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Populations of species of Pilosella in ruderal habitats in the city of Prague:
frequency, chromosome numbers and mode of reproduction
Populace druhů rodu Pilosella na ruderálních stanovištích v Praze: frekvence výskytu, počty chromozomů
a způsob rozmnožování
Veronika K ř i š ť á l o v á1, Jindřich C h r t e k2,3, Anna K r a h ul c o vá2,
Siegfried B r ä u t i g a m4& František K r a h u le c2*
1Department of Ecology, Faculty of Environmental Sciences, Czech University of Life Sci-
ences, Kamýcká 129, CZ-165 21 Prague 6 – Suchdol, Czech Republic, e-mail:
vkostalova@gmail.com; 2Institute of Botany, Academy of Sciences of the Czech Republic,
Zámek 1, CZ-252 43 Průhonice, Czech Republic, e-mail: frantisek.krahulec@ibot.cas.cz;
3Department of Botany, Faculty of Science, Charles University in Prague, Benátská 2, CZ-
128 01, Prague, Czech Republic, e-mail: jindrich.chrtek@ibot.cas.cz; 4Senckenberg
Museum für Naturkunde Görlitz, Postfach D-300154, 02806 Görlitz, Germany
* corresponding author
Křišťálová V., Chrtek J., Krahulcová A., Bräutigam S. & Krahulec F. (2010): Populations of species
of Pilosella in ruderal habitats in the city of Prague: frequency, chromosome numbers and mode of
reproduction. – Preslia 82: 437–464.
Populations of Pilosella (Hieracium subgenus Pilosella) at ruderal localities were investigated in an
urban area (Prague City) with respect to their distribution, variation in DNA ploidy level/chromo-
some number and mode of reproduction. The following species, hybridogenousspecies or hybrids
(with ploidy level/chromosome number and mode of reproduction) were found: P. aurantiaca,
P. caespitosa (4x, 5x), P. cymosa subsp. vaillantii (5x), P. officinarum (2n = 36, sexual; 2n = 54, sex-
ual; 2n = 63), P. piloselloides subsp. bauhinii (2n = 45, 54; both apomictic), P. piloselloides subsp.
praealta (5x; apomictic), P. brachiata (4x; sterile), P. densiflora,P. flagellaris,P. floribunda,
P. erythrochrista,P. glomerata (5x; apomictic), P. leptophyton (5x; apomictic), P. rothiana (4x,
apomictic), P. setigera,P. visianii (4x; apomictic), P. ziziana (4x, apomictic) and the previously
undescribed hybridogenous type P. piloselloides ×P. setigera (5x, apomictic). Pilosella visianii is
reported from the Czech Republic for the first time. New habitats resulting from highway construc-
tion are suitable for Pilosella species. Many previously rare types, such as P. rothiana, can colonize
these habitats and spread, not only locally, but also throughout the whole country.
K eywo r ds: chromosome numbers, DNA ploidy level, Pilosella, reproductive mode
Introduction
Plant species richness in cities is usually greater than in surrounding areas due to the
greater habitat diversity there, which is a result of more variable land use and enrichment
by alien species (e.g. Sukopp & Werner 1983, Pyšek & Pyšek 1990, Kühn et al. 2004). The
distribution of many species changes considerably in time due to the temporary nature of
many habitats, succesional changes, and various kinds of disturbances. Detailed invento-
ries of species diversity are thus of great importance for understanding species and com-
munity dynamics in human settlements.
In the genus Pilosella Vaill. there are both diploid (sexual) and polyploid (sexual or
facultatively apomictic) perennials, whose seed is dispersed by wind, and which fre-
Preslia 82: 437–464, 2010 437
quently hybridize and commonly spread vegetatively by means of stolons (Krahulcová et
al. 2000, Fehrer et al. 2007). These plants prefer open habitats, such as grasslands, heaths,
herbaceous fringes and rocky outcrops. Diploids are usually confined to more natural,
sometimes relic habitats. Polyploid species, both basic and intermediate (hybridogenous)
species often occur in man-made habitats, which have a more or less extensively disturbed
vegetation cover and where competition is low, e.g. railways, roadsides and slopes along
roads (e.g. Heinrichs 1998, Dunkel et al. 2007). The number of suitable localities in the
rural landscape is decreasing, mostly due to either fertilization and subsequent increase in
competition from more productive species or the abandonment of grasslands and conse-
quent succession. However, these tendencies are based only on casual observations. They
are documented very rarely at a landscape level, e.g. by Heinrichs & Gottschlich (1996)
and Gottschlich et al. (2004, 2006).
For this reason, it was decided to study the diversity of Pilosella in an urban landscape.
After a pilot study, all the morphologically distinguishable types were collected at selected
localities and their chromosome numbers/DNA ploidy level and mode of reproduction (sexu-
ality vs. apomixis) determined. Both these parameters are of crucial importance to Pilosella
and information on them may help clarify the evolutionary potential of particular species/types
and indicate why some of them are thriving, whereas others are in danger of disappearing.
A detailed population study might also elucidate the hybridization processes (generally com-
mon in this genus), the fate of hybrid progeny and the role of basic (non-hybridogenous) spe-
cies and habitat condition in determining the structure of mixed populations. The data set will
enable future students to evaluate more precisely the changes in the distribution and frequency
of particular species and changes in population composition. For that reason all species occur-
ring at the localities studied are listed, even if their mode of reproduction and chromosome
numbers/DNA ploidy level were not studied. However, this contribution definitively does not
provide a comprehensive list of Pilosella localities in the area studied.
Remarks on the taxonomic concept
For a long time, Pilosella was treated in central European literature as a subgenus within
Hieracium (Nägeli & Peter 1885; Zahn 1922–1930). In fact, we consider it, in agreement
with many other authors, as an independent genus, but in our previous papers we used the
traditional classification as a subgenus, because several names were not valid within
Pilosella. The full list of species in the recent paper by Bräutigam & Greuter (2007) is used
in this study. However, in all cases where the respective species is given in the Flora of
Czech Republic (Chrtek 2004) we refer to names within Hieracium.
Area studied
The city of Prague is situated in the center of the Bohemian basin, occupies 496 km2and is
at an altitude of 177 m in the Vltava valley in the north to 399 m near Zličín on the west.
