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Neilreichia 7: 119–155 (2015)
Pulsatilla styriaca (Ranunculaceae) is
a new species for the Bulgarian flora,
and conspecific with P. subslavica
Alexander Renate & Manfred
1 Department of Dendrology, Faculty of Forestry, University of Forestry, 10 Kliment Ochridski
2 Herbarium
GJO
, Abt. Botanik, Studienzentrum Naturkunde, Universalmuseum Joanneum
3 Department of Botany and Biodiversity, University of Vienna, Rennweg 14, 1030 Wien,
Abstract: Pulsatilla styriaca (P. halleri subsp. styriaca), up to now considered endemic to
Styria within Austria, is reported as new for the Bulgarian flora. Earlier, it had been identified as
P. halle ri s. str. (P. halleri subsp. halleri) because it is clearly different from P. rhodopaea (P. halleri
subsp. rhodopaea) distributed in Bulgaria. The only three populations situated in western Sredna
Gora (W Balkan mts.) are small and have been monitored during the period 1998–2013. Population
sizes are decreasing and the species thus endangered in Bulgaria. By morphological (phytographi-
cal) evidence, using features of traditional Pulsatilla taxonomy, the differences between the Bulgar-
ian and Styrian populations of P. styriaca, in respect to the variation amplitude, proved to be negli-
gible. This taxon, however, turned out to be conspecific with P. subslavica distributed endemically
in Slovakia. This is demonstrated by a comparative survey of Slovak specimens attributed to this
species and specimens of P. styriaca from Styria. Consequently, P. st yriaca is no longer endemic
to Styria and Austria but exhibits a highly disjunct distribution range covering western central Slo-
vakia, eastern Austria (Styria) and western Bulgaria. Bulgarian, Slovakian and Austrian habitats of
this species are compared und the conservation status is discussed.
Key words: Pulsatilla styriaca; Pulsatilla halleri subsp. styriaca; Pulsatilla halleri agg.;
Pulsatilla subslavica; Pulsatilla slavica; Pulsatilla rhodopaea; Pulsatilla grandis; new species of
Bulgaria; disjunct distribution range; endemism; Bulgaria; Slovakia; Styria
Zusammenfassung: Pulsatilla styriaca (Ranunculaceae) ist eine neue Art der
bulgarischen Flora und konspezifisch mit P. subslavica
Pulsatilla styriaca (P. halleri subsp. styriaca) wurde als drei kleine, benachbarte Populationen in
der westlichen Sredna Gora (Balkangebirge) gefunden. Die Sippe war ursprünglich als P. halleri
(P. halleri subsp. halleri) identifiziert worden, da sie sich deutlich von der in Bulgarien längst be-
kannten P. rhodopaea (P. halleri subsp. rhodopaea) unterscheidet. Sie stimmt jedoch mit den stei-
rischen Exemplaren der P. styriaca – in Anbetracht der Variationsbreite – völlig überein. Die Popu-
lationsgrößen nehmen von 1996 bis 2013 ab, sodass sie in Bulgarien gefährdet ist. Phytographische
Vergleiche anhand der in der Pulsatilla-Taxonomie üblichen Merkmale zeigten, dass P. styriaca
mit der bisher als slowakischen Endemiten betrachteten P. subslavica konspezifisch ist. Somit ist
P. st yriaca nicht länger ein steirischer und österreichischer Endemit, sondern weist ein deutlich
disjunktes Areal auf, das von der mittleren Slowakei über das östliche Österreich (Steiermark) bis
ins westliche Bulgarien reicht. Die Habitate der P. styriaca (s. lat.) in Bulgarien, der Slowakei und in
Österreich werden verglichen und der Naturschutzstatus erörtert.
120 &
Introduction, Materials and Methods
Objective
During floristic investigations in the western Sredna Gora by the first author, when
studying the locality of Anthemis argyrophylla – a critically endangered and protected
species, local endemic and tertiary relict –, a population of a Pulsatilla was found which
was preliminarily identified as Pulsatilla halleri (s. str.) (Anemone halleri) (
2008). Later, two more small populations were found, a survey on the literature and the
herbarium collections in Bulgaria and in Austria was performed and several Floras were
consulted: & (1970), & (1974: 221),
T & (1993), (1996), & al. (2008). By this, evidence
accumulated that these Bulgarian populations belong to P. styriaca, a species so far
considered endemic to Austria.
The habitat of the Bulgarian locality was identified according to & al.
(2005) and the Interpretation Manual for the habitats in the European Union, Eur 15.2.
(2002). The code followed
NATURA
2000 (
HD
-Code), European Nature Information
System (
EUNIS
database v. 2) and “Classification of the Palaearctic habitats” (
PAL
.
CLASS
), version 1996.
The second author, curator of the phanerogams in the state herbarium of Styria,
compared those Bulgarian specimens with plants not only of P. styriaca from Styria but
also with specimens of P. subslavica from Slovakia, because the new Bulgarian popula-
tions seem to look as closely to P. styriaca as to P. subslavica. So, the questions arose,
(1) whether both these species might be conspecific, separated just by a geographic
disjunction, and (2) whether and to what extent the three entities differ in those features
traditionally used in the taxonomy of Pulsatilla.
Materials
Pulsatilla “styriaca” fide A. Tashev from Bulgaria: 4 specimens (2 flowering, 2 fruiting;
herbarium A. Tashev, Sofia), used for diagnostic comparison with herbarium material in
the Bulgarian herbaria
SOM
,
SO
and
SOA
as well as in the Viennese hebaria W,
WU
,
WHB
and
W F BVA
. – Pulsatilla styriaca from Styria: 66 specimens (42 flowering, 15
fruiting, 9 vegetative;
GJO
), used for diagnostic comparison. – Pulsatilla subsla vica
from Slovakia: 62 specimens (18 flowering, 35 fruiting, 11 vegetative;
SAV
), used for
diagnostic comparison. – Pulsatilla slavica from Slovakia: ca. 30 specimens checked
(
SAV
). – Pulsatilla halleri s. str.: ca. 10 specimens checked (
GJO
,
GZU
). – Pulsatilla
rhodopaea from Bulgaria: 3 specimens checked (herbarium A. Tashev, Sofia).
Methods
For comparison of the Bulgarian populations with Pulsatilla styriaca and with P. sub-
slavica 27 features, most of them used by (1958) are analyzed on the basis of 132
specimens and the data arranged in Table 1. That student of Walter Zimmermann’s Pul-
satilla research group ( 1935–1939, 1958, & 1957)
Pulsatilla styriaca, new for Bulgaria 121
emphasized strong variation in all the taxa of P. halleri group and recommended the use
of mean values. Following her, our second author calculated arithmetic means as well.
Taxonomy of Pulsatilla halleri group
In their monograph of the genus Pulsatilla, & (1957) include
P. styriaca together with P. halleri, P. velezensis, P. taurica, P. slavica, P. grandis, and
P. vulgaris in their subsect. Vulgares within sect. Pulsatilla. This narrow species con-
cept is followed by, e.g., (1982) and & (1973).
(1958) in her comprehensive and detailed study sinks P. halleri, P. styriaca, P. slavica,
and P. taurica to subspecific rank summarizing them within P. halleri s. lat. and adding
subsp. macedonica and subsp. rhodopaea. This wider species concept is followed by, e.g.,
& (1970), & al. (1989), and & (1993).
The present paper, however, follows, for the time being, the narrow species concept:
At species rank, these taxa all together (excluding P. vulgaris and P. grandis), according
also to several more modern authors (e.g., & 1973) form P. hal-
leri agg.
