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Taraxacum sect. Palustria ( Compositae, Cichorieae ) in Bulgaria revised, with three new species

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Three new species of Taraxacum sect. Palustria (Compositae, Cichorieae) are described from Bulgaria, viz. T. abruptilobum, T. basilicum and T. rumelicum. The new species, on the basis of detailed, standardized descriptions and illustrations, are compared with similar species of this section. All the seventeen species of T. sect. Palustria in Bulgaria are agamospermous. An identification key to the members of T. sect. Palustria in Bulgaria is given. © 2017, Botanischer Garten und Botanisches Museum Berlin-Dahlem. All rights reserved.
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Taraxacum sect. Palustria (Compositae, Cichorieae) in Bulgaria revised, with
three new species
Author(s): Jan Štěpánek & Jan Kirschner
Source: Willdenowia, 47(2):155-165.
Published By: Botanic Garden and Botanical Museum Berlin (BGBM)
https://doi.org/10.3372/wi.47.47207
URL: http://www.bioone.org/doi/full/10.3372/wi.47.47207
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Willdenowia
Annals of the Botanic Garden and Botanical Museum Berlin-Dahlem
JAN ŠTĚPÁNEK1 & JAN KIRSCHNER1*
Taraxacum sect. Palustria (Compositae, Cichorieae) in Bulgaria revised, with three new
species
Version of record first published online on 13 July 2017 ahead of inclusion in A ugust 2017 issue.
Abstract: Three new species of Taraxacum sect. Palustria (Compositae, Cichorieae) are described from Bulgaria,
viz. T. abruptilobum, T. basilicum and T. rumelicum. The new species, on the basis of detailed, standardized descrip-
tions and illustrations, are compared with similar species of this section. All the seventeen species of T. sect. Palustria
in Bulgaria are agamospermous. An identification key to the members of T. sect. Palustria in Bulgaria is given.
Key words: Mediterranean, Balkan Peninsula, Bulgaria, Asteraceae, Compositae, Cichorieae, Taraxacum, Taraxacum
sect. Palustria, taxonomy, new species
Article history: Received 22 December 2016; peer-review completed 28 March 2017; received in revised form
9June 2017; accepted for publication 14 June 2017.
Citation: Štěpánek J. & Kirschner J. 2017: Taraxacum sect. Palustria (Compositae, Cichorieae) in Bulgaria revised,
with three new species. – Willdenowia 47: 155 165. doi: https://doi.org/10.3372/wi.47.47207
Introduction
The Taraxacum flora of Bulgaria and adjacent regions has
been dealt with by the present authors since the late 1980s
(Kirschner & Štěpánek 1993a, 1999b, 1999a; Štěpánek &
Kirschner 2014; Štěpánek & al. 2011; Zeisek & al. 2015),
with an ultimate goal of a critical, reasonably complete
treatment of Taraxacum F. H. Wigg. in the series of Flora
of Bulgaria. In particular, we focused our study on marsh
dandelions, T. sect. Palustria (H. Lindb.) Dahlst. Our first
expeditions to Bulgaria revealed eight species of this sec-
tion (four newly described, Kirschner & Štěpánek 1993a).
Later on, we monographed T. sect. Palustria (Kirschner &
Štěpánek 1998) and revealed another six new species in
the Bulgarian material. The present contribution completes
our knowledge of T. sect. Palustria in Bulgaria on the ba-
sis of additional and/or newly cultivated material. Another
three new species are described so that 17 taxa referred to
T. sect. Palustria are now known from Bulgaria, all with
agamospermous reproduction (see Material and methods).
Any taxonomic treatment of Taraxacum comparable
to modern standards should follow several principles.
They were summarized by Richards (1973), Kirschner
& Štěpánek (1996) and Ge & al. (2011), and reflect the
peculiar features and processes known in dandelions, par-
ticularly the coexistence of agamospermy and sexuality,
complex hybridity and polyploidy, the low level of struc-
tural morphological dierentiation and the high number of
mutually similar and mostly hybridogenous species. The
principles derived from the above features include (1) dif-
ferent kinds of species are to be recognized on the basis of
the extent of variation and modes of reproduction, (2) dis-
tribution of sexuality is to be explored, (3) variation within
a family of siblings should be studied for each taxon (to
1 Institute of Botany, Academy of Sciences, Zámek 1, CZ-25243 Průhonice, Czech Republic; *e-mail: jan.kirschner@ibot.cas.cz
(author for correspondence); jan.stepanek@ibot.cas.cz
156 Štěpánek & Kirschner: Taraxacum sect. Palustria in Bulgaria revised
detect autonomous aberrants or facultative sexuality), and
(4) the study should be started at the lowest variation level
(within and among populations). We attempted to follow
the above principles in the present paper.