Within the Czech Republic, the climate is warm, with an average annual temperature of
9.4 °C in the town center to 7.8 °C at Ruzyně. Winter is usually without or with very irreg-
ular snow cover. The geology in the area of the city is diverse, with extremely acid lydites
and sandstones to extremely base rich rocks, such as lime stones and marls. Their age is
Proterozoik to Mesozoik, but there are deep loess deposits in many places, which cover
438 Preslia 82: 437–464, 2010
bedrock and suppress its influence. Bedrock has a greater influence at many places along
railways and highways, especially where they are situated in trenches. On the other hand,
the embankments are often made from completely different material, the extreme being
pure sand. It is impossible to characterize the composition briefly. More information about
the environmental conditions, history and vegetation cover can be found, e.g. in books by
Moravec et al. (1991) and Kovanda (1995).
Within the city of Prague, the number of new habitats is increasing because of the build-
ing of new highways and an extensive road network. Along these new roads there are areas
with different habitat conditions (differing especially in exposition and for that reason also
in moisture, temperature etc.). These habitats mostly consist of open soil surfaces, which
are quickly colonized by seedlings and populations. Pilosella species are especially good
invaders of these habitats, because their seeds are dispersed by wind and the air currents
caused by traffic.
Material and methods
Plant material
Living plants were collected at 49 localities (mostly along roads and railways, seeAppen-
dix 1, Fig. 1), with most collected in 2002 and 2003, but occasionally also in other years in
this decade. They were transplanted to the experimental garden of the Institute of Botany
in Průhonice and used for determining chromosome number/ploidy level and mode of
reproduction. Voucher specimens of cultivated plants together with numerous specimens
collected in the field are deposited in the herbarium PRA. Both the taxonomic concepts
and nomenclature follow Bräutigam & Greuter (2007).
Determination of chromosome number, DNA ploidy level and mode of reproduction
Chromosomes were counted in root-tip meristems of pot-grown plants following the
method described in Krahulcová & Krahulec (1999). DNA ploidy level (Suda et al. 2006)
was determined using flow cytometry, following the method of Krahulcová et al. (2004).
As an internal standard karyologically examined plants cultivated in the experimental gar-
den were used.
Mode of reproduction was determined by comparing the seed set of open pollinated and
emasculated (cut) capitula (Gadella 1984, Krahulcová & Krahulec 1999). The plants, which
did not produce any achenes when open pollinated, were considered to be sterile, those of
which the open pollinated capitula produced achenes, but not the emasculated ones were
considered to be sexual. The plants on which the emasculated capitula produced achenes
were considered to be apomictic (agamospermous). For P. ziziana and P. piloselloides –
P. setigera we used the FCSS method (Matzk et al. 2000, Krahulcová & Rotreklová 2010).
Isozyme analysis
Isozyme analysis was used to estimate the genotype (isozyme phenotype) structure of
P. visianii, which is a rather common but previously unrecorded species for the Czech
Republic. Methods used are described in detail by Krahulec et al. (2004); two systems
were used in 2004, aspartate aminotrasferase (AAT) and esterase (EST). Esterases are
Křišťálová et al.: Pilosella species in ruderal habitats 439
highly variable and are valuable for clone determination in the genus Pilosella (Krahulec
et al. 2004). For estimating P. rothiana and P. visianii clones four systems (AAT, 6-PGDH,
LAP and EST) were used in 2010.
Overview of species
Altogether, five basic species (one with two subspecies) and 11 intermediate species were
found at 49 localities (see Appendix 1 and Fig. 1). They are briefly commented on below.
Figures in brackets following locality number indicate number of plants studied.
Basic species
Pilosella aurantiaca (L.) F. W. Schultz & Sch. Bip. subsp. aurantiaca
(syn.: H. aurantiacum L.)
This species was found only once (loc. 28) and has not been studied. No hybrids of
P. aurantiaca were found along the new roads.
Pilosella caespitosa (Dumort.) P. D. Sell & C. West (syn.: H. caespitosum Dumort.)
Localities: 9, 12, 24, 30, 47.
DNA ploidy level 4x: loc. no. 24 (10 plants); loc. 47 (3 plants).
DNA ploidy level 5x: loc. no. 24 (1 plant).
Pilosella caespitosa occurs rarely in the area studied. The estimated ploidy levels corre-
spond to two clones known from Central Europe (Fehrer et al. 2005).
440 Preslia 82: 437–464, 2010
Fig. 1. – Map of Prague indicating the localities studied.
Pilosella cymosa subsp. vaillantii (Tausch) S. Bräutigam & Greuter (syn: Hieracium
cymosum subsp. cymigerum Peter)
Locality: 47
DNA ploidy level 5x: locality no. 47 (2 plants).
Numerous chromosome counts are published for H. cymosum, ranging from diploids to
hexaploids (see Rotreklová et al. 2005 for references). However, they only rarely directly
refer to P. cymosa subsp. vaillantii (H. c. subsp. cymigerum): 2n = 4x (Fehrer et al. 2005)
for plants from the Krušné hory Mts, Czech Republic) and 2n = 4x, 2n = 54 (Marhold et al.
2007) for those from the Nízke Tatry and Veľká Fatra Mts, Slovakia. Chrtek (2004) gives
tetra– and pentaploids of this subspecies in the Czech Republic.
Pilosella officinarum Vaill. (syn.: Hieracium pilosella L.)
Localities: 6, 9, 10, 12, 15, 21, 24, 25, 31, 32, 33, 37, 38, 42, 44, 45.
DNA ploidy level: 4x (46 plants); locality no. 6 (5), 9 (11 plants, 2n = 36), 10 (9), 15 (5), 21 (2), 24 (1), 25 (2), 32
(2), 44 (1), 45 (8).
Mode of reproduction, sexual (44 plants): locality no. 6 (5), 9 (11), 10 (9), 15 (5), 21 (2), 25 (2), 32 (2), 45 (8).
DNA ploidy level 6x: locality no.12 (6 plants, 2n = 54, Fig. 2).
Mode of reproduction, sexual: locality no.12 (1 plant).
2n = 7x = 63: locality no. 12 (1 plant, Fig. 2); mode of reproduction not specified.
The flow cytometry histogram for all three cytotypes (tetraploid, hexaploid and heptaploid) is presented in Fig. 3.
Pilosella officinarum is the second most common species found in the area studied
(recorded at 16 localities). For this species in Prague, three ploidy levels were detected.