Pulsatilla halleri group (= P. halleri s. lat., P. halleri agg., “Spec. coll. [= ‘Sammel-
art’] Halleri” sensu & 1957 without presenting any diagnosis)
is, however, difficult to delimit from P. vulgaris group (P. vulgaris agg., “Spec. coll.
Vulgaris”), that is why both are united by & (1974: 216) as
Table 1: Excer pt of Table 9 in & (1957: 168), listing differential characters
(translated from German by authors R.H. and M.A.F.). — Tab. 1: Auszug aus Tab. 9 in &
(1957: 168). Liste der Unterscheidungsmerkmal zwischen den beiden Arten.
P. styriaca
P. slavica sensu
&
(1957)
Diameter of flower 7 cm 8 cm
Length of involucrum 3 cm 3.5 cm
Number of involucrum lobes 16 18
Scape (below involucrum) length at anthesis 7 cm 8 cm
Width of leaf lobes 1 cm 1.2 cm
Number of leaf lobes about 50 fewer than 50
Further traits basal leaf segments
usually slightly petiolulate;
catadromous segment of
2nd order slightly separated
leaves often with 1 pair of
segments only;
leaf segments relatively
strongly connate
Distribution endemic to Styria endemic to Slovakia: region
122 &
“P. vulgaris-Artengruppe”. (1958) does not provide any description of her P. hal-
leri (s. lat.). &’s (1993: 266) description mentions: “Stems 3–12 cm (up
to 45 cm in fruit). Basal leaves persistently and often densely lanate, 1-pinnate with
3–5 segments, the terminal long stalked, segments pinnatifid; lobes oblong-lanceolate;
cauline leaves united below, sericeous. Flowers 5.5–8.5 cm in diameter, campanulate,
erect or suberect, dark violet; perianth segments usually straight, acute, 2–3 times as
long as the stamens.” – & (1970) characterize “P. halleri ”
(comprising subsp. halleri and subsp. rhodopaea) in the species key by basal leaves
1-pinnate or rarely 2-pinnate with 3–5 basal segments. – &
(1974: 217, 220) characterize their “P. halleri ” (comprising subsp. halleri and subsp.
styriaca) by leaf segments (2–)6–11(–22) mm wide; basal leaves not fully developed
when flowering, strongly hairy and shining silky when young, later on villous, segment
tops with a conspicuous hair tuft; leaves comparatively weakly pinnate; tepals usually
6 and 3–5 cm long, outer ones usually longer, mainly purple-violet; top of styles violet;
fruitlets 5–5.5 mm long, the style (in fruit) 35–40 mm long and strongly hairy; chromo-
somes: 2n = 32. For most of these features no comparable counterparts are provided in
P. grandis and P. vulgaris.
This species group inhabits dry stony places in bushes, exclusively on limestone.
Its distribution includes central (Alps, western Carpathians) and southeastern Europe
(Balkan Peninsula, Crimea). In Bulgaria the species group reaches the easternmost
peripheral part of its distribution area ( 1958: 3, & 1970:
108, & 1993: 266). The species group, therefore, belongs to the Alpine-
Carpathian-Balkan floristic element (“Alp-Carp-Bal”). The present distribution of Pul-
satilla rhodopaea (= P. halleri subsp. rhodopaea) in Bulgaria – the only taxon within
Pulsatilla halleri group reported before – includes Middle Rhodopes and eastern Stara
planina ( & 1970, 1975, 1992,
2002, & 2003, & 2006). According to
(& 2006) its vertical distribution covers the range 300–1400 m a. s. l.,
and according to & (2003) P. rhodopaea occurs within the
range 500–1500 m a. s. l.
Pulsatilla styriaca (Pritz.) Simonk. in Magyar Bot. Lapok 5 (5–7): 178 (1906) (see Figs.
1–4).
Anemone halleri var. styriaca Pritz. in Linnaea 15: 575 (1841)
Anemone styriaca (Pritz.) Hayek in Oesterr. Bot. Z. 52: 477, 479 (1902)
Pulsatilla halleri subsp. styriaca (Pritz.) Zämelis in Acta Horti Bot. Univ. Latv. 1 (2):
104 (1926)
Pulsatilla slavica G. Reuss, Slov.: 5 (1853), non sensu&
(1957) (see Figs. 5–7).
P. halleri subsp. slavica (G. in Acta Hort Bot. Univ. Latv. 1 (2): 104
(1926)
Pulsatilla styriaca, new for Bulgaria 123
Anemone wahlenbergii Szontagh in Verh. k. k. Zool.-Bot. Ges. Wien 13: 1082 (1863),
nom. illeg.
Pulsatilla wahlenbergii (Szontagh) Baenitz (1895), nom. illeg.
P. halleri subsp. slavica var. wahlenbergii Aichele & Schwegler in Feddes Repert.
Spec. Nov. Regni Veg. 60: 1–230 (1957)
P. slavica var. wahlenbergii (Szontagh) Dostál in Folia Mus. Rerum Nat. Bohemiae
Occid., Bot. 21: 6 (1984), nom. illeg.
Pulsatilla subslavica Futák ex K. Goliášová in Biológia, A (Bratislava) 36: 867–868
(1981) (see Figs. 8–11).
= P. slavica auct., non G. Reuss
= P. slavica subsp. slavica var. slavica sensu & (1957) and
(1958), typo excl.
= P. slavica subsp. subslavica (Futák) Dostál in Folia Mus. Rerum Nat. Bohemiae
Occid., Bot. 21: 6 (1984)
Fig. 1: Herbarium specimens of fruiting Pulsatilla styriaca from Styria (Austria), Peggauer Wand N of
Graz, leg. E. Korb, 20 June 1908, and leg. K. Fritsch, 3 June 1902 (W). — Abb. 1: Herbarbelege von fruch-
tenden Pulsatilla styriaca aus der Steiermark (Österreich), Peggauer Wand N von Graz, leg. E. Korb,
20. Juni 1908 und leg. K. Frit sch, 3. Juni 1902 (W).
124 &
Fig. 2: Herbarium specimens of flowering and vegetative Pulsatilla styriaca from Styria (Austria), near
Peggau, leg. Eu. v. Pittoni (
GJO
). — Abb. 2: Herbarbelege einer blühenden und einer vegetativen Pulsa-
tilla styriaca aus der Steiermark (Österreich), bei Peggau, leg. Eu. v. Pittoni (
GJO
).
Pulsatilla styriaca, new for Bulgaria 125
Fig. 3: Herbarium specimen of flowering and postfloral Pulsatilla styriaca from Styria (Austria), rocks
near Gratwein, leg. Fürstenwärther (
GJO
). — Abb. 3: Herbarbelege eines blühenden und eines postflora-
len Exemplars der Pulsatilla styriaca aus der Steiermark (Österreich), Felsen bei Gratwein, leg. Fürsten-
wärther (
GJO
).
126 &
Fig. 4: Herbarium specimens of flowering and fruiting Pulsatilla styriaca from Styria (Austria), Peggau
(
GJO
). — Abb. 4: Herbarbelege blühender und fruchtender Pulsatilla styriaca aus der Steiermark (Öster-
reich), Peggau (
GJO
).
Pulsatilla styriaca, new for Bulgaria 127
Fig. 5: Herbarium specimen of fruiting Pulsatilla slavica from Slovakia, leg. Hubová, Kmêtová and
Futák (
SAV
). — Abb. 5: Herbarbeleg einer fruchtenden Pulsatilla slavica aus der Slowakei, leg. Hubová,
Kmêtová and Futák (
SAV
).
128 &
Fig. 6: Herbarium specimen of fruiting Pulsatilla slavica from Slovakia, leg. Fabianková (SA
V
). —
Abb. 6: Herbarbeleg einer fruchtenden Pulsatilla slavica aus der Slowakei, leg. Fabianková (
SAV
).