Material and methods
The material for the present study comes from our expedi-
tions to Bulgaria and from achene samples of other col-
lectors. We would like to acknowledge the enormous help
of the late Bogdan Kuzmanov during our field and her-
barium studies. The material is preserved in the herbarium
PRA, of the Institute of Botany, Academy of Sciences,
Průhonice, Czech Republic. It represents one of the larg-
est Taraxacum collections in the world; duplicates will
be deposited in SOM (see Index Herbariorum at http://
sweetgum.nybg.org/science/ih/). Most of the material was
also cultivated at the experimental garden of the Institute
of Botany, Academy of Sciences, Průhonice, Czech Re-
public, following the methods published in Kirschner &
Štěpánek (1993b).
All the plants included in the present study have aga-
mospermous reproduction. The mode of reproduction
was studied in cultivation using methods described in
Kirschner & al. (2006).
As a rule, more than one species grow side by side,
up to five species according to our observations. It is
therefore important that plants are collected and/or other-
wise documented taking this diversity into account. Only
well-developed plants are suitable for identification. Ide-
ally, plants in full flower should be evaluated, and their
achenes obtained in cultivation. Alternatively, the oldest
capitulum in a plant may be collected with its scape and
kept in water in a small vessel for up to ten days to get
ripe achenes.
Results and Discussion
Taraxacum sect. Palustria in Bulgaria
Standard descriptions were published for all the Bulgar-
ian species of Taraxacum sect. Palustria (Kirschner &
Štěpánek 1993a, 1998), with the exception of T. abrupti-
lobum, T. basilicum and T. rumelicum, the three species
that are described here as new. We therefore provide a
dichotomous identification key for all species, and full
descriptions are published for the new species only.
The character diversity of the Bulgarian members of
Taraxacum sect. Palustria is relatively low, and some
important character states are missing from that territory
(e.g., yellow stigmas, absence of pollen, outer phyllar-
ies all patent). They belong to five species groups only,
and T. sect. Palustria exhibits a relatively low diversity
near its southern limit. The majority of species, ac-
cording to the monograph of the section (Kirschner &
Štěpánek 1998), belong to the groups centred in Greece
and Bulgaria, only rarely extending to other regions: the
T. subalpinum group (seven species) and the T. apicula-
toides group (four species). The T. scaturiginosum group
(represented by two species in Bulgaria) is widespread
in the E Mediterranean and the Black Sea region, reach-
ing Iran in the east. Taraxacum scaturiginosum is a taxon
with one of the largest distribution ranges in this section;
it extends from Albania in the west through the Balkan
Pen insula, Crimea and Turkey to Armenia, Iraq and Iran
in the east. There is another group with higher diversity
in the central Mediterranean and extending to the Alps
and farther north to Central Europe, the T. tenuifolium
group, with two species in Bulgaria. The only species
with a large geographical range reaching its southern
limit in Bulgaria is the variable and widespread (Battjes
& al. 1992) T. vindobonense s.lat. belonging to the group
bearing the same name and diverse in the Pannonian re-
gion and the narrower Central Europe.
The members of Taraxacum sect. Palustria in Bul-
garia occupy habitats typical of the section: meadows in
the vicinity of springs, wet pastures, subhalophilous sites
and flood-meadows. As regards the possible gaps in the
knowledge of this section in Bulgaria, we should men-
tion wet meadows along the Danube River and the lower
streams of its tributaries in the Danube Lowlands. This
lowland region was not properly explored, and some taxa
not listed below might occur there.
Bulgarian representatives placed in species groups
Taraxacum subalpinum group
T. abruptilobum Kirschner & Štěpánek, sp. nov.
T. ambitiosum Kirschner & Štěpánek
T. basilicum Kirschner & Štěpánek, sp. nov.
T. melancholicum Kirschner & Štěpánek
T. obuncum Kirschner & Štěpánek
T. strictum Kirschner & Štěpánek
T. suspectum Kirschner & Štěpánek
Taraxacum apiculatoides group
T. apiculatoides Małecka
T. lentum Kirschner & Štěpánek
T. sophiae Kirschner & Štěpánek
T. turfosiforme Kirschner & Štěpánek
Taraxacum scaturiginosum group
T. scaturiginosum G. E. Haglund
T. subudum Kirschner & Štěpánek
Taraxacum vindobonense group
T. rumelicum Kirschner & Štěpánek, sp. nov.
T. vindobonense Soest, s.lat.
Taraxacum tenuifolium group
T. glaucolivaceum Kirschner & Štěpánek
T. refectum Sonck
157Willdenowia 47 – 2017
Key to species of Taraxacum sect. Palustria in Bulgaria
1. Outer phyllaries with abaxial surface ± evenly dark
coloured (dark green, blackish green, deep olivaceous
green), with a very narrow (usually 0.1 0.3 mm
wide) whitish or membranous border . . . . . . . . . . . 2
Outer phyllaries with abaxial surface broadly (at
least 0.5mm wide) bordered paler green, light green-
ish or whitish (usually also with a narrow membra-
nous margin) ............................ 10
2. Scapes glabrous or initially with a few araneous hairs
and later glabrescent . . . . . . . . . . . . . . . . . . . . . . . . 3
Scapes densely or sparsely araneous with persistent
hairs .................................... 4
3. Outer phyllariesbroadly ovate to ovate (3.5 5 mm
wide); achenes 3.6 4.3 mm long; beak 8 11 mm
long ......................................