The most common were populations of sexual tetraploid plants (2n = 4x = 36). Krahulcová
et al. (2009) report tetraploid (2n = 4x = 36) and pentaploid (2n = 5x = 45) plants at
Vysočany, loc. 31; both of which were sexual. A rich population of hexaploid (2n = 54)
plants, undoubtedly identical with those previously collected in the canyon of the Vltava
river (Mráz et al. 2008), was discovered at one locality. One heptaploid plant (2n = 7x =
63) was detected within this hexaploid population; it is probably a 2n + n hybrid between
tetraploid and hexaploid cytotypes. There was only a single plant of this heptaploid and it
was not recollected during repeated visits to this locality (T. Urfus, pers. comm.). In this
respect it is similar to the individual heptaploid plants found in Slovakia (Mráz et al. 2008).
The mode of reproduction of this single heptaploid plant was not determined as the plant
grew badly and did not survive cultivation in the experimental garden.
Pilosella piloselloides subsp. bauhinii (Schult.) S. Bräutigam & Greuter (syn.: Hieracium
bauhinii Schult.)
Localities: 1, 3, 4, 5, 6, 7, 8, 9, 10, 12, 13, 14, 15, 16, 17, 18, 19, 20, 22, 23, 24, 25, 26, 28, 29, 30, 31, 34, 35, 36, 40,
41, 44, 47.
DNA ploidy level 5x (35 plants): locality no. 1 (1 plant, 2n = 45, Fig. 3), 3 (5 plants, 2n = 45), 5 (3), 8 (2), 9 (3
plants, 2n = 45), 10 (1), 22 (5), 24 (1), 25 (3 plants, 2n = 45), 26 (2), 28 (2 plants, 2n = 45), 44 (4), 47 (4).
Mode of reproduction apomictic (34 plants): locality no. 3 (5), 5 (5), 6 (2), 8 (2), 9 (3), 10 (1), 22 (5), 25 (3), 26 (2),
28 (2), 44 (4).
DNA ploidy level 6x (32 plants): locality no. 10 (1), 14 (7 plants, 2n = 54, Fig. 3), 15 (6 plants, 2n = 54), 17 (5), 19
(2 plants, 2n = 54), 30 (10 plants, 2n = 54).
Mode of reproduction, apomictic (32 plants): locality no. 10 (1), 14 (8), 15 (6), 17 (5), 19 (2), 30 (10).
The flow cytometry histogram for both cytotypes is presented in Fig. 3.
Křišťálová et al.: Pilosella species in ruderal habitats 441
442 Preslia 82: 437–464, 2010
Pilosella *bauhinii is the most frequently recorded Pilosella species in ruderal habitats
within the area studied (34 localities). Penta- and hexaploids seem to be spatially separated
as no mixed populations were found. All 66 plants analyzed were apomictic. The rather
high frequency of pentaploids (recorded at altogether 11 localities) corresponds well with
previously published data from the Czech Republic and Germany (for a review see
Rotreklová 2004), where this cytotype seems to prevail. Both penta- and hexaploids are
reported from Prague, Vysočany (loc. no. 31) (Rotreklová et al. 2002) together with
apomictic tetraploids and a single heptaploid plant (Krahulcová et al. 2009).
Křišťálová et al.: Pilosella species in ruderal habitats 443
Fig. 2. – Microphotographs of somatic metaphases of selected Pilosella species and explanatory drawings.
1–P. officinarum (2n = 63, heptaploid cytotype), 2 – P. piloselloides subsp. bauhinii (2n = 45, pentaploid
cytotype), 3 – P. officinarum (2n = 63, heptaploid cytotype), 4 – P. piloselloides subsp. bauhinii (2n = 54,
hexaploid cytotype), 5 – P. officinarum (2n = 54, hexaploid cytotype).
444 Preslia 82: 437–464, 2010
Fig. 3. – Flow cytometry histograms for selected Pilosella species. (1) Three peaks corresponding to ploidy levels
detected in P. officinarum. All three cytotypes were analyzed simultaneously. (2) Two peaks corresponding to
ploidy levels detected in P. piloselloides subsp. bauhinii. Both cytotypes were analyzed simultaneously. (3) Two
peaks demonstrating the higher DNA content in the tetraploid P. brachiata (Bra) compared to its parental
tetraploid P. officinarum (Pi).
Pilosella piloselloides subsp. praealta (Gochnat) S. Bräut. & Greuter (syn.: Hieracium
praealtum Gochnat)
Localities: 11, 12, 18, 43.
DNA ploidy level 5x: locality no. 12 (4 plants, 2n = 45).
Mode of reproduction, apomictic: locality no. 12 (4 plants).
Populations (max. 10 individuals) of Pilosella *praealta were found only at three localities.
Intermediate species
Pilosella brachiata (DC.) F. W. Schultz & Sch. Bip. (P. piloselloides <P. officinarum;
syn.: Hieracium brachiatum Bertol. ex DC.)
Localities: 6, 10, 23, 24, 31.
DNA ploidy level 4x: locality no. 6 (2 plants, 2n = 36), 10 (1 plant, 2n = 36), 24 (1 plant).
Mode of reproduction, sterile: locality no. 6 (2 plants).
According to its morphology, P. brachiata is an intermediate hybrid between
P. officinarum and P. *bauhinii in this area. Because P. *bauhinii has a higher Cx-value
(monoploid DNA content) than P. officinarum, the tetraploid hybrid, P. brachiata also has
a higher DNA content (2C-value) than its tetraploid parent, P. officinarum (Fig. 3 – cf.
Suda et al. 2007). Out of the nine localities at which both parental species were present,
P. brachiata was detected at four of them. Several cytotypes of P. brachiata were previ-
ously reported occurring in a hybrid swarm at Vysočany (loc. no. 31 in this paper), namely
pentaploid, heptaploid, octoploid and several aneuploid cytotypes (Rotreklová et al. 2002,
Krahulcová et al. 2009). Pilosella brachiata is evidently a complex of recent hybrids,
which in the Czech Republic are represented by a number of cytotypes, ranging from
tetraploid to octoploid (for discussion see Rotreklová et al. 2002, 2005). The pentaploids
are recorded as mostly apomictic (Rotreklová et al. 2005). At locality no. 31 the plants
have diverse modes of reproduction (Krahulcová et al. 2009).