Pulsatilla styriaca, new for Bulgaria 129
Fig. 7: Herbarium specimen of fruiting Pulsatilla slavica from Slovakia, leg. Fabianková (
SAV
). —
Abb. 7: Herbarbeleg einer fruchtenden Pulsatilla slavica aus der Slowakei, leg. Fabianková (
SAV
).
130 &
Fig. 8: Herbarium specimen of a vegetative Pulsatilla styriaca (“P. subslavica”) from Slovakia, leg. &
det. as P. slavica, later rev. as P. subslavica by J. Futák (
SAV
). — Abb. 8: Herbarbeleg einer vegetativen
Pulsatilla styriaca (“P. subslavica”) aus der Slowakei, leg. & det. als P. slavica, später rev. als P. subsla-
vica von J. Futák (
SAV
).
Pulsatilla styriaca, new for Bulgaria 131
Fig. 9: Herbarium specimens of vegetative Pulsatilla styriaca (“P. subslavica”) from Slovakia (
SAV
).
— Abb. 9: Herbarbelege von vegetativen Pulsatilla styriaca (“P. subslavica”) aus der Slowakei (
SAV
).
132 &
Fig. 10: Herbarium specimens of vegetative Pulsatilla styriaca (“P. subslavica”) from Slovakia, from the
same locality as the specimen in Fig. 14, but with narrower leaf segments, leg. & det. as P. slavi ca, later rev.
as P. subslavica by J. Futák (
SAV
). — Abb. 10: Herbarbelege von vegetativen Pulsatilla styriaca (“P. sub-
slavica”) aus der Slowakei, von derselben Fundstelle wie das Exemplar in Abb. 14, aber mit schmäleren
Laubblattabschnitten, leg. & det. als P. slavica, später rev. als P. subslavica von J. Futák (
SAV
).
Pulsatilla styriaca, new for Bulgaria 133
Fig. 11: Herbarium specimens of vegetative Pulsatilla styriaca (“P. subslavica”) from Slovakia, from the
same locality as specimens in Fig. 13, but with wider leaf segments, leg. & det. as P. slavica, later rev. as
P. subslavica by J. Futák (
SAV
). — Abb. 11: Herbarbelege von vegetativen Pulsatilla styriaca (“P. sub -
slavica”) aus der Slowakei, von derselben Fundstelle wie die Exemplare in Abb. 13, aber mit breiteren
Laubblattabschnitten, leg. & det. als P. slavica, später rev. als P. subslavica von J. Futák (
SAV
).
134 &
Differential characters between P. styriaca, P. subslavica and P. slavica
according to taxonomic and floristic literature
& (1993: 266) as P. halleri subsp. styriaca: “Like subsp. [halleri] [i. e.:
plants usually more than 5 cm at anthesis; primary divisions of the basal leaves usually
5; lamina with fewer than 50 lobes, more or less lanate], but the lamina of the basal
leaves 5–11 cm. 2n = 32.” P. subslavica is not mentioned even in synonymy.
(1996: 88) describes the leaves of P. styriaca as 1-pinnate with 7–20 lobes,
6–11 mm wide.
& (1974: 221): P. halleri subsp. styriaca in contrast to
subsp. halleri: Plants in flower usually about 10 cm high (subsp. halleri: usually below
10 cm); tepals usually longer than 35 mm (h.: usually shorter than 35 mm); cauline
leaves at average 35–45 mm long (h.: 25–30 mm), hairs on flower and leaf 4–5 mm
(h.: on upper side of basal leaves after flowering and at fruiting time about 4 (2–4) mm
long). – The authors draw attention to the fact that “subsp. styriaca” is (because of gene
exchange) genetically more close to the geographically adjacent P. grandis than to the
geographically and altitudinally distant “subsp. halleri”, and that be a similar situation
like the contacts between P. grandis and subsp. slavica in the Carpathians.
(1902: 479) writes: The characters of “Anemone Styriaca” are intermediate
between “A. Halleri” und “A. grandis”: The leaves are distinctly pinnate with usually
three pairs of segments of 1st order, but the terminal segment is often sessile and there-
fore not distinctly separated from the uppermost segments of 1st order; the lobes are
wider than in A. grandis and, when young, densely silky villous, and when old still
amply hairy, particularly the petioles are fairly strongly hairy even when old. There is
no difference in the flower between all three species.
(1908: 371, as Anemone stiriaca) in his Styrian Flora mentions: Stems up to
30 cm high, patently villous; basal leaves long petiolate, imparipinnate with usually two
(rarely one) pairs of segments [pinnae], terminal leaflet usually sessile and palmately
3–7-fid with deeply dentate segments, the lateral ones usually bifid with multifid deeply
dentate segments, segments of last order [lobes] 4–8 mm wide; young leaves silky vil-
lous, when adult hairs densely appressed; flower erect. – Pulsatilla halleri is not treated
because absent from Styria.
& (1957: 168, 178–182) present detailed descriptions well
comparable with those of P. slavica and P. halleri. Differences between P. styriaca
and P. slavica, in our opinion, are minute and by far not sufficient for accepting two
different species (Tab. 2). Their summary (on p. 181) that P. styriaca is “well separable”
from the other species “morphologically as well as before all geographically” seems to
us justified only in respect to “geographically”.
Pulsatilla slavica, according to in & (1957), comprises two
varieties: var. slavica (basal leaves with 2–3 narrow segment pairs) and var. wahlen-
1 misprint on p. 175: “laciniis foliorum latis” should read – according to the German text – “… angustis”
Pulsatilla styriaca, new for Bulgaria 135
bergii (basal leaves with 1–2 segment pairs). Pulsatilla slavica var. slavica is said to
hold an intermediate position between P. slavica var. wahlenbergii and P. styriaca.
(1958) emphasizes the differences between “P. Halleri subsp. styriaca” and
“P. Halleri subsp. slavica”: Table 2. Also her values show that “styriaca” holds an inter-
mediate position between “slavica” and “wahlenbergii”.
It is important to note that the mean values in Tables 2 and 3 are rather insignificant
because variance in all values is generally very high as clearly shown especially by
(1958) who, in her statistical tables, presents the amplitudes as well.
Table 2: Differential characters mentioned by (1958: 34–35, 41) between three infraspecific
taxa of her “P. halleri”: P. styriaca and the two races within “P. slavica” (s. lat.). — Tab. 2: Unterschei-
dungsmerkmale zwischen den drei infraspezifischen Taxa der „P. halleri“ nach (1958: 34–35,
41): P. st yr iaca und die beiden Rassen innerhalb ihrer „P. slavica“ (s . l at .).
subsp. styriaca
subsp. slavica
var. slavica
= P. subslavica sensu
(1981, 1985)
and (1982)
subsp. slavica
var. wahlenbergii
= P. slavica sensu
(1981, 1985)
and (1982)
Plant size low height: max.:
14 cm in flower,
34 cm in fruit
18 cm: higher in flower,
but lower in fruit
17 cm: higher in flower,
but lower in fruit
Leaf length at
begin of flowering 4.0 cm 3.2 2.0
Hairs on involucrum
and flower short longer longer
Leaf indument,
especially in fruit strong less strong less strong
Ratio tepal
width/length 0.32, i. e., narrower 0.37, i. e., wider 0.40, i. e., wider
Inner tepals usually shorter
than outer
not shorter? not shorter?
Ratio leaf
width/length 1.3 1.2 1.4
Number of
leaf lobes 47 55 37
2-pinnate leaves 10% no no
Width of lobes 38 mm 37 mm 50 mm
Petiolule of lowest
leaf segments 4.8 mm 3.5 mm 1.4 mm
Length of lowest
rhachis “internode” 26 mm 27 mm 16 mm
136 &
Table 3: Comparison of diagnostic features in Styrian Pulsatilla styriaca, Bulgarian P. styriaca and Slovakian P. subslavica (= P. styriaca in our opinion).