. . T. abruptilobum Kirschner & Štěpánek, sp. nov.
Outer phyllaries lanceolate to ovate-lanceolate
(2.8 3.7 mm wide); achenes 4.2 5 mm long; beak
7 8mm long . . . T. strictum Kirschner & Štěpánek
4. Achenes sparsely spinulose above; beak 10 12mm
long . . . . . . . . T. ambitiosum Kirschner & Štěpánek
Achenes densely spinulose above; beak 7 9( 10)mm
long ..................................... 5
5. Outer phyllaries 8 15 . . . . . . . . . . . . . . . . . . . . . . . 6
Outer phyllaries 15 22 . . . . . . . . . . . . . . . . . . . . . . 8
6. Achenes 5 5.6mm long; cone 1.1 1.5mm long . .
...............T. lentum Kirschner & Štěpánek
Achenes 3.7 4.3mm long; cone 0.7 1mm long . . 7
7. Outer phyllaries loosely appressed to erect, imbri-
cate; beak 7.5 9mm long . . . . . . . . . . . . . . . . . . . .
............. T. obuncum Kirschner & Štěpánek
Outer phyllaries tightly to loosely appressed, not im-
bricate; beak 8.5 10mm long . . . . . . . . . . . . . . . . .
............ T. suspectum Kirschner & Štěpánek
8. Outer phyllaries loosely appressed to ± patent,
8 11mm long ..............................
. . . . . T. basilicum Kirschner & Štěpánek, sp. nov.
Outer phyllaries tightly appressed, 5.2 8mm long . .
........................................ 9
9. Achenes 3.7 4 mm long, body abruptly narrowing
into cone 0.8 1mm long; outer phyllaries ± imbri-
cate . . . . .T. melancholicum Kirschner & Štěpánek
Achenes 4.5 4.9mm long, body ± gradually narrow-
ing into cone 1.1 1.2mm long; outer phyllaries not
imbricate . . . . . . T. subudum Kirschner & Štěpánek
10. Pappus c.8.5mm long . . . . . . . . . . . . . . . . . . . . . . .
. . . . T. rumelicum Kirschner & Štěpánek, sp. nov.
Pappus 5 7mm long . . . . . . . . . . . . . . . . . . . . . . 11
11. Outer phyllaries variously loosely appressed . . . . . .
.......... T. turfosiforme Kirschner & Štěpánek
Outer phyllaries tightly appressed, rarely some (1 or
2) phyllaries suberect or with arcuate apex . . . . . 12
12. Cone 0.7 0.9( 1)mm long . . . . . . . . . . . . . . . . . . . .
................. T. vindobonense Soest, s.lat.
Cone 1 1.5mm long . . . . . . . . . . . . . . . . . . . . . . 13
13. Leaves deeply divided into long, acute triangular seg-
ments; distal margins of segments and interlobes usu-
ally with several thin, acute teeth . . . . . . . . . . . . . . .
.............. T. scaturiginosum G. E. Haglund
Leaves undivided, subentire or with small, patent teeth,
or shallowly lobed, rarely with lateral segments; lobes
or segments entire ......................... 14
14. Achenes 5.5 6.9mm long; cone 1.4 2mm long . .
..........................T. refectum Sonck
Achenes 4.5 5.2mm long; cone to 1.5mm long . .
....................................... 15
15. Beak 8 9mm long . . . . . . . . . . . . . . . . . . . . . . . . . .
.............. T. sophiae Kirschner & Štěpánek
Beak to 6.5mm long . . . . . . . . . . . . . . . . . . . . . . 16
16. Leaves grey-green or glaucous green, narrowly lin-
ear-oblanceolate, entire or remotely shallowly sinu-
ate-dentate; outer phyllaries with border 0.8 1.2mm
wide . . . . T. glaucolivaceum Kirschner & Štěpánek
Leaves paler mid-green, narrowly oblanceolate in
outline, usually deeply lobed to sinuate-lobulate; out-
er phyllaries with border 0.5 0.9mm wide . . . . . . .
..................... T. apiculatoides Małecka
Taraxacum abruptilobum Štěpánek & Kirschner, sp.
nov. – Fig. 1, 2.
Holotype: Bulgaria, Haskovo District, “ad compitum vi-
arum publicarum ca 3 km ad meridiem a pago Poljanovo
versus”, 2 May 1990, K. Sutorý, cultivated as JŠ 6031
(PRA no. det. 28597; isotypes: BRNM, PRA no. det.
