian (1997) studied the
delimitation and distribution of sections and
subsections of Centaurea subgenus on the basis
of morphological criteria. In Agababian’s
studies, Centaurea subg. Centaurea was subdi-
vided into three sections and seven subsections
including two new sections (sect. Vicentina M.
V. Agab., sect. Africana M. V. Agab) and four
new subsections (subsect. Ruthenicae M. V.
Agab., subsect. Aralocaspicae M. V. Agab.,
subsect. Turkestanicae M. V. Agab., subsect.
Iranicae M. V. Agab.).
The Euro+Med genera of Cardueae with
their relevant synonyms have recently been
published (Greuter 2003). In this work the
previous Centaurea sect. Centaurea has been
segregated from the Centaurea genus and
named as Rhaponticoides Vaill. Those taxo-
nomic and nomenclatural changes follow the
recommendations of Garcia-Jacas et al. (2000,
2001), based on the results of molecular
phylogenies. This statement was supported by
Hellwig, who gave detailed information on the
ecogeographical history of Cardueae-Centau-
reinae (Asteraceae) in the Mediterranean
region (Hellwig 2004). Furthermore, he
reported, that since Centaurea s.1. is not a
monophyletic taxon, several groups will be
segregated as separate genera. The name
Rhaponticoides was used by Vaillant for a
large and artificial genus of 29 or 30 species,
characterized by blunt, non-pungent phyllaries
of homogeneous consistency (Greuter et al.
2005). This name was not adopted by subse-
quent authors until Greuter (2003).
The genus Rhaponticoides Vaill. (Compos-
itae: Carduae-Centaureinae) comprises 32 spe-
cies occurring from Portugal and Morocco in
the West to Mongolia in the East (Hellwig
2004). Most of them are either narrow endem-
ics or have very disjunct areas (Wagenitz
1986). There are only a few numbers of species
such as R. ruthenica in a large continuous area.
In July 2003, while conducting field-work
with plant ecologist José Luis Hierro as part of
NSF project entitled ‘‘Mechanisms for suc-
cessful invasions: Yellow Starthistle (Centau-
rea solstitialis) in California, Central
Argentina and Turkey’’, the author found an
uncommon, very striking Rhaponticoides
around the Bakacak pass in the province of
Antalya. In the course of identifying some
critical taxa, the Rhaponticoides of the
Bakacak pass population came close to R.
mykalea, but it was also clearly different from
R. mykalea in a number of important features.
More material was gathered from the same
locality in 2004 and 2005. Detailed studies on
this plant and the related species have been
made during the stay of author in Berlin. Here
this plant is described as a new species,
bringing a total number of Rhaponticoides
species known from Turkey to six.
Materials and methods
All studies were based on the plant material
collected by author and the herbarium specimens
of Rhaponticoides kept at the following herbariums:
B, BULU, E, and GAZI. Pollen samples of
R. hierroi and R. mykalea were obtained from
author’s collections, which are numbered Eren 186/
03 and Eren 366/04 respectively. Pollen grains were
prepared for scanning electron microscopy (SEM)
according to the ‘‘Micro-Acetolyse’’ method
modified by Erdtman (1966). The pollen grains
were suspended in a drop of water, directly
transferred with a fine pipette to a metallic stub
Ö. Eren: The genus Rhaponticoides Vaill. (Asteraceae) in Turkey
using double-sided cellotape, and coated with
15 nm gold/palladium in a Low Voltage Cool
Sputter Coater (EMITECH K 550). The SEM
examination was carried out with a Philips SEM
515.
Results
Representatives of the genus Rhaponticoides
in Turkey.
1. Rhaponticoides hierroi Ö. Eren sp. nova
Holotype: Turkey, C3 Antalya, between
Antalya and Saklıkent road, below Bakacak
pass, Boyunduruk battı, 1100 m, rocky slope,
open Pinus brutia Ten. forest, limestone,
17.7.2004, Eren 252/04 & Şirin (AYDN; Iso-
types: B, E, GAZI, HUB). – Figs. 1, 2.
Diagnosis. Ab affini R. mykalea differt
caule sulcato, involucris late umbonato-ovoi-
deis vel subglobosis, parte proximali bractea-
rum pallide virentium appressa, cartilaginea
albescente sed distali subpatula, cochlearifor-
mi-concava, flosculis cum tubo antherarum
pallide flavis.