Numbers in square brackets are specimens studied; bold numbers in round brackets are arithmetic means. Literature data in the 4th and 6th columns. Lobes
= ultimate (smallest) segments of a leaf; pedicel = internode between involucrum and flower; scape = internode below involucrum; involucrum = whorl of
anaphylls below the flower = “stem leaves”; rhachidula = petiolulus. — Tab. 3: Vergleich diagnostischer Merkmale zwischen steirischer Pulsatilla styriaca,
bulgarischer P. styriaca und slowakischer P. subslavica (= P. styriaca unserer Meinung). Zahlen in eckigen Klammern: untersuchte Exemplare; Zahlen in
runden Klammern im Fettdruck: arithmetische Mittelwerte. Daten aus der Literatur in der 4. und 6. Spalte. Lobes = Laubblattzipfel; pedicel = Internodium
(Stängelabschnitt) zwischen Hochblatthülle und Blüte; scape = Stängelabschnitt unter der Hochblatthülle; involucrum = Hochblatthülle (= „Stängelblätter“);
rhachidula = Blättchenstiel.
Character parameter P. styriaca from Styria
P. styriaca from
Bulgaria
P. halleri subsp. styriaca
data by
= P. styriaca
P. subslavica from
Slovakia
P. var. slavica
data by
= P. subslavica
Number of specimens
studied
66 4 about 100 62 about 100
Height of flowering plant
[mm]
90–260 (154) [42] 140–160 (150) [2] 70–200 (135) 68–205 (144) [17] 70–240 (175)
Heigth of stem in fruit
[mm]
155–410 (305) [15] 340–350 (345) [2] – 210–495 (334) [35] –
Length of leaves (
LB
) [mm]
when first flower appears
0–110 (45) [42]
8
LB
19%
6
LB
20–40 = 14%
28
LB
= 67%
0–0 (0) [2]
2
LB
100%
(leaf buds!)
10–70 (40) 0–55 (16) [16]
10
LB
63%
3
LB
20–40 = 19%
3
LB
= 19%
10–60 (32)
Length of leaf lamina [mm] 28–115 (68) [23] 105–115 (110) [2] 50–180 (83) 50–130 (82) [45] 50–180 (91)
Width of leaf lamina [mm] 40–160 (88) [23] 120–125 (123) [2] 49–220 (110) 55–150 (99) [45] 60–180 (109)
Number of lobes 30–65 (47) [23] 37–64 (51) [2] 15–100 (47) 27–90 (49) [45] 20–130 (55)
Number of 1st-order-
segments
1–3 pairs (1.9) [21]
1 pair: 4× = 19%
2 pairs: 16× = 76%
3 pairs: 1× = 4.8%
2–3 pairs (2.5) [2]
2 pairs: 1× = 50%
3 pairs: 1× = 50%
– 1–3 pairs (2.2) [45]
1 pair: 1× = 2.2%
2 pairs: 32× = 71%
3 pairs: 12× = 27%
–
Degree of pinnate leaf
division (1× = 1-pinnate,
2× = 2-pinnate,
3× = 3-pinnate)
1–2× (1.5) [23]
1×: 12× = 52%
2×: 11× = 48%
2–2× (2) [2]
2×: 2× = 100%
2–4 (3.3) 1–3× (1.9) [45]
1×: 7× = 16%
2×: 37× = 82%
3×: 1×= 2.2%
3–4 (3.5)
Pulsatilla styriaca, new for Bulgaria 137
Width of 1st-order-segments
[mm]
5–17 (10) [23] 8–10 (9) [2] 5–22 (11) 4–20 (11) [45] 5–15 (10)
Length of petiole [mm] 50–210 (122) [23] 90–160 (125) [2] 50–300 (154) 70–240 (151) [45] 50–250 (139)
Length of lowest rhachis
“inter-nodium” [mm]
4–40 (21) [23] 35–50 (43) [2] 0.1–70 (26) 10–45 (24) [45] 0.1–80 (27)
Length [mm] of lowest
rhachidula “internodium”
0–10 (1.1) [23] 3–3 (3) [2] 0–24 (1) 0–13 (1.4) [45] 0–4 (0.5)
Width of terminal lobe of
lowest segments [mm]
1.5–4 (2.8) [23] 4–4 (4) [2] 2–9 (3.8) 2–6 (3.5) [45] 2–6 (3.7)
Length of petiolule of lowest
1st-order-segment [mm]
0–12 (3.2) [23] 5–14 (9.5) [2] 0–25 (4.8) 0–15 (2.5) [45] 0–21 (3.5)
Opposite 1st-order-segments 22 [out of 23] 96% 1 [out of 2] 50% – 11 [out of 45] 24% –
Alternate 1st-order-segments 1 [out of 23] 4.4% 1 [out of 2] 50% 1.5% 34 [out of 45] 76% 0%
Upper section of rhachis
present
15 [out of 23] 65% 2 [out of 2] 100% 70% 30 [out of 45] 67% 45%
Length of leaf hairs [mm] 2–4 (2.3) [23] 3–3 (3) [2] 3 0–2.5 (1.6) [45] 2.5
Length of scape (in fruit)
[mm]
60–150 (94) [15] 110–110 (110) [2] 50–200 (100) 40–190 (121) [35] 50–220 (105)
Length of involucrum
(at anthesis [mm]
20–50 (34) [42] 26–38 (32) [2] 20–57 (36) 20–37 (31) [18] 25–65 (45)
Length of involucrum hairs
[mm]
3–5 (4) [57] 5–6 (5.8) [4] 3–6 (4) 2–6 (4.3) [53] 4–10 (6)
Length of pedicel [mm] 90–315 (210) [15] 230–240 (235) [2] 120–400 (240) 115–370 (213) [35] 120–330 (210)
Length of tepals [mm] 25–50 (37) [42] 42–48 (45) [2] 20–54 (38) 30–45 (38) [18] 30–65 (45)
Width of tepals [mm] 9–18 (13) [42] 12–12 (12) [2] 8–23 (13) 9–19 (14) [17] 8–23 (16)
Length of tepal hairs [mm] 2–4 (2.7) [39] 4–5 (4.5) [2] 1–6 (2.5) 1.4–4 (2.6) [17] 2–8 (5)
Length of anthers [mm] 1–1.5 (1.1) [20] 2–2 (2) [1] – 1–1.8 (1.3) [16] 1.3–2.2 (1.8)
138 &
Pulsatilla slavica (s. str.) sensu Futák (P. halleri subsp. slavica var. wahlenbergii)
differs by its distinctly wider leaf segments as shown by photographs in (1958:
Taf. 1. Abb. 69) and the drawing by & (2011: 104).
& (1993: 266) characterize P. halleri subsp. slavica by “primary
divisions of the basal leaves usually 3, sessile; lamina with usually fewer than 50 lobes,
more or less lanate”. – Pulsatilla subslavica is missing as a taxon as well as a synonym.
(1958: 45–47) describes P. halleri subsp. slavica var. wahlenbergii (= P. sla-
vica sensu Futák) in her keys to the infraspecific taxa of P. halleri: Second leaf rhachis
“internodium” usually missing, segments usually wider than 15 mm, tepals usually longer
than 42 mm, ungainly (“plump”) with width/length 42.2 in average, anthers suborbicular.
In contrast, P. halleri subsp. slavica var. slavica (= P. subslavica) is described by
second leaf rhachis “internodium” usually present, segments usually narrower than
15 mm, terminal leaf segment narrower than 4 mm (wider in styriaca), leaf lamina at
least 9 cm long in average (shorter in styriaca), relatively narrow (in comparison to sty-
riaca), segments (5–)10(–15) mm wide, with more than 50 lobes, tepals usually longer
than 42 mm, to “slender” width/length of tepals 35.5 in average, anthers “oblong”.