28546, distributed also as Taraxaca Exs. no. 1016). – Ex-
siccates: Taraxaca Exs. no. 1016.
Diagnosis Plantae agamospermae, foliis profunde
lobatis, pinnatisectis, lobis lateralibus saepissime paten-
tibus lingulatis, phyllariis exterioribus adpressis, deinde
laxe adpressis, numerosis, subimbricatis, obscure brun-
neo-viridibus marginibus distinctis albo-membranaceis
angustissimis, antheris polliniferis, stigmatibus obscure
luteo-griseis, acheniis dense spinulosis, in pyramidem
tenuem cylindricam 0.8 1(– 1.1) mm longam abrupte
abeuntibus, rostro 8 11mm longo.
Description Herbs medium-sized, sometimes quite
tall, usually 14 25 cm tall. Plant base almost gla-
brous. Leaves variably erect-patent, usually 8 19 ×
1.5 4.5cm, light (brownish) green, unspotted, almost
glabrous, sometimes araneous on midvein; leaf blade
narrowly oblanceolate to narrowly elliptic in outline,
pinnatisect; terminal segment variable, usually 1 3 ×
1 2.5 cm, narrowly triangular to triangular, often tri-
lobed with ± patent basal lobules and distal subsegment
lingulate, acute to subacute, terminal segment with dis-
tal margin concave, ± undulate, entire, sometimes with
a single incision, proximal margin ± straight, entire; lat-
eral segments (3 or)4 6(or 7) pairs, relatively short, usu-
ally 7 20mm long, 6 12mm wide at base, ± narrowly
deltoid-triangular, usually ± patent, in outer leaves also
158 Štěpánek & Kirschner: Taraxacum sect. Palustria in Bulgaria revised
± recurved, acute, proximal margin slightly concave to
straight, often with a distinct tooth at base, distal margin
variable, entire or with a single tooth; interlobes rela-
tively broad, usually 10 15 × 2 8mm, green or faintly
brownish to brown-purplish, usually dark-bordered, en-
tire or with a single tooth, rarely dentate; midvein green,
proximally sometimes slightly purplish; petiole narrow
or narrowly winged, 2 5( 6) cm long, deep purple.
Scapes ± equalling leaves, purplish at base, otherwise
pale green, glabrous, initially with a few araneous hairs
below capitulum, later glabrescent. Capitulum 3 4cm
wide, ± slightly convex, deep yellow. Involucre c.8mm
wide, rounded at base. Outer phyllaries 14 19, broadly
ovate to ovate, with an attenuate and obtuse apex, 6 7 ×
3.5 5mm, appressed, loosely so after anthesis, slightly
imbricate, ± evenly brown-green, blackish when dry,
darker distally, with an abrupt transition into whitish
membranous border 0.1 0.2mm wide, margin irregu-
larly ciliate. Inner phyllaries 10 11mm long, dark oli-
vaceous green. Outer ligules flat, striped dark olivaceous
grey, later also suused purplish, ligule teeth blackish;
inner ligules canaliculate, teeth reddish to dark yel-
low. Stigmas discoloured, yellowish grey-green, outer
pubescence with apically dark hairs. Pollen abundant,
irregular in size. Achenes medium light (stramineous)
greyish brownish, 3.6 4.3 × 0.9 1.1mm, body medium
densely to subsparsely spinulose in distal ¼, other-
wise ± smooth, spinules short, ± broad, body ± abruptly
narrowing into a thin, cylindric to subcylindric cone
0.8 1( 1.1) mm long, usually with minute spinules at
base; beak (8 )10 11 mm long; pappus 5.5 6.5 mm
long, light brownish white. Agamospermous. Triploid:
2n = 24 (det. V. Jarolímová, 2000). Flowering from April
to May; fruiting in May.
Ecology and distribution — Wet grasslands and pastures.
Thracian plain (Polyanovo). Known from a single macro-
locality in Bulgaria only.
Anities Taraxacum abruptilobum belongs to the
group of T. subalpinum Hudziok and T. noterophilum
Kirschner & al. (Kirschner & Štěpánek 1998) and is quite
close to T. ambitiosum, particularly in outer phyllary fea-
tures, but the latter has a dierent character of achenes:
much less densely spinulose, with a subgradual transition
of body into a longer, usually 1.3 1.5 mm-long cone.
Also the leaf shape is useful to tell the two taxa apart.
Another taxon similar to T. abruptilobum is the Turkish
Fig. 1. Taraxacum abruptilobum sp. nov. – A: general habit (scale bar = 5cm); B: achenes (scale bar = 1mm); C: outer involucral
phyllary (scale bar = 1mm). – Del. J. Štěpánek according to cultivated plants of JŠ 6031.
