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Abstract

A species‐level taxonomic revision of Atocion and Viscaria, based on a recent phylogenetic study, is presented. Atocion includes six species (A. armeria, A. compactum, A. lerchenfeldianum, A. reuterianum, A. rupestre and A. scythicinum), and Viscaria includes three species (V. alpina, V. asterias and V. vulgaris). The highest species diversity is found on the Balkan Peninsula. Species descriptions based on a study of morphological characters and generated using the Prometheus description model are provided and are also available in the Sileneae online database. Complete synonymy, notes on the ecology and geographical distribution of the taxa and an identification key are provided. Lectotypes for nine names (Lychnis helvetica, L. suecica, Silene armeria var. angustifolia, S. berdaui, S. lituanica, S. compacta, S. orientalis, S. reuteriana and Viscaria media) and a neotype for one name (S. lerchenfeldiana) are assigned. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210.
Taxonomic revision of Atocion and Viscaria (Sileneae,
Caryophyllaceae)
BOŽO FRAJMAN1*, MIKAEL THOLLESSON2and BENGT OXELMAN3
1Institute of Botany, University of Innsbruck, Sternwartestraße 15, A-6020 Innsbruck, Austria
2Department of Systematic Botany, Evolutionary Biology Centre, Uppsala University, Norbyvägen
18D, SE-75236 Uppsala, Sweden
3Department of Biology and Environmental Sciences, Göteborg University, Box 461, SE-405 30
Göteborg, Sweden
Received 16 November 2012; revised 5 July 2013; accepted for publication 20 July 2013
A species-level taxonomic revision of Atocion and Viscaria, based on a recent phylogenetic study, is presented.
Atocion includes six species (A. armeria,A. compactum,A. lerchenfeldianum,A. reuterianum,A. rupestre and
A. scythicinum), and Viscaria includes three species (V. alpina,V. asterias and V. vulgaris). The highest species
diversity is found on the Balkan Peninsula. Species descriptions based on a study of morphological characters and
generated using the Prometheus description model are provided and are also available in the Sileneae online
database. Complete synonymy, notes on the ecology and geographical distribution of the taxa and an identification
key are provided. Lectotypes for nine names (Lychnis helvetica,L. suecica,Silene armeria var. angustifolia,
S. berdaui,S. lituanica,S. compacta,S. orientalis,S. reuteriana and Viscaria media) and a neotype for one name
(S. lerchenfeldiana) are assigned. © 2013 The Linnean Society of London, Botanical Journal of the Linnean
Society, 2013, 173, 194–210.
ADDITIONAL KEYWORDS: Europe – Lychnis – Middle East – Silene – taxonomy – typification.
INTRODUCTION
Atocion Adans. and Viscaria Bernh. are small genera
of tribe Sileneae (Caryophyllaceae), closely related to
Heliosperma (Rchb.) Rchb. and Eudianthe (Rchb.)
Rchb. (Erixon & Oxelman, 2008; Frajman, Eggens &
Oxelman, 2009a). A recent phylogenetic study
(Frajman, Heidari & Oxelman, 2009b) has shown that
relationships among the taxa as traditionally inferred
on the basis of morphology (e.g. Rohrbach, 1868;
Williams, 1896; Chowdhuri, 1957) and presented in
Table 1 do not reflect the evolutionary history of these
groups. Before the first study of Sileneae based on
DNA sequences (Oxelman & Lidén, 1995), Viscaria
and Atocion were never thought to be closely related.
Viscaria (excluding V. asterias (Griseb.) Frajman) was
often classified independently or merged with Lychnis
L., which in turn has recently been sometimes clas-
sified in Silene L. (e.g. Greuter, 1997). Atocion taxa
have almost invariably been classified in Silene
(exceptions being Rafinesque, 1840; Oxelman et al.,
2001a; Tzvelev, 2001), mostly as members of two,
allegedly closely related sections (Table 1). A common
characteristic of Viscaria and Lychnis is a capsule
dehiscing by a number of teeth equal to the number
of styles (five). However, fully ripe capsules of V. vul-
garis Bernh. sometimes have ten teeth (Oxelman &
Lidén, 1995). In contrast, in Silene the capsules
dehisce by twice as many teeth as the number of
styles (six or ten; e.g. Chowdhuri, 1957). Phylogenetic
studies (e.g. Oxelman & Lidén, 1995; Oxelman et al.,
2001a; Popp & Oxelman, 2004; Erixon & Oxelman,
2008; Frajman et al., 2009a) have shown that taxa
with five carpels are found at different positions in
phylogenetic trees for Sileneae, indicating that
changes in carpel number have happened several
times in the evolutionary history of Sileneae. There
are also morphological characters that separate Vis-
caria from Lychnis, for example, the capsule dehis-
cence of Viscaria is primarily loculicidal, whereas that
*Corresponding author. E-mail: bozo.frajman@uibk.ac.at
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Botanical Journal of the Linnean Society, 2013, 173, 194–210.
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210194
Table 1. Morphology-based classifications of Atocion and Viscaria in important Sileneae-related taxonomic literature
Linnaeus
(1753) Otth (1824)
Boissier
(1867)
Rohrbach
(1868,
1869–70)
Williams
(1896)
Neumayer
(1923)
Ascherson &
Graebner
(1929)
Pax &
Hoffmann
(1934)
Chowdhuri
(1957)
Chater et al.
(1993), Chater
(1993)
Greuter
(1995)
Atocion armeria Silene L. Silene sect.
Atocion Otth
Silene ser.
Compactae
Boiss.
Silene ser.
Compactae
Silene ser.
Compactae
Silene Silene ser.
Compactae
Silene sect.
Dichasiosilene
Rohrb. subsect.
Compactae
(Boiss.) Gürke
Silene sect.
Compactae
(Boiss.)
Schischk.
Silene sect.
Compactae
Silene sect.
Compactae
Atocion rupestre Silene Silene sect.
Rupifraga
Otth
Silene ser.
Brachyanthae
Rohrb.
Silene ser.
Brachyanthae
Silene ser.
Brachyanthae
Silene sect.
Dichasiosilene
subsect.
Brachyanthae
(Rohrb.) Gürke
Silene sect.
Rupifraga
Silene sect.
Rupifraga
Silene sect.
Rupifraga
Atocion
lerchenfeldianum
Silene sect.
Stachymorpha
Otth
Silene ser.
Decumbentes
Boiss.
Silene ser.
Brachyanthae
Silene ser.
Brachyanthae
Silene Silene ser.
Brachyanthae
Silene sect.
Dichasiosilene
subsect.
Brachyanthae
Silene sect.
Rupifraga
Silene sect.
Rupifraga
Silene sect.
Rupifraga
Atocion
compactum
Silene sect.
Atocion
Silene ser.
Compactae
Silene ser.
Compactae
Silene ser.
Compactae
Silene Silene ser.
Compactae
Silene sect.
Dichasiosilene
subsect.
Compactae
Silene sect.
Compactae
Silene sect.
Compactae
Silene sect.
Compactae
Atocion
reuterianum
–– Silene ser.
Compactae
Silene ser.
Compactae
Silene ser.
Compactae
–– Silene sect.
Dichasiosilene
subsect.
Compactae
Silene sect.
Compactae
––
Viscaria alpina Lychnis L. Lychnis sect.
Viscaria
(Bernh.)
Otth
Viscaria
Bernh.
Viscaria Viscaria Lychnis Viscaria Viscaria sect.
Liponeurum
Nyman
Lychnis Lychnis Silene sect.
Viscaria (DC.)
Greuter
Viscaria asterias –– Silene ser.
Armerioideae
Boiss.
Silene ser.
Compactae
Silene ser.
Compactae
Silene Silene ser.
Compactae
Silene sect.
Dichasiosilene
subsect.
Compactae
Silene sect.
Compactae
Silene sect.
Compactae
Silene sect.
Compactae
Viscaria vulgaris Lychnis Lychnis sect.
Viscaria
Viscaria Viscaria Viscaria Lychnis Viscaria Viscaria sect.
Viscaria
Lychnis Lychnis Silene sect.
Viscaria
REVISION OF ATOCION AND VISCARIA 195
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210
of Lychnis is septicidal (for additional differences see
Oxelman & Lidén, 1995: table 4). The phylogenetic
relationships in Atocion (Frajman et al., 2009b) do not
support recognition of two sections, as in Chowdhuri
(1957) and others (Table 1).
Most of the taxa belonging to Viscaria and Atocion
are native to Europe. The range of V. alpina (L.)
G.Don extends to north-eastern North America, and
some Atocion taxa occur in the Middle East. Atocion
armeria (L.) Raf. and V. vulgaris are sometimes cul-
tivated and can be found as occasional and adventive
garden escapes even outside of their native ranges
(e.g. Morton, 2005). There is extensive taxonomic
literature available about the European taxa (e.g.