Pulsatilla subslavica (Figs. 8–11) comprises populations with leaves the segments
of which are narrower than in P. slavica s. str. (= sensu 1982) but wider than in
P. grandis. Most of the herbarium specimens studied by the second author, in respect to
this character, exhibit the same variation amplitude like in P. styriaca. Almost all spec-
imens of P. slavica s. str., however, show distinctly wider segments than in P. styriaca.
In the key by (1982: 111), P. subslavica, P. slavica, and P. grandis are com-
pared in parallel:
Pulsatilla subslavica: Basal leaves not more than 2-pinnate or -pinnatisect, lobes
7–12 mm wide; segment pairs usually 3.
Pulsatilla slavica: Basal leaves 1-pin nate, lobes 12–25 mm wide; segment pairs 1 or 2.
Pulsatilla grandis: Basal leaves 2–3-pinnate or -pinnatisect, lobes 2–6(–7) mm
wide; segment pairs 4 or 5.
The description of P. subslavica in (1982) reads: Basal leaves at anthesis
usually not developed, reaching their definite shape towards end of summer. Leaf lam-
ina ± broadly ovate in outline, length like in P. slavica [no length given there], tomen-
tose when young, later dispersedly hairy, unpaired pinnate with usually 4 pairs of leaf-
lets, the lower leaflets sometimes 2-pinnate at base, terminal leaflet usually stalked,
3-partite, segments ± 3-lobed, lateral leaflets opposite, very rarely alternate, sessile,
rarely stalked [petiolulate], 3–5 times rather irregularly pinnatisect to pinnatilobe, lobes
7–12 mm wide. Otherwise like P. slavica. (Translated from Slovak.)
(1981) has investigated in detail that P. slavica shows abundant intro-
gressive hybridization to P. grandis as well as to P. subslavica (see also 1982:
119–129 and & 2011: 98 –99, 104–107). Pulsatilla grandis, in Slo-
vakia, is disjunctly distributed with centres in
SW
Slovakia and E Slovakia. Pulsatilla
slavica has its distribution centre in eastern central Slovakia, reaching to western Tatra
mts. in Poland. Pulsatilla subslavica is endemic to Slovakia, mainly distributed in west-
Pulsatilla styriaca, new for Bulgaria 139
ern central Slovakia, adjacent to and overlapping with the range of P. g randis in western
and in eastern parts of the country. For large regions both P. subslavica and P. slavica
are growing sympatrically and forming hybrids.
Other closely related taxa are P. rhodopaea and P. halleri (P. taurica, P. macedo-
nica, and P. velezensis being omitted here):
Pulsatilla rhodopaea (= P. halleri subsp. rhodopaea): according to &
(1993: 266) rarely taller than 5 cm at anthesis; lamina of basal leaves with 50–100 lobes,
primary segments usually 5, often petiolate, densely lanate. Balkan Peninsula. (See Figs.
12 and 13.) – According to (1958: 46): Leaf lobes usually more than 65, lower-
most rhachis “internodium” usually longer than 4 cm, primary segments (6–)11 (–24) mm
wide. – Description by &
gradsko).” Subsp. halleri:.” (Lower
internodes [= leaf rhachis segments] of the basal leaves longer than 2 cm; leaf segments
60–70. Central Rodops (Asenovgrad district). Subsp. halleri: … shorter than 2 cm; …
segments 40–45. Not present in our country [Bulgaria].)
In Bulgaria, P. rhodopaea occurs in eastern Stara planina and in the central Rho-
dopes (see Figs. 12 and 13). Pulsatilla slaviankae, included in P. rhodopaea by “Flora
Europaea” ( & 1993), is endemic to Slavianka mts. (
SW
edge of Bul-
garia) according to & (1970).
Pulsatilla halleri s. st r. (= P. halleri subsp. halleri) differs from P. styriaca accord-
ing to & (1993: 266): Plants usually taller than 5 cm in anthesis, lamina
of basal leaves 3–7 cm long, primary segments usually 5, sessile, usually with fewer than
50 lobes, more or less lanate. – According to (1958: 46): Plant at anthesis usually
lower than 10 cm, hairs on leaves 4–5 mm long& 1974: 221
further characters: hairs on upper side of leaves after anthesis and in fruit about 4 mm
long, rarely 2–4 mm), lobes usually fewer than 45, lowest petiolule longer than 2 mm,
tepals shorter than 35 mm, stem leaves (involucral leaves) 25–30 mm on average; hairs
on flowers usually shorter than 4 mm.
SW
Alps up to S Switzerland.
Pulsatilla grandis is rather close, though not included in P. halleri s. lat. (= P. hal-
leri agg.). According to & (1993: 264–265) P. vulgaris s. lat. (i. e. in-
cluding P. grandis) differs from P. halleri s. lat. by basal leaves 3- or 4-pinnatisect,
with 7–9 primary segments, plants at first sericeous, becoming glabrous. The trait of
P. grandis that leaves appear after flowers is common with P. halleri s. lat. – According
to & (1974: 217) P. grandis differs from P. halleri s. lat. (i. e.
including styriaca) by leaf lobes (2–)4(–11) mm wide and involucrum lobes ca. 20 in
P. grandis vs. leaf lobes (2–)6–11(–22) mm wide in P. halleri s. lat. – (1958:
34–35), in her character table of different taxa within P. halleri (s. lat.), includes P. gran-
dis. The figures there do not show any conspicuous differences against P. halleri s. lat.
1 included in subsp. rhodopaea by & (1993)
140 &
Fig. 12: Herbarium specimen of fruiting Pulsatilla rhodopaea from the central Rhodopes in Bulgaria,
Abb. 12: Herbarbeleg einer fruch-
tenden Pulsatilla rhodopaea
Wand], A. Ta shev, 16. April 1999.
Pulsatilla styriaca, new for Bulgaria 141
Fig. 13: Herbarium specimen of Pulsatilla rhodopaea from the central Rhodopes in Bulgaria, near the
village Dobrostan, Pinus pallasiana forest, 1235 m a. s. l., A. Tashev, 1 May 2008. — Abb. 13: Herbarbeleg
der Pulsatilla rhodopaea aus den Zentral-Rhodopen (Bulgarien), nächst dem Dorf Dobrostan, Pinus-
pallasiana-Wald, 1235 msm, A. Tashev, 1. Mai 2008.
142 &
Comparison of P. styriaca with P. subslavica
Table 3 shows virtually no significant differences between P. styriaca from Styria and
from Bulgaria and P. subslavica from Slovakia. Before all, the large variation ampli-
tude within all three “taxa” is obvious. The first-glance impression that all these speci-
mens belong to the same species is corroborated by detailed measuring of the character
parameters traditionally considered taxonomically significant. (Figs. 1–4, 8–11)
The leaves show conspicuous ontogenetic heterophylly: The number of lobes in-
creases strongly from the first leaves up to adult leaves developed only much after an-
thesis, towards fruiting time. (1958) mentions that lobe number increases by
doubling from one to the following leaf. Usually specimens are collected when in flow-
er, that is why study of herbarium material is difficult. Comparisons suffer also from
the fact that we had only few specimens from Bulgaria because of the poor populations
there.
Habitats of P. subslavica in Slovakia
Dry, grassy places on calcareous ground from the colline (ca. 350 m s. m.) up to the
montane zone (1000 m s. m.). Coenology: alliances Seslerio-Asterion alpini, Seslerio-
Festucion duriusculae, Quercion pubescentis-petraeae, Erico-Pinion, Festucion valesia-
cae and suballiance Cephalanthero-Fagion ( 1982: 128).