159Willdenowia 47 – 2017
T. pseudopulchrum Kirschner & Štěpánek (Kirschner &
Štěpánek 1998). Again, T. pseudopulchrum diers from
T. abruptilobum in its achenes with a very short cone
(0.5 0.7mm long) and shorter beak; its outer phyllar-
ies are indistinctly bordered, and its leaf lateral lobes are
much more densely dentate.
Etymology The adjectival epithet refers to the abruptly
incised lateral segments of the leaves.
Specimens examined B: Haskovo District,
near the crossroads c.3km S of Polyanovo, 2 May 1990,
K. Sutorý, cultivated as JŠ 4763, collected 1992 (PRA no.
det. 28548); ibid., cultivated from achenes of JŠ 4763 as
6031, collected 1998 (BRNM, PRA no. det. 28546,
distributed also as Taraxaca Exs. no. 1016).
Taraxacum basilicum Štěpánek & Kirschner, sp. nov.
Fig. 3, 4.
Holotype: Bulgaria, Sofia region, Gorni Lozen, summit
grasslands of Lozenska planina, 1000 1100m, 29 May
1988, J. Kirschner, cultivated as 3088 (PRA no. det.
28593; isotypes: PRA no. det. 29010, also Taraxaca Exs.
no. 1012). – Exsiccates: Taraxaca Exs. no. 1012 1014.
Diagnosis Plantae mediocres, primo adspectu Tara-
xaco obunco et T. copidophyllo anes, sed phyllariorum
exteriorum marginibus pallidis carentibus, acheniis mul-
to longioribus, pappo sordide albido discrepant.
DescriptionHerbs medium-sized, usually 12 18cm
tall. Plant base densely brownish arachnoid. Leaves
erect-patent, usually 6 14 × 1.3 4.5 cm, light oliva-
ceous green, adaxially often suused bronze, unspotted,
sparsely araneous to glabrate; leaf blade elliptic to nar-
rowly oblanceolate in outline, pinnatisect; terminal seg-
ment dominant, usually 1.3 3 × 1.5 4cm, triangular to
narrowly so, approaching helmet-shaped, apex subacute,
distal margin ± concave to slightly sigmoid, entire, basal
lobules subrecurved, acute, proximal margin convex to
sigmoid, entire, whole terminal segment ± sagittate; lat-
eral segments 2 or 3(or 4) pairs, proximal ones smaller,
slightly recurved, deltoid-triangular, sometimes bird-
wing-shaped, acuminate, distal margin ± convex to ± sig-
moid, entire, or with a few minute teeth, proximal margin
sigmoid to ± concave, entire; interlobes long, relatively
narrow, usually 3 11 × 2 4mm, not darker coloured, en-
tire, less often with a single thin, long, acute tooth or sev-
eral filiform teeth, margin ± brown-purple; midvein dis-
Fig. 2. Taraxacum abruptilobum sp. nov. – Holotype specimen (PRA).
160 Štěpánek & Kirschner: Taraxacum sect. Palustria in Bulgaria revised
tally pale, proximally pinkish to purplish (brown); petiole
narrowly to medium broadly winged, usually 2 3.5cm
long, greyish purple. Scapes equalling to ± overtopping
leaves, scattered floccose-araneous, densely so below ca-
pitulum, light green. Capitulum relatively large, 3 4cm
wide, slightly convex, (dirty) yellow. Involucre dark oli-
vaceous, flat to ± truncate at base, c.8mm wide. Outer
phyllaries 16 20, loosely appressed to almost patent
(appressed at base, distally arcuate to sigmoid-patent),
not imbricate, narrowly lanceolate to lanceolate, 8 11
× 2 3.3mm, lowermost ones with margin at base with
1 3 narrow, fimbriate teeth to 0.4mm long, adaxially
paler green and often suused bluish purplish, abaxially
dark blackish olivaceous green (brownish black-green
when dry), often suused violet in distal ½, border not
developed or border only in proximal part, reduced to a
very narrow (to 0.1mm wide), dark membranous fringe,
apex ciliate, flat to callose. Inner phyllaries 12 13mm
long, dark olivaceous green. Outer ligules slightly canal-
iculate, more distinctly so near apex, striped dark pur-
plish grey-black outside, apical teeth black-purple; inner
ligule teeth light purple. Stigmas dark, grey-green, with
black pubescence outside. Anthers polliniferous; pollen
irregular in size. Achenes light grey-brownish to light oli-
vaceous grey-brown, 4.4 5.3 × 1 1.1mm, body coarsely
spinulose and ribbed, either throughout (outer achenes)
or in distal (inner ones), spinules short, longest ones
(to 0.4mm long) just below cone, these sometimes coa-
lescing in squamules, erect-patent, body subgradually
narrowing into a ± thick, subcylindric cone 0.8 1.4mm
long, often with a few spinules at base; beak (6 )7 9mm
long; pappus light brownish white, 6.5 7.5 mm long.
Agamospermous. Flowering from April to May; fruiting
from May to June.