Ascherson & Graebner, 1929; Stojanoff, 1939; Chater,
1964; Böcher, 1977; Meusel & Mühlberg, 1978–1979;
Chater, Walters & Akeroyd, 1993; Greuter, 1995;
Greuter, 1997; Jonsell & Kurtto, 2001; Kurtto &
Wesenberg, 2001). The plant descriptions in the above
literature often differ and sometimes disagree. They
are usually geographically restricted and incompara-
ble, as they do not take the same characters into
account.
Much less is known about Atocion taxa from the
Middle East: A. scythicinum (Coode & Cullen)
Frajman from eastern Turkey is to our knowledge
only known from the type specimen (Coode & Cullen,
1967). There are few data available about A. reuteri-
anum (Boiss. & Blanche) Frajman, a taxon endemic to
Lebanon (Mouterde, 1966), and A. compactum (Fisch.
ex Hornem.) Tzvelev, distributed from the south-
eastern Balkan Peninsula to Iran (Coode & Cullen,
1967; Rechinger et al., 1988; Greuter, 1997).
The aim of the present study is a taxonomic revi-
sion of the two genera. Inferred phylogenetic trees
(Frajman et al., 2009b) identify hypothetical mono-
phyletic groups and help us in taxon delimitations.
The main focus is species-level taxonomy, but we also
discuss infraspecific taxa. Data from literature (e.g.
Stojanoff, 1939; Böcher, 1963; Meusel & Mühlberg,
1978–1979; Greuter, 1997; Jonsell & Kurtto, 2001;
Kurtto & Wesenberg, 2001) and results of critical
morphological study of herbarium material are used
to generate consistent species descriptions, applying
the Prometheus description model (Pullan et al.,
2005). The aim is to provide comparable descriptions
for Sileneae. An advantage of such a system is the
possibility to continuously update the available infor-
mation, for example when more data are gathered or
new taxonomic decisions are made, as the information
is available online (http://www.sileneae.info). A second
aim of our study is to detect possible morphological
characters differentiating Viscaria and Atocion, and
them together from other Sileneae. We also discuss
the geographical distribution and ecology of the taxa
and make a critical evaluation of the nomenclature.
An identification key for Atocion and Viscaria is also
provided.
MATERIAL AND METHODS
LITERATURE REVIEW
The original descriptions (e.g. Baumgarten, 1816;
Grisebach, 1843; Boissier, 1856; Huber-Morath,
Coode & Cullen, 1967) and several regional and
national floras (e.g. Coode & Cullen, 1967; Meusel &
Mühlberg, 1978–1979; Melzheimer, 1986; Chater
et al., 1993; Greuter, 1997; Jonsell & Kurtto, 2001;
Kurtto & Wesenberg, 2001) were checked in order to
create a list of morphological characters that poten-
tially differentiate the taxa studied. The list was
completed on the basis of field experience and study of
herbarium material and then used to extend the
Sileneae-specific character list established by F.
Eggens, M. Thollesson & B. Oxelman (unpubl. data)
in the X303 database based on the Prometheus
description model (Pullan et al., 2005).
FIELDWORK AND HERBARIUM MATERIAL
Specimens of A. armeria,A. lerchenfeldianum
(Baumg.) M.Popp, A. rupestre (L.) Oxelman, V. alpina,
V. asterias and V. vulgaris s.l. were collected during
excursions to the Pyrenees, Alps, Balkan Peninsula
and Caucasus in 2003–2012. Morphological charac-
teristics and ecology were observed in the field. Her-
barium material from BM, BRNM, C, CGE, CL, E,
G-BOIS, GH, GOET, K, KRAM, LE, LINN, LJM,
LJU, LW, O, P, PRC, S, SAV, SKO, SOM, UPS, W and
WU (abbreviations from Thiers, 2013), comprising
>500 specimens, was checked.
SPECIES DESCRIPTIONS AND IDENTIFICATION KEY
Variation of character states was observed on plants
in the field and on herbarium specimens. Flower
characters were mostly observed and measured under
a stereomicroscope using dissected flowers from her-
barium specimens. Variation observed was compared
and complemented with data from the literature and
used to generate species descriptions, employing the
Prometheus description model (Pullan et al., 2005)
with angiosperm ontology, using a template modified
for the use in Sileneae (F. Eggens, M. Thollesson & B.
Oxelman, unpubl. data), which was adjusted for our
purposes. The (morphological) terminology used is
mostly well defined in the original angiosperm ontol-
ogy (Pullan et al., 2005). ‘Basal leaves’ in our study
refers to the rosulate leaves at the base of the stem.
The characteristics of stem leaves were observed
approximately in the middle of the stem. All flower
characters were observed on hermaphroditic flowers,
196 B. FRAJMAN ET AL.
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210
as female flowers, which occur occasionally in Sile-
neae, usually have shorter calyces, anthophores and
petals. Petal colour was in most cases observed in the
field. Chromosome numbers and distribution data are
summarized from the literature. Habitat information
is based on field observations, herbarium labels and
literature. Descriptions, information about specimens
and some specimen images are available at the Sile-
neae database at http://www.sileneae.info.
An identification key for Atocion and Viscaria is
based on the morphological revision and species
descriptions. Whenever possible, several characters
are used to discriminate the (groups of) taxa. Even if
in some cases the character states are overlapping,
such characters can still be valuable in the identifi-
cation process.
RESULTS AND DISCUSSION
MORPHOLOGY
It is difficult to find morphological synapomorphies to
separate Atocion and Viscaria from other Sileneae.
Oxelman & Lidén (1995) and Oxelman et al. (2001a)
discussed some diagnostic features of Atocion and
Viscaria, but plants with similar characteristics can
also be found in Silene.Atocion and Viscaria are
mostly glabrous (except ciliate leaf bases in Viscaria),
and most taxa have viscid upper portions of distal
internodes (but not V. alpina,A. lerchenfeldianum
and A. rupestre). Bracts are mostly scarious or green
with a scarious margin. Oxelman et al. (2001a) also
listed inconspicuous calyx veins as diagnostic for
Atocion and Viscaria, but in some Atocion taxa
they are rather prominent (e.g. A. compactum and
A. armeria), as in V. asterias. Most of the taxa have
pink to purple or reddish petals; they are paler in
A. lerchenfeldianum and mostly white in A. rupestre.
The main character differentiating Atocion from
Viscaria is life form. Viscaria taxa are perennials with
creeping rhizomes, whereas Atocion comprises
annuals to short-lived perennials with taproots that
can, however, be slightly woody in perennial individu-
als. Atocion taxa also have a glaucous coating on the
surface of all above-ground vegetative parts, which is
not present in Viscaria. Glaucous here refers to a
greyish or bluish waxy coating or bloom that is easily
rubbed off; in the terminology of the Prometheus
description model (Pullan et al., 2005) it is defined as
a grey– or blue–green colour. Another differentiating
character between Atocion and Viscaria is the inflo-
rescence type. Viscaria has thyrsoid inflorescences,
i.e. panicle-like clusters with cymose, simultaneously
flowering lateral and terminal partial inflorescences
(dichasia), which are more (V. alpina) or less (some
populations of V. vulgaris) compact and head-like. In
V. asterias, the inflorescence is head-like with densely
clustered elements and the underlying inflorescence
type is difficult to assess. The inflorescence type in
Atocion is a terminal compound dichasium, which can
be rather dense and head-like, as in A. compactum.
Sometimes lateral (axillary) inflorescences are
formed, but always after the terminal inflorescence is
already developed. Whereas the pistil is generally
composed of three carpels in Atocion (four or five
styles are occasionally found), Viscaria taxa have
either five (V. alpina,V. vulgaris) or three (V. asterias)
carpels. Atocion and Viscaria taxa have the chromo-
some number 2n= 24 as do most of the European
members of Silene (e.g. Goldblatt & Johnson,
1979–2009; Degraeve, 1980; Chater et al., 1993).
TAXONOMIC REVISION
In general, subspecific and varietal names are
included in the synonymy, whereas the forms are
mostly ignored. Many of the synonyms listed have
been rarely cited, the most notable exceptions
being Rohrbach (1869, for Atocion and V. asterias;
Rohrbach, 1869–70, for other Viscaria taxa) and
Ascherson & Graebner (1929).
TAXONOMIC TREATMENT
ATOCION Adans., Fam. Pl. 2: 254. 1763 – Type:
Atocion armeria (L.) Raf.