Pulsatilla styriaca in Bulgaria
The comparison of the herbarium specimens of P. styriaca from Bulgaria (Figs. 14–17)
and Austria (W,
WU
,
WHB
,
W F BVA
) by the first author proved their complete identity
(Figs. 1–4, 15–17 ).
Bulgaria, western Sredna Gora (Ihtimanska Sredna Gora):
(1) North-west from Belovo, northern from the Maritza river bed, between Momi-
na Klisura village and Belovo. The slope is of south-eastern exposition and the rock is
GM
-57;
26 Apr. 1998, coll. Alexander Tashev (
SOM
164107).
Same23
GM
-57; 11 May 2008, coll.
Alexander Tashev (W 2009-0003619; http://herbarium.univie.ac.at/database/detail.php
?
ID
=136481).
The locality is situated in the central Rhodopean low-mountaineous climatic district
of the Transitional continental climatic sub-region of the European Continental climatic
region (& 1963; 2005). According to the forest regionalization, it
is located in the Thracian forest region, lower plain and hilly oak forests ( &
al. 1979), and – according to the regionalization of vegetation in Bulgaria – in the belt of
Pulsatilla styriaca, new for Bulgaria 143
Fig. 14: Flowers of Pulsatilla styriaca in western Sredna Gora; A. Tashev, 24 Feb. 2008. — Abb. 14:
Blühendes Individuum von Pulsatilla styriaca im westlichen Balkan-Gebirge; A. Tashev, 24. Feb. 2008.
Fig. 15: Specimen of Pulsatilla styriaca with fully developed leaves close to a tree of Ostrya carpini-
folia; western Sredna Gora; A. Tashev, 6 June 2008. — Abb. 15: Exemplar der Pulsatilla styriaca mit voll
entwickelten Laubblättern, neben einer Hopfenbuche (Ostrya carpinifolia); westliches Balkan-Gebirge;
A. Tashev, 6. Jun i 2008.
144 &
Fig. 16: Herbarium specimen of fruiting Pulsatilla styriaca from western Sredna Gora in Bulgaria, near
Belovo, 450 m a. s. l.; A. Tashev, 10 May 2009. — Abb. 16: Herbarbeleg einer fruchtenden Pulsatilla styri-
aca vom westlichen Balkan-Gebirge, bei Belovo, 450 m s. m.; A. Tashev, 10. Mai 2009.
Pulsatilla styriaca, new for Bulgaria 145
Fig. 17: Herbarium specimen of fruiting Pulsatilla styriaca from western Sredna Gora in Bulgaria, near
Belovo, 385 m a. s. l.; A. Tashev, 11 May 2009. — Abb. 17: Herbarbeleg einer fruchtenden Pulsatilla styri-
aca vom westlichen Balkan-Gebirge, bei Belovo, 385 m s. m.; A. Tashev, 11. Mai 2009.
146 &
xerophyte and mesoxerophyte, microthermic and mesothermic vegetation in the xero-
thermic oak belt and in hilly plains ( 1991). According to the floristic regional-
ization of the country it is in the floristic region western Sredna Gora ( 1966).
The habitat was identified as “Arborescent matorral with Juniperus spp.” (
HD
-Code
5210;
PAL
.
CLASS
.: 32.131) or “[Juniperus oxycedrus] arborescent matorral” (
EUNIS
:
F5.1311;
PAL
.
CLASS
.: 32.1311).), which is a habitat of European importance (“Interpre-
tation Manual for the habitats in the European Union”, Eur 2002).
The habitat is situated at the lower part of a 25° slope. The bedrock is limestone and
the soil is stony, shallow, poor and very dry. The soil type is rendzic leptosols. The orig-
inal secondary plant community dominated by Quercus pubescens with participation
of Ostrya carpinifolia (Fig. 15) and Fraxinus ornus, where P. styriaca was found, had
reached its final stage of degradation. Therefore, in 1955 it was afforested by Black Pine
(Pinus nigra) and single individuals of Scots Pine (Pinus sylvestris). At the moment
there are still several survived individuals or small groups of Black Pine in the lower
part and very few Scots Pine individuals. Also, single trees of Quercus pubescens, Qu.
frainetto, Qu. dalechampii, Ostrya carpinifolia, Fraxinus ornus, Pistacia terebinthus,
Pyrus pyraster participate in the tree composition. There are also small groups or single
individuals of shrubs: Juniperus deltoides, Carpinus orientalis, Paliurus spina-christi,
Cotinus coggygria, Coronilla emerus subsp. emeroides, Rhamnus rhodopaeus, Prunus
spinosa, Rosa obtusifolia, Amelanchier ovalis etc.
The herbaceous layer is formed by more than 50 species, the most typical being
Chrysopogon gryllus, Poa badensis, Stipa pennata, Koeleria nitidula, Carex humilis,
Teucrium polium, Jurinea consanguinea, Achillea clypeolata, Anthemis argyrophylla,
Convolvulus cantabrica, Erysimum diffusum, Onosma taurica, Minuartia rhodopaea,
Inula ascherssoniana, Sedum kostovii, Silene flavescens, Scorzonera austriaca, Sal-
via argentea, Stachys recta, Hypericum rumeliacum, Anthyllis vulneraria subsp. poly-
phylla, Thesium ramosum (Th. arvense), Helianthemum nummularium, Fumana pro-
cumbens, Cytisus (Corothamnus) procumbens, Ononis pusilla, Globularia bisnagarica
(G. aphyllanthes) etc. A clear trend to more dry conditions during the vegetation period
was observed during the years of monitoring, and most probably this is the reason of the
many dried individuals of Juniperus deltoides. It is also possible that the proximity of
the Pulp and paper plant “Belana” has also influenced the process.
Full inventory of the individuals of P. styriaca was performed on 7 May 1998, and
partial inventory on 6 May 2000 and 11 May 2003. The plants grow at south-eastern and
south-western exposition and there is a small group in the gully that separates them. The
total area where the individuals grow was roughly estimated to about 20,000 m².
Most individuals were recorded on the south-eastern slope, at altitude 385 m and geo-
Sixty-four
individuals were recorded in 1998 and 40 of them had generative stems. There were 17
micro-groups consisting of more than 1 individual – 1 group of 5 individuals, 2 groups
1 taxonomy and nomenclature of the taxa mentioned follow (2002)
Pulsatilla styriaca, new for Bulgaria 147
of 6 individuals, 3 groups of 3 individuals and 2 groups of 2 individuals each. The
remaining 20 individuals were solitarily distributed. In 2000 the total number of indi-
viduals was 59 and 5 of them had generative stems. There were 14 groups consisting of
more than 1 individual: 1 group of 8 individuals, 1 group of 5 individuals, 1 group of
3 individuals, and 9 groups of 2 individuals each. The remaining 16 individuals grew
solitary. In 2003 the population consisted of 39 individuals and only 1 had a generative
stem. There were 9 groups of more than 1 individual – 1 group of 5 individuals, 2 groups
of 4 individuals and 6 groups of 2 individuals each. The remaining 14 individuals grew
solit a r y.
These observations show how the number of individuals on the south-eastern slope
has decreased considerably during a 5-year period from 64 to 39, and the individuals
with generative stems from 40 to 1. The same is valid also for the spot in the gully, where
the number of individuals decreased two times. During the period of monitoring, many
damages caused by domestic animals were found, especially on the generative stems.
(2) In the gully, under the canopy of trees of tertiary relic Ostrya carpinifolia, at an
the group: six plants were recorded in 1998, all of them with generative stems. Only 3
individuals remained in 2003 and none had a generative stem.