Ecology and distribution — At most of its sites, Tara-
xacum basilicum grows in temporarily wet meadows
(vernal spring patches in meadows); the only known ex-
ception is the Vitosha locality with a slope permanently
soaked with spring water. It is apparently, so far as the
material goes, quite common in mountain ranges S of
Sofia (from Vitosha to Golo Bardo).
Anities Taraxacum basilicum, when compared with
the other members of T. sect. Palustria in Bulgaria, ap-
parently approaches the group of T. subalpinum (repre-
sented by seven species), and T. obuncum in particular.
T. basilicum diers from the latter in having substantially
longer achenes, much longer and pale brownish white
pappus, and longer outer phyllaries with almost absent
Fig. 3. Taraxacum basilicum sp. nov. – A: general habit (scale bar = 5cm); B: achenes (scale bar = 1mm). – Del. J. Štěpánek
according to cultivated plants of JŠ 3088.
161Willdenowia 47 – 2017
Fig. 4. Taraxacum basilicum sp. nov. – Holotype specimen (PRA).
162 Štěpánek & Kirschner: Taraxacum sect. Palustria in Bulgaria revised
borders. Taraxacum basilicum may be compared with the
group of T. copidophyllum Dahlst. of T. sect. Taraxacum.
The closest species, T. copidophyllum is characterized
by much shorter achenes with short cone, dierent leaf
shape (particularly the terminal segment and denticulate
distal margins to proximal leaf lateral segments).
Etymology The epithet is a Latin adjective, basilicus
(-a, -um), meaning kingly or noble.
Specimens examined — B: Golo Bardo Mts,
along path between alpine chalets of Orlite and Slavej,
19 May 1989, B. Kuzmanov 89109, cultivated as T 221
(PRA no. det. 26015, distributed also as Taraxaca Exs. no.
1014); Sofia, Ljulin planina, near the monastery, 29 May
1988, J. Kirschner (PRA no. det. 26014); Sofia, Gorni
Lozen, summit part of Lozenska planina, 1000 1100m,
29 May 1988, J. Kirschner, cultivated as 3087 (PRA
no. det. 26013, distributed also as Taraxaca Exs. no.
1013); ibid., cultivated as JŠ 3088 (PRA no. det. 26010,
distributed also as Taraxaca Exs. no. 1012); ibid., culti-
vated as 3089 (PRA no. det. 26009, distributed also
as Taraxaca Exs. no. 1013); Sofia, Lozenska Planina, wet
slopes above Gorni Lozen, May 1990, J. Kirschner, cul-
tivated as JK 1071 (PRA no. det. 26012); Sofia, Vitoša
Mts, meadows along path SW of alpine chalet of Aleko,
1600 m, 3 Jun 1988, J. Kirschner, cultivated as T102
(PRA no. det. 26011).
Taraxacum rumelicum Štěpánek & Kirschner, sp. nov.
– Fig. 5, 6.
Holotype: Bulgaria (southern), Rodopi Mts, Čepelare,
Mursalica (= E part of Perelik Mts), Široka Läka, wet
meadows at W foot of Mt Goljam Sněžnik (2188m),
between Lednicata and Mugla, c. 2 km NE of Mugla,
41°37'N, 24°31'E, 1700m, 23 Jul 1998, J. Štěpánek, R.
Bělohlávková, V. D. Vladimirov & D. Petkova, cultivat-
ed from root sample no. 41 as 6829/C (PRA no. det.
28595; isotypes: PRA no. det. 28555, and 15 duplicates
distributed as Taraxaca Exs. no. 1015). – Exsiccates:
Taraxaca Exs. no. 1015.
Diagnosis Plantae mediocres vel subrobustae, primo
adspectu Taraxaco subudo et T. vindobonensi similes,
sed acheniis et pyramide longioribus, pappo multo lon-
giore, phyllariis involucralibus exterioribus latioribus
discrepant.