Annual, biennial or perennial, glaucous and mostly
glabrous herbs with a taproot, rarely (A. lerchenfeldi-
anum) rhizome. Stem mostly erect, sometimes
KEY TO ATOCION AND VISCARIA IN SILENEAE
To key out Atocion and Viscaria from other Sileneae, the key published by Oxelman et al. (2001a) can be used, after
modifying couplet ten in the key and adding an additional couplet after that:
10. Plant hairy ......................................................................................................................... Silene
10*. Plant glabrous, leaves sometimes ciliate ...................................................................................... 10b
10b. Plant rhizomatous with head-like, densely congested inflorescences, subtended by conspicuous bracts. Petals
carmine, entire. Anthophore (5.0–)6.0–8.5(−9.0) mm long. Upper portion of the upper stem internodes glutinous
....................................................................................................................... Viscaria (V. asterias)
10b*. Plant different ................................................................... 11 (as in the key of Oxelman et al., 2001a)
REVISION OF ATOCION AND VISCARIA 197
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210
procumbent, decumbent or ascending; upper portion
of the upper internodes sometimes glutinous. Leaves
glabrous, rarely sparsely pubescent, sometimes with
ciliate lower portion. Basal leaf rosette often withered
at maturity. Inflorescence cymose (regular dichasia),
lax or capitate and (densely) congested. Lower-portion
bracts opaque, upper-portion bracts usually scarious.
Anthophore mostly 4–11 mm long, sometimes (A. rup-
estre) shorter than 1.5 mm. Calyx often purple,
cylindrical–clavate, sometimes (A. rupestre) campanu-
late. Calyx primary veins ten, not or slightly raised
and slightly anastomosing. Petals bright to purple
KEY TO SPECIES OF ATOCION AND VISCARIA
The most discriminating characters in the key are underlined.
1. Perennials with creeping rhizomes. Glaucous coating on the plant surface not present. Inflorescence thyrsoid
(panicle-like clusters with cymose, simultaneously flowering lateral and terminal partial inflorescences–dichasia),
sometimes compact and head-like. Styles five or three (Viscaria) ............................................................ 2
1*. Annuals to short-lived perennials with taproots. Glaucous coating on the plant surface present. Inflorescence a
terminal compound dichasium, sometimes dense and head-like. Styles three (Atocion) ............................... 4
2. Styles three. Petals carmine. Anthophore (5.0–)6.0–8.5(−9.0) mm long. Calyx teeth acuminate. Inflorescence
head-like, densely congested, subtended by conspicuous bracts. Middle stem leaves wider than 10 mm. Upper
portion of the upper stem internodes glutinous. Central Balkan Peninsula. ......................... Viscaria asterias
2*. Styles five. Petals purple to pink, rarely white. Anthophore 0.8–5.0(−5.5) mm long. Calyx teeth acute to rounded.
Inflorescence not head-like, lax to sparsely congested, not subtended by conspicuous bracts. Middle stem leaves up
to 10 (12) mm wide. Upper portion of the upper stem internodes glutinous or not glutinous. Europe, north-eastern
North America, Greenland. .............................................................................................................. 3
3. Plant (15–)25–80(−100) cm high. Upper portion of the upper stem internodes glutinous. Inflorescence usually not
congested. Calyx (6–)7–15(−16) mm long. Calyx primary veins prominent, slightly raised. Petal limb entire or
slightly emarginate, lobes up to 1 mm long. Coronal scales (0.5–)1.0–3.5 mm long. Capsules (6–)7–9(−10) mm long.
Europe, alien in North America................................................................................... Viscaria vulgaris
3*. Plant (2–)5–30(−40) cm high. Upper portion of the upper stem internodes not glutinous. Inflorescence usually
sparsely congested. Calyx 4.0–6.5(−7.5) mm long. Calyx primary veins not prominent, not raised. Petal limb
emarginated to half-length bifid, lobes (0.5–)1.0–3.0 mm long. Coronal scales up to 0.5 mm long or absent.
Capsules (3–)4–5 mm long. Europe, north-eastern North America, Greenland. ....................... Viscaria alpina
4. Usually perennial. Stem 0.5–1.5 mm wide, upper portion of the upper internodes not glutinous. Inflorescence
cymose, lax. Anthophore 0.6–6.5(−7.5) mm long. Petals white to bright pink. Calyx 3.5–11.0(12.0) mm long, stamen
filaments 2.5–7.5 mm long. Capsules (3–)4–6 mm long. Plant from Europe. .............................................. 5
4*. Usually annual to biennial, rarely perennial. Stem 1–10 mm wide, upper portion of the upper internodes glutinous.
Inflorescence cymose, lax or densely congested. Anthophore 5–11(−12) mm long. Petals purple–pink, rarely white.
Calyx (11–)12–20 mm long, stamen filaments 7.5–12.0 mm long. Capsules (6–)7–10 mm long. (Calyx, stamens and
capsules might be shorter in A. scythicinum, which is, however, absent from Europe). ............................... 6
5. Stem mostly erect, rarely ascending. Anthophore 0.6–1.5 mm long. Calyx campanulate, 3.5–5.5 mm long. Petals
white, rarely pink. Petal limb slightly emarginate. Coronal scales elliptic, 0.2–0.5 mm long. Scandinavia, Pyrenees,
Sierra Nevada, Sierra Cantabrica, Massif Central, Corsica, Alps, Apennines. ........................ Atocion rupestre
5*. Stem procumbent, decumbent or ascending. Anthophore (3.0–)4.0–6.5(−7.5) mm long. Calyx cylindrical–clavate,
(6.5–)7.0–11.0(−12.0) mm long. Petals bright pink, rarely white. Petal limb entire, rarely slightly emarginated.
Coronal scales linear–triangulate, (0.6–)0.8–1.5(−1.7) mm long. South-eastern Europe.Atocion lerchenfeldianum
6. Inflorescence lax to slightly congested, flower pedicels 2–15(20) mm long. ................................................ 7
6*. Inflorescence densely congested, capitate, flower pedicels 0–2(3) mm long. ............................................... 8
7. Anthophore (6–)7–9(−10) mm long. Calyx (11–)12–17(−18) mm long. Basal leaves soon withering. Europe,
north-eastern Anatolia, alien in Japan, India, South and North America. ............................ Atocion armeria
7*. Anthophore 5–7 mm long. Calyx 10–12 mm long. Basal leaves persistent. Turkey: Rize. .... Atocion scythicinum
8. (20–)30–80(−120) cm high biennial, rarely annual or perennial. Distinct stem internodes (6–)8–12(−14), stems
rarely branching in lower portion, sometimes in upper portion, branches not twiggy, without solitary flowers.
Bracts subtending the inflorescence 10–30 mm long, (3–)5–15 mm wide. Petal claw (4.5–)5.0–8.5(−9.0) mm long.
Capsules 6–8 mm long, 2.5–3.5 mm wide. South-eastern and eastern Europe, Turkey, Armenia, Georgia, Iran, Iraq
........................................................................................................................... Atocion compactum
8*. (60–)80–100(−120) cm high biennial or perennial. Distinct stem internodes three to six, stems sometimes
branching in lower portion, often in upper portion, branches usually twiggy, often with solitary flowers. Bracts
subtending the inflorescence 4–10 mm long, 2–4 mm wide. Petal claw 8–10 mm long. Capsules 8–10 mm long,
4–6 mm wide. Lebanon ......................................................................................... Atocion reuterianum
198 B. FRAJMAN ET AL.
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210
pink or white (A. rupestre). Petal limb distinct from
claw, mostly entire, sometimes slightly emarginate, in
A. rupestre emarginate. Coronal scales two, promi-
nent. Anthers pink to violet, in A. rupestre white.
Gynoecium styles three. Capsules robust, opaque,
oblong– to ovoid–ellipsoid, dehiscing by six teeth.
Seeds approximately reniform, with slightly papillate
testa. 2n= 24.
1. ATOCION ARMERIA (L.) Raf., Autik. Bot.: 26. 1840
Silene armeria L., Sp. Pl.: 420. 1753 Lychnis
armoraria Scop., Fl. Carniol., ed. 2, 1: 310. 1771,
nom. illeg. (‘armoraria’ is probably just a different
spelling of the Linnaean epithet) Cucubalus fas-
ciculatus Lam., Fl. Franç. 3: 27. 1779, nom. illeg.
Silene glauca Salisb., Prodr. Stirp. Chap. Allerton:
302. 1796, nom. illeg. Silene latifolia Gray, Nat.
Arr. Brit. Pl. 2: 647. 1821, nom. illeg., non Poir.
Silene variegata H. Leod. ex Fenzl, Adv. Bot. in
Del. Sem. Hort. Vindob.: 9. 1851, pro syn., nom.
inval. – Type: Herb. Linn. no. 583.49 (LINN!),
lectotype designated by Talavera & Muñoz Gar-
mendia in Anales Jard. Bot. Madrid 45: 435. 1989.
=Atocion armeroides Raf., Autik. Bot.: 29. 1840 –
Type not designated (specimen not at DWC, P, PH or
WIS!).
=Silene armeria var. sparsiflora Schur, Enum. Pl.
Transsilv.: 105. 1866 – Type: ‘Auf Nagelflüe bei Tal-
matsch’ Juli (holotype, LW!).
=Silene armeria var. serpentini Beck in Glasn.