Description. Perennial herb to 210 cm tall,
arising from a woody rootstock. Stem single or
a few, erect, sulcate, glabrous, c. 7 mm diam. at
base, divided above into few 1-headed
branches. Leaves firm, most of them in a basal
rosette, blade glabrous, margins and nerves
shortly scabridulous. Basal leaves either
undivided, elliptic-ovate, petiolate, 28–32 5–
6 cm (incl. petiole), petiole up to 12 cm long or
mostly bipinnatisect, petiolate, 35–60  14–
20 cm (incl. petiole) with 5–7 pairs of primary
segments; all segments ending in a distinct
mucro, secondary segments linear-lanceolate,
6–11  2:5–4 cm; pinnatifid, terminal segment
lanceolate 7:5–10  5–7 cm, pinnatisect or
pinnatilobed. Lower and middle cauline leaves
similar as the bipinnatisect basal ones but
smaller. Upper stem leaves pinnatisect, sessile,
smaller toward the apex of stem, segments
entire, apex acute, with a distinct mucro.
Involucre glabrous, broadly ovoid to almost
spherical 4–5 3:5–5 cm, umbonate at base.
Phyllaries multiseriate, coriaceous, pale green,
15
proximally white and cartilaginous, with
narrow (1–1.5 mm long) membranous margin;
margin entire, flat to undulate, becoming
brownish, all phyllaries conspicuously longitu-
dinally striate marked with blackish-green
lines, outer ones spoon shaped, cucullate, 10–
25 8–18 mm, inner oblong or linear-lanceo-
late, cucullate, 25–40  4–12 mm. Florets pale
yellow, c. 5 cm, distinctly 5 veined, marginal
ones scarcely radiant, sterile, ligules filiform up
to 10 mm long, anther tube pale yellow,
subexserted to exserted. Achenes cylindrical,
9:5–11  4–5 mm, brown, yellowish and glossy
in basal parts, laterally compressed, trans-
versely wrinkled. Pappus double, multiseriate,
scabrous, dirty white becoming brownish,
innermost bristles of outer pappus up to
16 mm, inner pappus uniseriately arranged
2–3 mm.
Etymology. In honour of the Argentinean
Plant Ecologist José Luis Hierro at the
University of Montana, United States, with
whom this interesting new species was first
collected in 2003.
Additional specimens (Paratypes). Type
locality, 17.7.2003, Eren 186/03 & Hierro
(AYDN, B); ibid 05.8.2003, Eren 252/03 &
Şirin (fruting; AYDN); ibid., 30.7.2005, Eren
213/05 & Çinbilgel (AYDN, EGE).
Phenology. Rhaponticoides hierroi flowers
in late June to early August, when the first
fruiting specimens were collected, and sets
fruits until October.
Distribution and site conditions. As a very
local endemic of the Western Taurus, R. hierroi
is hitherto only known from type locality and
has a surprisingly narrow altitudinal range
(1100–1150 m). It is a distinct East Mediterra-
nean mountain plant that grows on limestone
in open Pinus brutia forest (Fig. 3). Associated
species include Quercus coccifera L., Vicia
villosa Roth subsp. eriocarpa (Hausskn.)
P. W. Ball, Phlomis grandiflora H.S. Thomp-
son, Opopanax hispidus (Friv.) Griseb., Turge-
nia latifolia (L.) Hoffm., Picnomon acarna (L.)
Cass., Carduus nutans L., Coronilla varia L.
subsp. varia, Colutea melanocalyx Boiss. &
Heldr. subsp. melanocalyx, Dianthus caloceph-
16 Ö. Eren: The genus Rhaponticoides Vaill. (Asteraceae) in Turkey

Fig. 1. R. hierroi in its natural habitat, photograph by Özkan Eren

alus Boiss., Melilotus officinalis (L.) Lam., Rosa
canina L., Salvia tomentosa Mill., Styrax
officinalis L., Dactylis glomerata L. subsp.
hispanica (Roth) Nymán, Valeriana dioscoridis
Sm., Echinops ritro L., Centaurea solstitialis L.
subsp. solstitialis, Anthemis kotschyana Boiss.
var. kotschyana, Silene gigantea L. var. gigan-
tea, Onobrychis caput-galli (L.). Lam., Cistus
creticus L. and Gladiolus anatolicus (Boiss.)