Description Herbs usually medium-sized, ± subrobust,
most often 12 25cm tall. Plant base medium densely
brownish arachnoid. Leaves variably erect-patent, usual-
ly 9 16 × 1.5 3cm, light subglaucous green, not spotted,
glabrous or subglabrous; leaf blade linear-oblanceolate to
linear-oblong in outline, pinnatipartite to pinnatisect; ter-
minal segment medium-sized to ± large, usually 1.2 4×
1 3cm, triangular or narrowly so, acute to subacumi-
nate, distal margin slightly convex to ± sigmoid, entire,
or with a single shallow acute incision, basal lobules su-
brecurved or patent, quite often asymmetric, whole ter-
minal segment therefore sometimes ± sagittate; proximal
margin ± straight or subconcave, entire; lateral segments
3 or 4 pairs, deltoid-triangular to narrowly triangular, re-
curved, acute, usually 5 19mm long, 7 15mm wide at
base, from a very broad base ± gradually narrowing into a
long, narrowly triangular, acuminate apex, distal margin
± straight or subconvex or subsigmoid, ± entire, proxi-
mal margin similar, entire; interlobes long and relatively
narrow, usually 5 12 × 3 4(– 6) mm, green or distally
with narrowly bordered brownish purple, with ± raised
margins, entire or with a few small, narrow teeth; mid-
vein proximally pinkish, distally light green; petiole nar-
rowly winged, greyish light purple. Scapes overtopping
leaves, usually 10 22cm long, slightly suused purplish
at base, distally green, purple to bronze below capitulum,
very sparsely araneous to floccose-araneous. Capitulum
large, 3.5 4.5 cm wide, ± flat, deep yellow. Involucre
rounded at base, c.9mm wide, not pruinose. Outer phyl-
laries 20 26, slightly imbricate, loosely appressed with
erect apex, outermost (1 or 2) ones linear, middle ones
ovate to ovate-lanceolate, uppermost ones ± lanceolate,
7 8 × (2 –)3.5 5 mm, often with an elongated obtuse
apex, flat, suused purple in distal ⅓, with a variably
broad, dark green to black-green middle part (0.3 2mm
wide), with a gradual transition into variably broad, pale
olivaceous to whitish green marginal part bordered by a
0.1 0.4mm-wide, pale membranous, distally ciliate mar-
gin. Inner phyllaries 13 14mm long, of equal width, ±
dark olivaceous green, apex suused pink-brown. Outer
ligules flat, striped dark olivaceous grey outside, apical
teeth grey-black; inner ligules subcanaliculate, with api-
cal teeth dirty yellow. Stigmas exserted, relatively dark,
yellowish grey-green, with a black pubescence outside.
Pollen abundant, irregular in size. Achenes light grey-
ish stramineous-brown, 5.1 5.8 × 1.2 1.3mm, achene
body medium densely and shortly spinulose in distal ¼,
± gradually narrowing into a cylindric to subcylindric,
narrow cone 1 1.5mm long; beak 7.5 9mm long; pap-
pus conspicuously long, c.8.5mm long, pale yellowish
white. Agamospermous. Flowering from April to May;
fruiting from May to June.
Ecology and distribution Taraxacum rumelicum grows
in wet meadows around mountain springs. It is known
from a rich population at the type locality only.
Anities Taraxacum rumelicum is a quite robust plant,
relatively close to T. subudum, but diering from it pri-
marily in darker, more numerous outer phyllaries and
longer achenes with much longer pappus. Also the group
of T.paucilobum Hudziok and T. vindobonense Soest (as
defined in Kirschner & Štěpánek 1998) is to be compared
with T. rumelicum. If we disregard the small, slender
plants of T. paucilobum and its allies, T. vindobonense
and a few taxa closely related to it have much shorter
163Willdenowia 47 – 2017
achenes and cones, substantially shorter pappus, and a
narrower shape of middle outer phyllaries.
Etymology — The epithet derives from Eastern Rumelia,
the historical Balkan region where the species occurs.
Fig. 5. Taraxacum rumelicum sp. nov. – A: general habit (scale bar = 5cm); B: achene (scale bar = 1mm); C: outer involucral phyl-
laries (scale bar = 1mm). – Del. J. Štěpánek according to cultivated plants of JŠ 6829/C.
164 Štěpánek & Kirschner: Taraxacum sect. Palustria in Bulgaria revised
Fig. 6. Taraxacum rumelicum sp. nov. – Holotype specimen (PRA).
165Willdenowia 47 – 2017
Willdenowia
Open-access online edition www.bioone.org/loi/will
Online ISSN 1868-6397 · Print ISSN 0511-9618 · Impact factor 0.680
Published by the Botanic Garden and Botanical Museum Berlin, Freie Universität Berlin
© 2017 The Authors · This open-access article is distributed under the CC BY 4.0 licence
Acknowledgements
We are grateful to the late B. Kuzmanov and to K. Sutorý
for plant material. Thanks are due to V. D. Vladimirov
and D. Petkova for the organization of the expedition in
Bulgaria, and to R. Bělohlávková for field assistance.
The work was supported by the long-term research and
development project of the Institute of Botany, Acad-
emy of Sciences, Průhonice, Czech Republic, no. RVO
67985939, and the Flora of Bulgaria project. In addition,
Bohumil Trávníček (Olomouc, Czech Republic) and an
anonymous reviewer are thanked for their comments on
an earlier version of this paper.
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... Last, we can give a few names that are listed for Bulgaria erroneously: Taraxacum sect. Palustria was newly revised, and seventeen taxa were recognized (Štěpánek & Kirschner 2017). Another group that received a great attention was T. sect. ...
... Taraxacum sect. Palustria (Lindberg 1908: 17) Dahlstedt (1921: 37), see van Soest (1965), Sonck (1985aSonck ( , b, 1986Sonck ( , 1993, Kirschner & Štěpánek (1993aKirschner & Štěpánek ( , 1998, Štěpánek & Kirschner (2017). Taraxacum sect. ...
