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ISSN 2226-3063 e-ISSN 2227-9555
Modern Phytomorphology 11: 51–89, 2017
doi: 10.5281/zenodo.834040
© The Author(s) 2017. Published by Novikoff A.V., State Natural History Museum NAS of Ukraine on behalf of Modern
Phytomorphology. This is an open access article under the Creative Commons BY-NC-ND license (http://creativecommons.org/
licenses/by-nc-nd/4.0/) freely available on https://phytomorphology.org/ .
Introduction
In spite of long history of investigation,
taxonomy of the genus Aconitum L. is still
much discussed and remains unresolved. Many
taxonomic problems arise from the notorious
hybridity within the genus and subsequent
origin of number of putative hybridogenic
taxa, which form a morphological continuum
between the parental taxa (Kadota 1987;
Starmühler 2001). Another problem is that
dierent morphological traits of monkshoods
were considered for genus taxonomy by
dierent authors, and therefore very oen
taxonomic ranges vary signicantly for the
same taxonomic units (Götz 1967; Kmeťová
1982; Skalický 1990; Kadota 1987; Voroshilov
1990; Tamura 1995; Tzvelev 2001). Recent
revisions based on morpho-geographical
concept of species (Davis & Haywood 1963)
and concept of infraspecic dierentiation
(Skalický 1982) led to unication and progress
RESEARCH ARTICLE
Genus Aconitum in Slovakia: a phenetic approach
Andriy Novikov *
State Natural History Museum NAS of Ukraine, Teatralna str. 18, 79008 Lviv, Ukraine; novikoffav@gmail.com
* Andriy Novikov formerly known as Andrew Novikoff starting from 2017 changed spelling variant of his name in
accordance to legislation limitations
Received: 23.04.2017 | Accepted: 28.06.2017 | Published: 23.07.2017
Abstract
Comprehensive investigation on herbarium material was realized and taxonomical structure
of the genus Aconitum in Slovakia has been ascertained on the base of phenetic analysis.
Three new taxa (i.e. A. degenii subsp. degenii, A. moldavicum subsp. hosteanum, and
A. moldavicum nothosubsp. confusum stat. nov.) were rediscovered for this region, and for
one of them the new status was proposed. Morphological variation of A. anthora and its
subdivision onto infraspecific taxa (subsp. anthora and subsp. jacquinii) are discussed. The
new key for identification and maps of distribution are represented.
Keywords: Aconitum, Slovakia, morphology, phenetics, distribution, taxonomy
52 Novikov A.
Modern Phytomorphology 11, 2017
in taxonomy of the genus Aconitum in countries
of Carpathian region (Mitka 2003, 2008a;
Starmühler & Mitka 2001; Starmühler 2002;
Mitka & Szajna 2009; Boroń 2010; Boroń etal.
2011; Noviko & Mitka 2011; Sutkowska etal.
2013; Mitka etal. 2013, 2016; Noviko et al.
2016; Wacławska-Ćwiertnia & Mitka 2016).
Accordingly to Flora of Slovakia (Kmeťová
1982), genus Aconitum here includes 7 taxa
(A. lycoctonum L. em. Koelle, A. moldavicum
Hacq., A. variegatum L., A. lasiocarpum
(Rchb.) Gáyer, A. rmum Rchb. subsp.
rmum, A. rmum subsp. moravicum Skalický,
and A. anthora L.). Kmeťová (1982) noted
that in ora of Slovakia four subspecies of
A. lycoctonum (subsp. vulparia (Rchb.) Schinz
et ell., subsp. puberulum (Ser.) Gáyer, subsp.
thalianum (Wall.) Gáyer, and subsp. penninum
(Ser.) Gáyer) are sometimes distinguished
mostly on the base of variation in pubescence.
For example, subdivision of A. lycoctonum
on subsp. vulparia and subsp. puberulum was
accepted by Domin (1935), and subdivision
of A. lycoctonum on subsp. vulparia, subsp.
puberulum, and subsp. penninum – later by
Dostál (1958). But Kmeťová (1982) did not
accept such partition, arguing that considered
traits make a morphological continuum in
A. lycoctonum. She also noted that sometimes
A. lasiostomum Rchb. is reported from
adjacent regions, perhaps this species does
not occur in Slovakia. For A. moldavicum,
Kmeťová (1982) reported var. australe
(Rchb. ex Baumg.) Grinţ with villous carpels
(“schlpatými mechúrikmi”) from Eastern part
of Slovakia. In the same time, she suggested
that A. moldavicum subsp. hosteanum (Schur)
Graebner et P. Graebner in Aschers. et
Graebner most probably does not occur here,
although it was reported for Czechoslovakia by
Domin (1935) and Dostál (1958).
Domin (1935) and Skalický (1966)
distinguished 3 species of aconites from
Variegatum group (i.e. A. variegatum, A. gracile
Rchb. and A. dominii Sillinger), which also
were accepted by Dostál (1958), however in
range of subspecies – subsp. variegatum, subsp.
gracile (Rchb.) Gáyer, and subsp. dominii
(Sillinger) Dostál. consequently. Moreover,
in Skalický (1982) only A. variegatum is
mentioned. Kmeťová (1982) mentioned
in Flora of Slovakia three subspecies of
A. variegatum (subsp. gracile, subsp. dominii,
and subsp. kotulae), but concluded that they
make morphological intermediations.
Kmeťová (1982) also mentioned that besides
of A. rmum, for Slovakia A. tauricum Wulf.
(Domin 1935; Skalický 1966) or A. napellus L.
subsp. tauricum (Wulf.) Gáyer (Dostál 1958)
are sometimes considered. Finally, she noted
that A. anthora is highly variable taxon and
delimitation of var. jacquinianum G. Beck has
poor taxonomic background.
Starmühler (2002) published the key for
identication of Slovakian aconites with 14
taxa including 6 hybrids (A. lycoctonum subsp.
lycoctonum, A. moldavicum subsp. moldavicum,
A. anthora subsp. anthora, A.variegatum subsp.
variegatum var. variegatum, A. × pawlowskii
Mitka et Starmühl., A. lasiocarpum subsp.
kotulae (Pawł.) Starmühl. et Mitka, A.×lengyelii
Gáyer nothosubsp. lengyelii (= A. × berdaui
Zapał. nothosubsp. berdaui in accordance to
Mitka (2003)), A. × lengyelii nothosubsp.
walasii Mitka (= A. × berdaui nothosubsp.
walasii (Mitka in Starmühler et Mitka) Mitka
in accordance to Mitka (2003)), A. rmum
subsp. rmum, A. rmum subsp. moravicum,
A. rmum subsp. maninense (Skalický)
Starmühl., A. rmum subsp. rmum × A. rmum
subsp. maninense, A. rmum nothosubsp.
paxii Starmühl., and A. rmum nothosubsp.
zapalowiczii Starmühl.). In addition, Mitka
(2003, 2008a) later reported for Slovakia
5 more taxa – A. × cammarum L. em. Fries,
A. × triste (Rchb.) Gáyer (also mentioned
for Czechoslovakia by Domin (1935)),
A. lasiocarpum subsp. lasiocarpum, A.×gayeri
Starmühl. and A. × hebegynum DC.
Taking into account mentioned
publications, the genus Aconitum in ora
of Slovakia could be represented at least by
19 taxa. In this paper I provide results of
investigations on morphological variability and
taxonomic structure of this genus in Slovakia
on the base of analysis of herbarium material
and published works.
Genus Aconitum in Slovakia: a phenetic approach 53
Modern Phytomorphology 11, 2017
Material and methods
Herbarium investigations were conducted
in 2012–2013 in project frames of National
Scholarship Programme of the Slovak Republic
entitled “Taxonomy and chorology of the genus
Aconitum L. (Ranunculaceae) in Slovakian ora”.
Preliminary results of this study (supposed
taxonomic structure of the genus in Slovakia)
were published in 2013 (Noviko 2013) and
deposited online at hp://slovaconitum.
myspecies.info/ (Noviko 2013–2017).
However, in 2017 obtained data were revised
and reevaluated, all of taxonomic and phenetic
studies were redone.
In general more than 3000 herbarium
vouchers were analyzed from 7 main
repositories (Append. 1): Institute of Botany
SAS (SAV, Bratislava), Comenius University
in Bratislava (SLO), Slovak National Museum
(B, Bratislava), Slovak University of
Agriculture in Nitra (NI), Technical University
in Zvolen (ZV), Pavol Jozef Šafárik University
(KO, Košice), and Museum of Tatra National
Park (TNP, Tatranská Lomnica). From total
number, 970 specimens were included into
chorological database; 450 specimens were
included in phenetic database and tested
with 115 morphological traits (Tab. 1). For
dierent taxonomic groups (Lycoctonum,
Anthora, Cammarum and Aconitum) inside
the genus, dierent combinations of treats
were applied for advanced partial analyses
(Tab. 1). Additionaly, main recent literature
sources (Mitka & Starmühler 2000; Mitka
2003, 2008a) were analyzed for mention of
Aconitum specimens from Slovakia nested in
herbaria abroad. Finally, the maps from Flora
of Slovakia (Kmeťová 1982) were digitized
and extracted landmarks were applied in our
research too (Append.2).
For statistical analysis the Past 3.14 was
used (Hammer et al. 2001). For correlation
statistics linear r (Pearson’s) was applied.
For multivariate analysis UPGMA and
Ward’s clustering methods were applied in
combination with Euclidian index of similarity
and N = 100 bootstrap support. For ordination
non-metric multidimensional scaling was
applied in combination with Euclidian index of
similarity.
Distribution maps were built in QGIS 2.18
(QGIS Development Team 2017) with WGS84
(EPSG: 4326) geographical coordinate system
and aer that transformed in J-TSK_Krovak
(EPSG: 102065) projection system, which
is commonly applied in Slovakia for beer
visualization (Annoni et al. 2001). Maps from
Kmeťová (1982) were imported to QGIS as
independent geoti layers with Polygonal 1
WGS 84 transformation mode, and aer that
separated layers with landmarks corresponding
to points on that maps were generated.
For taxonomic verication IPNI (2017)
and e Plant List (2017) databases were
used. For developing of website, the free CMS
Scratchpads 2.0 was applied (Smith et al. 2012).
e raw datasets of treats applied for phenetic
analyses (S20, S21, S22, S23, S24, S25, S26,
S27, S28) and database of analyzed specimens
with generated coordinates of locations (S29
& S30) are freely available in Zenodo and I
kindly welcome researchers to use them for any
investigation purposes.
Results
General analysis was performed for 450 samples
and full dataset of 115 morphological traits
(Tab. 1). Cluster analysis (both UPGMA and
Ward’s methods) showed good delimitation
of main taxonomic groups (i.e. sections and
subgenera) in the genus Aconitum, however
in most cases did not allow discrimination
of taxa on species and subspecies levels, as
well as of some hybrid taxa (Figs. 1 & S1).
Multidimensional scaling showed good
delimitation of group Lycoctonum by rst two
axes however did not show any good limits
between this and other groups on second
scaerplot. Groups Anthora, Cammarum,
Acomarum and Aconitum were signicantly
overlapped (Fig. S2). erefore I provided
further statistical analyses on separated
taxonomic groups representing sections with
exclusion of uninformative traits.
54 Novikov A.
Modern Phytomorphology 11, 2017
Nr Group of traits Trait Lycoctonum Anthora Cammarum Aconitum
1 Helmets glabrous +* +* +*
2 sparsely pubescent +* + +
3 moderately pubescent +* + +
4 densely pubescent +* + +
5 with short curved trichomes +* + +*
6 with protruding straight trichomes + +* +*
7 with mixed trichomes + +*
8 Carpels glabrous +* +* +*
9 ventral pubescent +*
10 dorsal pubescent +* +* +*
11 sparsely pubescent +* + +
12 densely pubescent +* + +
13 with short curved trichomes +* + +*
14 with protruding straight trichomes +* + +
15 Stamens glabrous + + +
16 sparsely pubescent + + +
17 densely pubescent + +
18 All pubescent +
19 Part of them pubescent + +
20 Upper part of
pedicels
glabrous +* +*
21 pubescent only at receptacle
22 sparsely pubescent
23 moderately pubescent + +
24 densely pubescent + +
25 with short curved trichomes + +*
26 with protruding straight trichomes + + +*
27 with mixed trichomes + +*
28 Lower part of
pedicels
glabrous +* +*
29 sparsely pubescent +
30 moderately pubescent + +
31 densely pubescent + +
32 with short curved trichomes + +*
33 with protruding straight
trichomes
+ + +*
34 with mixed trichomes + +
35 Bracteoles glabrous + +*
36 ciliate (pubescent along
margins only)
+
37 sparsely pubescent + +
Table 1. Initial set of analyzed morphological traits. For partial analyses: + – analyzed for full set of treats of separated
taxonomic groups; * – analyzed for limited set of traits selected for each of separated taxonomic groups.
Genus Aconitum in Slovakia: a phenetic approach 55
Modern Phytomorphology 11, 2017
Nr Group of traits Trait Lycoctonum Anthora Cammarum Aconitum
38 Bracteoles moderately pubescent + +
39 densely pubescent + +
40 with short curved trichomes + +*
41 with protruding straight trichomes + + +*
42 Inflorescence
axes
glabrous +* +*
43 sparsely pubescent + +
44 moderately pubescent + + +
45 densely pubescent + + +
46 with short curved trichomes + + +*
47 with protruding straight trichomes + +*
48 with mixed trichomes +
49 Stems glabrous + + +
50 sparsely pubescent + + +
51 moderately pubescent + + + +
52 densely pubescent + + +
53 with short curved trichomes + +
54 with protruding straight trichomes + + +
55 with mixed trichomes +
56 Petioles glabrous + + +
57 sparsely pubescent + +
58 moderately pubescent + + +
59 densely pubescent + + +
60 with short curved trichomes + +
61 with protruding straight trichomes + + +
62 with mixed trichomes +
63 Adaxial surface
of leaf laminae
glabrous + + +
64 pubescent along veins +
65 sparsely pubescent + + +
66 moderately pubescent +
67 densely pubescent +
68 with short curved trichomes + +
69 with protruding straight trichomes + +
70 Abaxial surface
of leaf laminae
glabrous + + +
71 pubescent along veins + + +
72 sparsely pubescent + +
73 with short curved trichomes + +
74 with protruding straight trichomes + +
75 Perianth color pale yellow +
76 yellow +* +
Table 1. Continued.
56 Novikov A.
Modern Phytomorphology 11, 2017
Table 1. Continued.
Nr Group of traits Trait Lycoctonum Anthora Cammarum Aconitum
77 Perianth color dirty blue +*
78 violet +*
79 blue +
80 variegate +
81 Helmet shape bean-shaped +
82 crescent +*
83 hemispherical + +* +*
84 conical +* +*
85 elongated conical +*
86 cylindrical
87 Nectaries not reaching the top of helmet +* +
88 reaching the top of helmet + +
89 spurs ecapitate / capitate +
90 spurs semi-spiral /
curvedbackward bent
91 spurs coiled
92 spurs deflected bent
93 claws adjoined +
94 claws outstanding +
95 Carpels 3 + +
96 5 + +
97 Stamens without protuberances +* +
98 part of them with protuberances +* +
99 all of them with protuberances +
100 Bracteoles reaching the receptacle +* +
101 narrower than pedicels + +
102 as wide as pedicels + +
103 wider than pedicels + +
104 entire +
105 divided +
106 Pedicels longer than flowers + +
107 Inflorescences simple + + + +
108 condensed compound + + + +
109 ramified compound + + + +
110 Shoots orthotropic + +
111 semirosulate
112 nonrosulate
113 Leaves leaf to internode index > 3 + +
114 Perianth deciduous after blooming
115 Life forms polycyclic monocarpous
Genus Aconitum in Slovakia: a phenetic approach 57
Modern Phytomorphology 11, 2017
Lycoctonum group
152 specimens and 50 morphological traits
selected among total 115 (Tab. 1, pluses) were
analyzed. Neither cluster analysis (Figs. S3 &
S4) nor multidimensional scaling (Fig. S5) did
not show appropriated results, randomly mixing
specimens from A. moldavicum, A. lycoctonum
and A.×triste (A.lycoctonum × A.moldavicum).
is explains by fact that these taxa are mainly
dierentiated by color of perianth, which is from
dirty-blue to purple-violet in A. moldavicum,
from whitish to yellow in A. lycoctonum, and
yellowish-bluish in A. × triste (Skalický 1990;
Mitka 2008a). In all three species, helmet is
usually covered by short curved trichomes.
But density of such pubescence signicantly
varies in A. moldavicum, where helmet can
be even entirely glabrous or with only few
sparsely distributed trichomes. While in studied
specimens of A. × triste and A. lycoctonum
helmets were always densely covered by curved
trichomes.
In A. lycoctonum and A. × triste carpels
can be either glabrous or covered by short
curved trichomes. In A. moldavicum character
of carpels’ pubescence plays the key role for
delimitation of subspecies. In A. moldavicum
subsp. moldavicum carpels are glabrous. In
A. moldavicum subsp. hosteanum carpels are
entirely covered by short curved trichomes.
ere are also several other hybrids
distinguished on the base of perianth color
and pubescence of carpels. For example, in
A. moldavicum nothosubsp. simonkaianum
(Gáyer) Starmühl. (A. lasiostomum ×
A. moldavicum subsp. hosteanum) carpels are
villous – entirely covered by protruding long
trichomes (Starmühler 1998; Mitka 2008a).
In A.× porcii Starmühl. (A. moldavicum subsp.
moldavicum × A. moldavicum nothosubsp.
simonkaianum) helmet is purple-violet; carpels
are with mixed (villous at the base and curved
at upper part) pubescence (Starmühler 1998;
Mitka 2008a). In A. × baumgartenianum
Simonk. (A. lasianthum Simonk. ×
A. moldavicum) helmet is yellowish-bluish,
glandular pilose and eglandular pubescent;
carpels are entirely villous, with long
protruding trichomes (Mitka 2008a). A hybrid
between A. moldavicum subsp. hosteanum and
A.moldavicum subsp. moldavicum with “carpels
at the backside and/or on the suture pilose” is
also mentioned by Mitka (2008a).
However in recent personal communication,
W. Roensteiner (formerly W. Starmühler)
noticed that he unsuccessfully tried to
dierentiate subtaxa among A. moldavicum on
the base of carpels pubescence. So he concluded
that “the pubescence of the carpels, which is very
precious in other species, has no taxomonical
value in A. moldavicum”. In particular, he treats
A. moldavicum subsp. hosteanum as an endemic
of Southern Carpathians distributed only in
the highest altitudes and distinguished from
A. moldavicum subsp. moldavicum by dierent
pubescence of tepals. However, A. moldavicum
subsp. hosteanum was found in Poland and
Ukraine even in lower altitudes of subcarpathian
region and growing in mixed populations with
A. moldavicum subsp. moldavicum (Mitka 2003,
2008a, 2008b; Noviko & Mitka 2011; Nov iko
et al. 2016). Zapałowicz (1908) mentioned
A. hosteanum Schur from Czywczyny and
Gorgany Mts. in Ukraine. Tzvelev (2001)
also applied type of carpels pubescence for
dierentiation of A. hosteanum among other
taxa from Lycoctonum group and noted that it is
distributed in Eastern Carpathians. So there is
dierent understanding of taxonomic limits in
A. moldavicum.
It is important to note that both Starmühler
(2002) and Mitka (2008a) reported for Slovakia
A. lycoctonum subsp. lycoctonum, although
Kmeťová (1982) and some other researchers
(Domin 1935; Dostal 1958; Skalický 1966,
1985; Tutin et al. 1993) mentioned for this
region A. lycoctonum subsp. vulparia. In
preliminary published results (Noviko 2013)
I used the name A. lycoctonum subsp. vulparia,
however here for uniformity this taxon is
considered as A. lycoctonum subsp. lycoctonum
following Starmühler (1997, 2002) and Mitka
(2003, 2008a). Moreover, Starmühler (1997)
neotypied A. lycoctonum subsp. vulparia and
chose the illustration from Reichenbach (1823–
1827) with straight pubescence of the pedicel
and the tepals. Such straight type of pubescence
58 Novikov A.
Modern Phytomorphology 11, 2017
Fig. 1.
Ward’s dendrogram in
combination with Euclidian
distances for full dataset
of treats and samples.
Blue – group Lycoctonum;
black – group Aconitum;
grey – group Acomarum;
green–group Cammarum,
red–group Anthora.
Genus Aconitum in Slovakia: a phenetic approach 59
Modern Phytomorphology 11, 2017
Fig. 1.
Continued.
60 Novikov A.
Modern Phytomorphology 11, 2017
is also seen in Reichenbach (1840), however
contradicts with taxonomic interpretation
of Czech and Slovakian researchers (Domin
1935; Dostal 1958; Skalický 1966, 1985) as
well as revision of Tzvelev (2001). In particular,
Tzvelev (2001), in opposition to Starmühler
(1997), characterized A. vulparia by short
curved trichomes covering inorescence axes
and pedicels, and A. lycoctonum – by straight
trichomes.
Hence, I performed cluster analysis for
14 separated traits mentioned as diagnostic
(Tab. 1, asterisks), which gave good results, but
with low bootstrap support. In case of UPGMA,
cluster moldavicum has been joined with cluster
lycoctonum (Fig. S6), but in case of Ward’s
method of clustering (Fig. 2), the cluster
moldavicum was aached with cluster hosteanum.