Zemaljsk. Muz. Bosni Hercegovini 19: 20. 1907 –
Type: ‘Bei Žepcˇe auf Serpentin [Na serpentinu kod
Žepcˇa]’, 7.8.1896 (holotype, PRC!).
=Silene armeria var. angustifolia Rchb. ex Zapał.,
Consp. Fl. Gallic. Crit. 3: 138. 1911 (First published
as nom. nud. by Rchb. in Icon. Fl. Germ. Helv. 6: 51.
1844.) Note: on the herbarium label Wierzb. is cited
as the author of the name. – Type: ‘Bannat: auf
wüsten, steinigen Waldplätzen bei Orawicza und
Csiklova, in der zweiten Region.’, leg. Dr. Wierzbicki,
Aug. 1841, Flora Germanica Exsiccata no. 2287 (S
10–1484!), lectotype designated here. Several dupli-
cates (isolectotypes) are, according to Stafleu &
Cowan (1976–83), deposited in the following herbaria
(abbreviations according to Thiers, 2013): B, BM, BP,
BR, BREM, CGE, GFW, GOET, GRO, H, HAL, JE,
KIEL, L, LE, LJM!, M, MANCH, MPU, NCY, O, OXF,
PRC, PH, S, TU, U, W and WRSL.
=Silene berdaui Zapał., Consp. Fl. Gallic. Crit. 3:
139–140. 1911 S. armeria L. var. berdaui (Zapał.)
Kulcz. in Szafer (ed.), Fl. Polska 2: 192. 1921 – Type:
Poland, ‘na polach mie˛dzy zborzem koło Bielan’ (on
fields among cereals, near Bielany), leg. Berdau,
August 1849 (KRAM 106680!), lectotype, designated
here; ‘Janów’ prope Leopolim, leg. Król, 9.8.1874
(syntype, KRAM 106578!).
=Silene lituanica Zapał., Consp. Fl. Gallic. Crit. 3:
138. 1911 Atocion lituanicum (Zapał.) Tzvelev (as
lithuanicum’) in Novosti Sist. Vyssh. Rast. 33: 96.
2001 Atocion armeria (L.) Raf. var. lithuanicum
(Zapal.) Niketic´ & Stevanovic´ in Arch. Biol. Sci., Bel-
grade 59: 388. 2007 – Type: ‘Polesie Litewskie:
Porzecze’, leg. Rehman, 1882 (KRAM 107384!), lecto-
type, designated here; ‘Polesie volhynskii: Rokitno’,
leg. Rehman, (syntype, KRAM 107382!). Other syn-
types cited in the protologue, but specimens not seen:
‘In Polesia lituanica: Swieta Wola’ (leg. Rehman),
‘Welesnica’ (leg. Twardowska), and ‘in Polesia vol-
hyniensi: Karpilówka’ (leg. Rehman).
Description: Plant (7–)15–60(−75) cm high annual or
biennial, rarely perennial. Root a taproot, 1.5–
4.0(−5.0) mm wide. Stem erect, glaucous, 1–3(−5) mm
wide, rarely branching in lower portion, sometimes in
upper portion, glabrous, upper portion of the upper
internodes glutinous. Distinct internodes four to eight
(−13), lowermost (1.0–)1.5–2.5(−3.0) cm long, upper-
most (3–)4–8(−9) cm long. Basal leaves glaucous, gla-
brous, lower portion sometimes ciliate, oblanceolate to
obovate, 1.5–5.0(−10.0) cm long, 3–15 mm wide,
obtuse, base subcordate or slightly connate–perfoliate,
sessile. Stem leaves glaucous, glabrous, lower portion
rarely ciliate, linear, ovate–oblong or ovate to ovate–
lanceolate, (1.0–)2.5–6.0(−7.5) cm long, (7–)15–30(−35)
mm wide, obtuse or acute, base connate–perfoliate,
sessile. Inflorescence cymose, mostly lax. First flower
pedicel 0.2–1.5(−2.0) cm long. Lower-portion bracts
opaque, ovate, 5–20 mm long, (2–)3–12 mm wide,
acute, base rounded. Upper-portion bracts usually
scarious, lanceolate, 2–5(−7) mm long, (0.5–)1.0–
1.5(−2.0) mm wide, acute, base rounded. Flowers
diurnal. Anthophore glabrous, (6–)7–9(−10) mm long.
Calyx glaucous–purple, glabrous, cylindrical–clavate,
(11–)12–17(−18.5) mm long, 2.5–3.5(−4.0) mm wide.
Calyx primary veins ten, slightly raised and slightly
anastomosing. Calyx teeth mostly homogeneous, tri-
angulate, 0.7–1.4 mm long, 0.7–1.8 mm wide, obtuse
or acute, margin hairs absent or rarely present.
Corolla upper and lower surface purple–pink or rarely
white. Petal claw glabrous, 5.5–10 mm long, 0.8–
1.6 mm wide. Petal limb 3.5–6.5(−7.0) mm long,
(1.5–)2.0–5 mm wide, sometimes slightly emarginate.
Coronal scales two, linear–triangulate, (1–)2–3(−4)
mm long, apex acute. Stamen filaments 7.5–12.0 mm
long, glabrous. Anthers exserted, pink to violet. Gynoe-
cium styles three, exserted. Fruit stalks erect, gla-
brous, first fruit stalk 0.2–1.5(−2.0) cm long. Fruits
capsules, robust, opaque, oblong–ellipsoid, (6–)7–
9(−10) mm long, 3–4 mm wide. Seeds approximately
reniform, 0.50–0.65 mm wide, 0.4–0.6 mm high. Hilum
slightly sunken, side flat, groove present. Testa slightly
papillate. 2n= 24.
REVISION OF ATOCION AND VISCARIA 199
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210
Distribution: Southern, central and eastern Europe,
from the Pyrenees to the Balkan Peninsula (Jalas &
Suominen, 1986) and north-eastern Anatolia (Coode
& Cullen, 1967). In central and northern Europe it is
often considered as an established alien (Jalas &
Suominen, 1986, but see Jonsell & Kurtto, 2001).
Atocion armeria is widely cultivated as an ornamen-
tal and frequently escapes into natural habitats. It is
reported from, for example China, Japan, India,
South America, and eastern and western North
America (Meusel & Mühlberg, 1978–1979; Zhou,
Lidén & Oxelman, 2001; Morton, 2005).
Habitat: Atocion armeria prefers rather acidic soils,
often mineral-rich, mostly on schistose or clayey
ground, including ophiolithic soils like serpentines
(Greuter, 1997). It mostly grows in gravelly dry
meadows and rocky outcrops, open forests, forest
margins and clearings, and also in vineyards, fields
and ruderal places as an escape from gardens, where
it is grown as an ornamental.
Taxonomic remarks: Because of the pronounced vari-
ability across the distribution range, several infraspe-
cific taxa of A. armeria have been described (e.g.
S. armeria var. sparsiflora Schur, S. armeria var. ser-
pentini Beck), but these are probably environmentally
induced modifications. The same is true for
A. armeroides Raf., described from the eastern USA,
that was said to have narrower leaves and often few
flowers (Rafinesque, 1840). Atocion lituanicum
described from Polesia and Volhynia (nowadays
including eastern Poland, north-western Ukraine,
Belarus and southern Lithuania) is said (Zapałowicz,
1911) to differ from A. armeria by its always erect
stem, which is densely branched in the upper part,
with fleshy leaves forming a rosette at the base, and
the upper stem leaves are lanceolate with rounded–
truncate base and a revolute margin; the flowers and
capsules are a bit longer than in A. armeria, the petal
lamina is obovate, never obcordate–cuneate. Silene
berdaui from Poland is said (Zapałowicz, 1911) to
differ from A. armeria by being always annual, with
relatively short and tender stems, small leaves
shorter than internodes, which are narrowed at the
base and not amplexicaul, inflorescences with only
few flowers which are smaller than in A. armeria and
the calyx veins anastomosing in the upper part. Both
taxa described by Zapałowicz morphologically fall into
the range of variation of A. armeria, based on the
measurements and observations of a large sample
from different parts of the distribution area, and
therefore do not deserve recognition.
2. ATOCION COMPACTUM (Fisch. ex Hornem.) Tzvelev
in Novosti Sist. Vyssh. Rast. 33: 97. 2001 (pub-
lished on 30 March 2001, pre-dates the combina-
tion by Oxelman in Nordic J. Bot. 20: 517
published on 20 May 2001) Silene compacta
Fisch. ex Hornem., Hort. Bot. Hafn. 1: 417. 1813
(First published as nom. nud. by Fisch., Cat. Jard.
Pl. Gorenki ed. 2: 60. 1812) Silene purpurea
Knowles & Westc., Fl. Cab. 1: 87. 1837, nom. illeg.
Silene armeria L. var. compacta (Fisch. ex
Hornem.) B.Fedtsch. & Flerow, Fl. Evropeiskoi
Ross.: 396. 1911 – Type: ‘Silene compacta h.h., Hb.