Pilosella leptophyton (Nägeli & Peter) P. D. Sell & C. West (P. piloselloides subsp.
bauhinii >P. officinarum; syn.: Hieracium leptophyton Nägeli & Peter)
Localities: 9, 11, 12, 20, 23, 31.
DNA ploidy level 5x: locality no. 9 (15 plants, 2n = 45).
Mode of reproduction, apomictic: locality no. 9 (14 plants).
The plants examined come from a mixed population of tetraploid sexual P. officinarum
and pentaploid apomictic P. *bauhinii. This is the first report of pentaploid P. leptophyton
from the Czech Republic, previously only one heptaploid cytotype was recorded (Suda et
al. 2007).
Pilosella visianii F. W. Schultz & Sch. Bip. [Pilosella piloselloides subsp. piloselloides
and P. p . subsp. praealta >Pilosella officinarum; syn.: H. visianii (F. W. Schultz & Sch.
Bip.) Schinz & Thell.)] (Figs 4–6)
Localities: 2, 7, 8, 11, 12, 14, 17, 18, 19, 24, 27, 44.
DNA ploidy level 4x (41 plants): locality no. 11 (5 plants, 2n = 36), 12 (11 plants, 2n = 36), 14 (5), 17 (2), 19 (2
plants, 2n = 36), 24 (7), 27 (8 plants, 2n = 36).
Mode of reproduction, apomictic (33 plants): locality no. 11 (5), 12 (11), 14 (5), 17 (2), 19 (2), 27 (8).
Křišťálová et al.: Pilosella species in ruderal habitats 445
In the city of Prague, P. visianii was found at 11 localities and seems to be one of the most
common hybridogenous species. Three chromosome numbers are reported for P. visianii,
i.e. 2n = 4x = 36 and 2n = 7x = 63 (Bräutigam & Schuhwerk 2002, 2005) and 2n = 5x = 45
(P. v. subsp. fallaciniforme, Schuhwerk & Lippert 2002). This species is not reported from
the Czech Republic (Chrtek 2004). It is rare throughout its whole geographical range (the
Alps and adjacent areas, Germany, southeast to Balkan Peninsula; Zahn 1987, Gottschlich
1987, Bräutigam & Schuhwerk 2005, Schuhwerk & Fischer 2003). It occupies an inter-
mediate position between P. piloselloides (subsp. piloselloides or subsp. praealta) and
P. officinarum, but resembles more the former parent. Plants from Prague are 15–45 cm
tall, mostly without stolons and rarely with short underground rhizomes. Key identifica-
tion characters are loose inflorescences (the inflorescence make up 1/5–1/3 of the total
plant height) with usually 5–30 heads (involucral bracts are of an intermediate length
between those of P. piloselloides and P. officinarum, i. e. 7–9 mm) and numerous stellate
hairs on the lower surface of leaves but none on the upper surface). The plants from Prague
seem to be rather uniform in morphology and most of them match the descriptions of
plants from other parts of the geographical area. On the other hand, a few of the plants
(localities 14, 19 and 24) differ in having more deeply branched inflorescences (the inflo-
rescence makes up more than half of the plant height). These plants might at first be identi-
fied as P. arida (Freyn) Soják (H. aridum Freyn), the second intermediate species between
P. piloselloides (subsp. piloselloides and subsp. praealta and P. officinarum), which is
more closely related to P. officinarum. However, isozyme analysis of 34 plants, including
specimens with deeply branched inflorescences from six localities (10, 11, 14, 17, 19, 27),
showed no intra- and inter-population variation. To be certain, this analysis was repeated
446 Preslia 82: 437–464, 2010
Fig. 4. – Pilosella piloselloides subsp. praealta, (left) and P. visianii (center and right).
Křišťálová et al.: Pilosella species in ruderal habitats 447
Fig. 5. – Pilosella visianii.
448 Preslia 82: 437–464, 2010
Fig. 6. – Pilosella visianii.
with a set of plants collected in 2010: five and ten plants were collected at two distant
localities, nos. 19 and 24, respectively. The plants with the different morphology all have
the same genotype (isozyme phenotype). Hence, all these plants belong to one successful
clone, which is apomictic. Plants with a deeply branched inflorescence are thus in our
opinion aberrant forms of P. visianii.
Remark: There are zymograms (isozyme phenotypes) in Košťálová’s diploma thesis
(Košťálová 2004: 83, 84). She uses the name Hieracium anchusoides (Arv.-Touv.) St.-
Lag. (Pilosella anchusoides Arv.-Touv.) for this type, which is a parallel hybrid species
resulting from hybridization between P. ziziana and P. officinarum. It differs from
P. visianii in the presence of stellate hairs on the upper leaf surface. Morphology of both
types considered here as P. visianii is illustrated in Figs 4–6.
Pilosella glomerata (Froel.) Fr. (P. caespitosa –P. cymosa; syn.: Hieracium glomeratum
Froelich)
Localities: 10, 19, 24, 26, 39, 46.
DNA ploidy level 5x: loc. 24 (1 plant).
Mode of reproduction, apomictic: locality no. 46 (6 plants).
This species is recorded for the Czech Republic as tetra- and pentaploid and always
apomictic (e.g. Krahulec et al. 2004, 2008). Because apomictic reproduction allows the
spread of successful clones, as has been reported for example from Germany (Gottschlich
et al. 2006), it is expected that this species will spread further in the Prague area.
Pilosella rothiana (Wallr.) F. W. Schultz & Sch. Bip. (P. echioides >P. officinarum; syn.:
Hieracium rothianum Wallr.) (Fig. 7)
Localities: 3, 14, 30, 40, 46, 47, 48.
DNA ploidy level 4x: loc. 46.
Mode of reproduction, apomictic: locality no. 3 (1 plant), 14 (1), 30 (3), 46 (1).
The populations of P. rothiana consisted of apomictic plants with no evidence/presence of
one of the parental species (P. echioides) at any locality. Pilosella echioides occurs in the
Prague area, but is strictly confined to rocky outcrops in the Vltava canyon and never colo-
nizes secondary habitats (Peckert 2002). Because of their homogeneous morphology, all
plants seem to belong to one genotype that has spread throughout Prague. Plants belong-
ing to the same genotype (more precisely, isozyme phenotype) as those occurring at loc.
no. 47 were found in summer 2009 at Odolena Voda, close to highway (14°24'18.6"E,
50°13'38.3"N, close to locality no. 47) and also at Modrý Důl (15°42'46.0"E,
50°42'45.5"N), in the Krkonoše Mts at an altitude of 1013 m. This indicates that this geno-
type is successful and even able to colonize habitats at high altitudes.