In both cases, cluster triste was joined with cluster
lycoctonum, and their samples were mixed on plot
of multidimensional scaling (Fig. S7). Cluster
lycoctonum was divided on two subclasters
dierent by pubescence of carpels (glabrous
in samples lycoct: 1554 – lycoct: 1556 or curved
pubescent in samples lycoct: 3142 – lycoct: 995),
however with low bootstrap support (Figs. 2
& S6). Cluster moldavicum is subdivided on
several subclusters on the base of intensity of
pubescence of helmet and variation of perianth
color (dirt-blue or violet). Simultaneously,
cluster hosteanum is divided on the base of
variation in helmet and carpels pubescence, and
coloration of perianth too.
At SAV herbarium I found one specimen
of A. moldavicum with carpels covered on
dorsal sides by protruding villous trichomes
and reported also from Romania (Grințescu
1953). Preliminary I identied this specimen
as A. moldavicum nothosubsp. simonkaianum
(Noviko 2013), however A. moldavicum
nothosubsp. simonkaianum is characterized by
protruding villous pubescence of entire carpels.
Taking into account reports of Grințescu (1953)
and Mitka (2008a) representing diagnostic
features of mentioned taxa, name proposed
by Grințescu (1953), and following current
taxonomic hierarchy in Lycoctonum group, I
believe that this specimen should be described
as A. moldavicum nothosubsp. confusum
(Grinț.) A. Novikov. Due its hybrid origin and
low number of analyzed material, this specimen
does not occupy any stabile position neither
Fig. 2. Ward’s dendrogram in combination with Euclidian distances for Lycoctonum group with dataset of
14 diagnostic traits.
Genus Aconitum in Slovakia: a phenetic approach 61
Modern Phytomorphology 11, 2017
on dendrogram (Figs. 2, S3, S4, S6) nor on the
multidimensional scaerplots (Figs. S5 & S7).
Correlation analysis reveals the only
very strong dependence (Pearson’s r = 0.85)
– between yellow coloration of perianth
and dense pubescence of helmet, what
characterizes the most of specimens from
cluster lycoctonum.
Hence, Aconitum subgen. Lycoctonum (DC.)
Peterm. sect. Lycoctonum DC. in ora of Slovakia
is represented by 5 taxa: A. lycoctonum subsp.
lycoctonum, A. moldavicum subsp. moldavicum,
A. moldavicum subsp. hosteanum, A. moldavicum
nothosubsp. confusum and A.×triste.
A. lycoctonum subsp. lycoctonum is well
represented in herbaria of Slovakia (150
vouchers revealed). It is mostly distributed in
mountain regions in western and central parts
of Slovakia, and also has one documented
location from eastern region near the border
with Ukraine (Fig. 7). A. moldavicum subsp.
moldavicum is represented by 195 herbarium
vouchers. It is scaered predominantly in
eastern and central mountain regions of
Slovakia, however it has also several locations
from western part of country (Fig. 8).
Similarly, A. moldavicum subsp. hosteanum
(74 investigated vouchers) is represented
in eastern and central regions of Slovakia,
however it is not going so far to the west
(Fig. 8). A. moldavicum nothosubsp.
confusum was collected just once and more
than 50 years ago (Spišská Nová Ves; leg.
Májovský J., 23.08.1955; SAV). Collector
indicated on the label just a city Spišská Nová
Ves, which is located in Hornádska Kotlina,
however did not indicate any other details
(Fig. 3). A. × triste was also collected only once
in 2005 (Meliata, vlhke lesy; leg. KarasováE.,
16.06.2005; SAV), and it is represented only
by 5 vouchers in SAV herbarium (Fig. 8).
Before this, A. × triste was reported for Slovakia
by Skalický (1990) and whereupon by Mitka
(2008a), however without indication of strict
locations. Both hybrids, A. moldavicum
nothosubsp. confusum and A. × triste, seem
to be very rare for Slovakia and require further
eld expeditions.
A. moldavicum nothosubsp. confusum
(Grinț.) A. Novikov, stat. nov. (Fig. 3)
(A. moldavicum subsp. hosteanum ×
A. moldavicum subsp. moldavicum)
Basionym: A. moldavicum var. confusum
Grinț. in Săvulescu, Flora Repub. Pop. Române
2: (677), 1953.
Typu s: Reg.: Craiova: in pratis ad pagum
Baia de Fier. (Grințescu 1953).
Diagnostic characters: Perianth from
dirty-blue to violet, carpels villous (with long
protruding trichomes) only on dorsal side or/
and suture.
Geographical distribution: In Carpathians
and adjacent regions. Rarely in the Western
Carpathians and slightly oen in the Eastern
Carpathians; oen in the Małopolska Upland
(Poland), Opillya and Voronyaky (Ukraine)
(Mitka 2008a).
Anthora group
Aconitum subgen. Anthora (DC.) Peterm. sect.
Anthora DC. is represented in ora of Slovakia
by single species, A. anthora, which is however
characterized by high morphological variation.
In general I had analyzed 152 specimens of
A. anthora s.l., from which 66 specimens were
included in further phenetic analysis. Among
115 total features I dened 26, which had
variation for A. anthora and used them for next
data processing (Tab. 1, pluses). Cluster analysis
on the base of Ward’s method revealed two clear
groups – anthora and jacquinii (Fig. S8) with
high bootstrap support. However UPGMA
analysis showed that group anthora is not
homogenous and consists from two main
subclusters – jacquinii1 and jacquinii2 (Fig. 4).
Cluster anthora is characterized by curved short
trichomes covered helmet, pedicels, bracteoles,
and inorescence branches. Cluster jacquinii1
is represented by samples with protruding
villous trichomes on helmet, bracteoles and
pedicels (in some observations pedicels in
lower part had mixed cover consisting from
curved and protruding trichomes), and
with mixed trichomes on inorescence axes.
Cluster jacquinii 2 (samples jacquin: 2280
62 Novikov A.
Modern Phytomorphology 11, 2017
Fig. 3. A. moldavicum nothosubsp. confusum (Grinț.) A. Novikov, stat. nov. in SAV herbarium.
Genus Aconitum in Slovakia: a phenetic approach 63
Modern Phytomorphology 11, 2017
and jacquin: 2346) is characterized by mixed
pubescence of helmet and pedicels, protruding
pubescence of bracteoles, and exclusively
curved pubescence of inorescence axes.
Outstanding position of the cluster jacquinii2
is also well seen on plot of multidimensional
scaling (Fig. S9).
Correlation analysis revealed that curved
pubescence of helmet is very strongly related to
curved pubescence of pedicels (Pearson’s r = 1),
bracteoles (Pearson’s r = 1), and inorescence
axes (Pearson’s r = 0.80). In the same time,
protruding pubescence of the helmet is strongly
correlated with protruding pubescence of
pedicels (for pedicels above bracteoles Pearson’s
r = 1, and for pedicels below bracteoles Pearson’s
r = 0.86), protruding pubescence of bracteoles
(Pearson’s r = 0.80), and mixed pubescence
of inorescence axes (Pearson’s r = 1). Mixed
pubescence of helmet is very strongly correlated
with mixed pubescence of pedicels (above
bracteoles Pearson’s r = 1, and below bracteoles
Pearson’s r = 0.81). Pubescence type of carpels,
stems and petioles do not show any (even
moderate) correlations, as well as the shape
of helmet, colour (yellow or pale yellow) of
perianth and structure of inorescence.
erefore, I can conclude that pubescence
type of helmet, pedicels, bracteoles and
inorescence are the most signicant features
illuminating 3 main morphotypes of A. anthora
in Slovakia, while the shape of helmet and
perianth colour (I do not consider here
A. pseudanthora Błocki ex Pacz. with bluish
or variegate perianth) are uninformative traits.
Moreover, in cluster jacquinii 1 the variation
from entire protruding to mixed pubescence
of pedicels was observed, but there was no
such variation in group anthora with always
and entirely curved pubescent pedicels. Even
on generalized dendrograms (Figs. 1 & S1),
jacquinii samples are well delimited from other
anthora samples with bootstrap support 96
(UPGMA) or 91 (Ward’s method), with only
two samples jacquin: 2280 and jacquin: 2346
occupying intermediate position between two
main groups. Cluster jacquinii 2 (samples
jacquin: 2280 and jacquin: 2346) most probably
represent putative hybrid between A. anthora
subsp. anthora and A. anthora subsp. jacquinii,
which, however, requires detailed studies.
Taking into consideration that these
plants grow in mixed populations (e.g.,
jacquin: 2280 was collected by J. Májovský in
Fig. 4. Ward’s dendrogram in combination with Euclidian distances for Anthora group with full dataset of traits.
jacquinii 2
jacquinii 1
anthora
64 Novikov A.
Modern Phytomorphology 11, 2017
1955 together with anthor: 2276, anthor: 2278
and anthor: 2282), they can be considered as
subspecies. Hence, cluster jacquinii 1 can be
recognized as A. anthora subsp. jacquinii, and
cluster anthora – as A. anthora subsp. anthora.
Before, due to the taxonomic confusion,
I erroneously reported for Slovakia A. anthora
subsp. eulophum (Noviko 2013), which does
not occur here. A. eulophum Rchb. was reported
by Reichenbach (1819, 1820) from Alps region
including Liguria (France) and Helvetia (part
of modern Switzerland), and is also oen
mentioned for Ukraine as independent taxon
(Paczoski 1927; Visjulina 1953; Chopyk 1978;
Andrienko & Peregrym 2012) or synonym
of A. anthora (Tzvelev 2001; Didukh et al.
2004). Division of A. anthora onto subspecies
for ora of Czechoslovakia was reported by
Domin (1935), who recognized A. anthora
subsp. eu‑anthora and A. anthora subsp. jacquinii
(Rchb.) Domin.
e problem with classication of A. anthora
and related taxa has a long history starting from
description by Linnaeus (1753). Reichenbach
(1819, 1820, 1838, 1840) recognized several
taxa in section Anthoroideae Rchb. viz
A. anthora, A. jacquinii Rchb., A. decandolii
Rchb., A. anthoroideum Rchb., A. eulophum
Rchb. and A. nemorosum Rchb. He applied
complex morphological analysis for delimitation
of taxa, including pubescence of oral parts and
shape of helmet. In particular, for Carpathians
Reichenbach (1820) mentioned A. jacquinii,
which, as he noted, diers by unique shape of
helmet with elongated rostrum. In this paper
Reichenbach (1820) also mentioned that
A. jacquinii has glabrous helmets, what is
very unusual for Anthora representatives from
Carpathian region, and what I saw just once, in
isolated refugial population in Stinka (Ukrainian
Carpathians). However later Reichenbach
(1840) published monograph containing
illustration of A. jacquinii with villous helmet
(Lám. LXXVII) and illustration of A. anthora
γ Jacquinianum Rchb. (Lám. C) with glabrous
owers. Some of later researchers accepted level
of species (Zapalowicz 1908; Paczoski 1927;
Visjulina 1953) or level of subspecies Domin
(1935) for this taxon, but in all cases authors
mentioned plants with helmet covered by
protruding hairs contrasting with A. anthora
(or A. anthora subsp. anthora) having curved
pubescence of helmet. Same division is also
accepted by Tzvelev (2001), who recognized
A. anthora with pedicels and inorescence axes
covered exclusively by short curved trichomes,
and A. jacquinii – by straight and mixed
trichomes.
In our research, helmet shape showed high
variation, however type of helmet pubescence
seems to be a good discriminative trait for
recognition of infraspecic taxa in Anthora
group.
In general, A. anthora subsp. anthora is
more widely represented in ora of Slovakia.
I found 139 vouchers of A. anthora subsp.
anthora in Slovakian herbaria sparsely collected
from almost all territory of country (Fig. 9).
A. anthora subsp. jacquinii there is represented
only by 9 vouchers and occurs only in two main
sites, one of which (Dreveník, obec Levoča)
it shares with putative hybrid (represented by
only 2 specimens in Slovakian herbaria) and
other parental taxon – A. anthora subsp. anthora
(Fig. 9).
Cammarum group
Aconitum subgen. Aconitum sect. Cammarum
DC. is one of the most diverse group in sense
of morphological variability and taxonomic
representativeness in Slovakia. I used 121
specimens for analysis of 68 morphological
traits (Tab. 1, pluses). UPGMA clustering
did not showed any appropriated results
(Fig. S10). Clustering by Ward’s method
separated two main clusters representing
mostly A. variegatum and A. lasiocarpum
samples. However A. degenii Gáyer and
A. × pawlowskii were randomly nested in these
two main clusters (Fig. S11) and scaered on
multidimensional plot (Fig. S12). erefore
I performed analysis on limited number of
15 traits, which are considered diagnostic for
this group (Tab. 1, asterisks). is allowed
obtaining beer results, but some samples of
A. lasiocarpum subsp. lasiocarpum were nested
among A. degenii specimens, while samples of
Genus Aconitum in Slovakia: a phenetic approach 65
Modern Phytomorphology 11, 2017
A.×pawlowskii – among A. lasiocarpum subsp.
lasiocarpum (Figs. 5 & S13). On the plot of
multidimensional scaling, A. × pawlowskii was
placed between its parental taxa A. variegatum
and A. lasiocarpum, while A. degenii occupied
marginal position (Fig. S14). e reason
of such divergence of A. degenii samples was
illuminated later – only cluster of degen: 1173 and
degen: 1175 containing samples from Eastern
part of Slovakia truly represents A. degenii,
while other samples were misidentied and
belong to A. rmum subsp. maninense.
Correlation analysis showed dependence
of glabrous helmet with sutural pubescence of
carpels (very strong, Pearson’s r = 0.94), glabrous
pedicels (strong, Pearson’s r = 0.84 for upper
and Pearson’s r = 0.75 for lower part of pedicels),
glabrous inorescence axes (strong, Pearson’s
r = 0.65), what is typical for A. variegatum.
ere was also moderate positive correlation
(Pearson’s r = 0.59) between glabrous helmet
and nectarines not reaching the helmet top, what
also characterizes A. variegatum. A. variegatum
has elongated conical helmet, while the shape
of helmet in A. lasiocarpum varies. In the same
time, A. degenii has oen hemispherical or
slightly elongated helmet and glabrous carpels,
what was reected in strong positive correlation
(Pearson’s r = 0.7) between hemispherical
helmet and glabrous carpels in my analysis too
(Tab. 1).
Sect. Cammarum in ora of Slovakia is
represented by 6 taxa: A. variegatum subsp.
variegatum, A. lasiocarpum subsp. lasiocarpum,
A. lasiocarpum subsp. kotulae, A. ×pawlowskii
(A. lasiocarpum × A. variegatum), A. degenii
subsp. degenii and A. × gayeri (A. degenii ×
A. lasiocarpum).
A. degenii subsp. degenii is reported
here for the rst time for ora of Slovakia.
All other taxa were already reported before
(Mitka & Starmühler 2000; Starmühler 2002;
Mitka 2003). Mitka (2003) also reported
A. × hebegynum (A. degenii × A. variegatum)
for ora of Slovakia, however this taxon was
later rediscovered and nested in A. degenii
subsp. degenii as var. intermedium (Zapał.) Mitka
(Ilnicki & Mitka 2011).
In total, I found in Slovakian herbaria 190
specimens of A. variegatum subsp. variegatum
collected mostly from central Slovakia, however
few samples also were collected from eastern
Fig. 5. Ward’s dendrogram in combination with Euclidian distances for Cammarum group with dataset of
15 diagnostic traits.
66 Novikov A.
Modern Phytomorphology 11, 2017
regions (Fig. 10). A. lasiocarpum subsp.
lasiocarpum is represented by 18 (Fig. 11), and
A. lasiocarpum subsp. kotulae – by 11 vouchers
collected from central and eastern regions of
Slovakia (Fig. 11). A.×pawlowskii was found in
herbaria only twice (Fig. 12). A. degenii subsp.
degenii in analyzed collections is represented
only by 2 vouchers (Fig. 12), and A. × gayeri
– by single specimen hosted in SAV (Fig. 12).
However Mitka (2003) reported 1 more
voucher of A. × gayeri from Bukovske Vrchy,
which is deposited in herbarium of Jagiellonian
University in Kraków (Rabia Skała, 1150 m,
J. Mitka, P. Bochenek, J. Terray, 23.08.1995;
K).
A. lasiocarpum, A. × pawlowskii, A. degenii
and A. × gayeri are rare for ora of Slovakia. Last
two taxa are distributed here on the edge of their
natural range.
Acomarum group
Aconitum subgen. Aconitum nothosect.
Acomarum Starmühler in ora of Slovakia
is represented by cultivated intersectional
hybrid A. × cammarum (A. napellus [?] ×
A. variegatum). A. × cammarum was reported
for Czechoslovakia by Skalický (1982), and
also mentioned by Domin (1935) and Dostál
(1958) as A.×stoerckianum (=A. napelus subsp.
neomontanum (Wulfen) Gáyer × A. variegatum
subsp. euvariegatum Dostál). I did not include
this taxon neither in morphological nor in
chorological analyses, because it is ornamental
plant and therefore there is low number of
specimens (only 17 were observed) deposited in
herbaria (Fig. 13). Sometimes A. × cammarum
can be found escaped into the wild nature.
Skalický (1966) also reported for
Czechoslovakia A. × exaltatum Bernh. ex Rchb.
(A. plicatum Köhler ex Rchb. subsp. plicatum
× A. variegatum subsp. variegatum), which
is morphologically close to A. × cammarum
(Mitka 2003). However, there are no mentions
of this hybrid exactly from Slovakia.
Starmühler (2002) mentioned for
Slovakia two more taxa for this nothosection,
however without details on their distribution
– A. × lengyelii Gáyer nothosubsp. lengyelii
(A. rmum subsp. rmum × A. variegatum
subsp. variegatum) and A.×lengyelii nothosubsp.
walasii Mitka (A. rmum subsp. moravicum
× A. variegatum subsp. variegatum). Later
these two nothotaxa were also mentioned
in Starmühler & Mitka (2001), but only for
territory of Poland and are not listed for Slovakia.
In personal communication W. Roensteiner
also noted that these nothotaxa are highly
possible to be nd in Slovakia, since both
parental taxa are present there.
In Mitka (2003) A. × lengyelii nothosubsp.
lengyelii and A. × lengyelii nothosubsp.
walasii were rediscovered as A. × berdaui
Zapał. nothosubsp. berdaui and A. × berdaui
nothosubsp. walasii (Mitka in Starmühler et
Mitka) Mitka, respectively. Mitka (2003) listed
only 2 specimens of A. × berdaui nothosubsp.
berdaui from Slovakian Tatra Mts. (Wycieczka
w Tatry Spiskie w r. 1878. Aconitum napellusL.,
Podspady nad Jaworzynką; leg. A. Rogalski,
25.07.1878.) and Slovenský Raj (Plantae
exsiccatae regni Hungariae, Comit. Szepes, in m.
Hollókö pr. Sztraczena; G. Lengyel, 01.08.1901;
BP) (Fig. 13). In personal communication
J. Mitka also provided me with two more
recently rediscovered locations – one in Tatras
(Tatras, Javorova Dolina; leg. A. Sutkowska,
18.08.2009; herb. J. Mitka) and other in Malá
Fatra (Malá Fatra, Terchová; leg. J. Mitka,
31.07.2009; K 0464764). However,
Mitka (2003) does not mention A. × berdaui
nothosubsp. walasii for Flora of Slovakia.
During our revision of Slovakian herbaria, I did
not nd specimens of A. × berdaui, and therefore
this nothospecies is provided here on the base of
reports of Starmühler (2002) and Mitka (2003)
only. Presence and distribution of A. × berdaui
nothosubsp. walasii in Slovakia still require
conrmation in eld.
Aconitum group
I performed cluster analysis for 104 specimens
and 82 morphological treats (Tab. 1, pluses),
which revealed division of Aconitum subgen.
Aconitum sect. Aconitum in ora of Slovakia
on two main clusters – rmum and moravicum
(Figs. S15 & S16). Cluster rmum includes
Genus Aconitum in Slovakia: a phenetic approach 67
Modern Phytomorphology 11, 2017
A. rmum subsp. rmum. Cluster moravicum
consisted mostly from A. rmum subsp.
moravicum samples, but also included randomly
dispersed samples of A. rmum subsp.
maninense, A. rmum nothosubsp. paxii and
A. rmum nothosubsp. zapalowiczii (Fig. S17).
All samples of A. rmum subsp. maninense
had localized position together, which also
conrmed by multidimensional scaling,
however they were nested inside the cluster
moravicum. erefore, I performed analysis
for limited number of 23 morphological traits
considered as diagnostic for Aconitum group
and obtained trees were beer ing accepted
taxonomic division – there were independent
and clearly distinguished clusters of A. rmum
subsp. rmum, A. rmum subsp. moravicum and
A. rmum subsp. maninense, however hybrids
(A. rmum nothosubsp. paxii and A. rmum
nothosubsp. zapalowiczii) were still overlaped
with other taxa (Figs. 6, S18 & S19).
For this limited number of 23 morphological
traits I provided correlation analysis (Tab. 1,
asterisks), which showed very strong positive
correlation between glabrous helmet and
glabrous pedicels (Pearson’s r = 0.9 for pedicels
above bracteoles and Pearson’s r = 0.88 for
pedicels below bracteoles), glabrous bracteoles
(Pearson’s r = 0.88) and glabrous inorescence
axes (Pearson’s r = 0.88), which are typical for
A. rmum subsp. rmum. Similarly, this analysis
showed very strong positive correlation between
curved pubescent helmet and curved pubescent
pedicels (Pearson’s r = 0.93 for pedicels above
bracteoles and Pearson’s r = 0.83 for pedicels
below bracteoles), curved pubescent bracteoles
(Pearson’s r = 0.91) and curved pubescent
inorescence axes (Pearson’s r = 0.87), what
characterizes A. rmum subsp. moravicum.