Stapf . Most likely R. hierroi is a ruderal plant
growing in forest clearings. It is probably
unable to tolerate shade and low soil pH since
Ö. Eren: The genus Rhaponticoides Vaill. (Asteraceae) in Turkey 17

Fig. 2. R. hierroi. 1 flowering capitula; 2 achene and pappus; 3 outer sterile floret and inner fertile one; 4 series
of involucral bracts from outer to inner; 5 undivided basal leaf; 6 bipinnatisect basal leaf

it is rarely found under the P. brutia. The
vegetation formed by R. hierroi and its associ-
ates has not yet been classified phytosociolog-
ically, but stand can be grouped into the Cisto-
Micromerietea Oberd. 1954 class.
Recommended IUCN threat category. It is
rather remarkable that R. hierroi has man-
aged to survive in such a localised area. At
present the rapid development of the area
threats its habitat. The area is easily accessi-
ble by road, subjected to heavy grazing
pressure, and limits with arable fields and
settlement houses; all of which imply a high
risk of extinction. Moreover, because of its
18 Ö. Eren: The genus Rhaponticoides Vaill. (Asteraceae) in Turkey
Fig. 3. Distribution map of Rhaponticoides species in Turkey. : R. hierroi; : R. amplifolia; j: R. mykalea; d:
R. amasiensis; m: R. pythiae; : R. iconiensis, ´: Unknown
striking appearance, it is collected at flower-
ing time by local people for companies that
sell it as a souvenir. Hence, I strongly
recommend placing R. hierroi under the
IUCN category ‘‘Critically Endangered
(CR)’’ (IUCN 2001), because the estimated
area of occupancy is less than 10 km2 and it
is only known from a single locality. Accord-
ing to field observations, it is estimated that
the total number of individuals of R. hierroi
does not exceed 80 in its single locality. This
very small number does not secure the
unhindered survival of the species for a long
time. Fruits of this stenoendemic plant should
be collected and stored in a seed bank.
2. Rhaponticoides (subsect. Centaurea) ampli-
folia (Boiss. & Heldr.) M. V. Agab. &
Greuter ” Centaurea amplifolia Boiss. &
Heldr. in Boiss., Diagn. Ser 2(3): 68. 1856.
Specimens examined: A2 Istanbul, Aydos
hill, on the slopes, 02.07.1974, Tuzlacı 30115
(E).
3. R. (subsect. Ruthenicae M. V. Agab.) am-
asiensis (Bornm.) M. V. Agab. & Greuter
” Centaurea amasiensis Bornm. in Feddes
Repert. 3: 54. 1906.
Specimens examined: A5/A6 Amasya, in
montis subalpinis, Abadschi-dagh, 800–900 m,
3.7.1889, Bornmüller 1100 (Holotype, B); A5
Amasya, Merzifon, 1904, Manissadjiani s.n.
(E); B3 Eskisehir, Sündiken Dağ, Bozdağ
Deresi, ca. 1100 m, 21.06.1973, Ekim 456 (E).
4. R. (subsect. Ruthenicae M. V. Agab.) pythiae
(Azn. & Bornm.) M. V. Agab. & Greuter ”
Centaurea pythiae Azn. & Bornm. in Mitt.
Thür. Bot. Ver. N. F. 37: 48. 1927.
Specimens examined: A2 Istanbul, near
‘‘Thermas Pythiae’’ not far from Yalova,
30.6.1905, Aznavour s.n. (Holotype, B); A2
Istanbul, Yalova, 12 km südl., od ber Heilbä-
der, 120 m, 28.6.1954, H. Demiriz 2301 (B).
5. R. mykalea (Hub.-Mor.) M. V. Agab. &
Greuter ” Centaurea mykalea Hub.-Mor.
in Bauhinia 6: 370. 1979.
Specimens examined: C1 Aydın, 7 km
nördlich von Davutlar, zwischen Selçuk und
Davutlar, 30 m, 27.7.1993, Nydegger 47546
(Topotype, B); C1 Aydın, between Kuşadası
and Davutlar, roadside, 35 m, 14.05.2005,
Eren 89/5, Erdağ & Kırmacı (sterile, AYDN);
ibid., 02.07.2005, Eren 163/5, Erdağ & Kırmacı
(AYDN); B3 Isparta, Uluborlu, Dam, slopes,
open forest clearing, 1200 m, 30.7.1998, G.