Liebm.’ (C, 193, III, 6!), a voucher of the specimen
cultivated in the Botanical Garden in Copenhagen
(‘h.h.’ – hortus hauniensis), lectotype, designated
here.
Note: Greuter (1997) lists the following syntypes,
which are, however, not cited in the protologue:
Russia, Caucasia: ‘ad latera montis Beschtau’ and ‘in
Iberiae lapidosis circa metallifodinas Achtalenfes’,
leg. Marschall von Bieberstein. Bieberstein in Fl.
Taur.-Caucas. 1: 339. 1808, considered the plants
from the Caucasus as belonging to Silene armeria.
Later, Bieberstein (in Fl. Taur.-Caucas. 3: 306. 1819)
listed his S. armeria from Fl. Taur.-Caucas. 1 in syn-
onymy with S. compacta.
=Silene hypanica Klokov in Bot. Zhurn. (Kiev) 5:
20. 1948 Atocion hypanicum (Klokov) Tzvelev in
Novosti Sist. Vyssh. Rast. 33: 97. 2001 – Type:
Ukraine, ‘distr. Amanijev, prope illapsu riv. Bakshala
in Hypanim, sub rupibus graniticis’, leg. J. Paczoski,
4.7.1913 (holotype, LE!). Note: according to M.
Fedoronchuk (herbarium LE, pers. comm.), ‘4 VII
1913’ should be ‘7 VII 1913’, attribtuable to a mis-
print in the protologue. In the protologue, another
specimen is cited from the same locality (collected on
30 July 1930), conserved in CWB. As the first speci-
men cited is clearly designated as a type (‘typus’),
the latter, at CWB, is a paratype.
=Silene vandasii Nábeˇlek in Spisy Prir. Fak.
Masarykovy Univ. 35: 43–44. 1923 – Type: ‘Kurdis-
taniae Turcicae distr. Hakkiari: ad pagum Hasitha
dit. Gulamerik, alt. c. 1700 m’, leg. Nábeˇlek,
17.6.1910 (holotype, SAV 4255!).
Note: Silene bella Clarke, Travels Eur. Asia & Africa
1, Appendix 5: 746. 1810, described from Crimea, was
listed by Ledebour (1842) and Rohrbach (1868) in
synonymy with A. compactum, but according to
Rubcov (1972), neither A. compactum nor any other
Atocion occurs in Crimea. Also Silene orientalis Mill.
in Gard. Dict., ed. 8., 10. 1768, was in the past listed
in the synonymy with A. compactum (e.g. Rohrbach,
1868; Ascherson & Graebner, 1929), but the specimen
Lychnis orientalis, longifolia nervosa, flore purpuras-
cente (BM 000939426!) from the Chelsea Physic
Garden, where Miller was superintendent, is actually
Silene noctiflora L. ‘Lychnis orientalis, longifolia
200 B. FRAJMAN ET AL.
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210
nervosa, flore purpurascente Tourn. Cor. 24’ is cited in
the synonymy with S. orientalis Mill. We here desig-
nate the specimen BM 000939426 as the lectotype of
S. orientalis Mill.
Description: Plant (20–)30–80(−120) cm high biennial,
rarely annual or perennial. Root a taproot, 3–10 mm
wide. Stem erect, glaucous, 2–10 mm wide, rarely
branching in lower portion, sometimes in upper
portion, glabrous, upper portion of the upper inter-
nodes glutinous. Distinct internodes (6–)8–12(−14),
lowermost 0.5–2.5 cm long, uppermost (2–)3–7(−8) cm
long. Basal leaves glaucous, glabrous, approximately
oblanceolate, 1–8 cm long, 5–15 mm wide, obtuse or
rarely acute, base slightly connate–perfoliate, sessile.
Stem leaves glaucous, glabrous, ovate to ovate–
lanceolate, (1.5–)2.5–8.5 cm long, (5–)10–30(−35) mm
wide, obtuse or rarely acute, base connate–perfoliate,
sessile. Inflorescence densely cymose and capitate,
congested. First flower pedicel 0–0.3 cm long. Lower-
portion bracts opaque, ovate, 10–30 mm long, (3–)5–
15 mm wide, acute, base rounded. Upper-portion
bracts usually scarious, ovate–lanceolate, 2–6 mm
long, 1–2 mm wide, acute, base rounded. Flowers
diurnal. Anthophore glabrous, (6–)7–11(−12) mm
long. Calyx glaucous–purple, glabrous, cylindrical–
clavate, (13–)14–18(−20) mm long, 2.5–3.5(−4.0) mm
wide. Calyx primary veins ten, slightly raised and
slightly anastomosing. Calyx teeth mostly homogene-
ous, triangulate–ovate, 0.8–1.8 mm long, 1.0–1.8 mm
wide, obtuse or acute, margin hairs absent or rarely
present, glandular hairs rarely present. Corolla upper
and lower surface purple–pink or rarely white. Petal
claw glabrous, (4.5–)5.0–8.5(−9.0) mm long, 0.7–
1.6 mm wide. Petal limb 3.5–7.5 mm long, 1.0–
3.0(−3.5) mm wide, sometimes slightly emarginate.
Coronal scales two, linear–triangulate, (1.5–)2.0–
3.5(−4.0) mm long, apex acute. Stamen filaments 7.5–
11.0 mm long, glabrous. Anthers exserted, pink to
violet. Gynoecium styles three, exserted. Fruit stalks
erect, glabrous, first fruit stalk 0–0.3 cm long. Fruits
capsules, robust, opaque, oblong-ellipsoid, 6–8 mm
long, 2.5–3.5 mm wide. Seeds approximately reni-
form, 0.50–0.65 mm wide, 0.40–0.55 mm high. Hilum
slightly sunken, side flat, groove present. Testa
slightly papillate. 2n= 24.
Distribution: South-eastern and eastern Europe (Mac-
edonia, Bulgaria, Greece, Romania, Ukraine; Jalas &
Suominen, 1986; Micevski, 1993), Turkey, Armenia,
Georgia, Iran, Iraq (Coode & Cullen, 1967; Rechinger
et al., 1988).
Habitat: Atocion compactum has a similar ecology to
A. armeria. It grows on siliceous substrate (schist or
granite) in open forests, gravelly places, rock ledges
and cliffs and at field and road margins and other
ruderal places.
Taxonomic remarks: Silene vandasii, described from
Turkey, was said to differ from A. compactum in being
perennial and in having acute calyx teeth (Nábeˇlek,
1923). Coode & Cullen (1967) and Rechinger et al.
(1988) considered it as a perennial variant of A. com-
pactum, whereas Lazkov (2003) retains its independ-
ent status. As annual/biennial forms can have
acute calyx teeth, we consider it as a synonym of
A. compactum.
Chater et al. (1993) considered A. hypanicum a
variant of A. compactum. According to Klokov (1948),
it differs from A. compactum in having herbaceous
(not membranaceous) involucral bracts, i.e. upper
cauline leaves, shorter bracts and coronal scales,
acuminate (not obtuse) calyx teeth and brown (not
black) seeds. The length of the bracts and coronal
scales falls into the variation range of A. compactum,
and ripe seeds in A. compactum (as in other Atocion
spp.) are dark brown, not black, with a greyish layer
that quickly erodes (seeds that are not ripe are
brownish). The type specimen of A. compactum has
only partly (in some inflorescences) membranaceous
involucral bracts, but there are also specimens from
Caucasus, with herbaceous bracts, as is the case
throughout the range of the species (B. Frajman, pers.
observ.). This character therefore seems of no taxo-
nomic value and we treat A. hypanicum as synonym
of A. compactum.
The phylogenetic study by Frajman et al. (2009b)
indicates that accessions of A. compactum from the
Caucasus and Ukraine are different from accessions
from other parts of the distribution area (plastid DNA
sequences, ITS, RPD2a,RPD2b), but the geographical
sampling was rather limited. Moreover, in the plastid
DNA tree the accessions of A. compactum are found in
three different positions and the pattern seen likely
resulted through plastid capture (Frajman et al.,
2009b). Accessions from the Caucasus and Ukraine
are grouped together, and the ones from Turkey form
their own clade, whereas the accession from Macedo-
nia is positioned in A. armeria. A phylogenetic and
morphological study with more extensive geographi-
cal sampling of A. compactum would be helpful to
clarify whether some genetically and morphologically
delimited geographical entities could be recognized.