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... However, the problem also concerns species within taxonomically problematic and species-rich genera with morphologically similar or cryptic species, such as Rubus, Hieracium, Campanula, Orobanche, Stipa, Oxytropis, etc. (Wolanin et al. 2016, 2020Aleksić et al. 2018;Piwowarczyk et al. 2019;Chen et al. 2020;Nobis et al. 2020a;Baiakhmetov et al. 2021;Kosiński et al. 2021;Trávníček et al. 2021;Havlíček et al. 2022;Szeląg 2022;Vintsek et al. 2022;Wang et al. 2022;Nobis et al. 2023). Globally distributed dandelions (Taraxacum, Asteraceae) can also be included in this group (Uhlemann 2002(Uhlemann , 2016Scott and Rich 2013;Štěpánek and Kirschner 2017;Marciniuk et al. 2018;Kirschner et al. 2020Kirschner et al. , 2022. ...
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On the basis of allozyme and cultivation data, and of additional herbarium material, a taxonomic and nomenclatural revision ofTaraxacum sect.Piesis A.J. Richards exKirschner etŠtěpánek is provided. The section is made up of halophilous, sexually reproducing taxa. InT. stenocephalum Boiss. etKotschy,T. pindicum Kirschner etŠtěpánek, sp. nov., andT. perenne Kirschner etŠtěpánek, sp. nov., a tetraploid chromosome number has been recorded, representing the only known case of sexuality at the tetraploid level in the genus. The complex ofT. stenocephalum, includes some geographically and morphologically extreme populations treated as subspecies: subsp.gumusanicum (Soest)Kirschner etŠtěpánek, comb. nov., subsp.magnum Kirschner etŠtěpánek, subsp. nov., and subsp.daralagesicum (Schischk.)Kirschner etŠtěpánek, comb. nov. In addition toT. bessarabicum (Hornem.)Hand.-Mazz., a widely distributed Eurasian species,T. stenocephalum, a complex centred in Transcaucasia and Anatolia, andT. pachypodum H. Lindb., a North African endemic, four new species are described:T. salsum Kirschner etŠtěpánek, sp. nov., a diploid endemic confined to E Crimea,T. perenne Kirschner etŠtěpánek, sp. nov., a tetraploid sexual species known only from SW Crimea,T. pindicum Kirschner etŠtěpánek, sp. nov., a remarkable tetraploid endemic to the Pindos Mts., Greece, andT. salsitatis Kirschner, Štěpänek etYirdirimli, sp. nov., an Anatolian diploid species. Furthermore, a hybrid betweenT. salsum andT. bessarabicum from Crimea (documented on the basis of allozyme data elsewhere) is given a binomial,T. xmesohalobium Kirschner etŠtěpánek, nothosp. nov.
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The genus Taraxacum is readily divided into “primitive” and “advanced” forms, on morphological and cytological criteria. It is thought that the genus arose in the west Himalayas during the Cretaceous and that apomixis arose at an early stage by means of polyploidy, precocious embryony and asynapsis in the female meiosis. Advances of sexual primitive types and “precursor” types west into Europe were paralleled by the spread of arctic-alpine types into many regions of the world. During the Pleistocene the precursor types are thought to have generated the widespread advanced section Ceratophora, which gave rise to many of the advanced species after the last glacial period by hybridizing with primitive and precursor sexuals, thus “fixing” a hybrid swarm as apomicts.
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Modes of evolution of species classified within different sections inTaraxacum involve diverse processes, viz. primary divergence of an ancestral sexual diploid, hybridization between a tetraploid apomict and a diploid sexual hybrid, differentiation of an advanced apomictic taxon at one ploidy level, hybridization between a sexual tetraploid and a sexual diploid, formation of a polyploid series from an apomictic ancestor of a lower polyploidy level, and remote hybridization between an autumn-flowering ancestral diploid and a spring-flowering derivative diploid or apomict. Various reproduction systems of the plants involved, different environments and different timing of the processes contribute to a very varied nature of the species groups.
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An introduction is presented to a taxonomic evaluation of rich cultivated and herbarium material of the genusTaraxacum from the Caucasus. Sources of the material, a survey of localities, and a preliminary account of the sections and species groups ofTaraxacum described from the Caucasus are given. The introduction is followed by a monograph ofT. sect.Porphyrantha, a section shown (contrary to the original conception and some literature data) to be endemic to the Caucasus. Six new species are described belonging to the section divided in two subsections, subsect.Porphyrantha and subsect.Haemantha, viz.T. pseudoporphyranthum, T. vepallidum, T. haemanthum, T. ignivomum, T. exstinctum andT. deminutum. Lectotypes are selected forT. porphyranthum Boiss. andT. pseudoroseum Šiškin.