Kaynak 10221B (BULU, GAZI); ibid.,
03.08.2004, Eren 366/04 & Şirin (AYDN, B,
herb. Parolly).
6. R. (subsect. Ruthenicae M. V. Agab.) iconi-
ensis (Hub.-Mor.) M. V. Agab. & Greuter
Ö. Eren: The genus Rhaponticoides Vaill. (Asteraceae) in Turkey
” Centaurea iconiensis Hub.-Mor. in Bau-
hinia 7: 77.1981.
Specimens examined: C4 Konya, 22 km
östlich Seydişehir, zwischen Seydişehir und
Bozkır, Kanalböschung, 1050 m, 30.7.1992,
Nydegger 46895 (Topotype, B).
The necessary detailed key to the Rhapon-
ticoides species occur in Flora of Turkey area is
proposed below. The characters given in the
key mainly consider the herbarium material
studied as well as the descriptions in the
references cited above.
1. Florets 2 cm long, pink, phyllaries with
scarious erose (ca. 5 mm long) appendage,
outer pappus <8:5 mm long ...……
…………….……...............……R. amplifolia
1¢. Florets 3 cm long, yellow, phyllaries
without appendage only with very narrow
(0.3–1.5 mm long) scarious margin, outer
pappus 10 mm..............................................2
2. Stem 80 cm, basal leaves bipinnatipartite,
ovate-oblong, 16–20 cm long, segments lin-
ear 0.5–1.5 mm broad ……….R. iconiensis
2¢. Stem 90 cm, basal leaves pinnatipartiate,
pinnatifid or bipinnatisect 20 cm long, seg-
ments linear-lanceolate or lanceolate broad-
er than 5 mm..............................................3
3. Basal leaves pinnatipartite, lateral segments
sharply dentate, involucre 3.5 cm
long............................................................4
3¢. Basal leaves mostly bipinnatisect, lateral
segments laxly serrate-dentate, involucre
4 cm long
4. Involucre small 2.5 long, stem foliated
on upper part, leaves with elongated
terminal segments to 5–6 cm long
........................................……R. amasiensis
4¢. Involucre larger 3 cm long, leaves con-
centrated on lower part of stem, upper part
of stem naked, terminal leaf segments not
elongated and smaller in size
................................................... R. pythiae
5. Outer involucral bracts broadly ovate,
obtuse, margin entire, florets gold,
anther tube bright sulphur
........................................……....R. mykalea
19
5¢. Outer involucral bracts spoon-shaped,
cucullate pale green proximally white and
cartilaginous, with narrow membranous,
entire, flat to undulate margin, becoming
brownish, florets and anther tube both pale
yellow…….…...............................R. hierroi
Discussion
The genus Rhaponticoides represents an inter-
esting topic for taxonomic studies because of
its geographical distribution, particular pollen
of Centaurium type, and recent reconsidera-
tion of its placement within the Centaureinae
from the derived group of the basal one.
Agababian’s important studies (1997) on the
subgenus Centaurea, other studies dealing with
the morphology, anatomy and molecular tax-
onomy of the Centaureinae group and recog-
nition of Vaillant’s names paved the way for
the later work of Greuter (2003), who draw
new taxonomic conclusions and split the
polyphyletic Centaurea s.1. into more natural
genera corresponding to the sections of Cen-
taurea. The main morphological and palyno-
logical characters (e.g. involucral bracts with
several dark blackish-green nerves near apex,
multiseriate pappus, transversely wrinkled ach-
enes and Centaurium type pollen) place the
studied plant clearly into Rhaponticoides Vaill.
General appearance and combination of char-
acters make its taxonomic position close to R.
mykalea (Hub.-Mor.) M. V. Agab. & Greuter.
Rhaponticoides hierroi shares with R. mykalea
bipinnatisect leave shape, tall habit, large
capitula and long florets. In addition the
pollen of R. mykalea is an excellent match of
R. hierroi (Fig. 4). Both have tricolporate
pollen type, surface pattern is echinate-micro-
perforate and colpus surface granulate. How-
ever, R. hierroi is easily distinguished from R.
mykalea by a number of highly distinctive
features, especially in the regard to those of
involucral and floral characters. The main
differential characters used to distinguish tech-
nically these two species are compiled in
Table 1.