3. ATOCION LERCHENFELDIANUM (Baumg.) M.Popp in
Nordic J. Bot. 20: 517. 2000, publ. 2001 Silene
lerchenfeldiana Baumg., Enum. Stirp. Transsilv. 1:
398–399. 1816. Described from Romania (Transsyl-
vania): ‘in summis alpium jugis ad Distr. Fagaras
ibidem inter rupium fissures versus Warfulo Goltz’
REVISION OF ATOCION AND VISCARIA 201
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210
leg. Baumgarten, 1799; ‘in alpibus Huyadensibus
ac quidem la Warfulo Paringuluj 1812 iterum
observata’. As the plant was just observed (‘obser-
vata’) at Warfulo Paringuluj, the specimen from
Fagaras is a holotype. As it is likely that the
holotype has been lost (it is not in the Baumgarten
herbarium in Cluj, CL), we here designate a
neotype. – Type: ‘In Alpe Rolz.’, leg. Baumgarten,
Flora Germanica Exsiccata no. 1995 (S 09-42550!),
neotype designated here. Several duplicates (iso-
neotypes) are according to Stafleu & Cowan
(1976–83) deposited in the following herbaria
(abbreviations according to Thiers, 2013): B, BM,
BP, BR, BREM, CGE, GFW, GOET, GRO, H, HAL,
JE, KIEL, L, LE, LJM!, M, MANCH, MPU, NCY,
O, OXF, PRC, PH, S, TU, U, W and WRSL.
=Silene macedonica Formánek in Verh. Naturf.
Vereins Brünn 32: 183. 1893, publ. 1894 S. lerchen-
feldiana Baumg. var. macedonica (Formánek) Bornm.
in Bot. Jahrb. Syst. 59: 409. 1925 – Type: Macedonia,
‘Plasnica et Pusta rjeka pl.’, leg. Formánek, 10.8.1893
(PR), lectotype designated by Greuter in Strid & Tan
(eds.), Fl. Hellenica 1:291. 1997; isotype, WU!.
=Silene lerchenfeldiana Baumg. var. rhodopea
Formánek in Verh. Naturf. Vereins Brünn 36: 97.
1898 – Type: Bulgaria, Rodopi Mountains, ‘Katlovi- et
Milevi Skali’, leg. Formánek, 1897. Syntypes, accord-
ing to Greuter (1997) possibly stored at BRNM, but
according to K. Sutorý (herbarium BRNM, pers.
comm.) not there; also not at PR!
Description: Plant (3–)5–25(−30) cm high perennial.
Rhizome (1.5–)2.0–4.0(−5.0) mm wide. Stem procum-
bent, decumbent or ascending, glaucous, 0.5–1.5 mm
wide, rarely branching in lower and upper portion,
glabrous, upper portion of the upper internodes not
glutinous. Distinct internodes four to seven, lowermost
1.0–2.5 cm long, uppermost 1.5–3.5 cm long. Basal
leaves glaucous, glabrous or rarely sparsely pubescent,
lower portion sometimes ciliate, oblanceolate, (1.0–)
1.5–6.0 cm long, 2–8(−12) mm wide, obtuse or acute,
base tapering or slightly connate–perfoliate, sessile.
Stem leaves glaucous, glabrous, lower portion rarely
ciliate, ovate–lanceolate to lanceolate, 1.0–3.5 cm long,
(2–)3–6(−7) mm wide, obtuse or acute, base connate–
perfoliate, sessile. Inflorescence cymose, mostly lax.
First flower pedicel 0.2–0.8(−1.5) cm long. Lower-
portion bracts opaque, linear–lanceolate, 2–6(−8) mm
long, 0.5–2.0 mm wide, acute, base rounded. Upper-
portion bracts usually scarious, linear, 1–3 mm long,
0.3–1.0(−1.5) mm wide, acute, base rounded. Flowers
diurnal. Anthophore glabrous, (3.0–)4.0–6.5(−7.5) mm
long. Calyx purple–glaucous, glabrous, cylindrical–
clavate, (6.5–)7.0–11.0(−12.0) mm long, 2.5–3.5 mm
wide. Calyx primary veins ten, slightly raised and
slightly anastomosing. Calyx teeth mostly homogene-
ous, triangulate–ovate, 0.8–2.0 mm long, 1.0–2.5 mm
wide, rounded, margin hairs absent. Corolla upper and
lower surface bright pink or rarely white. Petal claw
glabrous, 2.8–5.0(−5.5) mm long, 0.7–1.3 mm wide.
Petal limb (2.5–)3.0–4.5(−5.5) mm long, (0.8–)1.0–
2.0(−2.2) mm wide, sometimes slightly emarginate.
Coronal scales two, linear–triangulate, (0.6–)0.8–
1.5(−1.7) mm long, apex acute. Stamen filaments 3.5–
7.5 mm long, glabrous. Anthers exserted, pink.
Gynoecium styles three, exserted. Fruit stalks erect,
glabrous, first fruit stalk 0.2–0.8(−1.5) cm long. Fruits
capsules, robust, opaque, ovoid–ellipsoid, (3.0–)
3.5–4.5(−5) mm long, 1.8–3.0(−3.5) mm wide. Seeds
approximately reniform, 0.5–0.7 mm wide, 0.40–
0.55 mm high. Hilum slightly sunken, side flat, groove
present. Testa slightly papillate. 2n= 24.
Distribution: South-eastern Europe: Romania, Serbia,
Macedonia, Bulgaria, Greece (Jalas & Suominen,
1986).
Habitat: Rock crevices, mainly on siliceous
substrates.
Taxonomic remarks: We consider a white-flowered
variant (S. macedonica) and a geographical variant
from the Rodopi Mountains (var. rhodopea) as con-
specific with A. lerchenfeldianum and follow Chater
et al. (1993) and Greuter (1997) in this respect.
4. ATOCION REUTERIANUM (Boiss. & Blanche)
Frajman in Taxon 58: 822. 2009 Silene reuteri-
ana Boiss. & Blanche in Boissier, Diagn. Pl.
Orient., ser 2, 5: 54–55. 1856 – Type: Lebanon, ‘ad
Solima’, leg. Blanche, August 1847 (G-BOIS!), lec-
totype, designated here; ‘inter El Meroudj et Beck-
faia’, leg. Gaillardot, 5.7.1853 (syntype, G-BOIS!).
Description: Plant (60–)80–100(−120) cm high bien-
nial or perennial. Root a taproot, 6–10 mm wide.
Stem erect, glaucous, 4–8 mm wide, sometimes
branching in lower portion, branching in upper
portion, glabrous, upper portion of the upper inter-
nodes glutinous. Distinct internodes three to six,
lowermost (1.5–)2.0–5.0(−6.0) cm long, uppermost
5–10(−20) cm long. Basal leaves glaucous, glabrous,
oblanceolate to ovate–lanceolate, 3–7(−8) cm long,
5–14 mm wide, obtuse or acute, base rounded, slightly
connate–perfoliate, sessile. Stem leaves glaucous, gla-
brous, ovate to ovate–lanceolate, 2.5–5.0 cm long,
7–15 mm wide, obtuse or rarely acute, base subcor-
date and connate–perfoliate, sessile. Inflorescence
cymose and capitate, congested. First flower pedicel
0–0.2 cm long. Lower-portion bracts opaque, ovate,
4–10 mm long, 2–4 mm wide, acute, base rounded.
202 B. FRAJMAN ET AL.
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210
Upper-portion bracts usually scarious, ovate–
lanceolate, 2–6(−10) mm long, 1.0–2.5 mm wide,
acute, base rounded. Flowers diurnal. Anthophore
glabrous, 8–11 mm long. Calyx glaucous, glabrous,
cylindrical–clavate, 16–20 mm long, 2.5–4.5 mm wide.
Calyx primary veins ten, slightly raised and slightly
anastomosing. Calyx teeth mostly homogeneous,
triangulate–ovate, 1.5–1.8 mm long, 1–2 mm wide,
obtuse or rounded, margin hairs absent. Corolla
upper and lower surface purple–pink. Petal claw
glabrous, 8–10 mm long, 1–2 mm wide. Petal limb
4–5 mm long, (1.5–)2.0–4.0 mm wide, sometimes
slightly emarginate. Coronal scales two, linear–
triangulate, 2–3 mm long, apex acute. Stamen fila-
ments 10–12 mm long, glabrous. Anthers exserted,
pink. Gynoecium styles three, exserted. Fruit stalks
erect, glabrous, first fruit stalk 0–0.2 cm long. Fruits
capsules, robust, opaque, ovoid–ellipsoid, 8–10 mm
long, 4–6 mm wide. Seeds approximately reniform,
side flat, groove present. Testa slightly papillate.
Distribution: Lebanon (Mouterde, 1966).
Habitat: Sandy meadows in mountainous regions
(Mouterde, 1966).
5. ATOCION RUPESTRE (L.) Oxelman in Nordic J. Bot.
20: 517. 2000, publ. 2001 Silene rupestris L., Sp.
Pl.: 420. 1753 Cucubalus saxatilis Lam., Fl.
Franç. 3: 30. 1778, nom. illeg. Oncerum rupestre
(L.) Dulac, Fl. Hautes-Pyrénées: 257. 1867 Silene
alpestris Willd. ex Nyman, Consp. Fl. Eur. 1: 92.