Very strong positive correlation was found also
for pilose pubescence of helmet and pilose
pubescence of pedicels (Pearson’s r = 1 both
for pedicels above and below the bracteoles),
and pilose pubescence of inorescence axes
(Pearson’s r = 0.86); as well as strong positive
correlation (Pearson’s r = 0.64) – for pilose
pubescences of helmet and bracteoles, what
in general characterizes A. rmum subsp.
maninense. Finally, strong positive correlation
(Pearson’s r = 0.7) was also shown for mixed
pubescence of helmet and mixed pubescence
of pedicels above the level of bracteoles, what
Fig. 6. Ward’s dendrogram in combination with Euclidian distances for Aconitum group with dataset of 23 diagnostic
traits.
68 Novikov A.
Modern Phytomorphology 11, 2017
identies A. rmum nothosubsp. paxii. is
analysis did not reveal any strong or even
moderate correlations for shape of helmet,
which can vary from crescent to slightly conical
even in the same inorescence.
Starmühler (2002) and Starmühler &
Mitka (2001) reported for Slovakia also hybrid
between A. rmum subsp. rmum × A. rmum
subsp. maninense with glabrous perianth and
pilose pedicels, discovered on the base of single
specimen deposited in herbarium of Natural
History Museum in Sibiu, Romania (Hungaria,
comitatus Szepes, Montes Bélaenses, in
valle Drechslerhäuschen [Dolina siedmich
prameňov valley] sub monte Stirnberg, alt.
cca. 1400–1500 m, solo calc.; 01.09.1907; leg.
E.G. Nyárády; SIB). However in Slovakian
herbaria I did not found such specimens.
In total I found and analyzed 152 specimens
of A. rmum in Slovakian herbaria. e most
represented there is collection of A. rmum
subsp. rmum counting 121 herbarium
vouchers mostly from central Slovakia
(Fig. 14). A. rmum subsp. moravicum is
represented by 26 vouchers (Fig. 14). I found
just few herbarium vouchers of A. rmum
subsp. maninense in SLO, SAV and B
herbaria (Fig. 14). Six more vouchers of
A. rmum subsp. maninense are mentioned
by Mitka (2003) from Vysoké Tatry and
Stražovské vrchy, and deposited in herbaria SIB
(Natural Histori Museum in Sibiu, Romania),
PRC (Charles University in Prague, Czecz
Republic) and K (Jagiellonian University
in Kraków, Poland). Moreover, Mitka et al.
(2015) noted that A. rmum subsp. maninense
is characterized by high genetic divergence and
probably is a relict taxon isolated from other
representatives of Aconitum group.
Morphologically A. rmum subsp.
maninense is similar to A. degenii subsp. degenii
and can be easily misidentied on the base of
protruding glandular cover of oral parts.
A. rmum nothosubsp. paxii was represented
only by 2 herbarium vouchers, and A. rmum
nothosubsp. zapalowiczii – only by 3 specimens,
one of which is mounted on the same herbarium
sheet with A. rmum subsp. rmum (Fig. 14).
Hence I can conclude that Aconitum subgen.
Aconitum sect. Aconitum in ora of Slovakia is
represented by 5 A. rmum subspecies and one
unnamed hybrid.
Key for identification
1. Helmet cylindrical, about 2.5–3 times higher
than wide; nectaries do not reach the top of
helmet and have coiled spurs; leaf laminas
incised less than ½, with very broad segments;
semirosulate plants with rhizomes .................... 2
1*. Helmet from crescent to elongated conical,
but its height does not exceed 2.5 of width;
nectaries reach or not the top of helmet, their
spurs from semi-spiral curved to ecapitate; leaf
laminas incised more than ½, with segments
from broad to narrow linear; nonrosulate plants
with tubers ............................................................. 6
2. Perianth from white-yellowish to yellow;
helmet densely covered by short curved
trichomes; carpels glabrous or pubescent by
short curved trichomes ..........................................
A. lycoctonum subsp. lycoctonum
2*. Perianth has other coloration ...................... 3
3. Perianth yellowish-bluish; helmet densely
covered by short curved trichomes; carpels
glabrous or pubescent by short curved
trichomes ............................................... A.×triste
3*. Perianth from dirty-blue to purple-violet;
helmet more or less covered by short curved
trichomes or, rarely, glabrous ............................. 4
4. Carpels glabrous ....... A.moldavicum subsp.
moldavicum
4*. Carpels pubescent ......................................... 5
5. Carpels entirely covered by short curved
trichomes .... A.moldavicum subsp. hosteanum
5*. Carpels have long protruding trichomes
only on dorsal side or/and suture ........................
A. moldavicum nothosubsp. confusum
Genus Aconitum in Slovakia: a phenetic approach 69
Modern Phytomorphology 11, 2017
6. Perianth from yellowish to yellow, persists
aer blooming; helmet hemispherical or slightly
elongated, sometimes with protruded front
part; nectaries reach the top of helmet, with
semi-spiral curved spurs; leaves deeply incised,
with very narrow linear segments; tubers from
ovate to napiform, sometimes divided ............. 7
6*. Perianth usually dark-blue, but sometimes
violet, lilac, white or variegate, deciduous
aer blooming; nectaries reach or not the top
of helmet, with spurs from backward bent to
ecapitate; leaves incised near ½–⅔, with more
or less broad segments; tubers from ovate to
napiform ................................................................. 9
7. Helmet, pedicel, and bracteoles covered by
short curved trichomes; carpels covered by
curved or protruding trichomes; inorescence
axes covered by short curved trichomes ............
A. anthora subsp. anthora
7*. Pubescence type diers ................................ 8
8. Helmet, pedicel, and bracteoles covered
by protruding trichomes; carpels covered
by protruding trichomes; inorescence axes
covered by mixed curved and protruding
trichomes .............. A.anthora subsp. jacquinii
8*. Helmet and pedicels covered by mixed
curved and protruding trichomes; bracteoles
and carpels covered by protruding trichomes;
inorescence axes covered by curved trichomes
........... A.anthora subsp. anthora × A.anthora
subsp. jacquinii
9. Helmet from hemispherical to elongated
conical; nectaries usually do not reach the
top of helmet and have hooked or backward
bent spurs; tubers ovate with short stolones
or napiform; plants usually high with ramied
inorescences .................................................. 10
9*. Helmet from crescent to hemispherical;
nectaries reach the top of helmet and have
capitate or ecapitate spurs; tubers napiform;
plants usually low with compact or dense
ramied inorescences ...................................... 18
10. Helmet elongated conical, oen with
pronounced rostrum; tubers ovate with
stolones ................................................................ 11
10*. Helmet from hemispherical to slightly
elongate; tubers ovate or napiform ................. 14
11. Helmet usually glabrous, rarely covered by
short solitary trichomes; pedicels and brateoles
glabrous; carpels have trichomes only along
ventral sutures; inorescence axes glabrous ......
A. variegatum subsp. variegatum
11*. Helmet, pedicels (at least above the
bracteoles) and bracteoles covered by
protruding glandular (pilose) trichomes ...... 12
12. Pedicels and bracteoles entirely covered by
protruding glandular trichomes; inorescence
axes covered by protruding pilose trichomes;
carpels entirely pilose .............................................
A.lasiocarpum subsp. lasiocarpum
12*. Pedicels covered by protruding glandular
trichomes only above the bracteoles, below
the bracteoles they are glandular and/or
eglandular pubescent, sparsely glandular
pilose or even glabrous. In very rare cases
pedicels are entirely glabrous. Inflorescence
axes have different cover, from sparsely pilose
to glabrous .................................................... 13
13. Carpels entirely pilose ........ A.lasiocarpum
subsp. kotulae
13*. Carpels have trichomes only along ventral
sutures ......................................... A.×pawlowskii
14. Helmet glabrous or only sparsely covered
by protruding trichomes; carpels sterile; tubers
napiform ............................................................... 15
14*. Helmet entirely covered by short protruding
trichomes; pedicels and bracteoles covered by
short protruding trichomes (sometimes less
covered or glabrous in lower part of pedicels);
tubers ovate ......................................................... 17
15. Helmet glabrous; carpels glabrous and
sterile; pedicels have trichomes just near the
70 Novikov A.
Modern Phytomorphology 11, 2017
Fig. 7. Map of Aconitum lycoctonum distribution in Slovakia.
Fig. 8. Map of Aconitum moldavicum and A. × triste distribution in Slovakia.
Genus Aconitum in Slovakia: a phenetic approach 71
Modern Phytomorphology 11, 2017
Fig. 9. Map of Aconitum anthora distribution in Slovakia.
Fig. 10. Map of Aconitum variegatum distribution in Slovakia.
72 Novikov A.
Modern Phytomorphology 11, 2017
Fig. 11. Map of Aconitum lasiocarpum distribution in Slovakia.
Fig. 12. Map of Aconitum degenii, A. × gayeri and A. × pawlowskii distribution in Slovakia.
Genus Aconitum in Slovakia: a phenetic approach 73
Modern Phytomorphology 11, 2017
Fig. 13. Map of Aconitum × cammarum and A. × berdaui distribution in Slovakia.
Fig. 14. Map of Aconitum firmum distribution in Slovakia.
74 Novikov A.
Modern Phytomorphology 11, 2017
receptacle; bracteoles glabrous or covered by
solitary trichomes ..................... A. × cammarum
15*. Helmet glabrous or only sparsely covered
by protruding trichomes; carpels glabrous or
pilose along ventral sutures, sometimes with
solitary trichomes on dorsal side .................... 16
16. Helmet, pedicels, bracteoles and
inorescence axes glabrous ...................................
A.×berdaui nothosubsp. berdaui
16*. Helmet glabrous or only sparsely covered by
protruding trichomes; pedicels and bracteoles
covered by curved trichomes; inorescence axes
covered by short curved trichomes or almost
glabrous ...... A.×berdaui nothosubsp. walasii
17. Carpels glabrous ... A.degenii subsp. degenii
17*. Carpels densely covered by protruding
trichomes ............................................. A.×gayeri
18. Helmet glabrous .......................................... 19
18* Helmet covered by trichomes .................. 21
19. Pedicels, bracteoles, and inorescence axes
glabrous; carpels glabrous or rarely pubescent
on dorsal side ........... A.rmum subsp. rmum
19*. Pedicels covered by trichomes ................ 20
20. Pedicels and bracteoles covered by mixed
curved and protruding trichomes; inorescence
axes covered by mixed or curved trichomes;
carpels glabrous or slightly pubescent ................
A.rmum nothosubsp. zapalowiczii
20*. Pedicels at least in upper parts are covered
by protruding pilose trichomes; iorescence
axes glabrous; carpels glabrous or slightly
pubescent on dorsal side ....... A .rmum subsp.
rmum × A.rmum subsp. maninense
21. Helmet, pedicels, bracteoles and
inoresence axes covered by short curved
trichomes; carpels usually glabrouse, rarely
pubescent by solitary curved trichomes ..........
A.rmum subsp. moravicum
21*. Type of pubescence diers ...................... 22
22. Helmet, pedicels, bracteoles and
inorescence axes covered by protruding pilose
trichomes; carpels usually glabrous, rarely
covered by pilose trichomes ..................................
A.rmum subsp. maninense
22*. Helmet, pedicels, and bracteoles covered
by mixed curved and protruding trichomes;
inorescence axes covered by mixed or
curved trichomes; carpels glabrous or slightly
pubescent ........... A.rmum nothosubsp. paxii
Conclusions
is investigation showed that genus
Aconitum in Slovakia is represented by 21
taxa of species and infraspecic levels, and
probably contains two more unnamed hybrids
belonging in general to 3 subgenera. ree new
taxa for ora of Slovakia (A. degenii subsp.
degenii, A. moldavicum subsp. hosteanum
and A. moldavicum nothosubsp. confusum
stat. nov.) have been ascertained, for one
of which the new taxonomic status has been
proposed. General taxonomic structure of the
genus Aconitum in ora of Slovakia is next:
Subgen. Lycoctonum (DC.) Peterm.
Sect. Lycoctonum DC.
Ser. Lycoctonia Tamura et Lauene
A. lycoctonum L. em. Koelle
subsp. lycoctonum
A. × triste (Rchb.) Gáyer
A. moldavicum Hacq.
subsp. moldavicum
subsp. hosteanum (Schur)
Graebner et P. Graebner in
Aschers. et Graebner
nothosubsp. confusum (Grinț.)
A. Novikov, stat. nov.
Subgen. Anthora (DC.) Peterm.
Sect. Anthora DC.
A. anthora L.
subsp. anthora
subsp. jacquinii (Rchb.) Domin
subsp. anthora × subsp. jacquinii
Genus Aconitum in Slovakia: a phenetic approach 75
Modern Phytomorphology 11, 2017
Subgen. Aconitum
Sect. Cammarum DC.
Ser. Va r iegata Steinberg ex Starmühl.
A. variegatum L.
subsp. variegatum
Ser. To x ic u m (Rchb.) Mucher
A. lasiocarpum (Rchb.) Gáyer
subsp. lasiocarpum
subsp. kotulae (Pawł.) Starmühl.
et Mitka
A. degenii Gáyer
subsp. degenii
A. × gayeri Starmühl.
Nothoser. Toxigata Starmühl.
A. × pawlowskii Mitka et Starmühl.
Nothosect. Acomarum Starmühl.
A. × cammarum L. em. Fries
A. × berdaui Zapał.
nothosubsp. berdaui
nothosubsp. walasii (Mitka in
Starmühler et Mitka) Mitka
Sect. Aconitum
A. rmum Rchb.
subsp. rmum
subsp. moravicum Skalický
subsp. maninense (Skalický)
Starmühl.
nothosubsp. zapalowiczii Starmühl.
nothosubsp. paxii Starmühl.
subsp. rmum × subsp. maninense
Acknowledgements
I would like to thank to all people who helped
in realization of this project and organization
of my stay in Slovakia, especially to RNDr.
Kornélia Goliašová, RNDr. Pavol Mereďa and
Prof. RNDr. Karol Marhold from Institute of
Botany SAS.
I thank Prof., Dr. hab. Józef Mitka from
Institute of Botany of Jagiellonian University
in Kraków for years of help and support in my
research activities, and particularly for help
in this study with comments and for provided
additional localities. I also thank Dr. Walter
Roensteiner from Graz for help in improvement
of this paper, comments about distribution of
aconites in Slovakia and discussion on taxony.
I thank so much to RNDr. Ján Kliment from
Botanical Garden of Comenius University for
help in improvement of distribution database
and decryption of problematic labels, as well as
for critical comments and advices.
I am thankful for different kind of help
in realization of this investigation to Zuzana
Kyselová from Tatra National Park, Jaroslav
Vlcko from Technical University in Zvolen,
Lenka Mártonfiová from Pavol Jozef Šafárik
University, staff of herbarium of Comenius
University in Bratislava, staff of herbarium
of Slovak University of Agriculture in Nitra,
and staff of herbarium of Slovak National
Museum.
is study was supported by grant of National
Scholarship Programme of the Slovak Republic
for the Support of Mobility of Students, PhD
Students, University Teachers, Researchers and
Artists in 2012–2013.
References
Andrienko T.L., Peregrym M.M. 2012. Official lists of
regional rare plants of administrative territories of
Ukraine (reference book). Alterpress, Kyiv.
Annoni A., Luzet C., Gubler E., Ihde J. (eds) 2001.
Map projections for Europe. European Commision.
Joint Research Centre.
Boroń P. 2010. Taksonomia molekularna rodzaju
Aconitum (Ranunculaceae) w Karpatach Zachodnich
i Sudetach. Phd Thesis. Uniwersytet Jagielloriski,
Wydzial Biologii i Nauk o Ziemi, Kraków.
Boroń P., Zalewska-Gałosz J., Sutkowska A.,
Zemanek B., Mitka J. 2011. ISSR analysis points
to relict character of Aconitum bucovinense Zapał.
(Ranunculaceae) at the range margin. Acta Soc.
Bot. Pol. 80: 315–326.
Chopyk V.I. 1978. Rare and endangered plants
of Ukraine. The guide. Naukova Dumka, Kyiv.
(In Russian)
Davis P.H., Heywood V.H. 1963. Principles of
angiosperm taxonomy. Edinburgh University Press,
Edinburgh.
Didukh Ya.P., Zyman S.M., Burda R.I.,
Korotchenko I.A. 2004. Aconitum anthora L. s. str.
– Akonit protyotruyniy. In: Didukh Ya.P. (ed.), Ecoflora
of Ukraine. Vol. 2: 97–98. Phytosociocentre, Kyiv.
Domin K. 1935. Plantarum Čechoslovakiae enumeratio.
Preslia, Praha 13–15: 1–305.
Dostal J. 1958. Klíč k úplné květeně ČSSR.
Nakladatelství ČSAV, Praha.
76 Novikov A.
Modern Phytomorphology 11, 2017
Götz E. 1967. Die Aconitum variegatum-Gruppe und
ihre Bastarde in Europa. Feddes Repert. 76: 1–62.
Grințescu G. 1953. Aconitum L. In: Săvulescu T., Flora
Republicii Populare Române. Vol. 2: 460–511,
676–686.
Hammer Ø., Harper D.A.T., Ryan P.D. 2001. PAST:
Paleontological statistics software package for
education and data analysis. Palaeontol. Electron.
4 (1). http://palaeo-electronica.org/2001_1/past/
issue1_01.htm
Ilnicki T., Mitka J. 2011. Chromosome numbers in
Aconitum sect. Cammarum (Ranunculaceae) from
the Carpathians. Caryologia 64 (4): 446–452.
IPNI 2017. The International Plant Names Index. Published
on the Internet http://www.ipni.org/index.html
QGIS Development Team. 2017. QGIS Geographic
Information System. Open Source Geospatial
Foundation Project. http://www.qgis.org/
Kadota Y. 1987. A revision of Aconitum subgenus
Aconitum (Ranunculaceae) of East Asia. Sanwa
Shoyaku Co., Utsunomiya.
Kmeťová E. 1982. Aconitum L. In: Futák J., BertováL.
(red.), Flóra Slovenska. T. III: 76–97. Véda, Bratislava.
Linnaeus C. 1753. Species Plantarum. T. 1: 532.
Holmiae.
Mitka J. 2003. The genus Aconitum L.
(Ranunculaceae) in Poland and adjacent countries:
A pheneticgeographic study. Institute of Botany of
the Jagellonian University, Cracow.
Mitka J. 2008a. Aconitum moldavicum Hacq.
(Ranunculaceae) and its hybrids in the Carpathians
and adjacent regions. Roczniki Bieszczadzkie 16:
233–252.
Mitka J. 2008b.Tojad Hosta. Aconitum moldavicum
Hacq. subsp. hosteanum (Schur) Graebn. &
P. Graebn. In: Mirek Z., Piękoś-Mirkowa H.
(eds), Czerwona Księga Karpat Polskich. Rośliny
naczyniowe: 78–79. Kraków.
Mitka J., Starmühler W. 2000. Phenetic variability
of Aconitum lasiocarpum (Rchb.) Gáyer
(Ranunculaceae): extention of taxonomic and
gegrafic borders. Acta Soc. Bot. Pol. 62 (2):
145–155.
Mitka J., Szajna B. 2009. A phenetic study on Aconitum
× hebegynum DC. (Ranunculaceae) in the Eastern
Carpathians. Roczniki Bieszczadzkie 17: 253–266.
Mitka J., Boroń P., Sutkowska A. 2013. Holocene
history of Aconitum in the Polish Western
Carpathians and adjacent regions: long-distance
migrations or cryptic refugia? Mod. Phytomorphol.
3: 9–18. https://doi.org/10.5281/zenodo.161587
Mitka J., Boroń P., Wróblewska A., Bąba W. 2015.
AFLP analysis reveals infraspecific phylogenetic
relationships and population genetic structure of
two species of Aconitum in Central Europe. Acta
Soc. Bot. Pol. 84: 267–276.
Mitka J., Boroń P., Novikoff A., Wróblewska A.,
Binkiewicz B. 2016. Two major groups of
chloroplast DNA haplotypes in diploid and tetraploid
Aconitum subgen. Aconitum (Ranunculaceae) in
the Carpathians. Mod. Phytomorphol. 9 (Suppl.):
5–15. https://doi.org/10.5281/zenodo.159700
Novikoff A.V. 2013. Introduction to the taxonomy of the
genus Aconitum L. (Ranunculaceae) in the flora of
Slovakia. Proc. of V Internat. Sci. Conf. (Sumy, 23-25
May 2013). Vol. 1: 303-306. (In Ukrainian)
Novikoff A. 2013–2017. Genus Aconitum L.
(Ranunculaceae) in Slovakia. Published on the
Internet. http://slovaconitum.myspecies.info/
Novikoff A.V., Mitka J. 2011. Taxonomy and ecology of
the genus Aconitum L. in the Ukrainian Carpathians.
Wulfenia 18: 37–61.
Novikoff A., Mitka J., Kuzyarin A., Orlov O.,
Ragulina M. 2016. Some notes on the genus
Aconitum in Chornohora Mts. Mod. Phytomorphol.
9 (Suppl.): 35–73. https://doi.org/10.5281/
zenodo.159703
Paczoski J. 1927. Aconitum. In: W. Szafer (red.), Flora
Polska. T. 3: 19–25. PWN, Warszawa–Krakow.
Reichenbach H.G.L. 1819. Uebersicht der Gattung
Aconitum. Regensburg.
Reichenbach H.G.L. 1820. Monographia generis
Aconiti. Sumtibus F.C.G. Vogelii, Lipsiae.
Reichenbach H.G.L. 1823–1827. lllustratio specierum
Aconiti generis. Lipsiae.
Reichenbach H.G.L. 1838. Icones florae Germanicae
et Helveticae. Bd. I. Sumtibus Ambrosii Abel,
Lipsiae.