This hybridogenous species is tetraploid (rarely triploid) and apomictic (Rotreklová et
al. 2002, 2005, Suda et al. 2007).
Pilosella setigera Fr. (P. cymosa –P. echioides; syn.: Hieracium fallax Willd.) (Fig. 8)
Localities: 30, 40.
DNA ploidy level and/or chromosome number were not analyzed.
Mode of reproduction, apomictic: loc. 30 (3 plants).
Křišťálová et al.: Pilosella species in ruderal habitats 449
450 Preslia 82: 437–464, 2010
Fig. 7. – Pilosella rothiana.
Křišťálová et al.: Pilosella species in ruderal habitats 451
Fig. 8. – Pilosella setigera.
This hybridogenous species was detected only at two localities and is certainly not common
in the area studied. Co-occurrence of parental species was not observed. Due to absence of
plants of the parental species and its apomictic mode of reproduction, the expectation is that
this morphotype of P. setigera is fixed and can form apomictic populations in any suitable
place even if parental species are not present. No chromosome number is reported for this
species in the Czech Republic, but pentaploid plants are recorded in other countries (Chrtek
2004).
Pilosella ziziana (Tausch) F. W. Schultz & Sch. Bip. (P. cymosa –P. piloselloides subsp.
piloselloides and P. p. subsp. praealta; syn.: Hieracium zizianum Tausch) (Fig. 9)
Locality: 24, 47.
DNA ploidy level 4x: loc. number 24 (1 plant), 47 (2 plants).
Pilosella ziziana is a hybridogenous species morphologically between P. cymosa and
P. piloselloides (subsp. piloselloides and subsp. praealta). The parental species were not
observed either at the same localities or close by. This and the fact that this species is
apomictic indicate that this morphotype of P. ziziana is fixed and can form apomictic pop-
ulations anywhere within the area studied, like P. setigera mentioned above. At locality
no. 24 there were approximately 10–20 individuals. Triploid (apomictic) and hexaploid
plants are reported from France and tetraploid plants from Slovakia (Rotreklová et al.
2005). The tetraploid and pentaploids are also recorded occurring in Bavaria, Germany
and tetraploid plants in Italy (Schuhwerk & Lippert 2002).
Pilosella densiflora (Tausch) Soják (P. cymosa – P. piloselloides subsp. bauhinii; syn.:
Hieracium densiflorum Tausch) (Fig. 10)
Localities: 10, 16, 30, 40.
All the plants collected from the three closely situated localities in southern part of Prague
and one on the eastern margin of the town are morphologically homogeneous indicating
that P. densiflora is a stable hybridogenous species in this area. In the Czech Republic it is
recorded as tetraploid (and sexual) and pentaploid, and in other countries as triploid and
hexaploid (Chrtek 2004).
Pilosella erythrochrista (Nägeli & Peter) S. Bräutigam & Greuter [P. caespitosa –
P. piloselloides subsp. praealta and subsp. piloselloides; syn.: Pilosella arvicola (Nägeli
& Peter) Soják, Hieracium arvicola Nägeli & Peter].
Locality: 49
In the area studied, this hybridogenous species is only known from this locality. Tetraploid
plants are reported by Rotreklová et al. (2005) in the Czech Republic and tetra- and
pentaploids in other countries (Chrtek 2004).
Pilosella floribunda (Wimm. & Grabowski) Fr. (P. caespitosa >P. lactucella; syn.:
H. floribundum Wimm. & Grabowski)
Locality: 43.
This stabilized hybridogenous type, which occurs mainly in mountain meadows (Chrtek
2004), was also found within the area studied.
452 Preslia 82: 437–464, 2010
Křišťálová et al.: Pilosella species in ruderal habitats 453
Fig. 9. – Pilosella ziziana.
454 Preslia 82: 437–464, 2010
Fig. 10. – Pilosella densiflora.
Pilosella flagellaris (Willd.) Arv.-Touv. (P. caespitosa –P. officinarum; syn.: H. flagellare
Willd.)
Locality: 24.
This species is very probably a local hybrid as it was found at this locality along with both
parents.
Pilosella piloselloides – P. setigera (Fig. 11, 12)
Locality: 6, 24.
DNA ploidy level 5x: loc. 24 (13 plants).
Mode of reproduction, apomictic: loc. 24 (4 plants)
At loc. no. 24 (Fig. 11) hybrid plants were found, which seem to fit the combination of par-
ents P. piloselloides and P. setigera. Because it lacks a stolon the first parent was probably
the rare subspecies, P. p . subsp. praealta. On the other hand, similar plants with long sto-
lons were collected at locality no. 6 (Fig. 12), which indicates that P. p . subsp. bauhinii
was involved in the hybridization. Both these combinations remain to be described and
studied in greater detail.
Discussion
During this research on the genus Pilosella at ruderal localities within the city of Prague,
five basic species (P. officinarum, P. caespitosa, P. cymosa subsp. vaillantii, P. aurantiaca,
P. piloselloides subsp. bauhinii and P. p. subsp. praealta) and 12 intermediate species
(P. brachiata,P. leptophyton,P. ziziana,P. visianii,P. rothiana,P glomerata,P. setigera,
P. floribunda,P. flagellaris,P. densiflora,P. erythrochrista and undescribed hybrido-
genous type between P. setigera and P. piloselloides) were found. Seven of these taxa were
not previously reported from this area (Špryňar & Münzbergová 1998), viz. P. floribunda,
P. glomerata,P. leptophyton,P. piloselloides subsp. praealta,P. visianii,P. erythrochrista
and the undescribed hybridogenous type between P. setigera and P. piloselloides. Except
for the last three taxa, all the others are rather common in other parts of the country and as
a consequence their occurrence in the Prague area is not surprising, especially when one
considers their fast rate of spread. Several of the species are currently spreading through-
out the whole of Central Europe. For instance, P. glomerata, which occurs mainly at high
altitudes, has recently started spreading in Germany (Gottschlich et al. 2006). Pilosella
visianii which is rather common in Prague and could have been here for a long time but
overlooked or was introduced or originated here recently. Two other Pilosella species are
known from the Prague area, which do not occur in ruderal habitats, viz. P. lactucella, and
P. echioides (Špryňar & Münzbergová 1998). The number of intermediate species
reported from Prague is higher than recorded in this study, but most of them were found
more than 80 years ago (Špryňar & Münzbergová 1998). The old data may be unreliable
due to common misidentifications and different and not fully compatible concepts adopted
by past and present authors. Thus, a detailed comparison of the past and present diversity
was not undertaken as this would have required a detailed revision of herbarium speci-
mens, which was not possible in the present study.