1878, non Jacq., pro syn., nom. inval.– Type: Herb.
Linn. no. 583.50 (LINN!), lectotype designated by
Talavera & Muñoz Garmendia in Anales Jard. Bot.
Madrid 45: 445. 1989.
Note: de Lamarck (1778) listed both ‘Silene quad-
rifida Lin. Sp. 602’ and ‘Silene rupestris Lin. Sp. 602’
in the synonymy of Cucubalus saxatilis Lam., which
is probably a mixture of two taxa, Heliosperma alp-
estre (Jacq.) Griseb. and Atocion rupestre (L.)
Oxelman. Two specimens (P 00286979 and P
00286980) from Lamarck’s herbarium labelled as
Silene rupestris, almost certainly by Lamarck himself
(P. Corsi, pers. comm.), are actually H. alpestre and
A. rupestre, respectively.
=Silene kaulfussii Spreng., Pl. Min. Cogn. Pug. 2:
64. 1815 – Type not designated (specimen not in W!).
Description: Plant (2–)8–25(−30) cm high biennial or
perennial, rarely annual. Root a taproot, 2–5 mm
wide. Stem erect or ascending, glaucous, 0.5–1.5 mm
wide, sometimes branching in lower and upper
portion, glabrous, upper portion of the upper inter-
nodes never glutinous. Distinct internodes (3–)4–
6(−7), lowermost 0.5–2.0(−3.0) cm long, uppermost
(0.5–)1.0–2.5(−3.5) cm long. Basal leaves glaucous,
glabrous, lower portion sometimes ciliate, oblong–
lanceolate to oblanceolate, 0.5–3.0(−4.0) cm long,
1.5–2.5 mm wide, obtuse, base subcordate or slightly
connate–perfoliate, sessile. Stem leaves glaucous,
glabrous, lower portion rarely ciliate, ovate–
lanceolate to elliptic, (0.7–)1.0–2.5(−3.5) cm long,
3–9(−15) mm wide, obtuse or acute, base connate–
perfoliate, sessile. Inflorescence cymose, lax. First
flower pedicel 1.0–3.0(−3.5) cm long. Lower-portion
bracts opaque, ovate–lanceolate, 5–15 mm long,
2–6 mm wide, acute, base rounded. Upper-portion
bracts usually scarious, linear, 0.6–2.0 mm long, 0.3–
0.6 mm wide, acute, base rounded. Flowers diurnal.
Anthophore glabrous, 0.6–1.5 mm long. Calyx glau-
cous, glabrous, campanulate, 3.5–5.5 mm long,
2–3 mm wide. Calyx primary veins ten, not raised
and slightly anastomosing. Calyx teeth mostly homo-
geneous, triangulate–ovate, 0.5–1.5(−1.7) mm long,
0.8–1.5 mm wide, obtuse, margin hairs absent or
rarely present, glandular hairs rarely present.
Corolla upper and lower surface white or rarely
pink. Petal claw glabrous, 2.5–4.5 mm long, 0.5–
1.5 mm wide. Petal limb 2.0–4.0(−4.5) mm long,
(1.3–)1.5–2.5(−3.0) mm wide, slightly emarginate,
lobes oblong, slightly divergent, 0.2–0.7 mm long,
0.5–1.3 mm wide. Coronal scales two, elliptic, 0.2–
0.5 mm long, apex obtuse. Stamen filaments 2.5–
5.0 mm long, glabrous. Anthers exserted, white.
Gynoecium styles three, exserted. Fruit stalks erect,
glabrous, first fruit stalk 1.0–3.0(−3.5) cm long.
Fruits capsules, robust, opaque, ovoid–oblong, (3.5–
)4.0–6.0 mm long, 1.8–3.0(−3.5) mm wide. Seeds
approximately reniform, 0.50–0.65 mm wide, 0.40–
0.55 mm high. Hilum slightly sunken, side flat,
groove present. Testa slightly papillate. 2n= 24.
Distribution: Scandinavia (Norway, Sweden, Finland),
Pyrenees (Spain, France), Sierra Nevada, Sierra Can-
tabrica (Spain), Massif Central, Corsica (France),
Alps (France, Italy, Switzerland, Germany, Austria,
Slovenia), Apennines (Italy) (Jalas & Suominen,
1986).
Habitat: Limestone and mineral-poor, base-rich, but
relatively acidic soils in mountainous regions. A plant
of siliceous rock fissures and ledges and on open soil
in rocky outcrops; in Scandinavia also in open lichen-
rich pine forests (Meusel & Mühlberg, 1978–1979;
Jonsell & Kurtto, 2001).
6. ATOCION SCYTHICINUM (Coode & Cullen) Frajman
in Taxon 58: 822. 2009 S. scythicina Coode &
Cullen in Notes Roy. Bot. Gard. Edinburgh 28: 6.
1967 – Type: Turkey, Rize, Ikizdere: terraced hill-
REVISION OF ATOCION AND VISCARIA 203
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side, 700 m, flowers pink; leg. Stainton, 11.4.1960,
Stainton 8167 (holotype, E!).
Description: Plant 12–22 cm high biennial. Root a
taproot, 5 mm wide. Stem erect, glaucous, 3 mm wide,
sometimes sparsely branching in lower portion,
branching in upper portion, glabrous, upper portion of
the upper internodes glutinous. Distinct internodes
four to five, lowermost 1.5 cm long, uppermost 4.0 cm
long. Basal leaves glaucous, glabrous, oblanceolate,
4–6 cm long, 4–7 mm wide, acute, base tapering or
slightly connate-perfoliate, sessile. Stem leaves glau-
cous, glabrous, ovate–lanceolate, 2.5–4.0 cm long,
7–10 mm wide, acute, base connate–perfoliate,
sessile. Inflorescence cymose, slightly congested. First
flower pedicel 0.6 cm long. Lower-portion bracts
opaque, ovate, 17–32 mm long, 7–12 mm wide, acute,
base rounded. Upper-portion bracts usually scarious,
ovate–lanceolate, 3–5 mm long, 1.5–2.0 mm wide,
acute, base rounded. Flowers diurnal. Anthophore
glabrous, 5–7 mm long. Calyx glaucous–purple, gla-
brous, cylindrical–clavate, 10–12 mm long, 2.5–
3.0 mm wide. Calyx primary veins ten, slightly raised
and slightly anastomosing. Calyx teeth mostly homo-
geneous, triangulate–ovate, 0.8–1.0 mm long, 0.6–
1.0 mm wide, obtuse or acute, margin glandular hairs
sometimes present. Corolla upper and lower surface
pink. Petal claw glabrous, 7–8 mm long, 1.3–1.8 mm
wide. Petal limb 4–5 mm long, 2 mm wide, entire.
Coronal scales two, linear–triangulate, 1–2 mm long,
apex acute. Gynoecium styles three, exserted. Fruits
capsules.
Distribution: Known only from the type locality:
Turkey, Rize, Ikizdere (Coode & Cullen, 1967).
Habitat: Terraced hillside (Coode & Cullen, 1967).
Taxonomic remarks: Atocion scythicinum is in overall
habit similar to A. armeria, which is only reported
from the neighbouring province of Coruh (Coode &
Cullen, 1967). The phylogenetic position of A. scyth-
icinum in Atocion still needs to be investigated.
VISCARIA Bernh., Syst. Verz.: 261. 1800, nom. cons.
Steris Adans., Fam. Pl. 2: 255, 607. 1763, nom. rej.
(see Oxelman, Lidén & Jonsell, 2001b). – Type: Vis-
caria vulgaris Bernh.
Perennial, not glaucous and mostly glabrous herbs
with rhizomes. Stem mostly erect, sometimes ascend-
ing; upper portion of the upper internodes sometimes
glutinous. Leaves glabrous, sometimes with ciliate
lower portion. Basal leaf rosette mostly persistent.
Inflorescence cymose–paniculate (regular dichasia),
lax or capitate and (densely) congested. Lower-portion
bracts opaque or scarious, upper-portion bracts
usually scarious. Anthophore mostly 0.8–9.0 mm long.
Calyx purple–violet to green, clavate to campanulate
or cylindrical–clavate. Calyx primary veins ten, not
or slightly raised and slightly anastomosing. Petals
purple to pink or carmine, rarely white. Petal limb
distinct from claw, entire, slightly emarginated or
bifid. Coronal scales absent or two, prominent only in
V. vulgaris. Anthers pink to lilac or violet–brown in
V. asterias. Gynoecium styles three (in V. asterias)or
five. Capsules robust, opaque, ellipsoid to (ob)ovoid,
dehiscing by six (V. asterias) or ten teeth. Seeds
(orbicular–)reniform, with slightly to strongly papil-
late testa. 2n= 24.