(3) Two other spots of P. styriaca were found on the south-western slope. The first
-
uals were recorded in 1998 within a group of Juniperus deltoides and below Pinus nigra
canopy and three of them had generative stems. The second one was at 440 m a. s. l. and
under Pinus nigra canopy and near Fraxinus ornus. Nine of them had generative stems.
At the same place six individuals were found in 2003 – all of them with generative stems.
Subsequent observations up to 2007 confirmed the trend of decreasing population
size. In total 42 individuals having 58 generative stems were found in the locality during
the last inventory (24 Feb. 2008).
Conservancy of the Bulgarian sites of Pulsatilla styriaca
The species closest to P. styriaca in the Bulgarian flora, P. halleri, is considered to be of
high conservation importance in Bulgaria. It is listed in the Red Data Book of P. R. Bul-
garia ( 1984) in the category “rare species” and was protected in 1961 under
the name Anemone rhodopaea (“Journal of the Presidium of the National Assembly”, 63,
1961). Its protection status was confirmed by inclusion in the new list of the protected
species under its present name (“State Gazette”, 56, 1989). Finally, it was included in
Appendix 3 of the “Biodiversity Act of Bulgaria” (2002). In the Red Lists of the higher
plants in Bulgaria elaborated in 2005, Pulsatilla halleri was listed as “endangered” (
EN
)
species (& 2009). The species is included under the same status
in the new Red Data Book of the Republic of Bulgaria ( 2011).
148 &
The state of P. styriaca population in the locality has been destroyed during the last
decade, as shown by the results of the periodical observations. Several reasons could
be hypothesized: regular grazing by goats, which is rather common in the region; the
evident xerophytization of the climate during the monitoring period, and the negative
influence by the pulp and paper plant “Belana”, which is close to the locality. Also, high
situated. Therefore, it is proposed the locality to be given a status of protected site. This
necessity is underlined by the fact that most part of the locality of P. styriaca overlaps
with the unique locality of the critically endangered Anthemis argyrophylla.
Distribution and habitats of Pulsatilla styriaca in Styria
The habitats of both the Bulgarian and the Styrian sites of P. styriaca (Figs. 1–4) resem-
ble each other as to bedrock (calcareous) and altitude, while, evidently, accompanying
species are different. The Bulgarian habitat is a submediterranean Quercus pubescens
and Ostrya carpinifolia forest with a number of thermophilic balkan-illyric elements
(see above). The Pulsatilla sites in Styria are characterized by (1981: 112 and
1996: 88) as “lichte blaugrasreiche Rotföhrenwälder an steilen felsigen Hängen und an
Felswänden; submontan bis montan; auf Kalk und Dolomit, nur ausnahmsweise auch
auf benachbarte Tonschiefer übergehend, von 390 bis 1650 m” (Pinus sylvestris forests
rich in Sesleria caerulea (S. varia) on steep slopes on cliffs; submontane to montane,
on calcareous and dolomitic bedrocks, exceptionally on adjacent tonschiefer, 390 to
1650 m s. m.). According to (1951) is Pulsatilla styriaca a characteristic spe-
cies of the “relictic pine-forest with Sesleria”. – Habitat type according to
HD
-Code
(
FFH
-Richtlinie): carbonate-rock-steppes (
FFH
-
LRT
6210, 6240) of the Upper Mur val-
ley and the Grazer Bergland (calcareous mountain range north of Graz) and partly also
“Karbonat-Rotföhrenwälder” (Scots pine forests on carbonate bedrocks).
The Styrian habitats belong to
HD
-Code (
FFH
-Lebensraumtypen) Karbonat-
Fels trockenrasen (
HD
-Code 6210 p. p. = Naturnahe Kalktrockenrasen und deren Ver-
buschungsstadien, 6240 p.p. = Subpannonische Steppentrockenrasen) des Oberen
Murtales und Grazer Berglandes und teilweise auch zum Karbonat-Rotföhrenwald
(Erico-Pinetum sylvestris p. p.; no
HD
-Code).
(1958: 12–13; translated from German) mentions in a phytosociological
relevé:
Locality (1): “Vorberg der Gfäller Wand” near Mautern in Steiermark [Liesing val-
ley]: steep rocky slope with humus pockets, slope 50° S-
SE
, coverage 50%, rocks 40%,
open soil 10%, 1400 m s. m., 50 m below der timberline: Pulsatilla “Halleri subsp. sty-
riaca” [= P. styriaca], Sesleria caerulea (= S. varia), Carex humilis, Globularia cordi-
folia, Euphorbia cyparissias, Galium sp., Teucrium chamaedrys, Cerastium arvense,
Sempervivum montanum (in rock crevices only), Helianthemum sp., Achillea clavennae,
Thymus sp., Lotus corniculatus, Senecio doronicum.
Pulsatilla styriaca, new for Bulgaria 149
Mautern in Steiermark: Mesobrometum on slope of calcareous scree; strongly humose
with several calcareous stones, slope 20° S; coverage 95%; A-horizon ca 20 cm; relevé
area 15 m²: Pulsatilla Halleri subsp. styriaca, Bromus erectus, Festuca “longifolia”
[= ? ], Juniperus communis, Carex montana, Carex ornithopoda, Carex alba, Asperula
cynanchica, Helianthemum sp., Lotus corniculatus, Euphorbia cyparissias, Hippocre-
pis comosa, Prunella grandiflora, Polygonatum “officinale” [= P. odoratum], Potentilla
heptaphylla, Alectorolophus minor [= Rhinanthus minor], Biscutella laevigata, Trifo-
lium montanum, Teucrium chamaedrys, Orobanche teucrii, Plantago media, Plantago
lanceolata, Helleborus niger, Thuidium tamariscinum, Hylocomium rugosum.
(1981; translated from German) enumerates from a site near Stübing, Grat-
wein and Peggau (Mur valley north of Graz) as frequently accompanying: Tree- and
shrub layer: Pinus sylvestris, Viburnum lantana, Ligustrum vulgare, Amelanchier ova-
lis, Rhamnus cathartica; herb layer: Sesleria caerulea (= S. varia), Potentilla arenaria
[= P. incana], Carex humilis, Erysimum sylvestre, Seseli austriacum, Leontodon incanus,
Thymus praecox, Anthericum ramosum. – At the Peggauer Wand (near the town Peggau)
additionally: Arthemisia campestris, Scorzonera austriaca, Melica ciliata, Geranium
sanguineum, Thalictrum foetidum, Geranium rotundifolium und Minuartia setacea. –
Near the town Stübing additionally Euphorbia verrucosa, Pulsatilla pratensis subsp.
nigricans and Pulsatilla nigricans × P. styriaca (= P. ×weberi).
Discussion of the ecological and phytogeographic position of P. styriaca by
(1981: 79–80; translated from German): “The preference of xerothermic habitats on cal-
careous rocks in the Mur valley with a touch of subcontintental climate points to rela-
tionship with mit continentally spread taxa in eastern Europe and Asia Minor. Accord-
ing to (1973 and 1979), the phytochoria of species at the eastern edge of the
Alps are in accordance to the comparatively insignificant pleistocene glaciation of the
mountains at the eastern edge of the Alps and agree also with the benefits of locally
sunny, warm slopes in the steep valleys of calcareous mountains.” In such climatically
favoured habitats on calcareous bedrocks are also stands of Quercus pubescens.