20 Ö. Eren: The genus Rhaponticoides Vaill. (Asteraceae) in Turkey

Fig. 4. SEM micrographs of pollens (A-E) R. hierroi and (F-J) R. mykalea (see Wagenitz 1955)
Ö. Eren: The genus Rhaponticoides Vaill. (Asteraceae) in Turkey
Table 1. Differential characters of R. hierroi and R. mykalea
Character R. mykalea
Stem longitudinally striate
Involucre broadly ovate
Involucral bracts bright green, subcoriaceous, outer
phyllaries ovate, obtuse, margin
entire
Florets gold
Anther tube bright sulphur, subexserted
Inner pappus 1–1.5 mm
R. hierroi is clearly distinguished from
R. mykalea by the sulcate stem, broadly ovoid
to almost spherical involucre, florets and
anther tube both pale yellow, inner pappus
2–3 mm long, and finally, most remarkable
and unique within the genus, the pale green,
coriaceous, proximally white and cartilaginous
cucullate and spoon-shaped involucral bracts,
which become brownish marginally.
Although the spoon-shaped phyllaries in
R. hierroi recall the genus Rhaponticum Hill,
they are quite different if a special attention is
payed to their bracts and appendage. Rhapont-
icum comprises Centaureinae species in which
the phyllaries end in a spoon-shaped, well
developed entire or tardily lacerate, scarious
appendage whereas in Rhaponticoides hierroi
the bract itself forms the spoon shape and the
appendage consists in a narrow membranous
margin. This unique character within Rhapon-
ticoides might be derived or arose relatively
late, in which case it can be considered as
apomorphic. The latter character probably
facilitates dispersal by ants. The presence of
elaisomes has been well known in Rhapontico-
ides for a long time (Wagenitz and Hellwig
1996, Hellwig 2004).
The unique combination of characters
makes R. hierroi a distinct and easily identifi-
able species. Within sect. Centaurea, following
the subsectional delimitation of Agababian
(1997), the narrow membranous margin of the
phyllaries of R. hierroi accounts for its place-
ment within the subsect. Ruthenicae. This
21
R. hierroi
sulcate
broadly ovoid to almost spherical
pale green, coriaceous, proximally white
and cartilaginous, all phyllaries cucullate
and spoon shaped, margin entire, flat to
undulate, becoming brownish
pale yellow
pale yellow, subexserted to exserted
2–3 mm
result is consistent with the fact that the more
closely related species to R. hierroi, namely
R. mykalea, belong to this subsection too
(Agababian 1997).
It is reported by Agababian (1997) that the
Mediterranean representatives of this group
are relatively few and taxonomically well
defined and little variable, whereas the non-
Mediterranean group includes a great number
of either polymorphic or very local species
often connected by intermediate forms. It is
also believed by Agababian (1997) that the
western species are more ancient than eastern
taxa. While western representatives grow in
oak or coniferous forests the eastern taxa grow
in steppe-like vegetation. New finding of R. hi-
erroi from the pine forest clearings supports
this view.
I have not seen the imperfect specimen of
Birand & Karamanoğlu included by Wage-
nitz (1975) under R. amasiensis, which has
some clear differences when compared to R.
amasiensis such as mostly smaller parts,
conspicuously glaucous leaves with narrow
segments. This specimen may well belong to
one of the Transcaucasian species or more
likely may represent another undescribed
taxa.
I am highly indebted to Prof. Dr. W. Greuter for
translation of English diagnosis to Latin and for
providing working facilities at BGBM, particularly
for SEM studies. I would like to thank M. Lüchow
for SEM analysis. I wish to thank Prof. Dr. H.
Duman and Prof. Dr. G. Wagenitz for kindly
22 Ö. Eren: The genus Rhaponticoides Vaill. (Asteraceae) in Turkey
checking my specimen and confirming that this
species belongs to the undescribed taxa. I am very
grateful to Dr. M. Agababian and Dr. T. Raus for
useful discussion. My sincere thanks go to Dr. A.
Erdağ who prepared Fig. 2 and directors and
keepers of the herbaria (B, BULU, E, and GAZI)
for loan materials. Dr. José L. Hierro improved the
English of this manuscript.
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