Reichenbach H.G.L. 1840. Icones florae Germanicae
et Helveticae, simul Pedemontanae, Tirolensis,
Istriacae, Dalmaticae, Austriacae, Hungaricae,
Transylvanicae, Moravicae, Borussicae, Holsaticae,
Belgicae, Hollandicae, ergo Mediae Europae. Apud
Fridericum Hofmeister, Lipsiae.
Skalický V. 1966. Československé oměje (Aconitum L.).
Živa 52/14 (6): 205–207.
Skalický V. 1982. Notula systematicae, diagnosticae
et nomenclaturicae ad Aconitum generis
investigationem pertinentes. Preslia, Praha 54:
115–122.
Skalický V. 1985. Taxonomische und nomenklatorische
Bemerkungen zu den Gattungen Aconitum L. und
Pulsatilla Mill. Preslia, Praha 57: 135–143.
Skalický V. 1990. Rod Aconitum na Československu.
Zprávy Českoslov. Bot. Spol. 25 (2): 1–27.
Smith V.S., Rycroft S., Scott B., Baker E., Livermore L.,
Heaton A., Bouton K., Koureas D.N., Roberts D.
2012. Scratchpads 2.0: A virtual research
environment infrastructure for biodiversity data.
http://scratchpads.eu
Starmühler W. 1997. Typification of names in the genus
Aconitum. Feddes Repert. 108: 102–103.
Genus Aconitum in Slovakia: a phenetic approach 77
Modern Phytomorphology 11, 2017
Starmühler W. 1998. Ranunculaceae subtrib.
Delphiniinae exsiccatae, Fasc. 3 (Nr. 41–60).
Fritschiana 13: 1–10.
Starmühler W. 2001. Die Gattung Aconitum in Bayern.
Ber. Bayer. Bot. Ges. 71: 99–118.
Starmühler W. 2002. Ranunculaceae subtrib.
Delphiniinae exsiccatae, Fasc. 5 (Nr. 81–100).
Fritschiana 30: 1–12.
Starmühler W., Mitka J. 2001. Systematics and chorology
of Aconitum sect. Napellus (Ranunculaceae) and
its hybrids in the Northern Carpathians and Forest
Carpathians. Thaiszia 10: 115–136.
Sutkowska A., Boroń P., Mitka J. 2013. Natural
hybrid zone of Aconitum species in the
Western Carpathians: Linnaean taxonomy
and ISSR fingerprinting. Acta Biol. Cracov. Ser.
Bot. 55 (1): 114–126. https://doi.org/10.2478/
abcsb-2013-00015
Tamura M. 1995. Aconitum L. In: Engler A., Prantl K.
(eds), Die natürlichen Pflanzenfamilien: 274–291.
Aufl. Dunker & Humblot, Berlin.
The Plant List 2017. Version 1.1. Published on the
Internet. http://www.theplantlist.org/
Tutin T.G., Akeroyd J.R., Chater A.O. 1993.
AconitumL. In: Tutin T.G., Burges N.A., ChaterA.O.,
Edmondson J.R., Heywood V.H., Moore D.M.,
Valentine D.H., Walters S.M., Webb D.A. (eds),
Flora Europaea. 2nd ed. Vol. 1. Psilotaceae to
Platanaceae: 254–256. Cambridge University
Press, Cambridge, New York, Melbourne.
Tzvelev N.N. 2001. Borets – Aconitum L. In: TzvelevN.N.
(ed.), Flora of Eastern Europe. T. X : 55–66. Mir i
semja, Saint-Petersburg. (In Russian)
Visjulina O.D. 1953. Rid 299. Akonit – Aconitum L. In:
KlokovM.V., Visjulina O.D. (eds), Flora of Ukrainian
SSR. Vol. 5: 56–70. Publishing of AS of Ukrainian
SSR, Kyiv.
Voroshilov V.N. 1990. European species from genus
Aconitum L. Bull. Main Bot. Garden 158: 36–41.
(In Russian)
Wacławska-Ćwiertnia K., Mitka J. 2016. Typification of
Zapałowicz’s names in Aconitum section Aconitum.
PhytoKeys 58: 119–126. https://doi.org/10.3897/
phytokeys.58.7110
Zapałowicz H. 1908. Conspectus florae Galiciae
criticus. T. 2: 205–237. Akademia Umiejętności,
Kraków.
Appendix 1. Specimens examined in herbaria
Aconitum lycoctonum subsp. lycoctonum
SLOVAKIA: Muránsky hrad; leg. Hubová O., Kmeťová E., 06.06.1968; (SAV). — Vrch Riečky [Mt. Vrchriečky near
village Litmanová]; leg. Zahradníková K., 05.07.1972; (SAV). — Považský Inovec, Marhát, Aceti–Fagetum; 690 m a.s.l.;
leg. Michalko J., Magic D., 03.06.1965; (SAV). — Veľká Fatra: Blatnická dolina, na svahu SE od Kečerovky [Kačárová];
700 m a.s.l.; leg. Kováčiková, 20.09.1974; (SAV). — Veľká Fatra: Gaderská dolina; leg. Futák J., Jasičová M., 25.06.1963;
(SAV). — Oravský Podzámok: vrch Skalka; 530 m a.s.l.; leg. Futák J., 08.07.1959; (SAV × 2 vouchers). — Bánovce nad
Bebravou [?], Calamagrostis; 803 m a.s.l.; leg. Futák J., 12.07.1960; (SAV). — Manínska nížina, pri potoku, vápenec;
350 m a.s.l.; leg. Futák J., 05.06.1958; (SAV × 2 vouchers). — Orava: Podbieľ, Biela skala; leg. Futák J., Jasičová M.,
24.06.1963; (SAV). — Ostrý kameň; leg. Maglocky Št., 09.08.1967; (SAV). — Ľubochňa; leg. Futák J., Jasičová M.,
23.06.1963; (SAV). — Veľká Fatra: [Kráľova studňa], svetlina v bučine pod chatou; 1100 m a.s.l.; leg. Grebenščikov O.,
02.07.1953; (SAV). — Bánovce nad Bebravou [?], Fagetum; 750 m a.s.l.; leg. Futák J., 15.07.1960; (SAV). — Štubn.
Teplice, Kostelecká dol., svetliny; leg. Horák P., 13.07.1942; (SAV). — Smolenice – hrebeň. Malé Karpaty; leg.
Krippelová T., 21.06.1960; (SAV × 3 vouchers). — Strážovská hornatina, dolomit, bučina; 540 m a.s.l.; leg. Futák J.,
26.07.1961; (SAV × 4 vouchers). — Bánovce nad Bebravou [?], Kanisova skala, dolomit; 700 m a.s.l.; leg. Futák J.,
15.07.1960; (SAV). — Záruby, hřeben; 700 m a.s.l.; leg. Horák P., 07.1923; (SAV). — Tematínske kopce, Modriná; leg.
Michalko J., 03.07.1957; (SAV). — pri Jablonici, okraj lesa; leg. Krippelová T., 07.1967; (SAV). — Strážovská hornatina;
leg. Futák J., Hubová O., 21.05.1954; (SAV). — Malé Karpaty: vápencový hrebeň západne od zrúcaniny Ostrý kameň v
bučine; 500 m a.s.l.; leg. Kollár J., Zahradníková K., 12.06.1968; (SAV). — Stráž. vrchy, medzi obcami Mníchova Lehota
a Kostolné Mitice na hrebeni Maiková a Lapkov kameň, v lese pri ceste v bučine; 600 m a.s.l.; leg. Schidlay E., 01.08.1950;
(SAV). — Vlhké... pod vrcholom Javořiny; leg. ?, 19.07.1911; (SAV). — Revúca [Liptovské Revúce]. Vel. Fatra; 1000m
a.s.l.; leg. Nábĕlek V., 07.1936; (SAV). — Slovenský raj: Sokolia dolina, dolná časť; 600 m a.s.l.; leg. Hubová O., 16.08.1962;
(SAV). — Strečno; leg. Popovič, 21.06.1959; (SAV). — Tematínske kopce, pri potoku Kalištia za obcou Modrová; 220 m
a.s.l.; leg. ?, 31.05.1957; (SAV). — Lubina: Veľká Javorina [Biele Karpaty] – pri turist. chate – prenesené na pokusný
pozemok Bratislava–Patrónka (kultúra 3 roky); leg. Zahradníková K., 16.07.1963; (SAV). — Považský Inovec, kopce nad
Tematínskym Podhradím, vápenec, lúčina; 500 m a.s.l.; leg. Michalko J., Zahradníková K., 08.07.1959; (SAV). — Rajecké
Teplice, in monte Tlstá hora, in fageto; 450 m a.s.l.; leg. Schidlay E., 10.06.1947; (SAV). — Biele Karpaty, Vrbovce, les nad
žel. stanicou Vrbovce, pod vrchom Stráň (550,3 m.m.n.), pri červenej turistickej značke (cesta M. Kudeříkové); 550 m
a.s.l.; leg. Perný M., 12.06.1998; (SAV). — Strážovská hornatina, Strážov (k. 1240 m), lúčne porasty pri kóte; 1150 m
78 Novikov A.
Modern Phytomorphology 11, 2017
a.s.l.; leg. Kollár J., 21.07.1966; (B × 2 vouchers). — Pohoří Rokoše: Smrkový les pod vrcholem Rokoš; 950 m a.s.l.;
leg. Dvořák J., 14.06.1979; (B). — Pohoří Rokoše: in fageto sub cacumine montis Rokoš; 960 m a.s.l.; leg. Dvořák J.,
05.06.1975; (B). — Považský Inovec: dolina Kalištia na V od obce Modrová, pri lesnej ceste v rekr. stredisku; 240 m
a.s.l.; leg. Mucina L., 26.06.1974; (B). — Slovenský raj: Prielom Hornádu, vápence; 550 m a.s.l.; leg. Hajdúk J.,
11.06.1959; (B). — Malé Karpaty: Smolenice, les pod Havraňou skalou, tmavosivý vápenec; 400 m a.s.l.; leg.
VozárováM., 10.07.1996; (B × 2 vouchers). — Strážovské vrchy, dolina medzi V. Málenicou (909 m) a Rohatínom
(832,4 m), v bučine; 520 m a.s.l.; leg. Michalková V., 26.05.2001; (B). — Liptovská Osada – Podsuchá; 700 m a.s.l.;
leg. ?, 16.07.1972; (B). — Vel. Fatra: Čierny kameň –; 1400 m a.s.l.; leg. ?, 21.08.1968; (B). — Malé Karpaty: prope
vici Smolenice, montis Záruby in iugo, ad terra calcareum; 730 m a.s.l.; leg. Michalko J., 08.08.1965; (B). — Stará
Lehota – Kňažný vrch; leg. Maglocký Št., 13.08.1965; (B). — ober Harmanec; leg. ?, 09.07.1857; (B). — pri obci
Topoľčianky; leg. Drévlackanský Fr., 19.06.1958; (B × 2 vouchers). — Na Nitr. Javorine [Panská Javorina] miestami;
leg. Holuby J., 07.1864; (B). — Malé Karpaty,Trnava: in fageto sub arce Ostrý kameň prope pag. Biksard [today
Buková village]; 500 m a.s.l.; leg. Valenta V., 18.07.1940; (B × 2 vouchers). — Kremnica, Bartoška; leg. Zechentner, ?;
(B). — prope Banská Štiavnica – in declivibus borealis, orientalians montis Koelberg in fruticetis – solo humoso –
planta sociabilis; leg. Hlavaček A., 29.05.1936; (B). — Ružomberok: in saxosis calcareis m. Vysoký gruň supra pag.
Ľubochňa; 650 m a.s.l.; leg. Valenta V., 16.06.1971; (B). — Ilava: in convalle inter pagos Beluša et Mojtín; 340 m a.s.l.;
leg. Valenta V., 29.06.1936; (B). — Prievidza: in fruticetis ad marginum silvae in m. Revan ad viam Gajdel – Fačkov;
850 m a.s.l.; leg. Valenta V., 11.07.1935; (B). — Západné Tatry: kataster obce Pribylina, okres Liptovský Mikuláš,
vysokobylinné spoločenstvá okolo potôčkov v kroví; 850 m a.s.l.; leg. Horníčková J., 12.08.1982; (B). — Liptovské
Revúce, Čierny kameň; leg. Grodkovszký G., 13.07.1934; (B × 2 vouchers). — Veľká Fatra, Dedošova dolina, v bučine;
800 m a.s.l.; leg. Uhlířová J., 11.07.1994; (B). — Slovenský raj: Stratená, Dedinky, orientatio W, acclivitas 40°,
substratum CaCO3, terra rendzina; 900 m a.s.l.; leg. Hajdúk J., 31.07.1956; (B). — Biele Karpaty, Veľká Javorina; leg.
Záborský J., 20.08.1972; (SLO × 9 vouchers). — Biele Karpaty, Veľká Javorina, v lese; leg. Záborský J., 16.09.1972; (SLO).
— Zlatnícka dolina pri Skalici, v lese; leg. Záborský J., 01.08.1972; (SLO × 2 vouchers). — Sever. Biele Karpaty, Pruské;
leg. ?, 19.06.1989; (SLO × 3 vouchers). — Biele Karpaty, Veľká Javorina; leg. Záborský J., 14.09.1973; (SLO). — Skalica:
Zlatnícka dolina; leg. Záborský J., 24.07.1965; (SLO × 3 vouchers). — Juž. Biele Karpaty, Skalica: Zlatnícka dolina; leg.
Záborský J., 27.08.1984; (SLO × 2 vouchers). — Juž. Biele Karpaty, Skalica: Zlatnícka dolina; leg. Činčura F., Záborský J.,
02.09.1984; (SLO). — Násyp pri Harmanci; leg. Záborský J., 30.06.1970; (SLO). — Biele Karpaty, Veľká Javorina; leg.
Záborský J., 14.09.1973; (SLO × 3 vouchers). — Slovenský raj: Kyseľ; leg. ?, 27.06.1971; (SLO). — Biely Potok [part of
town Ružomberok]; leg. ?, 25.06.1979; (SLO). — Nemecká – Hradisko; leg. Háberová I., 05.06.1975; (SLO). —
Manínska úžina; leg. Háberová I., 04.06.1976; (SLO). — Palota; leg. Majzlanová E., 10.06.1977; (SLO). — Malá Fatra
– sedlo pod Kľakom; lesy v okolí chaty; leg. Činčura F., 13.07.1968; (SLO × 2 vouchers). — Veľká Fatra – lesy pri chate
pod Δ Úplaz, smerom od Harmanca; leg. Činčura F., 04.07.1968; (SLO). — Malé Karpaty – Ostrý kameň; leg. Filová M.,
08.07.1949; (SLO). — Harmanecká dolina; leg. Michalko J., 28.06.1952; (SLO). — Neresnícka dolina pri Zvolene – v
lesoch vedlá cesty; leg. Činčura F., 14.06.1967; (SLO × 4 vouchers). — Rokoš vs. od Bánovce nad Bebravou; leg. Futák J.,
04.07.1931; (SLO). — Veľká Fatra: dolina Bystrice pod Kráľovou studňou; leg. Futák J., 15.06.1947; (SLO). — Veľká
Fatra: Majerova skala; leg. Futák J., 15.06.1947; (SLO). — Kopce a lúky nad Ľubochňou; leg. Májovský J., 06.07.1951;
(SLO). — Kozel – Biela skala, les; leg. Hollá D., 11.07.1968; (SLO). — Veľká Fatra: Gaderská dolina; leg. Májovský J.,
21.06.1948; (SLO). — Malá Fatra; leg. Futák J., 11.06.1947; (SLO). — Harmanecká dolina; leg. Michalko J., 28.06.1952;
(SLO). — Veľká Fatra: Horný Jelenec; leg. Futák J., 14.06.1947; (SLO). — Bučiny nad Riečkou a dolina B.B. [Banská
Bystrica] vodovodu; leg. Májovský J., 29.06.1952; (SLO × 2 vouchers). — Zvolen; leg. Futák J., 07.1941; (SLO). —
Reváň; leg. Pantocsek J., 02.08.1902; (SLO). — Moravia Occidentalis. Ad connes Slovakiae: Javorníky, in pratis
“Javornické louky”; 1056 m a.s.l.; leg. Říčan G., 07.1924; (SLO × 1 voucher, ZV × 1 voucher). — Juhoslovenský kras; leg.
?, 27.06.1964; (SLO). — Neresnícka dolina pri Zvolene – v lesoch vedlľacesty; leg. Činčura F., 14.06.1967; (SLO × 2
vouchers). — Veľká Fatra: Tlstá; leg. Exkurzia, 22.06.1948; (SLO). — Veľká Fatra: Dedošová dolina; leg. Májovský J.,
21.06.1948; (SLO). — Rokoš: bučina na sev. svahu; 920 m a.s.l.; leg. Futák J., 28.07.1942; (SLO × 2 vouchers). — Malé
Karpaty: Záruby – Buková; leg. Medovič J., 25.08.1955; (SLO × 3 vouchers). — Malé Karpaty: Veterlín; leg. Filová M.,
04.07.1948; (SLO × 2 vouchers). — Malé Karpaty: Driny – Čelo; leg. Berta J., Medovič J., 24.08.1955; (SLO). —
Strážovská hornatina, Rajecké Teplice, hrebeň na Tlstú horu, les; leg. Hallonová, 04.06.1977; (SLO). — Strážovská
hornatina, Lietava, severný svah pod Lietavským hradom, les; leg. Hallonová, 17.06.1978; (SLO). — Remata – Roviny;
leg. Takáč, 17.07.1962; (NI). — Stráž. vrchy: Uhrovské Podhradie, sedlo Rokoš; 945 m a.s.l.; leg. Eliáš P. jun., 22.07.2003;
(NI). — Stráž. vrchy: Δ Baske, v lese pod chatou; leg. Eliáš P. jun., 23.07.2003; (NI × 2 vouchers). — M. Karpaty:
Δ Záruby, hojne okolo turist. chodníka; 755 m a.s.l.; leg. Eliáš P. jun., 21.07.2010; (NI). — Silická planina, 620 m,
“Fujanco”, Z, 15°; leg. ?, 17.05.1946; (NI). — Podsuchá – vzácne na úpätí stráne nad riekou pri letnom tábore; leg. ?,
10.07.1992; (NI). — Vápencové predhorie medzi Kysucou a Varínkou – Brodnianka – Tac–FQ, exp. V–Z, skl. 10–20°;
700 m a.s.l.; leg. Greštiak M., 13.06.1965; (ZV). — Homôľka; leg. Mikoláš V., 15.09.1984; (KO). — pod Tesnou rizňou
[Tesná rizňa – Malá Fatra]; leg. Mikoláš V., 18.08.1984; (KO). — Zádielska dolina; leg. Vojtúň A., 14.06.1968; (KO).
Genus Aconitum in Slovakia: a phenetic approach 79
Modern Phytomorphology 11, 2017
Aconitum moldavicum subsp. moldavicum
SLOVAKIA: Vernár; leg. ?; (SAV). — okres Bánovce nad Bebravou: Skupina Kňažného stola, medzi Ľutovom a
Timoradzou; 525 m a.s.l.; leg. Futák J., 05.07.1956; (SAV). — Zádielska dolina: na strmých skalách; 450 m a.s.l.; leg.
Schidlay E., 05.08.1954; (SAV). — bukový les v doline Čierneho Váhu; 800 m a.s.l.; leg. Futák J., 12.07.1957; (SAV × 2
vouchers). — na hrebeni vrchu Rígeľ medzi Čiernym Váhom a Svarínom, dolomit, smrečina; 850 m a.s.l.; leg. Futák J.,
11.07.1957; (SAV). — Slovakia …; 800 m a.s.l.; leg. Futák J., 19.08.1955; (SAV). — Veľká Fatra; 1180 m a.s.l.; leg.
Hubová O., 27.05.1975; (SAV). — Pusté pole; leg. Futák J., 21.08.1973; (SAV). — Veľká Fatra: Majerova skala; leg.
Nábĕlek V., 15.06.1947; (SAV). — Novoselica [Nová Sedlica] – Príkry – Stužica. Rezervácia [Bukovské vrchy, NP
Poloniny]; leg. Exkurzia, 16.07.1955; (SAV × 4 vouchers). — Okres Snina, Ruský Potok, Malý Bukovec; leg. Exkurzia,
16.07.1955; (SAV × 2 vouchers). — Nízke Tatry, ŠPR Ohnište, údolie potoka Šuštiačka; leg. Drábová J., Hrouda L.,
16.07.1986; (SAV). — Humenská; leg. Maglocký Št., 19.05.1971; (SAV). — Studňa [Muránska planina]; leg. Futák J.,
Zahradníková K., 08.07.1970; (SAV). — Vihorlat, Jovsa, Jovsianska hrabina (štvorec 7198); 162 m a.s.l.; leg. Mráz P.,
28.05.1998; (SAV). — Pusté pole; 1000 m a.s.l.; leg. Futák J., 13.07.1965; (SAV). — Muránska planina: Δ Vrbiarka; 740
m a.s.l.; leg. Šípošová H., Peniašteková M., 17.07.1980; (SAV). — Muránska planina; leg. Zahradníková K., 08.07.1970;
(SAV). — Zádiel, Zádielska dolina; leg. Kmeťová E., 11.06.1969; (SAV). — Korytnica, okraj krovia; 800 m a.s.l.; leg.