Křišťálová et al.: Pilosella species in ruderal habitats 455
456 Preslia 82: 437–464, 2010
Fig. 11. – Pilosella piloselloides × P. setigera.
Křišťálová et al.: Pilosella species in ruderal habitats 457
Fig. 12. – Pilosella piloselloides × P. setigera.
Of the basic and intermediate species, only P. officinarum is sexual, and both tetraploid
and hexaploid cytotypes of this species occur in Prague and its vicinity. This fully corre-
sponds with the recent study for the whole Czech Republic (Mráz et al. 2008). The mode
of reproduction of P. aurantiaca in the Prague area was not studied, but plants cultivated in
gardens and escaping from them are tetraploid apomicts (Chrtek 2004) with one common
clone occurring throughout the whole of central Europe (Fehrer et al. 2005). Half of the
intermediate species have P. officinarum as one of the parents. This fact clearly demon-
strates the exceptional importance of this species in the formation of the whole agamic
complex in Central Europe. Pilosella officinarum has a similar role in the Krkonoše Mts
(Krahulec et al. 2004) and Šumava Mts (Krahulec et al. 2008). In Central Europe, it is the
only widely distributed species, which is regularly sexual at ploidy levels higher than dip-
loid. It is common in both non-ruderal and ruderal habitats. The other sexual species,
P. echioides, is rather rare in the Prague area and the sexual tetraploid of P. piloselloides
subsp. bauhinii has not been found there. In general, apomictic species are at a great
advantage when colonizing new habitats. This is especially true for linear habitats, such as
roads and railways. One seed of a successful genotype can form a big population by seed
dispersal and clonal growth. Some intermediate species without clonal growth are also
successful, like P. visianii. The success of some of the hybridogenous species in the
Prague area can be demonstrated by the occurrence of P. rothiana (the identical isozyme
phenotype) not only in Prague and its close vicinity, but also in the Krkonoše Mts, at an
altitude higher than 1000 m, where it was probably introduced by cars (it grows only
within several meters of areas occasionally used for parking).
It is predicted that large-scale building activities, especially those planned for the
periphery of Prague will result in the creation of new habitats suitable for Pilosella species
and hybrids and extend their spread into the countryside. It would be interesting to follow,
which of them will be successful; whether those that already occur here, or a new taxon.
Easy hybridization within Pilosella favours the evolution of new taxa. New hybrids are
frequently reported, e.g. for the combination P. piloselloides and P. officinarum (the
hybrids correspond to P. brachiata).
Comparison of these results with rare data from the western part of Europe reveals sim-
ilar trends and colonization of new habitats by species of Pilosella. For example,
Gottschlich (1990) reports new localities for Hieracium fallax (= P. setigera) along rail-
ways in Basel, Gottschlich et al. (2004) mention H. caespitosum (P. caespitosa) and
H. glomeratum (P. glomerata) as recently colonizing habitats in Hessen. Gottschlich et al.
(2006) especially record H. caespitosum (= P. caespitosa), H. cymosum “nordische Sippe”
(very close to P. cymosa subsp. vaillantii), H. glomeratum (P. glomerata), H. rothianum
(P. rothiana) and H. zizianum (P. ziziana) as spreading into ruderal habitats; all of which
were also found in Prague.
Acknowledgements
We would like to express our thanks to all colleagues who helped us: Vlasta Jarolímová with counting andphoto-
graphing chromosomes, Pavel Trávníček for editing FCM histograms, Franz Schuhwerk and Günter Gottschlich
for identifying many plants, Institute of Botany for logistic support in the experimental garden. Ivana Plačková
and Adéla Macková helped with the isozyme study, Jan Wild, Jiří Machač and Zdenka Konopová with producing
some of the graphs. Olga Rotreklová and Patrik Mráz are acknowledged for their comments on earlier drafts.
458 Preslia 82: 437–464, 2010
Financial support of the Academy of Sciences of the Czech Republic (AV0Z6005908) and Czech Science Foun-
dation (grants GAČR 206/08/0890 to FK and GAČR 521/08/1131 to VK) is gratefully acknowledged.
Souhrn
Nově budovaná silniční tělesa s velkými terénními zářezy a různě exponovanými svahy poskytují řadu vhodných
stanovišť pro šíření rostlin. Jednou ze skupin, pro které je tento typ stanovišť extrémně vhodný, jsou jestřábníky
rodu Pilosella. V uplynulých letech byla tato skupina podrobně studována podél nových velkých silnic a podél
vybraných železničních tratí na území Prahy (několik lokalit bylo situováno těsně za hranicemi města). U řady se-
braných jestřábníků byla sledována ploidie či počet chromosomů a jejich reprodukční systém, který ukazuje, jaké
má daný základní (nehybridogenní) či hybridogenní druhmožnosti v další evoluci a šíření. Během tohoto studia
bylo nalezeno pět základních druhů: P. aurantiaca,P. caespitosa (4x, 5x), P. cymosa subsp. vaillantii (5x), P. of f i -
cinarum (2n = 36, sexuální; 2n = 54, sexuální; 2n = 63),P. piloselloides subsp. bauhini (2n = 45, 54; obě ploidní
úrovně apomiktické), P. piloselloides subsp. praealta (5x; apomiktické), dále byla nalezena i celá řada hybrido-
genních taxonů a zřejmě i primárních hybridů – P. brachiata (4x; sterilní), P. densiflora, P. erythrochrista,
P. flagellaris, P. floribunda, P. glomerata (5x), P. leptophyton (5x), P. rothiana (4x, apomikt), P. setigera,P. zizia-
na (4x) a P. visianii (4x; apomikt). Kromě uvedených druhů byl nalezen ještě dosud nepopsaný hybridogenní typ
vzniklý křížením P. piloselloides aP. setigera (5x, apomikt). Hybridogenní druh P. visianii je poprvé udáván
z území České republiky; na území Prahy patřil v této době k častěji se vyskytujícím. Podél nových silnic se
mohou šířit i typy donedávna vzácné, jako P. rothiana; tento hybridogenní druh již byl nalezen na těchto
stanovištích i mimo Prahu a její okolí.