Distribution in Styria
The distribution of P. styriaca in Styria (see also 1838, 1864, 1868,
1908, 1951, 1963, F. & H. 1967,
1981, & al. 1989, 2009) is verbally summarized by (1996:
88): “… endemisch im Grazer Bergland beiderseits der Mur von Gratkorn bis Grat-
wein bei Graz nordwärts bis zum Röthelstein und zur Roten Wand bei Mixnitz; weiter
nördlich gegen Aflenz im Thörlgraben, nördlich Leoben im Vordernberggraben und
einigen Seitentälern, besonders bei St. Peter-Freienstein sowie westlich von Leoben
1 taxonomy and nomenclature follow & al. (2008)
150 &
bis St. Michael und den südlichen Ausläufern des Reiting bei Kammern und Mautern.”
(See Fig. 18.)
Conservancy of Pulsatilla styriaca in Styria
& al. (1989: 178, 200) categorize P. styriaca in Styria as “vulnerable”
(rank 3). It is endangered by forestry managements, construction activities, quarries and
collecting; species protection and conservation of biotopes are recommended.
The “Rote Liste gefährdeter Pflanzen Österreichs” (Red List of threatened plants in
Austria) by & (1999) ranks P. styriaca as “gefähr-
det” (vulnerable = rank 3).
According to the Styrian Law for the Protection of Nature (“Steiermärkisches
Naturschutzgesetz”) of 1976 and the Prescription of Species Protection (“Artenschutz-
verordnung”) of 2007, Pulsatilla styriaca
Fig. 18: Distribution grid map of Pulsatilla styriaca in Styria: black circles: records since 1990, currently
confirmed; gray circles: records 1945–1989, currently not confirmed; white circles: records prior to 1945,
currently not confirmed; : 8853/2 (near Judenburg, 1963) and 8359/1 (Neuberg an der Mürz,
2009). – According to: Floristic Mapping of Austria (H. Niklfeld & L. Schratt-Ehrendorfer, Univer-
sity of Vienna). — Abb. 18: Rasterverbreitungskarte von Pulsatilla styriaca in der Steiermark: schwarze
Kreise: Meldungen seit 1990, aktuell bestätigt; graue Kreise: Meldungen 1945–1989, aktuell nicht bestä-
tigt; weiße Kreise: Meldungen vor 1945, aktuell nicht bestätigt; : Ansalbungen: 8853/2 (bei Judenburg,
1963) und 8359/1 (Neuberg an der Mürz, 2009). – Nach: Floristische Kartierung Österreichs
(H. Niklfeld & L. Schratt-Ehrendorfer, Universität Wien).
Pulsatilla styriaca, new for Bulgaria 151
leaf rosettes and underground plant parts must not be removed, whereas a single bundle
of flowers in one hand (“Handstrauß”) is allowed (“von den nicht geschützten Teilen der
Pflanzen ist die Entnahme von mehr als einem Handstrauß verboten”).
Results and Discussion
(1958: 3): “Experimentell sind die einzelnen Formen gut miteinander kreuzbar;
natürlicherweise verhindert aber die räumliche Isolation jeglichen Genaustausch (abge-
sehen von slavica [= subslavica]—Wahlenbergii [= slavica]). […] Jedes dieser Taxa hat
charakteristische Eigentümlichkeiten; das weist auf eine selbständige Geschichte hin.
Da alle Taxa relativ kleine Areale besiedeln, kann der zufällige Allelverlust, die sog.
‘genetic drift’, besonders leicht eintreten; auch Mutationen haben Sippendifferenzierun-
gen begünstigt. Die Rassen sind jedoch so polymorph, […] die Übergänge also gleitend
sind.” (In the experimental garden the taxa hybridize, in nature, however, geographical
isolation prevents gene flow between slavica [= subslavica] and Wahlenbergii [= sla-
vica]. […] Each taxon has characters of its own, indicating independent history. As all
these taxa inhabit relatively small areas, loss of alleles by genetic drift easily can happen,
together with mutations. The taxa are rather polymorphic, […] variation is continuous.”
(1985), in her investigation on the variability of P. slavica, P. grandis
and P. subslavica, states that P. subslavica exhibits, by morphological and anatom-
ical characters of the vegetative and generative organs, an intermediate position be-
tween P. slavica and P. grandis. She concludes that P. subslavica probably originated
by hybridization towards the end of Pleistocene, when both parents came into contact.
Today, P. slavica forms distinct and pure populations in northern Slovakia, mainly in
the Fatra mts. region but also further east; this species ranges from montane to subalpine
altitudes. Pulsatilla grandis shows equally pure populations in southern Slovakia rang-
ing up to eastern Slovakia east of Košice, where P. slavica is missing (maps by
1982: 125, 1985: 94, 160 and & 2011: 104). Its habitats
range from lowland (colline) to submontane altitudes. However, in all contact zones
of the parental species there is ample introgressive hybridization. Pulsatilla subslavica
shows an intermediate position in respect to its vertical range from colline to montane.
The strong similarity, i. e. the broad congruence in all characters considered of taxo-
nomic relevance leads us to the conclusion that P. subslavica and P. styriaca, including
the newly discovered Bulgarian populations must be treated as conspecific. Therefore,
the geographic range of this emended species is split into three regions: a small one
in the West (central Styria), another, even smaller one in the east (Ihtimanska Sredna
Gora in central Bulgaria) and a comparatively large one in central Slovakia which is,
however, likewise disrupted because consisting of a large area in western Slovakia and
several fragments further in the east. Thus, the whole species range is remarkably dis-
junct. The reason for this disjunction could be traced back to Quaternary period. Why
the species survived only in these splitted refugial area – ranging from Austria in the
152 &
west to Bulgaria in the east, remains unclear. Theoretically, however, polytopic origins
of P. styriaca s. lat. cannot be excluded and would need confirmation or exclusion by
genetic data.
As to the distribution of the Bulgarian populations, further studies in the region
should be performed, particularly because there exists information provided by local
people that this plant could be found also in other places in the region. There is high
probability of finding new localities in similar habitats. Also, the monitoring of those
rare and isolated Bulgarian populations must be continued.
Pulsatilla styriaca (s. lat.) seems to be an element within a network of closely related
taxa exhibiting geographical differentiation (different main areas) but phenetically close
and – if geography allows – connected by hybrid zones. In sympatric areas hybridization
seems to be abundant. Typical populations of these taxa, however, are rather distinct
and characterized by a set of – though variable – features (leaf shape, size of flower,
indumentum of plant, phenology); furthermore, there is also ecological differentiation
as to altitude (gradient from subalpine to colline: halleri s. str. – slavica – styriaca s. lat.
– grandis); P. halleri s. str. and P. rhodopaea represent dwarfy mountain ecotypes, evi-
dently. This is enough reason for several authors to attribute specific rank to these enti-
ties, because otherwise all of them would have to be lumped into one single polymor-
phic species. Distinction of two species groups (aggregates: &
1973) or two species (e. g. by & 1993) hardly seems acceptable to
us. Therefore we rather follow & (1974) proposing a large
“P. vulgaris Artengruppe” complying with P. sect. Pulsatilla subsect. Vulgares sensu
& (1957).
This network includes at least seven taxa, all of them tetraploid (see Fig. 19). (The
diploid species P. patens, P. pratensis, and P. montana are excluded.)
Fig. 19: Reticulate relations within the tetraploid Pulsatilla vulgaris s. latiss. group. — Abb. 19: Bezie-
hungen innerhalb der tetraploiden Gruppe der Pulsatilla vulgaris s. latiss.
vulgaris s. str.
oenipontana
grandis slavica
subslavica
styriaca
halleri s. str.
taurica
rhodopaea
macedonica
slaviankae
velezensis
phenetically close
phenetically close and
genetic contacts (gene flow, hybrids)
Pulsatilla styriaca, new for Bulgaria 153
Acknowledgements
We are thankful to Kornélia Goliašová for providing the loan of herbarium material from
SAV
and for
help in providing literature, and to Harald Niklfeld for additional information on the Styrian distribution
range of P. styriaca.
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