Schidlay E., 11.08.1941; (SAV × 2 vouchers). — Veľká Fatra, Horný Jelenec; leg. Futák J., 12.07.1965; (SAV). — Slizké;
410 m a.s.l.; leg. Kliment J., 10.06.1976; (SAV). — Striežovce; leg. Kliment J., 30.06.1976; (SAV × 2 vouchers). — Jelšava;
leg. ?, 20.05.1934; (SAV). — Liptovská Teplička ?; 600 m a.s.l.; leg. Michalko J., 17.07.1972; (SAV). — Stratenská
hornatina: roklina Suchá Belá / pri Hrabušiciach /; 600 m a.s.l.; leg. Hubová O., Tatarková G., 22.07.1966; (SAV). —
Stratenská hornatina: roklina Veľký Sokol; 700 m a.s.l.; leg. Hubová O., Tatarková G., 22.07.1966; (SAV). — vrch
Zámčisko pri Kvetnici, Tilio–Piceetum; leg. Michalko J., 13.06.1984; (SAV). — Muránska plošina: Cigánka, JZ od obce
Muráň, okraj lesa; 900 m a.s.l.; leg. Jasičová M., 19.06.1963; (SAV). — Vyšná Slaná, vrch Radzim; 920 m a.s.l.; leg.
HajdúkJ., 05.08.1956; (SAV). — Studňa, Mochnatá; leg. Futák J., Zahradníková K., 08.07.1970; (SAV). — Veľká Fatra,
vrch Krížna [+ sample from Čorna Riká in Podkarpatská Rus ?]; leg. Nábĕlek V., 07.1936; (SAV). — dolina Lesnica [?];
560 m a.s.l.; leg. Hubová O., 08.1962; (SAV). — Veľká Fatra: Gader, u potoka; leg. Schidlay E., 24.08.1936; (SAV). —
Šturec [Veľká Fatra], južný svah pri cestárskom domčeku; leg. Futák J., 12.07.1965; (SAV). — Slovenské rudohorie – vrch
Stolica – zadný Kyprov; leg. Kliment J., 11.08.1977; (SAV). — Vrch Riečky [Mt. Vrchriečky near village Litmanová]; leg.
Zahradníková K., 05.07.1972; (SAV). — Liptovský Hrádok, na okraji lesa medzi krovím, dolomit, N; 700 m a.s.l.; leg.
Futák J., 10.07.1957; (SAV). — Žiar [časť lúčanskej časti Malej Fatry]; 1010 m a.s.l.; leg. Futák J., 11.07.1957; (SAV). —
Slovakia orientalis, in silvis caeduis supra Kavečany prope oppidum Košice, solo calcareo; 500 m a.s.l.; leg. Hlavaček A.,
21.07.1950; (SAV). — Pieniny, Vysoké skalky, na SZ svahu pri hranici; leg. Michalko J., 22.08.1953; (SAV). — Slovenské
rudohorie, vrch Radzim; 920 m a.s.l.; leg. Hajdúk J., 05.08.1956; (SAV). — Ďurkovec [Bukovské vrchy Mts] (okres
Snina); leg. Májovský J., 05.10.1958; (SAV). — Spišská Nová Ves; leg. Májovský J., 23.08.1955; (SAV). — Riaba skala
(okr. Snina); leg. Májovský J., 04.10.1958; (SAV). — Kráľova Lehota, skaly nad stanicou; leg. Futák J., Jasičová M.,
Zahradníková K., 22.06.1863; (SAV). — Maša [Hnúšťa]; leg. Rošetská K., 12.07.1954; (SAV). — V lese na Šturci u pasu;
leg. Horák P., 13.07.1938; (SAV). — Malé Karpaty, Ostrý kameň; leg. Perný M., 23.10.1997; (SAV). — Muřánské
pohorie: in monte Veľká Stožka, in declivibus silvaticis; 1450 m a.s.l.; leg. Dvořák J., 23.07.1975; (B). — Nízke Tatry,
Liptovský Hrádok, vrch Zapač – východná krovinatá stráň v lese; 700 m a.s.l.; leg. Horníčková J., 07.1983; (B × 3
vouchers). — Slovenský raj: Stratená, Dedinky, acclivitas 40°, substratum Ca, Mg, CO3, terra rendzina; 1000 m a.s.l.; leg.
Hajdúk J., 17.07.1956; (B). — Slovenský raj: Dedinky, Hnilec, orientatio J, acclivitas ca. 35°, substratum calcareus,
terra rendzina, Abieto–Fagetum vegetatio; 1000 m a.s.l.; leg. Hajdúk J., 23.07.1955; (B). — Stratenská hornatina, pri
Dobšinskom hydrostave, kóta 921; leg. Hajdúk J., 19.07.1956; (B). — Slovenský raj: Dobšinská Maša, collis Skala,
umen Hnilec, orientatio V (E), substratum calcit + dolomit, terra rendzina čierna, vegetatio Piceetum; 1000 m a.s.l.; leg.
Hajdúk J., 27.07.1955; (B). — Slovenský raj: Hrabušice, Veľký Sokol, substratum calcit carb., terra litozem, orientatio
NW; 900 m a.s.l.; leg. Hajdúk J., 1956; (B). — Slovenský raj: Radzim, orientatio S (N), acclivitas ca. 40°, substratum
calcit, terra rendzina, vegetatio Abieto–Fagetum; 940 m a.s.l.; leg. Hajdúk J., 13.08.1955; (B). — Slovenský raj: Vernár,
dolomit, Pulsatillo–Pinetum; 1100 m a.s.l.; leg. Hajdúk J., 01.08.1955; (B). — Slovenské rudohorie, montes Muráňská
planina: in jugo saxoso ab arce Muráň ad orientem versus; 1040 m a.s.l.; leg. Vašák V., 18.07.1978; (B × 2 vouchers).
— hrad Muráň; leg. ?, 27.06.1972; (B). — Brzotín – les, výstup na Silickú planinu; leg. ?, 23.06.1972; (B). —
Bardejov; leg. Berganský J., 1923; (B). — Nízke Poloniny: in platis inter pag. Topoľa et Runina; 400 m a.s.l.; leg.
PišútI., 18.07.1962; (B × 2 vouchers). — Revúca, mont. Cigánka supra pag. Muráň; 800 m a.s.l.; leg. Valenta V.,
09.07.1970; (B). — Nízke Tatry: Tále, na okraji řídkého lesa pod silnicí; 750 m a.s.l.; leg. Dvořák J., 17.07.1972;
(B). — Baba bei Lučivná; leg. Grodkovszký G., 26.06.1933; (B). — Tisovec, Voniaca; leg. Vraný V., 06.06.1922;
(B). — N. Tatry, údolí Štiavnica; leg. Balthasar V., 07.1934; (B). — Muráňská vrchovina: Javorníková dolina,
vápencové svahy v horní části doliny; 850 m a.s.l.; leg. Dvořák J., 08.08.1980; (B). — Zádielská dolina; leg. ?,
19.07.1956; (B). — Muráň; leg. ?, 04.07.1962; (B). — Bardejov: Zabavastrasse; leg. Berganský J., 17.06.1922;
(B × 2 vouchers). — Bardejov: Baadgelände; leg. Berganský J., 18.08.1929; (B). — Bardejov: Kapitnoka; leg.
Berganský J., 25.06.1927; (B × 3 vouchers). — Bardejov: Gresak Kert; leg. Berganský J., 10.06.1922; (B × 8
80 Novikov A.
Modern Phytomorphology 11, 2017
vouchers). — Bardejov: Teglam; leg. Berganský J., 14.05.1922; (B × 3 vouchers). — Bardejov: fürdő erdö féle; leg.
Berganský J., 14.06.1922; (B × 2 vouchers). — Bardejov: Bartfeld; leg. Berganský J., 28.06.1922; (B). — Bardejov:
Alte Bleiche; leg. Berganský J., 30.05.1924; (B). — Bardejov: Moliterka; leg. Berganský J., 04.07.1927; (B × 3
vouchers). — Bardejov: Zabava; leg. Berganský J., 17.06.1922; (B). — Bardejov: vedľa potoka pri papierni; leg.
Berganský J., 1924; (B). — Bardejov: Moliterka; leg. Berganský J., 09.06.1922; (B × 2 vouchers). — Bardejov; leg.
Berganský J., 1925; (B × 3 vouchers). — Liptovský Hrádok, vrch Zapač, krovinaté úpätie, SV svah; 670 m a.s.l.; leg.
Horníčková J., 20.06.1973; (B). — Bardejov: Rurňa; leg. Berganský J., 12.06.1924; (B × 3 vouchers). — Bardejov:
Kohlengrund; leg. Berganský J., 02.05.1924; (B × 3 vouchers). — Bardejov: Popova hura Gelände; leg. Berganský J.,
25.05.1924; (B). — Slovenský raj: Dedinky, Gerava, Hnilec, orientatio W, acclivitas 25°, substratum CaCO3, terra
rendzina; 1000 m a.s.l.; leg. Hajdúk J., 17.07.1956; (B). — Ružomberok – Biele skaly; leg. ?, 07.1964; (B). —
Čergov – Bystrá, v bukovom lese na vrchole; leg. Májovský J., 20.06.1947; (SLO × 2 vouchers). — Sninský kameň, v
bučinách na sev. svahu, Aceri–Fagetum; leg. Májovský J., 10.07.1967; (SLO). — Δ Sokol, v bučinách nad dedinou, skala;
leg. Májovský J., 21.05.1968; (SLO). — Vihorlat [vrch], na hrebeni na okraji lesa pri chodníku; leg. Májovský J.,
23.06.1961; (SLO). — Bučiny nad Riečkou a dolina B.B. [banskobystrického] vodovodu; leg. Májovský J., 29.06.1952;
(SLO × 4 vouchers). — Čergov – Bystrá, na hrebeňoch v bukovom lese (pieskovce); 900 m a.s.l.; leg. Májovský J.,
20.06.1947; (SLO). — Čierny Váh: na lúke Stará Poľana; leg. Ferjanec J., 26.07.1965; (SLO). — Skalky nad Dubovou;
leg. Suchá D., 29.06.1952; (SLO). — Pohorie Čergov, Δ Bystrá, v bučine na hrebeni; leg. Májovský J., 20.06.1947; (SLO).
— Pieniny: Vysoké Skalky –; leg. exkurzia, 19.07.1954; (SLO). — Muráň; leg. Michalko J., 08.07.1952; (SLO). — Cestou
z Dlhej Lúky do Muráňa (okr. Revúca) [Muránska Dlhá Lúka]; leg. Opluštilová T., 08.1952; (SLO). — Muráň; leg.
FutákJ., 10.07.1944; (SLO). — Zádiel; leg. Futák J., 27.07.1949; (SLO × 2 vouchers). — Špania Dolina, Panský diel,
Baláže; leg. Michalko J., 27.06.1952; (SLO × 4 vouchers). — Čergov – Solisko, v bukovom lese; leg. Májovský J.,
24.06.1947; (SLO × 2 vouchers). — Nízke Tatry: pri Váhu medzi Lipt. Hrádkom a Lipt. Jánom, vápenec; leg. Futák J.,
16.08.1944; (SLO). — Spišská Nová Ves, ad stationem viae ferr. “Štiavník”; 600 m a.s.l.; leg. Suza J., 20.06.1931; (SLO).
— Flora Leutschoviensis [Levoča], Dolina; leg. Greschik V., 08.1887; (SLO). — Flora Tatrae Magnae, Leutschovie
[Levoča], Dolina; leg. Greschik V., 07.1942; (SLO). — Flora tatrae Magnae, Leutschovie [Levoča], Dolina; leg.
GreschikV., 06.1944; (SLO × 3 vouchers). — Flora Tatrae Magnae, Leutschovie [Levoča], in convalle “Dolina”; leg.
Greschik V., 06.1910; (SLO × 2 vouchers). — Kesmark [Kežmarok], Kleinwäldchen; leg. Greschik V., 06.1886; (SLO).
— Flora tatrae Magnae, Leutschovie [Levoča], in convalle “Dolina”; leg. Greschik V., 07.1920; (SLO). — Flora Tatrae
Magnae, Löcse [Levoča]: Löcsefüred [Levočské Kúpele]; leg. Greschik V., 06.1888; (SLO × 2 vouchers). — Čierny Váh
– Δ Cudzenica pri Malej Lehote, les; 750 m a.s.l.; leg. Vartíková E., 21.07.1972; (SLO × 2 vouchers). — Čierny Váh, lesy
po hájovňu Niž. Chmelinec, S exp.; 750 m a.s.l.; leg. Vartíková E., 18.07.1972; (SLO). — Nízke Tatry: Čierny Váh, Δ
Gregové; 1000 m a.s.l.; leg. Vartíková E., 03.07.1974; (SLO). — Nízke Tatry, Δ Vysoké; 900 m a.s.l.; leg. Vartíková E.,
04.07.1974; (SLO). — Nízke Tatry: Čierny Váh, Δ Gregové; 950 m a.s.l.; leg. Vartíková E., 03.07.1974; (SLO ×
2vouchers). — Nízke Tatry: Δ Muránik, Čierny Váh; 850 m a.s.l.; leg. Vartíková E., 09.07.1974; (SLO). — Korytnická
dolina – svetlé, trávnaté miesta na lesných svahoch vedľa cesty smerom na Donovaly; leg. Činčura F., 06.07.1968; (SLO ×
5 vouchers). — Veľká Fatra: Majerova skala; leg. Futák J., 15.06.1947; (SLO × 2 vouchers). — Slovakia …; leg. MájovskýJ.,
16.06.1949; (SLO). — Kopec medzi Kolbasovom a Uličom (okres Snina); leg. exkurzia, 14.07.1955; (SLO × 3 vouchers).
— Muránska planina; leg. Svobodová Z., 1970; (NI). — Rybník (okres Rimavská Sobota): les na vápencovom podklade
v doline potoka Drienek nad dedinou [now obec Revúca]; leg. Řehořek V., 31.05.1968; (NI × 2 vouchers). — Ľubietovský
Vepor: paseka smerom na Tri Vody; leg. Svobodová Z., 10.06.1968; (NI). — Slovenský kras: Rožňava, skalky; leg. Kostúl
[?], 06.1936; (NI). — Spišská Nová Ves, ad stationem viae ferr. “Štiavník”; 600 m a.s.l.; leg. Suza J., 20.06.1931; (NI). —
Banskobystrická vrchovina, Radvaňská jamka – Varta, Banská Bystrica. Svahy nad Jamkou.; 480 m a.s.l.; leg. Manica M.,
01.06.1960; (ZV). — u Sivce; leg. Mikoláš V., 13.07.1962; (KO). — Kysak, sev. svahy Hradu; leg. Mikoláš V., 29.05.1999;
(KO). — Veľká lúka [Muránska planina] – Cigánka; leg. Mikoláš V., 04.07.1984; (KO). — Turnianske sedlo – Erneho
jask.; leg. Mikoláš V., 19.06.1962; (KO). — Príkrá nad Spišskou Teplicou; leg. Paclová L., 28.06.1956; (TNP). —
Slovenský raj, proti Letanovskému mlynu; leg. Odložilíková L., 30.06.1957; (TNP). — bei Poprad; leg. Scherfel A.,
29.06.1979; (TNP).
Aconitum moldavicum subsp. hosteanum
SLOVAKIA: Košická kotlina, Zádiel; leg. Krippelová, 16.06.1966; (SAV). — Humenné: svahy nad potokom; leg.
Michalko J., 22.06.1959; (SAV). — Nízke Tatry: Skalky pri Ružomberku, v kroví. 2 r.; leg. Nagy G., 06.1969; (SAV).
— Nízke Poloniny, Runina – brehy “Pľaša potoka” pred obcou; 450 m a.s.l.; leg. JZ, 21.08.1970; (SAV). — Slovenské
rudohorie – vrch Stolica – zadný Kyprv; leg. Kliment J., 11.08.1977; (SAV). — Jovsa, les pri obci; leg. MichalkováE.,
17.06.1992; (SAV). — Liptovský Hrádok, na okraji lesa medzi krovím [?], dolomit, N; 700 m a.s.l.; leg. FutákJ.,
10.07.1957; (SAV). — v lese na vrchu Sokol [vrch v Žiari]; 950 m a.s.l.; leg. Futák J., 12.07.1957; (SAV). — Novosedlica
– Príkry – Stužica. Rezervácia [Bukovské vrchy, NP Poloniny]; leg. exkurzia, 16.07.1955; (SAV). — Slovenské rudohorie:
Hrušovo, vápence; 350 m a.s.l.; leg. Hajdúk J., 21.05.1968; (B × 3 vouchers). — Bardejov; leg. BerganskýJ., 20.06.1929;
(B). — Muráňská vysočina, Muráň; leg. Skřivánek V., 06.07.1951; (B). — Slovenský raj: Dobšinská Maša, collis
Genus Aconitum in Slovakia: a phenetic approach 81
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na skalu, umen Hnilec, substratum calcit, terra rendzine; 850 m a.s.l.; leg. Hajdúk J., 31.07.1956; (B). — Slovenské
rudohorie, montes Muráňská planina: in jugo saxoso ab arce Muráň ad orientem versus; 1040 m a.s.l.; leg. VašákV.,
18.07.1978; (B). — Nízke Poloniny: brehy potoka “Plaša” pri križovatke na Runinu; 450 m a.s.l.; leg. Gallo A.,
18.06.1970; (B). — Nízke Poloniny: in valle rivi inter pag. Topoľa et Runina, loco umbroso; 380 m a.s.l.; leg. PišútI,
18.07.1962; (B). — Muráň, lesy; leg. Blaný T., 18.06.1947; (B). — Baba ; leg. Grodkovszký G., 26.06.1933;
(B). — Bardejov; leg. Berganský J., 18.08.1929; (B). — Bardejov: Baadgelände; leg. Berganský J., 18.08.1929;
(B). — Bardejov: Zabavastrasse; leg. Berganský J., 17.06.1922; (B). —Zádielská dolina; leg. ?, 19.07.1956; (B).
— Hnilec, potok, vápencové skály, Z 20°; 900 m a.s.l.; leg. Marva V. [?], 06.07.1958; (B). — Bardejov: Zabavastrasse;
leg. Berganský J., 17.06.1922; (B × 10 vouchers). — Bardejov: Kľušov; leg. Berganský J., 1925; (B). — Bardejov:
Stationsgelände; leg. Berganský J., 12.06.1929; (B). — Bardejov: Baadgelände; leg. Berganský J., 18.08.1929; (B).
— Bardejov; leg. Berganský J., ?; (B × 14 vouchers). — Bardejov; leg. BerganskýJ., 1927; (B). — Bardejov: Zabava;
leg. Berganský J., 17.06.1922; (B). — Bardejov: Moliterka; leg. Berganský J., 20.06.1929; (B × 2 vouchers). —
Bardejov: Kohlengrund; leg. Berganský J., 02.05.1924; (B). — Bardejov: Gresak Kert; leg. Berganský J., 10.06.1922;
(B). — Slovenský raj: Hrabušice, Malý Sokol, horná časť, orientatio NW, acclivitas 40°, substratum Ca, Mg, CO3,
terra rendzina, vegetatio Acerion; 700 m a.s.l.; leg. Hajdúk J., 07.08.1955; (B). — Čergov – Bystrá, v bukovom lese na
vrchole; leg. Májovský J., 20.06.1947; (SLO). — Vihorlat [vrch], na hrebeni na okraji lesa pri chodniku; leg. MájovskýJ.,
23.06.1961; (SLO). — Kopce medzi Darou [obec Dara] a Starinou (okr. Snina); leg. Exkurzia, 14.07.1955; (SLO).
— Pieniny: Vysoké skalky; leg. exkurzia, 19.07.1954; (SLO). — Čergov – Šoltýsova hora v bukovom lese na vrchole;
leg.MájovskýJ., 23.06.1947; (SLO × 2 vouchers). — Cestou na Muráňsku planinu; leg. Opluštilová T., 08.1952; (SLO).
— Muráňska vysočina: Cigánka; leg. Futák J., 11.07.1944; (SLO). — Flora Tatrae Magnae, Leutschovie [Levoča], Dolina;
leg. GreschikV., 07.1890; (SLO). — Flora Tatrae Magnae, Leutschovie [Levoča], in convalle “Dolina”; leg. GreschikV.,
06.1910; (SLO). — Flora Leutschoviensis [Levoča], Dolina; leg. Greschik V., 07.1926; (SLO). — Flora Tatrae Magnae,
Löcse [Levoča]: Löcsefüred [Levočské Kúpele]; leg. Greschik V., 06.1888; (SLO). — Korytnická dolina – svetlé, trávnaté
miesta na lesných svahoch vedľa cesty smerom na Donovaly; leg. Činčura F., 06.07.1968; (SLO). — Hrabušice – Podlesok;
leg. exkurzia, 08.07.1955; (SLO). — Baba [vrch] nad Svitom; 750 m a.s.l.; leg. Odložilíková L., 13.07.1956; (TNP).
Aconitum moldavicum nothosubsp. confusum
SLOVAKIA: Spišská Nová Ves; leg. Májovský J., 23.08.1955; (SAV).
Aconitum × triste
SLOVAKIA: Meliata, vlhké lesy; leg. Karasová E., 16.06.2005; (SAV × 5 vouchers).
Aconitum anthora subsp. anthora
SLOVAKIA: Bukovské vrchy, Stinská, hraničný hrebeň; leg. Marhold K., 18.08.1988; (SAV). — Šivec [vrch]; leg. FutákJ.,
30.08.1950; (SAV). — Biela skala; leg. Futák J., 26.08.1950; (SAV × 2 vouchers). — Sokol pri Humennom; leg.
MájovskýJ., 27.08.1950; (SAV). — Juhoslovenský kras: Felsőhegy [Horný vrch] pri Zádieli; leg. Michalko J., Popovič,
30.07.1956; (SAV). — Zádielská dolina, na strmých skalnatých svahoch na dne, sreby na dne údolia; 350 m a.s.l.; leg.
Schidlay E., 05.08.1954; (SAV). — Vihorlat: [Humenský] Sokol, váp. skalky, Fageto–Quercetum; 300 m a.s.l.; leg.