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Křišťálová et al.: Pilosella species in ruderal habitats 461
Appendix 1. – List of localities studied (abbreviations of collectors: VK – Veronika Křišťálová; JC – Jindřich
Chrtek, AK – Anna Krahulcová and FK – František Krahulec)
1. Praha-Slivenec: Novořeporyjská highway, SW oriented slope next to the road bridge, N 50°01'11.2", E
14°20'7.3". Detected species: Pilosella piloselloides subsp. bauhinii. VK
2. Praha-Slivenec: Novořeporyjskáhighway, SW oriented slope on the right side, approximately 50 m behind the
bridge towards Barrandov, N 50°01'26.5", E 14°19'6.8". Detected species: P. visianii. VK
3. Praha-Slivenec: Novořeporyjská highway, SW oriented slope close to the bridge, N 50°01'26.4", E
14°19'36.2". Detected species: P. piloselloides subsp. bauhinii,P. rothiana. VK
4. Praha-Řeporyje: Novořeporyjská highway, SW oriented slope next to the exitStodůlky, towards to Řeporyje, N
50°02'31.74", E 14°16'13.56". Detected species: P. piloselloides subsp. bauhinii. VK
5. Praha-Řeporyje: S oriented slope in front of the Billa store, the crossing of Mukařovského and Jeremiášova
streets, N 50°02'24.78", E 14°19'36.18". Detected species: P. piloselloides subsp. bauhinii. VK
6. Praha-Řeporyje, Bavorská street, final stop of bus 219, the locality is situated right in front of the Auto-salon
Honda, SE oriented site, N 50°03'13.92", E 14°18’ 37.14". Detected species: P. officinarum,P. piloselloides
subsp. bauhinii,P. brachiata,P. piloselloides ×P. setigera. VK
7. Praha-Řeporyje: S oriented slope in Pekařská street, spot next to the Avia gas station, N 50°03’ 27.42", E
14°20’ 32.70". Detected species: P. piloselloides subsp. bauhinii,P. visianii. VK
8. Praha-Řeporyje: S oriented slope in Pekařská street close to the bakery Odkolek, on the left side of
Rozvadovská street towards the city center, N 50°03’ 22.20", E 14°20'25.50". Detected species:
P. piloselloides subsp. bauhinii,P. visianii. VK
9. Praha-Hrdlořezy: Za mosty street, W oriented slope close to the railway embankment, N 50°05'44.82", E
14°31'20.58". Detected species: P. caespitosa,P. officinarum,P. piloselloides subsp. bauhinii,P. leptophyton.
VK
10. Praha-Kyje: Railway station Praha-Kyje, SW oriented site on the right side of the bridge, N 50°05'46.14",E
14°32'49.68". Detected species: P. officinarum,P. piloselloides subsp. bauhinii,P. brachiata,P. densiflora,
P. glomerata. VK, JC.
11. Praha-Běchovice: East part of the railway station Praha-Běchovice, N 50°04'55.62", E 14°36'12.60".
Detected species: P. leptophyton,P. piloselloides subsp. praealta, P. visianii. VK, FK, JCH
12. Praha-Černý most: E oriented slope, beginning of the highway near the rail bridge, which crosses the highway
from Praha to Liberec, locality is on the right side toward Prague, Exit 1, Horní Počernice, N 50°06'52.44", E
14°35'16.56". Detected species: P. caespitosa,P. officinarum,P. piloselloides subsp. praealta,P. piloselloides
subsp. bauhinii,P. leptophyton,P. visianii. VK
13. Praha-Černý most: SW oriented slope close to Makro-store and KFC on the Ocelkova street (besides
Chlumecká street), N 50°06'35.82", E 14°34'32.64". Detected species:P. piloselloides subsp. bauhinii. VK
14. Praha-Braník: S oriented slope next to the crossing of V podzámčí and Na strži streets, N 50°02'11.76",
E14°26'38.70". Detected species: P. piloselloides subsp. bauhinii,P. rothiana,P. visianii. VK
15. Praha-Braník: S oriented mound along the highway called South highway, also crossing of V Podzámčí and
Na Strži streets, N 50°02'15.06", E 14°27'0.24". Detected species: P. officinarum,P. piloselloides subsp.
bauhinii. VK
16. Praha-Spořilov: SE oriented slope between the railway and Na nivách street, N 50°02'34.98", E 14°27'51.24".
Detected species: P. piloselloides subsp. bauhinii,P. densiflora. VK
17. Jirny: Highway D11, SE oriented slope next to the bridge, on the rightside of the highway (direction Prague),
N 50°07'21.96", E 14°42'13.32". Detected species: P. piloselloides subsp. bauhinii,P. visianii. VK
18. Praha-Hrdlořezy: Ca 0.3 km ESE of the railway station Praha-Libeň, N 50°06'01.2", E 14°30'17.9". Detected
species: P. piloselloides subsp. bauhinii,P. piloselloides subsp. praealta,P. visianii. VK
19. Praha-Veleslavín: Railway station Praha-Veleslavín, left side of track towards Praha-centre. N 50°5'25.20", E
14°20'25.80". Detected species: P. piloselloides subsp. bauhinii,P. glomerata,P. visianii. VK, JCH, FK
20. Praha-Staré Město: Masarykovo nádraží railway station, trackage, N 50°05'23.29", E 14°26'15.42". Detected
species: P. piloselloides subsp. bauhinii,P. leptophyton. VK
21. Řevnice: Beginning of the village of Lety on the way from the town of Řevnice, crossing of main road and
Zahradní street, SE oriented road side, N 49°55'12.24", E 14°14'58.02" Detected species: P. officinarum. VK
22. Praha-Radotín: Area close to the Radotín Cement mill, bus station opposite the mill. N 49°59'40.02", E
14°20'33.48". Detected species: P. piloselloides subsp. bauhinii. VK
23. Praha-Vysočany: SE oriented slope alongside the track, approx. 500 m behind the railway station Praha-
Vysočany towards the centre, N 50°06'50.59", E 14°29'32.20". Detected species: P. piloselloides subsp.