MichalkoJ., 26.08.1954; (SAV). — Slovenský raj: Skrik nad Letanovským mlynom, kroviny, exp. J.; 750 m a.s.l.; leg.
Hubová O., 22.08.1961; (SAV × 2 vouchers). — Slovenské rudohorie, vrch Radzim, exp. J., skaly; 945 m a.s.l.; leg.
HajdúkJ., 05.08.1956; (SAV). — Vihorlat: [Humenský] Sokol, váp. skalky, Fageto–Quercetum; 300 m a.s.l.; leg. MichalkoJ.,
26.08.1954; (SAV × 2 vouchers). — Skalka, Jelšava; leg. ?, 02.08.1953; (SAV). — Zádielská dolina, na skalách na dne
údolia; 420 m a.s.l.; leg. Schidlay E., 05.08.1954; (SAV). — Pohornádie: Skalky pri M. Vieske severne od Košíc; leg.
Michalko J., 19.09.1956; (SAV). — Zádielská rokle; 400 m a.s.l.; leg. Horák P., 14.07.1936; (SAV). — Zvolen; 450 m a.s.l.;
leg. Futák J., 20.08.1961; (SAV). — Kováčovské skály, Hronská Breznica; leg. Nábĕlek V., 09.1936; (SAV). — Zádielská
dolina; leg. Krč J., 29.07.1954; (SAV). — Slovenský raj: Tomášovský výhľad; leg. Zahradniková K., 20.08.1961; (SAV).
— Zádielská dolina; leg. Fabianková K., 12.09.1972; (SAV). — Malá Vieska – Trebejov; leg. Fabianková K., 23.08.1973;
(SAV). — Jelšava; 510 m a.s.l.; leg. Kliment J., 28.08.1977; (SAV). — Slovakia Orientalis, ad saxa calcarea in valle Hájska
dolina prope Háj apud opp. Košice; 350 m a.s.l.; leg. Hlavaček A., ?; (SAV). — Pohornádie: váp. skalky na Šivci pri Ružíne,
hrebeň; 780 m a.s.l.; leg. Michalko J., 23.08.1960; (SAV). — Bratislava, pestované v záhrade p. Černého (pochádza z
okolia Parkána [Štúrovo]); leg. Schidlay E., 13.09.1944; (SAV). — Juhoslovenský kras: Zádielská dolina. V spodnej časti
rokle; leg. Krippelová T., Zahradníková K., 26.09.1962; (SAV). — Zobor [vrch]; leg. Jasičová M., Fabianková K.,
12.10.1972; (SAV). — Nitra – Zobor [vrch]; leg. ?, 24.07.1966; (SAV). — Zádielská dolina; leg. Krippelová T.,
26.08.1966; (SAV). — Slovakia media, in fruticetis iugi montis Pešianska prope Kotionem viae ferroviae Hronská
Dúbrava, solo audes; 500 m a.s.l.; leg. Hlavaček A., 27.07.1956; (SAV). — Slovakia australis, distr. Plešivec, regio Slovenský
kras: ad margines sylvae deciduae collis Čertova diera prope Domica, solo calcareo; 400 m a.s.l.; leg. Deyl M., 03.09.1933;
82 Novikov A.
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(B). — Gemerské rudohorie: na južnom svahu kopca Cíger na V od obce Jelšava; 320 m a.s.l.; leg. Gallo A., 30.08.1969;
(B). — Kováčovske kopce; leg. ?, 09.09.1969; (B). — Humenské chrbáty: svah na Z od obce Oreské [okres
Michalovce]; 220 m a.s.l.; leg. Gallo A., 03.10.1969; (B). — Slovenské rudohorie: Stolické vrchy, Brdárka, Vyš. Slaná,
Radzim, orientatio medit., acclivitas 45°, substratum Ca, Mg, CO3, vegetatio Spiraetum mediae; 930 m a.s.l.; leg. Hajdúk J.,
13.08.1955; (B). — Slovenské rudohorie: Stolické vrchy, Brdárka, Vyš. Slaná, Radzim, orientatio SW, acclivitas 20°,
substratum Ca, Mg, CO3, terra rendzina, litozem, vegetatio Spiraetum mediae; 940 m a.s.l.; leg. Hajdúk J., 04.08.1957;
(B). — Slovenský raj: Dedinky, Biele Vody – collis Geravy, Baranie rohy, orientatio J (S), acclivitas 10°, substratum
calcit, terra litozem, vegetatio Festucetum pallentis; 1000 m a.s.l.; leg. Hajdúk J., 02.09.1956; (B × 2 vouchers). —
Muráň; leg. Blaný T., 25.08.1950; (B). — Nitra, pod Zoborom; leg. ?, 20.09.1959; (B × 4 vouchers). — Kovačovské
kopce, Kováčov; 320 m a.s.l.; leg. Valenta V., 25.08.1936; (B × 3 vouchers). — Revúca, Muráň; 925 m a.s.l.; leg.
ValentaV., 11.08.1937; (B). — Hronská Breznica; leg. Nábĕlek V., 09.1936; (B). — Muráňyi fensik [Muránska
planina]; leg. ?, 08.1953; (B). — Muráň; leg. ?, 25.08.1950; (B). — Slovensko již., Nitra, v lese na Zoboru; 350 m
a.s.l.; leg. Plaskavka J., 06.08.1936; (B). — Nitra, pod Zoborom; leg. ?, 20.09.1959; (B × 8 vouchers). — Gemerské
rudohorie: J svahy vrchu Dúbrava na S od obce Jelšava; 420 m a.s.l.; leg. Gallo A., 24.08.1970; (B). — Prierva Hornádu;
leg. Májovský J., 05.09.1960; (SLO). — Slovenský kras: Hrušov, vápencové svahy; leg. Májovský J., Záborský J.,
08.09.1960; (SLO). — Turniansky hradný vrch; leg. Májovský J., 15.09.1966; (SLO). — Turčianský hradný vrch; leg.
Májovský J., 15.09.1966; (SLO). — Hôrky a Vápenicé (vápence južne od vrchu Ploská); leg. Eliáš P., 02.09.1968; (SLO ×
2 vouchers). — Jelenec: gaštanica, v svetlom lese (dub, gaštan); leg. Májovský J., 07.10.1965; (SLO). — Tisovec, Δ
Čeremošná, v bučinách hrebeňa od kameňolomu na Δ Martinová, čistinky; 850 m a.s.l.; leg. Májovský J., Magic D.,
18.08.1970; (SLO × 2 vouchers). — Klátova Nová Ves, Skalka, posled. vrch na hrebeni J Δ Kostrín; 440 m a.s.l.; leg.
Kováčiková, 19.09.1972; (SLO). — Veľká Lodina, vrch Babiná, Quercetum pubescentis; leg. Futák J., 08.09.1946; (SLO).
— Kováčovské kopce [now pohorie Burda], medzi Kováčovom a Kamendinom; leg. Futák J., 22.08.1948; (SLO × 3
vouchers). — Zádielska dolina; leg. Futák J., 27.07.1949; (SLO). — Šivec, pri obci Ružín; leg. Futák J., 30.08.1950;
(SLO). — Dreveník, obec Levoča; leg. Májovský J., 27.08.1955; (SLO × 3 vouchers). — Kysak, Trebejov, Malá Vieska
[now Družstevná pri Hornáde]; leg. Bosáčková E., 11.09.1956; (SLO × 3 vouchers). — Margecany: Kurtova skala; leg.
Bosáčková E., 12.09.1956; (SLO). — Kysak; leg. Futák J., 10.09.1946; (SLO). — Dvorník pri obci Spišské Podhradie;
leg. Futák J., 11.09.1946; (SLO). — Sokol pri Humennom; leg. Májovský J., Michalko J., 14.09.1950; (SLO × 3
vouchers). — Zobor [vrch] pri Nitre, v lese; 350 m a.s.l.; leg. Futák J., 23.07.1942; (SLO). — medzi Budčou a
Dúbravou, okres? Zvolen; leg. Futák J., 18.07.1933; (SLO). — Slovakia …; leg. ?, 08.1941; (SLO). — Slovakia …; leg.
Futák J., 15.09.1948; (SLO). — Krásna Hôrka pri Rožňave; leg. Futák J., 24.08.1947; (SLO × 2 vouchers). —
Juhoslovenský kras, Hrušov, Ičast; leg. Futák J., 03.09.1949; (SLO). — Juhoslovenský kras, Hrušov, II čast; 580 m
a.s.l.; leg. Futák J., 03.09.1949; (SLO × 2 vouchers). — Dreveník; leg. ?, 08.1889; (SLO). — Muránska planina, Δ
Cigánka; leg. Záborský J., 25.09.1984; (SLO). — Δ Sokol – exp. S–V; leg. Hudáková, 20.08.1975; (SLO). — Δ Sokol
– skaly, exp. V–J; leg. Hudáková, 29.07.1975; (SLO). — Δ Podskalka, skaly, exp. V; leg. Hudáková, 07.08.1974; (SLO).
— Δ Podskalka, skaly, exp. V; leg. Hudáková, 18.08.1975; (SLO). — Δ Podskalka, skaly, exp. V; leg. Hudáková,
18.08.1975; (SLO). — Váp. skala nad Jasovom; leg. Jurko A., 20.08.1950; (SLO). — Mladina v horu nad ?; leg.
Záborský J., 10.08.1959; (SLO × 3 vouchers). — Vrch pri obci Veľká Lodina; leg. Futák J., 25.08.1947; (SLO). —
Ružín na záp. od Kysaku; leg. Futák J., 04.09.1946; (SLO × 3 vouchers). — Malá Lodina; leg. Futák J., 08.09.1946;
(SLO). — Belá hora; leg. Jurko A., 07.08.1959; (SLO). — Vihorlat: Sokol, rúbaň po Quercus, vápenec, J svah; leg.
Michalko J., 14.09.1950; (SLO × 6 vouchers). — Sokol pri Humennom; leg. Májovský J., Michalko J., 27.08.1950;
(SLO). — Vihorlat: Dupne [Dúpna jaskyňa], v Quercus; 300 m a.s.l.; leg. MichalkoJ., 11.09.1953; (SLO). — Nitra, Δ
Zobor; leg. Svobodová Z., 08.07.1955; (NI × 2 vouchers). — Pov. Inovec, Topoľčany: Závada, juž. svah Δ Vinište, v
riedkom poraste, pospolite asi 300 rastl.; 400 m a.s.l.; leg. Eliáš P. jun., 10.09.2004; (NI). — distr. Nitra: in locis
stepposis fruticosisque monti Zobor apud opp. Nitra, solo calcareo; 550 m a.s.l.; leg. OsvačilováV., 13.09.1953; (NI).
— distr. Nitra: in rupibus ad decl. septentr. sub summa montis Zobor supra opp. Nitra, solo andesitico; 550 m a.s.l.;
leg. Osvačilová V., 11.07.1953; (NI). — Kozlica – Jánova Ves; leg. Takáš, 14.09.1962; (NI). — Zádiel, hor. č.; leg.
Mikoláš V., 23.08.1984; (KO). — Plešivec – Hrad; leg. Mikoláš V., 12.09.1984; (KO). — Ružinské skaly; leg.
MikolášV., 20.08.1959; (KO). — Muránska planina – Hrad; leg. Mikoláš V., 24–25.08.1959; (KO × 2 vouchers). —
Turňa – pod hradom; leg. Vojtúň A., 04.08.1966; (KO). — Jasovská skala; leg. Vojtúň A., 20.08.1977; (KO). — Zádiel,
prvý balkón [Zádielska tiesňava]; leg. Vojtúň A., 22.09.1967; (KO). — Kurtova skala u Margecan; leg. PaclováL.,
25.07.1957; (TNP).
Aconitum anthora subsp. jacquinii
SLOVAKIA: Slovenské rudohorie, vrch Radzim, exp. J., skaly; 945 m a.s.l.; leg. Hajdúk J., 05.08.1956; (SAV). — Spišské
kotliny: Spišský hradný vrch; leg. Marciová, 19.08.1996; (SAV). — Slovenské rudohorie: Stolické vrchy, Brdárka, Vyš.
Slaná, Radzim, orientatio medit, acclivitas 40°, substratum Ca, Mg, MgCO3, terra rendzina, litozem, vegetatio Spiraetum
mediae; 940 m a.s.l.; leg. Hajdúk J., 26.08.1955; (B). — Dreveník, obec Levoča; leg. Májovský J., 27.08.1955; (SLO).
— in saxis meridionalis ad Dreveník; leg. Greschik V., 08.1910; (SLO). — in monte Dreveník; leg. Greschik V., 07.1910;
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(SLO). — Dreveník; leg. Vojtúň A., 26.09.1964; (KO). — Kirchdrauf – Drewenyik [Dreveník]; leg. Scherfel A.,
08.08.1924; (TNP). — in Garten, Drewenyik [Dreveník]; leg. Scherfel A., 08.1891; (TNP). — Drewenyik [Dreveník];
leg. Waldebrenner, ?; (TNP).
Aconitum anthora subsp. anthora × A. anthora subsp. jacquinii
SLOVAKIA: Dreveník, obec Levoča; leg. Májovský J., 27.08.1955; (SLO). — in monte Dreveník; leg. Greschik V.,
07.1910; (SLO).
Aconitum variegatum subsp. variegatum
SLOVAKIA: Slovenské rudohorie: Galmus, okr. Sp. N. Ves, pravá strana Belej, v úžľabine; leg. Hajdúk J., 22.08.1959;
(SAV). — Tatry. Faixová a Votrubova chata; leg. Horák P., 25–26.08.1937; (SAV). — Liptovská kotlina: Pribylina; leg.
Futák J., 18.09.1970; (SAV × 2 vouchers). — Slovenské rudohorie – vrch Stolica – zadný Kyprov; leg. KlimentJ.,
11.08.1977; (SAV). — Slovenský raj: Suchá Belá, vápencová roklina; leg. Zarzycki, Zahradniková K., 14.09.1970; (SAV).
— Čergov; leg. Flašíková, 21.05.1969; (SAV). — M. F. [Malá Fatra], Kľak , cesta na Čičmany; leg. ZahradniíováK.,
23.07.1975; (SAV). — Veľká Fatra: Blatnická dolina; leg. Zahradníková K., 27.08.1972; (SAV × 2 vouchers). — Belianske
Tatry, Kotlina – Skalné vráta; leg. Michalko J., Zahradníková K., 07.08.1972; (SAV). — Slovakia …; leg. PtačovskýJ.,
11.09.1954; (SAV). — Biela skala [in massif of Sivý vrch?], in sylve in alpestr; 1385 m a.s.l.; leg. Schidlay E., 09.09.1953;
(SAV). — Pusté Pole; leg. Peniašteková M., 21.08.1973; (SAV × 3 vouchers). — Nizke Tatry, sev.-vých. Pustia, Iľanovská
dolina; leg. Zahradníková K., 16.10.1962; (SAV × 2 vouchers). — Chočské pohorie, Prosiecka dolina; leg. ZahradníkováK.,
Jasičová M., 10.08.1966; (SAV). — Nízke Tatry, vch Siná; leg. Zahradníková K., Fabianková [Goliašová] K., 15.08.1972;
(SAV × 2 vouchers). — Západné Tatry, Biela skala; leg. Marhold K., 08.07.1986; (SAV). — Pilsko, Babia hora; leg.
Futák J., Magic D., 10.08.1964; (SAV). — pod vrcholom Klenovského Vepra (Slovenské rudohorie); leg. LetzR.,
23.08.1993; (SAV). — vrch Sitno; leg. Zahradníková K., 08.07.1972; (SAV). — Slovenský raj: Stratenská hornatina,
substrat calcit; 900 m a.s.l.; leg. Hajdúk J., 09.08.1956; (B). — Vys. Tatry; leg. Szép R., 20.08.1904; (B). —
Slovenský raj, Stratenské vrchy. Dobšiná. Andrejisko, hrebeň Andrejiská. Orientatio SN, acclivitas 30°, substratum kalcit,
terra rendzina, vegetatio Abieto–Fagetum; 1200 m a.s.l.; leg. Hajdúk J., 09.08.1956; (B). — Západné Tatry: Liptovský
Hrádok, krovitý breh – Džadovica; 680 m a.s.l.; leg. Horníčková J., 25.08.1976; (B). — Slovenský raj: Vernár, dolomit,
rendzina, Pinetum sylv.; 1000 m a.s.l.; leg. ?, ?; (B). — Slovenský raj: Hnilec, orientatio V(E), substratum calcit; 850m
a.s.l.; leg. Hajdúk J., 31.07.1956; (B). — Stratenská hornatina, Vernár, juž. exp.,vápence; 1000 m a.s.l.; leg. HajdúkJ.,
16.08.1956; (B). — Nízke Poloniny: brehy “Pľaša potoka” pri križovatke na Runinu; 450 m a.s.l.; leg. Gallo A.,
21.08.1970; (B). — Ružomberok: in convalle rivi Lubochňanka ad dom. tenat. Vyš. Tajch; 800 m a.s.l.; leg. ValentaV.,
16.08.1940; (B). — Bel. Tatry: Muráň; 1500 m a.s.l.; leg. Nábĕlek V., 07.1936; (B). — Bel. Tatry: in declivibus
supra pag. Tatranská Kotlina; 1000 m a.s.l.; leg. Valenta V., 08.08.1936; (B × 2 vouchers). — Vysoké Tatry: Bielovodská
dolina; 1700 m a.s.l.; leg. Drévlackanský Fr., 07.10.1962; (B). — Horná Nitra, cestou M. Kľakk – V. Kľak, na horských
lúkach; 1350 m a.s.l.; leg. Drévlackanský Fr., 14.08.1963; (B × 3 vouchers). — Dobšiná, pri jaskyni; leg. ?, 07.08.1962;
(B × 2 vouchers). — Slovenský raj: Stratenské vrchy, Dobšiná, Vernár, collis Kopanec, od cesty Stratená – Hrabušice
ca. 250 m, orientatio J (S), acclivitas ca 30°, substratum calcit, terra rendzina, vegetatio Cephalanthero–Fagetum; 1100m
a.s.l.; leg. Hajdúk J., 16.08.1956; (B). — Slovenský raj: Stratená, Vernár, collis Havrania skala, orientatio 35° J (S),
substratum calcit, terra rendzina, vegetatio Aceretum pseudoplatani; 1000 m a.s.l.; leg. Hajdúk J., 03.08.1955; (B). —
Slovenský raj: Stratenské vrchy, Stratená, collis Roveň (Havrania skala), substratum CaCO3, terra rendzina, vegetatio
Fagetum; 1000 m a.s.l.; leg. Hajdúk J., 24.08.1956; (B). — Slovenský raj: Stratená, collis Andrejisko, na vrchole kopca,
orientatio N, acclivitas 20°, substratum calcit, terra rendzina, vegetatio zmiešaný les; 1100 m a.s.l.; leg. Hajdúk J.,
01.08.1956; (B). — Slovenský raj, Stratenská hornatina, Dobšinská Maša, horná časť prehodý, orientatio S (N),
substratum calcit, terra rendzina, vegetatio Abieto–Fagetum; 900 m a.s.l.; leg. Hajdúk J., 31.07.1956; (B). — Slovenský
raj: Dobšiná, collis Andrejisko, od ľadovej jaskyne 2,8 km, orientatio S (N), acclivitas ca 30°, substratum calcit, terra
rendzina, vegetatio Cephalanthero–Fagetum; 1000 m a.s.l.; leg. Hajdúk J., 09.08.1956; (B). — Nízké Tatry: vápencové
svahy nad Trangošskou dolinou pod “Kozími chrbáty”; 1750 m a.s.l.; leg. Dvořák J., 11.07.1966; (B). — Nízké Tatry:
Prostredná dolina, na okraji olšiny poblíž Kyslé; 750 m a.s.l.; leg. Dvořák J., 15.07.1972; (B). — Stratená; leg. ?,
09.08.1949; (B). — Tisovec (Voniaca) ; leg. ?, 12.08.1948; (B). — Dobšiná, pri jaskyni; leg. ?, 07.08.1962; (B).
— Strážovská hornatina, Strážov (k. 1214 m), lúčne porasty pri kóte; 1150 m a.s.l.; leg. Kollár J., 21.07.1966; (B). —
Staškov, v záhrade; leg. ?, 26.08.1960; (B × 3 vouchers). — Slovenský raj, na pravom brehu Bieleho potoka; leg.
Vyparina St. [?], 16.08.1958; (B). — Slovenský raj, na pravom brehu Bieleho potoka, 1.5 km južne od Sokolovej
doliny; 600 m a.s.l.; leg. Vyparina St. [?], 16.08.1958; (B). — Horná Nitra, cestou M. Klák – V. Klák, na horských
lúkach; 1350 m a.s.l.; leg. Drévlackanský Fr., 14.08.1963; (B × 3 vouchers). — Veľká Fatra, in convalle uvii
Lubochňanka ad dom. tenat. Vyš. Tajch; 800 m a.s.l.; leg. Valenta V., 16.08.1940; (B). — Brezno; leg. Zechentner, ?;
(B). — Ružomberok – Biele skaly; leg. Hodoval V., ?; (B). — Cestou na Čergov; leg. Drévlackanský Fr., 09.08.1957;
84 Novikov A.
Modern Phytomorphology 11, 2017
(B × 5 vouchers). — Muránska planina: Muráň, hradný vrch, nádvorie hradu; leg. Kochjarová J., 06.08.1985; (B).
— Liptovská Osada – Podsuchá, Vel. Brankov, pod Kuruckou skalou; leg. ?, 16.07.1972; (B). — Vel. Fatra – Čierny
kameň; 1400 m a.s.l.; leg. [Horváthová ?], 21.08.1968; (B). — Nízké Tatry: Pusté Pole, na břehu menšího rybníka
poblíž hájovny; 850 m a.s.l.; leg. Dvořák J., 17.08.1976; (B). — Chočské pohorie: Veľký Choč; leg. ?, 06.1964; (B).
— Tisovec, Voniaca; leg. Blaný T., 12.08.1948; (B). — Veľká Fatra: Gaderská dolina, in fundo vallis, ad marginae
silvae; leg. Uhlířová J., 18.09.1993; (B × 2 vouchers). — Vysoké Tatry: Tatranské Matliare, SV od osady, dlhý les a
mokriny, podmáčaná smrečina; 850 m a.s.l.; leg. Vozárová M., 23.08.1995; (B). — Slovenský raj: ústie doliny Veľký
Sokol za hájovňou; 800 m a.s.l.; leg. Gallo A., 24.08.1971; (B). — Veľká Fatra: Tlstá, vrchol; leg. Bernátová [?],
02.09.1976; (B). — Belanské Tatry: cestou z Tatranskej Kotliny; leg. Májovský J., 04.09.1960; (SLO). — Muránska
planina, Δ Cigánka (hrad); leg. Záborský J., 25.09.1984; (SLO). — Dolina Kyseľ (Slovenský raj); leg. Hejná, Rošetzká,
21.08.1951; (SLO). — Dolina Kyseľ (Slovenský raj); leg. Hejna, Rošetzká, 21.08.1951; (SLO × 3 vouchers). — Δ Solisko,
na okraji lesa, pri hrebeňovom chodníku, v Nardetum; 1000 m a.s.l.; leg. Májovský J., 07.1955; (SLO × 4 vouchers). —
Oravská Polhora: vystup na Babiu horu?; leg. Futák J., 25.08.1964; (SLO). — Západ. Tatry: Roháčske plesá; leg.
ŠomšákL., 08.1972; (SLO). — Veľká Fatra: Bukovina [Bukovinka]; leg. Michalková V., 05.09.1962; (SLO). — Poprad;
leg. Šmarda J., Paclová L., 11.09.1956; (SLO). — Vysoké Tatry: Zadné Meďodoly; 1350 m a.s.l.; leg. Májovský J.,
20.09.1967; (SLO × 5 vouchers). — Vysoké Tatry: Zadné Meďodoly, pri lesnej ceste do Javoriny, v smrečine; 1350m
a.s.l.; leg. Májovský J., 20.09.1967; (SLO). — Skala na V. Fatre, vápence; 1100 m a.s.l.; leg. Futák J., 11.08.1942; (SLO).
— Záp. Beskydy: Oravská Lesná, Juzikovka; leg. Murín A., 25.07.1983; (SLO × 2 vouchers). — Jastub na skalkach v
lese; leg. Májovský J., 18.09.1947; (SLO). — Roszutec [Veľký Rozsutec]; leg. Futák J., 23.08.1943; (SLO). — Tatr.
Kotlina; leg. Futák J., 28.08.1954; (SLO × 2 vouchers). — Belianske Tatry: Ždiarska Vidla – Havran; leg. Májovský J.,
23.08.1957; (SLO × 2 vouchers). — Turzovka; leg. Halbová I., 20.08.1962; (SLO). — Lúky pod Vys. Tatrami, cestou
z Podbanska na Štrbské Pleso; leg. Pastýrik Ľ., Májovský J., 27.08.1941; (SLO × 2 vouchers). — Flora Leutschoviensis
[Levoča]; leg. Greschik V., 07.1890; (SLO). — Leutschovia [Levoča], Gr. Rehberg; leg. Greschik V., 07.1918; (SLO).
— Flora Leutschoviensis [Levoča], Dolina; leg. Greschik V., 07.1889; (SLO). — Flora Tatrae Magnae, Leutschovia
[Levoča]; leg. Greschik V., 07.1928; (SLO). — Flora Tatrae Magna, Leutschovia [Levoča], in silvae Kienwald; leg.
Greschik V., 07.1900; (SLO). — Flora Tatrae Magnae, Leutschovia [Levoča], in silvus elesioribus; leg. Greschik V.,
08.1928; (SLO). — Leutschovia [Levoča], in [Wäldern ?]; leg. Greschik V., 07.1918; (SLO). — Vysoké Tatry: Téryho
chata; leg. FutákJ., 18.08.1946; (SLO). — Vyšné Hágy, rašelinisko; leg. Futák J., 26.08.1954; (SLO × 2 vouchers). —
Juhoslovenský kras, Silická planina; leg. Futák J., 05.09.1946; (SLO). — Tatr. Kotlina; leg. Futák J., 28.08.1954; (SLO).
— Slovenský raj: na lúčke pod Lipovcom [planina Lipovec] nad potokom; leg. Ferjanec J., 04.09.1962; (SLO × 4
vouchers). — Čierny Váh: na lúke Stará Poľana; leg. Ferjanec J., 26.07.1965; (SLO). — Údolie Čierneho Váhu; leg.
Májovský J., 18.08.1957; (SLO × 7 vouchers). — Údolie Čierneho Váhu; leg. Májovský J., 18.08.1957; (SLO). —
Orava: Polhora kúpele; leg. JurkoA., 16.09.1955; (SLO × 6 vouchers). — Fatra Major: in valle “Žarnovická dolina”
non procul a p. Dolná Štubňa [now Turčianske Teplice], in declivibus; 600 m a.s.l.; leg. Brižický G., Červeňová Ž.,
18.08.1940; (SLO). — Veľká Fatra: Tlstá; leg. Hrivnáková, 20.08.1954; (SLO). — Čierny Váh – pri horárni Biely
Potok; 830 m a.s.l.; leg. Vartíková E., 15.09.1972; (SLO). — Jamnícka Dolina – v kosodrevine; leg. ?, 09. 1981; (NI).
— Lúčky [okres Ružomberok]; leg. Svobodová Z., 04.09.1973; (NI × 2 vouchers). — Muráňská vysočina: Veľká
Stožka; leg. Sillinger P., 11.08.1937; (NI). — Bielske Tatry: Drabina [near Kobylí vrch]; leg. Domin K., 22.08.1937;
(NI). — Bielske Tatry: Ivanka; leg. Domin K., 29.08.1937; (NI). — Kľak; leg. Takáč, 06.09.1962; (NI). — in monte
Kľak; leg. Brancsik K., 07.1900; (NI). — Pieniny (Vysoké Skalky), Stráňany; 1052 m a.s.l.; leg. Magic D., 19.08.1963;
(ZV). — v doline potoka nad obcou Olcnava; leg. Šmídt I., 20.08.1990; (KO × 2 vouchers). — úd. Kvačanské rokle;
leg. Mikoláš V., 19.09.1984; (KO). — sev. a sv. svahy pod Kľakom; leg. Mikoláš V., 15.08.1984; (KO). — Homôľka
(Lúč. MF); leg. Mikoláš V., 15.09.1984; (KO). — str.–sp. č. úd. od haj. Vonžovec – červ./žltá; leg. Mikoláš V.,
18.09.1984; (KO). — Vavrišovo; leg. ?, 19.08.1960; (KO). — Tokáreň; leg. Mikoláš V., 14.08.1958; (KO). —
Muránska planina – Hrad; leg. Mikoláš V., 24–25.08.1959; (KO × 6 vouchers). — úd. Dudliavky – Kamenné Vráta; leg.
Mikoláš V., 16.08.1984; (KO × 2 vouchers). — Olcnava, pri potoku v doline Raj, cca. 1km nad dedinou; leg. Šmídt I.,
20.08.1990; (KO). — Belanské Tatry, chodník Kopské sedlo – Zadné Meďodoly; leg. Najvarová Š., 30.08.1968;
(TNP). — Belanské Tatry, Dolina Siedmich prameňov, Lavinový žľab; 1450 m a.s.l.; leg. Odložilíková L., 12.08.1955;
(TNP). — Belanské Tatry, Dolina Siedmich prameňov, Lavínový žľab; 1500 m a.s.l.; leg. Odložilíková L., 01.09.1955;
(TNP × 3 vouchers). — Západné Tatry: Tichá Dolina, pod Štrkami; leg. ŠmardaJ., 25.08.1955; (TNP). — Belanské
Tatry, Zadné Meďodoly, nad Bránkou; 1210 m a.s.l.; leg. Paclová L., ŠoltésováA., 31.08.1989; (TNP × 2 vouchers).
— Belanské Tatry, nad chatou Plesnivec v Doline Siedmich prameňov; 1350 m a.s.l.; leg. Šoltés R., Šoltésová A.,
11.09,1974; (TNP). — Západné Tatry: Tichá Dolina, pod Štrkami; leg. Paclová L., 25.08.1957; (TNP). — Belanské
Tatry, Dolina Čiernej vody (Rakúskej); 900 m a.s.l.; leg. Odložilíková L., 14.08.1955; (TNP). — Belanské Tatry,
Červená hlina; 1370 m a.s.l.; leg. Odložilíková L., 21.08.1955; (TNP). — Belanské Tatry, Dolina Čiernej vody
(Rakúskej); 980 m a.s.l.; leg. Odložilíková L., 09.09.1955; (TNP). — Bad Lucsivna [Lučivná]; leg. Scherfel A.,
08.1891; (TNP). — Poprad; leg. Scherfel A.; (TNP).
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Aconitum lasiocarpum subsp. lasiocarpum
SLOVAKIA: Užská hornatina: Rabia skala , pieskovec, skalky, južný svah; 1190 m a.s.l.; leg. Michalko J., 08.1956; (SAV × 4
vouchers). — Vihorlat, vrch Vihorlat, Calamagrostidetum arundinaceae; 1050 m a.s.l.; leg. Michalko J., 26.07.1967; (SAV).
— pod vrcholom Stinskej; leg. Marhold K., 18.08.1988; (SAV). — vrch Vihorlat, tesne pod vrcholom; leg. Michalko J.,
27.08.1963; (SAV). — Slovenské rudohorie: Slovenský raj. Stratená, Dobšiná. Andrejisko. Orientatio S(N), acclivitas
35°, substratum calcit, terra rendzina, vegetatio Fagetum; 1100 m a.s.l.; leg. Hajdúk J., 09.08.1956; (B). — Slovenský
raj, Stratenská hornatina. Stratená, Strat. pila. Hnilec, Vodná nádrž. Orientatio S2, NW, acclivitas 50°, substratum CaCO3,
terra litozem, rendzina, vegetatio Piceetum calcicolum; 840 m a.s.l.; leg. Hajdúk J., 31.07.1956; (B). — Slovenský raj,
Stratenské vrchy. Dobšiná. Andrejisko, hrebeň Andrejiská. Orientatio SN, acclivitas 15°, substratum kalcit, terra rendzina,
vegetatio Abieto–Piceetum; 1200 m a.s.l.; leg. Hajdúk J., 09.08.1956; (B). — Nízke Beskydy: Osadné, nad Δ Čereniny
v bučine, pri hranici; leg. Májovský J., 29.09.1975; (SLO). — Vysoké Tatry: Kriváň; leg. Májovský J., 21.08.1957; (SLO).
— Vihorlat [vrch], lúky (typu Calamagrostion) na najvyššom vrchole, J; 1060 m a.s.l.; leg. Májovský J., Michalko J.,
10.09.1953; (SLO). — Slovakia; leg. ?, ?; (SLO × 3 vouchers). — Nízke Beskydy: Osadné, pri chodníku štatnej hranice
na Δ Čereniny, nízky zárast Salix caprea; 950 m a.s.l.; leg. Májovský J., 29.09.1975; (SLO). — Nízke Tatry, Slovenský raj:
Javorina nad Vernárem; leg. Sillinger P., 06.08.1931; (NI).
Aconitum lasiocarpum subsp. kotulae
SLOVAKIA: Pusté Pole; leg. Futák J., 21.08.1973; (SAV). — Užská hornatina: Rabia skala, pieskovec, skalky, južný svah;
1190 m a.s.l.; leg. Michalko J., 08.1956; (SAV × 2 vouchers). — Vihorlat, pod vrcholom vrchu Vihorlat; leg. Michalko J.,
27.08.1963; (SAV × 2 vouchers). — vrch Vihorlat, tesne pod vrcholom; leg. Michalko J., 27.08.1963; (SAV × 4 vouchers).
— Šíp [vrch], obec Stankovany; 1100 m a.s.l.; leg. ?, 23.08.1975; (B). — Čierny Váh: na lúke Stará Poľana; leg.
FerjanecJ., 26.07.1965; (SLO).
Aconitum × pawlowskii
SLOVAKIA: Čierny Váh: na lúke Stará Poľana; leg. Ferjanec J., 26.07.1965; (SLO). — Belianske Tatry: Ždiarska Vidla –
Havran; leg. Májovský J., 23.08.1957; (SLO).
Aconitum degenii subsp. degenii
SLOVAKIA: Slovakia orientalis, Pri chyžke u potoku Stužica; leg. Šmidt[?], 13.08.1964; (SAV × 2 vouchers).
Aconitum × gayeri
SLOVAKIA: medzi Čolom a Rabiou skalou, vo Fagete pod hranicou; leg. Šmidt[?], 15.08.1964; (SAV).
Aconitum × cammarum
SLOVAKIA: Bardejov; leg. Berganský J., 06.08.1928; (B × 7 vouchers). — Bardejov: Wachterhof; leg. Berganský J.,
25.07.1924; (B). — Bardejov: Roch–Vorgelände; leg. Berganský J., 09.10.1929; (B). — Pri železničnej stanici Čirč;
leg. ?, 30.07.1957; (B). — Sasinkovo, v zahrade; leg. ?, 30.06.1963; (B × 2 vouchers). — Vrátna [dolina]; leg.
Brančík K., 08.1904; (SLO). — Felka [Veľká, Tatry]; leg. Scherfel A., 10.08.1924; (TNP × 2 vouchers). — Felka eig.
Garten [Veľká, Tatry]; leg. Scherfel A., 14.08.1923; (TNP × 3 vouchers).
Aconitum firmum subsp. firmum
SLOVAKIA: Belianske Tatry; 1250 m a.s.l.; leg. Futák J., 14.07.1861; (SAV). — Slavkovský potok [Vysoké Tatry]; leg.
Grebenščikov O., 07.08.1954; (SAV). — Vysoké Tatry, Zelené pleso hezin; 1550 m a.s.l.; leg. Hubová O., 23.06.1969;
(SAV × 2 vouchers). — Vysoké Tatry, Velická dolina, Velické pleso; 1700 m a.s.l.; leg. Fabianková , 09.09.1974; (SAV).
— TANAP, Biely Váh dolina; 1350 m a.s.l.; leg. Futák J., 27.08.1957; (SAV). — Lúč. Malá Fatra, vrch Reváň; 1100m
a.s.l.; leg. Schidlay E., 20.08.1949; (SAV). — Belianske Tatry, Javorina, Zadné Koperšady (Zadné Meďodoly) v smrečine;
1400 m a.s.l.; leg. Schidlay E., 09.08.1946; (SAV). — Vihorlat: Popričný vrch, v sédolé; leg. Šramko E., 08.1953; (SAV).
— Pilsko, Babia hora; leg. Futák J., Magic D., 11.08.1964; (SAV). — Djumbier [Ďumbier], svahy; leg. Ptý [Ptačovský],
09.1928; (SAV). — Vihorlat: V ridoli potoka; leg. Šramko E., 08.1953; (SAV). — Vys. Tatry, Velická dolina; leg. FutákJ.,
08.08.1962; (SAV). — B.T., Široká dolina, dolná časť; 1500 m a.s.l.; leg. Flašíková, Hubová O., 02.08.1971; (SAV × 2
vouchers). — Bel. Tatry: Faixová; leg. Ptý, 22.07.1939; (SAV). — Bel.Tatry: pri Bielom plese (smerom na Belanskú
kopu); 1600 m a.s.l.; leg. Zahradníková K., Šípošová H., 19.08.1980; (SAV × 3 vouchers). — Vysoké Tatry: N svahy
86 Novikov A.
Modern Phytomorphology 11, 2017
Kežmarského štítu nad Zeleným plesom; 1600 m a.s.l.; leg. Zahradníková K., 07.08.1969; (SAV). — Vysoké Tatry: cestou
na Kriváň; leg. Zahradníková K., 15.08.1970; (SAV). — Vysoké Tatry: južný svah Jahňacieho štítu; 2000 m a.s.l.; leg.
ZahradníkováK., 09.08.1969; (SAV). — Vysoké Tatry: Kriváň; leg. Zahradníková K., Jasičová M., 11.08.1966; (SAV). —
Votrubova chata; leg. Ptý, 07.1941; (SAV). — V kleči u Popradského plesa; leg. Ptý, 07.1939; (SAV). — Záp. Beskydy:
Babia hora; leg. Zahradníková K., 21.07.1973; (SAV). — Vysoké Tatry: Furkotská dolina – Wahlenbergovo pleso – Bystré
sedlo; leg. Zahradniková K., 13.08.1970; (SAV). — Tatry. Faixová a Votrubova chata; leg. Horák P., 25–26.08.1937;
(SAV). — [Chočská] Fatra, Biela Skala; 1600 m a.s.l.; leg. Futák J., 24.08.1966; (SAV × 3 vouchers). — Slovakia borealis:
montes Vysoké Tatry, ad ripas lacus “Popradské pleso” in Pineto mughi, solo granitico; 1640 m a.s.l.; leg. Hlavaček A., ?;
(SAV). — Západné Tatry: Osobitá, Suchá dolina, pri potoku; 1000 m a.s.l.; leg. Futák J., 15.07.1965; (SAV × 2 vouchers).
— Vys. Tatry; leg. Szép R., 20.08.1904; (B). — Vys. Tatry; 2000 m a.s.l.; leg. ?, 07.1936; (B). — Vys. Tatry: ad
margines sylvae, Štrbské Pleso et Vyš. Hágy; 1200 m a.s.l.; leg. Valenta V. 01.08.1935; (B). — Nízké Tatry: pramenitý
svah pod vrcholem Krupovy hole; 1550 m a.s.l.; leg. Dvořák J., 07.08.1973; (B). — Vysoké Tatry; leg. Berganský J.,
28.08.1928; (B × 2 vouchers). — Vysoké Tatry; leg. Vaslora L. [?], 07.08.1952; (B). — Vysoké Tatry: Javorová
dolina; 1900 m a.s.l.; leg. Drévlackanský Fr., 12.10.1962; (B). — Štrbské pleso; leg. Černý J., 08.07.1924; (B). —
Slovenské rudohorie: Volovské vrchy, Slovenský raj. Dedinky, Mlynky, collis Kukla, umen Hnilec. Orientatio N, acclivitas
35°, substratum silik.+carb., vegetatio Mugedio–Aconitetea; 980 m a.s.l.; leg. Hajdúk J., 18.07.1956; (B). — Vysoké
Tatry: montis Satan; leg. Weber F., 08.1936; (B). — Vysoké Tatry: montis Končistá; 2450 m a.s.l.; leg. WeberF.,
08.1925; (B). — Bielské Tatry: montis Tokárna; 1200 m a.s.l.; leg. Weber F., 08.1936; (B). — Vysoké Tatry: ad lac.
Ľadové pleso; leg. Weber F., 08.1925; (B). — Bielské Tatry: montis Havran; leg. Weber F., 08.1936; (B). — Vysoké
Tatry: montis Vel. Vysoká; 2400 m a.s.l.; leg. Weber F., 08.1925; (B). — Tatra; leg. Scherfel A., ?; (B). — Belanské
Tatry: Tatr. Kotlina – “Zadné Meďodoly”; 1400 m a.s.l.; leg. Z., 07.08.1969; (B). — Záp. Tatr y: Bystrá dolina, ad ripam
rivi “Bystrý ”; 1300 m a.s.l.; leg. Dvořák J., 28.07.1976; (B). — Orava – Osobitá; leg. Magdolenová S., 08.1967; (B).
— Bielske Tatry; leg. ?, 07.08.1970; (B). — Žiarska dolina; leg. ?, 31.07.1942; (B). — Vysoké Tatry: Česká dolina,
kamenité svahy nad Českým plesom [now Ťažké pleso]; 1620 m a.s.l.; leg. Dvořák J., 07.09.1974; (B). — Záp. Tatry:
Bystrá dolina, ad ripam rivi “Bystrý”; 1300 m a.s.l.; leg. Dvořák J., 28.07.1976; (B). — Vysoké Tatry; 950 m a.s.l.; leg.
Kavka V., 15.07.1930; (B). — Vysoké Tatry: Kriváň; leg. Májovský J., 21.08.1957; (SLO). — Vysoké Tatry: cestou z
Popradského na Žabie plesá; leg. Gálisová E., 23.08.1956; (SLO). — Nízke Tatry: Ohnište; leg. Májovský J., 20.07.1957;
(SLO). — V. Tatry: Velická dolina; leg. Schidlay E., 10.10.1952; (SLO). — V. Tatry: Hincovo pleso; leg. HáberováI.,
16.07.1968; (SLO). — Záp. Beskydy: Babia hora; leg. Murín A., 27.07.1983; (SLO × 2 vouchers). — Rašelinisko
Mutné (okres Námestovo); leg. Májovský J., 30.07.1955; (SLO). — Kamenné vráta; leg. Futák J., 19.08.1946; (SLO ×
3 vouchers). — Belianske Tatry: hrebeň; leg. Futák J., 09.07.1946; (SLO). — Tatranská Lomnica: Kežmarskej Žľaby,
okolie; leg. Futák J., 08.07.1946; (SLO). — Popradské pleso; leg. Futák J., 23.08.1943; (SLO). — Zlomisková dolina,
Ľadové pleso, V. Tatry; leg. Futák J., 26.08.1943; (SLO). — Belianske Tatry: Javorina; leg. Futák J., 14.07.1946; (SLO). —
Belianske Tatry: Tristárska dolina; leg. Futák J., 14.07.1946; (SLO × 3 vouchers). — Vysoké Tatry: Litvorová kotlina; leg.