bauhinii,P. brachiata,P. leptophyton. VK
462 Preslia 82: 437–464, 2010
24. Jirny: Highway D11, S oriented slope next to the exit Jirny, left side, N 50°07'19.60", E 14°42'14.00".
Detected species: P. caespitosa,P. officinarum,P. piloselloides subsp. bauhinii,P. brachiata,P. flagellaris,
P. glomerata,P. visianii,P. piloselloides ×P. setigera. VK, JC, AK, FK
25. Praha-Řeporyje: Alongside the ramparts of Novořeporyjská highway (E 50), SW oriented site ca 100 m of the
bridge to Dalejská street, N 50°01'26.89", E 14°19'02.34". Detected species: P. officinarum,P. piloselloides
subsp. bauhinii. VK
26. Praha-Ruzyně: Railway station Praha-Ruzyně, SW oriented slight slope on the left side of the track towards
Praha centre, N 14°18'14.94", E 50°05'05.9". Detected species: P. piloselloides subsp. bauhini,P. glomerata.
VK
27. Praha-Řeporyje: Novořeporyjská highway, SW oriented slope on the right side, 200 m of exit 19 (direction
Řeporyje), N 50°01'30.6", E 14°17'54.9". Detected species: P. visianii. VK
28. Praha-Řeporyje: Novořeporyjská highway, SW oriented slope on the right side, 200 m of exit to Chráštany
(direction Řeporyje), N 50°02'29.40", E 14°16'12.50". Detected species: P. aurantiaca,P. piloselloides
subsp. bauhinii. VK
29. Praha-Prosek: Prosecké skály rocks, rocky spot in the park close to Na rozhraní street, N 50°07'01.1", E
14°29'08.9". Detected species: P. piloselloides subsp. bauhinii. VK
30. Praha-Spořilov: Grassy place at the crossing of streets 5. května and Jižní spojka, N 50°02'31.4", E
14°28'14.3". Detected species: P. caespitosa,P. piloselloides subsp. bauhinii,P. densiflora,P. rothiana,
P. setigera. VK
31. Praha-Vysočany: S oriented site alongside the railway ca 300 m W of the railway station Praha-Vysočany, N
50°06'38.87", E 14°29'03.71". Detected species: P.officinarum,P. piloselloides subsp. bauhinii,P.brachiata,
P. leptophyton. VK, JC, AK, FK.
32. Praha-Šeberov: Exit of the D1 highway, on the E side of the roundabout to Opatov and Šeberov, N 50°01'14",
E 14°30'34.8". Detected species: P. officinarum,P. glomerata. VK
33. Praha-Šeberov: SW oriented site of road mound, 3rd bus station in the village of Šeberov, against to football
ground, N 50°00'36.4", E 14°30'54.9". Detected species: P. officinarum. VK
34. Praha-Horní Počernice: Liberecká highway E65/R10 at the NW marginof the village, W oriented slope on the
right side of the road (direction Liberec), N 50°07'18.3", E 14°36'03.8". Detected species: P. piloselloides
subsp. bauhinii. VK
35. Praha-Opatov: Close surroundings of the Opatov metro station area, N 50°01'37.1", E 14°30'28.6". Detected
species: P. piloselloides subsp. bauhinii. VK
36. Rudná: Slopes along the highway (direction Plzeň), 0.3 km SW of the exit 5, 50°01'21.3" N, 14°12'04.5" E.
Detected species: P. piloselloides subsp. bauhinii. JC.
37. Praha-Hradčany: Corner of the streets Patočkova and Bělohorská, N 50°05'35.0", E 14°23'14.1". Detected
species: P. officinarum. VK
38. Praha-Butovice: Side ditch in the Dalejské údolí valley, N 50°02'41.6", E 14°23'12.8". Detected species:
P. officinarum. VK
39. Praha-Zličín: SE oriented grassy slope in angle (cross-road) of the streets Makovského and Jeremiášova, N
50°03'46.5", E 14°18'46.5". Detected species: P. glomerata. VK
40. Praha-Spořilov: SW oriented slope of Spořilovská street, close to the road bridge on Hlavní street, N
50°02'48.2", E 14°29'05.9". Detected species: P. piloselloides subsp. bauhinii,P. densiflora,P. rothiana,
P. setigera. VK, JC
41. Praha-Spořilov: W oriented slope close to the garages Kačerov, N 50°02'22.57", E 14°28'54.87". Detected
species: P. piloselloides subsp. bauhinii. VK
42. Praha-Kyje: S oriented grassy site where the streets Broumarská and Rožmberská cross. N 50°05'39.25", E
14°32'47.86". Detected species: P. officinarum. VK
43. Praha-Horní Počernice, left side of the highway direction Olomouc, 3rd km, near the bridge over highway. N
50°6'0", E 14°36'32". Detected species: P. piloselloides subsp. praealta,P. floribunda. VK
44. Praha-Horní Počernice: Highway D11, SE oriented site along the right side of the highway towards Prague, ca
1,5 km ENE of petrol station, N 50°06'41.16", E 14°38'58.37". Detected species: P. officinarum,
P. piloselloides subsp. bauhinii,P. visianii. VK
45. Praha-Hrdlořezy: Old orchard on the right side of the railway towards the station Praha-Libeň, area between
the streets Lísková and Morušová, N 50°05'53.5", E 14°31'22.86". Detected species:P. officinarum. VK
46. Praha-Hloubětín: Grassy place where the Kolbenova and Poděbradská cross. N 50°06'26.99", E 14°33'07.96".
Detected species: P. glomerata,P. rothiana. VK
47. Klíčany: Petrol station on the highway, N 50°12'26.3", E 14°26'08.1". Detected species: P. caespitosa,
P. cymosa subsp. vaillantii,P. piloselloides subsp. bauhinii,P. rothiana,P. ziziana. JC, AK, FK
Křišťálová et al.: Pilosella species in ruderal habitats 463
48. Praha-Dejvice: Podbaba, near the ruin Baba (below the street Nad Paťankou), N 50°07'07.1", E 14°23'’ 27.1".
Detected species: P. rothiana. JC.
49. Praha-Řeporyje: Ridge above the Dalejský potok brook,waste places along the path, ca 800 E of the chapel in
the village. N 50°02'00", E 14°19'15". Detected species: P. erythrochrista. JC
464 Preslia 82: 437–464, 2010