Vartíková E., 28.09.1971; (SLO × 2 vouchers). — Vysoké Tatry: circum l. d. “Zelené pleso”; leg. Nábĕlek V., 12.08.1940;
(SLO × 2 vouchers). — Belské Tatry: sev. svah Hlúpeho; leg. Futák J., 21.08.1943; (SLO). — Jánska dolina, JZ svahy
Ohnišťa, nad lesom na trávnatých svahoch a v suti; leg. Májovský J., 15.07.1967; (SLO × 2 vouchers). — Javorová dolina
pri Javorine vo V. Tatrách; leg. Futák J., 17.08.1943; (SLO). — Dolina Mlynica vo V. Tatrách; leg. Futák J., 25.08.1943;
(SLO × 4 vouchers). — Belianske Tatry: Zadné Koperšady; leg. Nábĕlek V., 12.08.1940; (SLO). — Kamenné vráta;
leg. Futák J., 19.08.1946; (SLO). — Žiarska dolina: hojne na prameniskách a popri potokoch v pásme kosodreviny; leg.
?, 16.08.1992; (NI). — montes Vysoké Tatry: in valle Zlomisko, solo granitico; leg. Osvačilová V., 14.08.1951; (NI).
— Belianske Tatry: Tristárska dolina; leg. Eliáš P. jun., 1998; (NI × 2 vouchers). — Račkova dolina; leg. Mikoláš V.,
08.1956; (KO). — Sliezsky Dom vo V.T. [horský hotel na prahu Velickej doliny vo Vysokých Tatrách]; leg. Vojtúň A.,
03.08.1969; (KO × 2 vouchers). — ud. Bocianky, str.–sp. časť; leg. Mikoláš V., 15.08.1984; (KO). — Račkova dolina;
leg. Mikoláš V., 08.1956; (KO). — Priehyba pod Kriváňom [vo Vysokých Tatrách]; leg. Vojtúň A., 16.07.1968; (KO ×
2 vouchers). — Západné Tatry: v kosodrevine pod Tomanovským sedlom; 1650 m a.s.l.; leg. Šoltésová A., 18.09,1974;
(TNP). — Bujačí Vrch – južný svah, v kosodrevine; 1800 m a.s.l.; leg. Šoltés R., Šoltésová A., 11.09,1974; (TNP). —
Belanské Tatry. Bujačí Vrch, medzi Malým a Veľkým Košiarom; 1850 m a.s.l.; leg. Šoltésová A., 17.09.1974; (TNP). —
Belanské Tatry. Bujačí Vrch, pod Veľkým Košiarom; 1850 m a.s.l.; leg. Šoltésová A., 17.09.1974; (TNP). — V. Tatry.
V.Studená dol.; leg. Paclová L., 29.08.1954; (TNP). — V. Tatry. Velická dol.; leg. Najvarová Š., 23.07.1968; (TNP). —
Vysoké Tatry: pri značkovanem chodníku zo Šalvejového prameňa na Zelené pleso; leg. Najvarová Š., 27.07.1967; (TNP).
— Velická dolina, Kvetnica; 1820 m a.s.l.; leg. Odložilíková L., 06.07.1954; (TNP). — Vysoké Tatry. Veľká Studená dol.;
1610 m a.s.l.; leg. Odložilíková L., 15.08.1954; (TNP). — Belanské Tatry, pod Siedmimi prameňmi; 1230 m a.s.l.; leg.
Odložilíková L., 08.08.1956; (TNP). — Mengsdorfer [Mengusovce]; leg. Scherfel A., 08.1923; (TNP × 2 vouchers). —
Felka [Poprad-Veľká] eig. Garten [Veľká, Tatry]; leg. Scherfel A., 08.1923; (TNP). — Felka eig. Garten [Veľká, Tatry];
leg. Scherfel A., 08.1923; (TNP). — Mengsdorfer [Mengusovce]; leg. Scherfel A., 13.07.1924; (TNP). — Salbaual; leg.
Scherfel A.; (TNP). — Gwostar [?]; leg. Scherfel A., 07.1880; (TNP).
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Aconitum firmum subsp. moravicum
SLOVAKIA: Malá Fatra, vrch Chleb; leg. Nábĕlek V., 29.08.1942; (SAV). — Malá Fatra, Šútovská dolina; leg.
ZahradníkováK., 22.07.1973; (SAV). — Vysoké Tatry, popri turistickom chodinku z Kriváňa na Tri studničky (po
zelenej značke); 2300 m a.s.l.; leg. Peniašteková M., 20.08.1987; (SAV). — Manínska úžina (v jej prvej – dolnej polovici);
leg. Schidlay E., 08.09.1945; (SAV). — Krivánska Malá Fatra: hrebeňovka V. Kriváň – Biele skaly – Suchý; 1500 m
a.s.l.; leg. Futák J., Jasičová M., Zahradníková K., 11.09.1964; (SAV × 2 vouchers). — Kriváňska Malá Fatra: Snilovská
dolina – pri potoku; leg. Letz R., 1994; (SAV). — Malá Fatra: Malý Kriváň, v porastoch kosodreviny; 1600 m a.s.l.;
leg. GrebenščikovO., 22.09.1954; (SAV). — Vysoké Tatry: Bielovodská dolina; 1700 m a.s.l.; leg. Drévlackanský Fr.,
07.10.1962; (B). — ad valle Studenec ad pedem m. Malý Kriváň; leg. Holuby J., 29.07.1896; (B). — [Krivánska]
Malá Fatra. Veľký Rozsutec; leg. Drévlackanský Fr., 11.07.1961; (B). — ad valle Vrátna; leg. Brancsik C., 09.1898;
(B). — Malá Fatra: in saxis montis R ásutec [Rozsutec] supra pag. Štefanová; leg. Weber F., 07.1935; (B). — Fatra
Minor: in declivibus saxosis m. Kriváň; 1500 m a.s.l.; leg. Brižický J., 27.08.1940; (SLO). — Malá Fatra: Kriváň; leg.
Činčura F., 19.07.1971; (SLO × 4 vouchers). — Rašelinisko Mutné (okres Námestovo); leg. Májovský J., 30.07.1955;
(SLO × 2 vouchers). — Kamenné vráta; leg. Futák J., 19.08.1946; (SLO). — Javorová dolina pri Javorine vo V. Tatrách;
leg. Futák J., 17.08.1943; (SLO). — Rozsutec – horná hranica lesa; leg. ?, 09. 1981; (NI). — Horné diery; leg. MikolášV.,
18.08.1984; (KO). — Horné diery (Kriv. MF); leg. Mikoláš V., 12.09.1984; (KO). — Vysoké Tatry: pri značkovanom
chodníku zo Šalvejového prameňa na Zelené pleso; leg. Najvarová Š., 27.07.1967; (TNP).
Aconitum firmum subsp. maninense
SLOVAKIA: Kamenné vráta; leg. Futák J., 19.08.1946; (SLO). — Manín [vrch]; leg. Futák J., 25.08.1971; (SAV). —
Súľov; leg. Futák J., 25.08.1971; (SAV). — Žilina. In saxosis calcareis in convalle Sedlovina et Kopaná ad Kunerad; 700m
a.s.l.; leg. Valenta V., 28.07.1937; (B).
Aconitum firmum nothosubsp. paxii
SLOVAKIA: Fatra Minor: in declivibus saxosis m. Kriváň; 1500 m a.s.l.; leg. Brižický J., 27.08.1940; (SLO). —Belianske
Tatry: Tristárska dolina; leg. Futák J., 14.07.1946; (SLO).
Aconitum firmum nothosubsp. zapalowiczii
TANAP, dolina Bieleho Váhu, pri potoku; 1250 m a.s.l.; leg. Futák J., 27.08.1857; (SAV). — Vys. Tatry, Velická dolina,
pri jazierku; 1825 m a.s.l.; leg. Futák J., Hubová O., 06.08.1962; (SAV). — V. Tatry: Hincovo pleso; leg. HáberováI.,
16.07.1968; (SLO).
Appendix 2. Specimens cited in published sources
Aconitum moldavicum subsp. moldavicum
SLOVAKIA: Spišská Nová Ves (ad stationem viae ferr. “Štiavník”), 600 m; leg. J. Suza, 20.06.1931; (K 003671);
source: Mitka (2008a). — Tomašovská Belá, nad potokiem w lesie bukowym; leg. K. Oklejewicz, 07.08.2004; (K);
source: Mitka (2008a). — Stratená, Fagetum; leg. J. Mitka, 18.07.2006; (K); source: Mitka (2008a). — Liptovské
Revúce, near torrent; leg. J. Mitka, 23.07.2006; (K); source: Mitka (2008a). — Liptovský Ján, Jánska dolina, ok. 800
m; leg. J. Mitka, 20.08.2007 (K); source: Mitka (2008a).
Aconitum variegatum subsp. variegatum
SLOVAKIA: Północne obrzeże wapienne, Kopieniec Wielki, górna cząść łąk Hali Koprowej; leg. H. & T. Tacik, 18.08.1966;
(KM); source: Mitka (2003). — Comit. Szepes, in apertis piceetorum “Gähe Leit” supra Tátrabarlagligent; leg.
A. Boros, 24.8.1938; (BP); source: Mitka (2003). — Kiczora od strony Dol. Międzyściennej, przerzedzony, skalisty
las, wapień, 1100-1200 m; leg. B. Pawłowski, 22.07.1939; (K); source: Mitka (2003). — Eisernes or, 1550 m; leg.
F.Pax, 21.08.1905; (BP); source: Mitka (2003). — wycieczka w Spiskie Tatry w 1878, Dolina między Hawranem a Nową
ku szczytowi Czamego Wierchu; leg. A. Rogalski, 28.07.1878; (KM); source: Mitka (2003).
88 Novikov A.
Modern Phytomorphology 11, 2017
Aconitum lasiocarpum subsp. lasiocarpum
SLOVAKIA: Bieszczady Zach.: Rabia skala, wśród zarośli z Rubus idaeus, 1150 m; leg. P. Bochenek, J. Mitka, 23.08.1995;
(K); source: Mitka (2003). — Stinska, na skraju Fagetum wsrod Rubus idaeus; P. Bochenek, leg. J. Mitka, 24.08.1995;
(K); source: Mitka (2003). — Stinka, 1093 m; leg. Z. Dolezalova, ?; (PRC); source: Mitka (2003). — Wyhorlat;
leg.C.Chyzer, 21.08.1879; (BP); source: Mitka (2003). — Comit. Gömör, in apertis silv. pedis montis Hanneshöhe
prope Ľadová jaskyňa, 900 m; leg. A. Boros, 03.09.1955; (BP); source: Mitka (2003).
Aconitum lasiocarpum subsp. kotulae
SLOVAKIA: Nizke Tatry, Tále; leg. J. Mitka, 19.08.2007; (K 0464740); source: Mitka, personal communication. —
Wyhorlat, ex herbario doctoris Cornelii Hyzer; leg. ?, 21.08.1879; (BP); source: Mitka & Starmühler (2000) and Mitka
(2003). — In monte Vihorlat, 1074 m; leg. F. Belšán, 25.07.1936; (PRC); source: Mitka & Starmühler (2000) and Mitka
(2003). — Rabia skala, 1150; leg. J. Mitka, P. Bochenek, J. Terray, 23.08.1995; (K); source: Mitka & Starmühler
(2000) and Mitka (2003). — Rabia skala, cota 1168; leg. J. Soják, 23.08.1963; (KM); source: Mitka & Starmühler
(2000) and Mitka (2003). — Slov. orient, montes Poloninské Karpaty, in monte Rabia skala supra pagum Nová Sedlica,
1160 m; leg. J. Soják, 20.08.1963; (PR); source: Mitka & Starmühler (2000) and Mitka (2003). — Slov. orient, montes
Poloniny, ad cacumen montis Čereniny supra pagum Zvala; 933 m; leg. J. Soják, 21.08.1963; (PR); source: Mitka &
Starmühler (2000) and Mitka (2003). — Na skale Rabia skala, trig. 1168 m; leg. J. Šourek, 18.07.1957; (PR); source:
Mitka & Starmühler (2000) and Mitka (2003). — Výslunna škála nad Runinou; 1150 m; leg. J. Šourek, 7.08.1957; (PR);
source: Mitka & Starmühler (2000) and Mitka (2003). — montes Poloniny, ad cacumen jugi inter pagos Zvala et Ruské,
c.1000 m; leg. J. Soják, 21.08.1963; (PR); source: Mitka & Starmühler (2000) and Mitka (2003). — Nízke Poloniny,
Runina, výslunné skály na Rabia skale, 1150 m; leg. J. Soják, 6.08.1957; (PR); source: Mitka & Starmühler (2000) and
Mitka (2003). — Dol. Koprowa [Kôprová dolina] – skraj lasu, ok. 1300 m, leg. B. Pawłowski, 19.8.1925; (KM-
Pawlowski 320228, Paratypus); source: Mitka & Starmühler (2000) and Mitka (2003).
Aconitum × pawlowskii
SLOVAKIA: Dol. Koprowa (górna), zarośla mieszane; leg J. Trela, 19.08.1825; (K); source: Mitka & Starmühler
(2000) and Mitka (2003). — Tatras, Kôprová dolina; leg. K. Zahradníková, M. Jasičová, 13.08.1966; (SAV); source:
Mitka & Starmühler (2000) and Mitka (2003). — Belianske Tatry, Lysá Poľana; leg. K. Zahradníková, M. Jasičová,
14.08.1966; (SAV); source: Mitka & Starmühler (2000) and Mitka (2003). — apud rivulorum Javorový potok supra
pag. Javorina (Tatranská Javorina), 1050 m; leg. J. Dostál, 15.08.1955; (PR); source: Mitka & Starmühler (2000) and
Mitka (2003). — Nízke Tatry, Dudlavá skala; leg. V. Kneblová, 08.09.1955; (PR); source: Mitka & Starmühler (2000)
and Mitka (2003). — Dobsinai jegbevlong szkecenei völgy [jégbarlang, Sztraczenai völgy?] leg. D. Filarszky, 23.08.1911;
(BP); source: Mitka & Starmühler (2000) and Mitka (2003). — Křižovatka; leg. A.G., 27.05.1970; (KO); source: Mitka
& Starmühler (2000) and Mitka (2003). — Hrad Mt.; leg. ?, 24.08.1959; (KO 13340); source: Mitka & Starmühler
(2000) and Mitka (2003). — Ûdolie Dudlavky (ad Kamenná vrata); leg. V. Mikoláš, 16.08.1984; (KO); source: Mitka &
Starmühler (2000) and Mitka (2003).
Aconitum × gayeri
SLOVAKIA: Rabia skala, 1150 m; leg. J. Mitka, P. Bochenek, J. Terray, 23.8.1995; (K); source: Mitka (2003).
Aconitum × cammarum
SLOVAKIA: Veľká Fatra, Donovaly; leg. J. Mitka, 27.07.2007; (K 0347248); source: Mitka, personal communication.
Aconitum × berdaui
SLOVAKIA: Tatras, Javorová dolina; leg. A. Sutkowska, 18.08.2009; (herb. J. Mitka); source: Mitka, personal
communication. — Malá Fatra, Terchová; leg. J. Mitka, 31.07.2009; (K 0464764); source: Mitka, personal
communication. — Wycieczka w Tatry Spiskie w r. 1878. Aconitum napellus L., Podspady nad Jaworzynką; leg. A . Rogalski,
25.07.1878; (KM 13348); source: Mitka (2003). — Plantae exsiccatae regni Hungariae, Comit. Szepes, in m. Hollókö
pr. Sztraczena; leg. G. Lengyel, 01.08.1901; (BP); source: Mitka (2003).
Aconitum firmum subsp. firmum
SLOVAKIA: pod szczytem Łomnicy; leg. H. Zapałowicz, 12.08.1908; (KM); source: Mitka (2003). — in subalpinis
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montium Magas-tatra supra Kešmark: Tokarna Zöltó; leg. L. Simonkai, 7-8.07.1890; (BP); source: Mitka (2003). —
Hinszko [lake Hinco]; leg. D. Filarszky, Kümmerle, 07.08.1909; (BP); source: Mitka (2003). — [Vysoké and Západné
Tatry Mts.] bélei Tiefergrund; leg. Magócsy, ?; (BP); source: Mitka (2003). — [Vysoké and Západné Tatry Mts.] ad lacum
Buckholz [Buchholz ?], 1950 m; leg. G. Lengyel, 04.08.1931; (BP); source: Mitka (2003). — im Kleinkohlbachtale,
Hohe Tatra; leg. A. Margiai, 28.7.1930; (M); source: Mitka (2003). — Breznóbánya, Zadny Uplaz; leg. S.Kupcok,
02.08.1904; (PR); source: Mitka (2003). — ad Javorinam, in alt. c. 900 m; leg. G. Lengyel, 17.07.1931; (BP); source:
Mitka (2003). — Alpenkessel zwischen Nový und Havran; leg. K. Ronniger, 09.07.1918; (W); source: Mitka (2003).
— Fatra: Hungaria, comitatus ad conn. Árva et Trencsén, montes Kriván Fatrae, in declivibus montis Roszudec, alt.
c.1300m, solo dolomitico; leg. E. Nyárády, 27.07.1907; (SIB); source: Mitka (2003). — Fatra, Terchová; leg. K. Domin,
01.08.1918; (PRC); source: Mitka (2003). — in rupestribus “Magas Fátra” loco “Bartya” [A. rmum Rchb., rev. G. Gáyer];
leg. L. Simonkai, ?.08.1904; (BP); source: Mitka (2003). — Comit. Árva, in pineti infra locus “Rohačsi savak“ – alpinum
liptoviensium, c. 1500 m [A. tatrae Borb., rev. G. Gayer 1913]; leg. A. Jávorka, 22.08.1911; (BP); source: Mitka (2003).
— [Roháče Mts.], Liptauer Alpen, Rohacs [Roháč], 1600 m [A. napellus L. var. tauricum (Rchb.) Ser., rev. De Rapaics];
leg. F. Pax, 14.08.1905; (BP); source: Mitka (2003). — Comit. Liptó, Tatra Inf., in valle Lucsany ad Deményfalven; leg.
G.Lengyel, 25.07.1928; (BP); source: Mitka (2003).
Aconitum firmum subsp. moravicum
SLOVAKIA: Montes Liptovské Tatry, in alpinis Osobitá, alt. c. 1580 m, solo calcareo; leg. M. Deyl, 07.07.1938; (PR);
source: Mitka (2003). — Roháče, u Zverovky; leg. Kopecký, 06.08.1968; (PR); source: Mitka (2003). — Hungaria,
comitatus Liptó, montes Tatrae Liptoviensae, in saxis “Javor” in parte superiore vallis Tycha, alt. cca. 1500 m, solo calc.;
leg. E. Nyárády, 01.08.1907; (SIB); source: Mitka (2003). — Siná, 1500–1550 m; leg. P. Sillinger, 27.07.1930; (PRC);
source: Mitka (2003). — in lapidosis in declivi merid. montis Chleb, cca. 1600 m, solo calcareo; leg. J. Soják, 04.08.1950
(PR); source: Mitka (2003). — Fatra – Kriváň [A. napellus L. var. multidum Koch. var. canescens (Schleich.) Rapaics, det.
Rapaics]; leg. ?, 13.08.1889; (BP); source: Mitka (2003). — [Malá] Fatra, Terchová; leg. K. Domin, 01.08.1918; (PRC);
source: Mitka (2003). — Slovakia, Malá Fatra, Révalov, Révalovská dolina, along torrent 1150–1200 m; leg. J. Mitka,
02.09.2001; (K); source: Mitka (2003).
Aconitum firmum subsp. maninense
SLOVAKIA: Galicia, ad pedem montis Tatrae Magnae, in valle Poduplaski, alt. c. 1000 m, solo granitico; leg. E. Nyárády,
27.07.1908; (SIB); source: Mitka (2003). — [Súľovské vrchy] Manínská soutěska, Súl’ov; leg. K. Domin, 01.08.1920;
(PRC); source: Mitka (2003). — Manín (SW od Súľovskych vrchov), ok. 400 m; leg. Skalický, Skalická, 27.07.1973;
(PRC); source: Mitka (2003). — Manín, Strážovské vrchy Mts., Manínska úžina, in oak-hornbeam forest Tilio‑
Carpinetum along a torrent; leg. J. Mitka, 03.09.2001; (K); source: Mitka (2003). — Slovakia, Manín, Strážovské vrchy
Mts., Manínska užina, on steep stone slope; leg. J. Mitka, 03.09.2001; (K); source: Mitka (2003).
Aconitum firmum nothosubsp. paxii
SLOVAKIA: Montes Liptovské hole, mons Červené vrchy, in locis lapidosis calcareis rupium Javorové skalky, alt. 1450m
[A. rmum (Rchb.) Neilr. mutatio – cum pilis adscendentibus copissimus, rev. V. Skalický]; leg. J. Dostál, 21.08.1930;
(PRC); source: Mitka (2003). — Liptovské hole, Tichá dolina; leg. P. Svoboda, 07.08.1935; (PR); source: Mitka (2003).
Aconitum firmum subsp. firmum × A. firmum subsp. maninense
SLOVAKIA: Belianske Tatry Mts.: Comitatus Szepes, Contes Bélaenses, in valle Drechslerhäuschen sub monte Stirnberg,
alt. c. 1400–1500 m; leg. E.G. Nyárády, 01.09.1907; (SIB); source: Mitka (2003).