1. VISCARIA ALPINA (L.) G.Don, Gen. Hist. 1: 415.
1831 Lychnis alpina L., Sp. Pl.: 436. 1753
Agrostemma alpina (L.) J.Forbes, Hort. Woburn.:
104. 1833 Liponeurum alpinum (L.) Schott,
Nyman & Kotschy in Analect. Bot.: 55. 1854
Silene liponeura Neumayer in Verh. Zool.-Bot. Ges.
Wien 72: 55. 1923 Steris alpina (L.) Šourková in
Novit. Bot. Inst. Horto Bot. Univ. Carol.: 27. 1976
– Type: Herb. Linn. no. 602.3 (LINN!), lectotype
designated by Ikonnikov in Novosti Sist. Vyssh.
Rast. 24: 80. 1987. Jonsell & Jarvis in Nordic J.
Bot. 14: 157. 1994 designated a Lapland specimen
in Paris no. 185 (LAPP) as lectotype, but their
choice is pre-dated by that of Ikonnikov.
=Lychnis frigida Schrank in Denkschr. K. Baier.
Bot. Ges. Regensburg 1(2): 25. 1818. Note: according
to Rohrbach (1869–70), L. frigida is synonymous with
V. alpina. – Type not designated (specimen not at M
nor REG!).
=Lychnis suecica Lodd. in Bot. Cab. 9: t. 881. 1824
Viscaria suecica (Lodd.) Sweet, Hort. Brit., ed 3: 66.
1839 Silene suecica (Lodd.) Greuter & Burdet in
Willdenowia 12: 190. 1982 – Type: illustration plate
881 in Loddiges, Bot. Cab. 9. 1824, lectotype desig-
nated here (no original material at K!). Note:
Agrostemma suecica Maund in Bot. Gard. 6: 576.
1835–36, has sometimes been listed as synonym to
L. suecica, but it is clear from Maund’s illustration
that he applied this name to a Dianthus species.
=Lychnis helvetica G.Don ex Loudon, Hort. Brit.:
186. 1830 – Type: illustration plate 394 of Lychnis
alpina in Curtis, Bot. Mag. 11: 394. 1797, lectotype
designated here (no original herbarium material at
BM!).
=Lychnis alpina var. americana Fernald in
Rhodora 42: 259. 1940 Viscaria alpina subsp. ameri-
cana (Fernald) Böcher in Biol. Skr. 11: 27. 1963
Lychnis alpina subsp. americana (Fernald) Feilberg
in Meddel. Gronland, Biosci. 15: 12. 1984 Steris
americana (Fernald) Ikonn. in Novosti Sist. Vyssh.
Rast. 24: 81. 1987 – Type: SE Canada, ‘Blomidon
204 B. FRAJMAN ET AL.
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210
Mountains, Bay of Islands, Newfoundland’, leg.
Fernald, Wiegand & Kittredge, 3395, 24.7.1910 (holo-
type, GH!).
=Viscaria alpina var. serpentinicola Rune in Acta
Phytogeogr. Suec. 31: 56. 1953 Lychnis alpina var.
serpentinicola (Rune) Kallio & Y.Mäkinen in Ann.
Univ. Turku, A, Biol. Geogr. Geol. 67: 63. 1982, pre-
dating the new combination by Ericsson in Svensk
Bot. Tidskr. 79: 115. 1985. – Type: ‘Suecia, Lapponia
lulensis: Purnu, in saxis serpentini montis Lingonber-
get’, leg. O. Rune, 5.7.1949 (holotype, UPS!).
=Viscaria alpina subsp. borealis Böcher in Biol.
Skr. 11: 27. 1963 Steris alpina subsp. borealis
(Böcher) Á.Löve in Phytologia 50: 171. 1982 Steris
borealis (Böcher) Ikonn. in Novosti Sist. Vyssh. Rast.
24: 82. 1987 – Type: Island, ‘Hafnarfjördur meridi-
onali’, leg. T. W. Böcher, 14. 7. 1962 (holotype, C!).
Description: Plant (2–)5–30(−40) cm high perennial.
Rhizome 1–3(−5) mm wide. Stem erect or sometimes
ascending, green, 0.5–2.5(−3.5) mm wide, rarely
branching in lower and upper portion, glabrous,
upper portion of the upper internodes not glutinous.
Distinct internodes (1–)2–4(−5), lowermost 0.3–2.5 cm
long, uppermost 0.5–6.5 cm long. Basal leaves green,
glabrous, lower portion sometimes ciliate, lanceolate–
spathulate to linear, (1.0–)1.5–6.0(−8.0) cm long,
(1.0–)1.5–4.0(−6.0) mm wide, acute, base slightly
connate–perfoliate, sessile. Stem leaves green, gla-
brous, lower portion sometimes ciliate, linear–
lanceolate to ovate–oblong, 1.0–4.0(−6.5) cm long,
1.5–6.0(−10.0) mm wide, acute or rarely obtuse, base
connate–perfoliate, sessile. Inflorescence cymose–
paniculate, usually congested. First flower pedicel
(0.1–)0.2–0.4(−0.6) cm long. Lower-portion bracts
opaque or scarious, ovate or lanceolate, (5–)8–20(−25)
mm long, (1.0–)1.5–3.0(−4.0) mm wide, acute or
obtuse, base rounded. Upper portion bracts scarious,
ovate or lanceolate, 2–8(−10) mm long, 0.5–2.0 mm
wide, acute or obtuse, base rounded. Flowers diurnal.
Anthophore glabrous, 0.8–2.0 mm long. Calyx mostly
purple–violet, glabrous, campanulate or ovoid, 4.0–
6.5(−7.5) mm long, (2.0–)3.0–4.5(−5) mm wide. Calyx
primary veins ten, not raised, obscure, slightly anas-
tomosing. Calyx teeth mostly homogeneous,
triangulate–ovate, 0.5–1.5 mm long, 0.8–2.0 mm
wide, rounded, margin hairs absent. Corolla upper
and lower surface pink, rarely white. Petal claw gla-
brous, 3.5–5.0(−5.5) mm long, 0.5–1.5 mm wide. Petal
limb (2.5–)3.5–5.0(−6.0) mm long, 1.5–3.5(−4.0) mm
wide, approximately bifid, lobes usually linear,
slightly divergent, (0.5–)1.0–3.0 mm long, 0.7–1.8 mm
wide. Coronal scales 0 or two, usually linear,
0–0.5 mm long, apex rounded to obtuse. Stamen fila-
ments 4.5–8.5 mm long, glabrous. Anthers exserted,
pink to lilac. Gynoecium styles five, exserted. Fruit
stalks erect, glabrous, first fruit stalk (0.1–)0.2–
0.4(−0.6) cm long. Fruits capsules, robust, opaque,
ovoid, (3–)4–5 mm long, 2.0–3.0(−3.5) mm wide. Seeds
orbicular–reniform, 0.5–0.7(−0.8) mm wide, 0.4–
0.6(−0.7) mm high. Hilum slightly sunken, side flat,
groove present. Testa slightly papillate. 2n= 24.
Distribution: Viscaria alpina is an amphi-atlantic
alpine plant (Hultén, 1958; Meusel & Mühlberg,
1978–1979), found in north-eastern North America
(Canada), Greenland, Iceland, Scandinavia (Norway,
Sweden, Finland), parts of north-western Russia,
Scotland (UK), Sierra Cantabrica, Pyrenees (Spain,
France), Alps (France, Switzerland, Italy, Austria)
and Apennines (Italy; Hultén, 1958; Jalas &
Suominen, 1986).
Habitat: Great variety of substrates, mostly acidic,
limestone-poor and dry ground, with a slight prefer-
ence for richer mineral soils. From hard gneiss and
rapakivi rock and poor sandy soils to rich mica schist
soils and serpentine gravel (Meusel & Mühlberg,
1978–1979; Kurtto & Wesenberg, 2001). In the Alps,
it grows mostly on dry, gravelly grasslands and rocky
outcrops (Meusel & Mühlberg, 1978–1979), and in
Scandinavia on grass heaths, gravely snowbeds, cliffs,
short-grass meadows, river banks, and apophytically
along roads, paths and in pastures (Kurtto &
Wesenberg, 2001).
Taxonomic remarks: Viscaria alpina is morphologi-
cally rather variable and several infraspecific taxa
have been recognized, being geographically and/or
ecologically correlated (e.g. Viscaria alpina subsp.
americana, subsp. borealis, var. serpentinicola).
However, the pattern of variation in the allozyme
study by Haraldsen & Wesenberg (1993) and a lack of
resolution in the phylogenetic study by Frajman et al.
(2009b) do not support these infraspecific taxa. Kurtto
& Wesenberg (2001) questioned the value of subdivi-
sion into subspecies and varieties, and we concur.
2. VISCARIA ASTERIAS (Griseb.) Frajman in Taxon 58:
822. 2009 Silene asterias Griseb., Spic. Fl.
Rumel. 1: 168. 1843 Saponaria asterias (Griseb.)
Griseb., Spic. Fl. Rumel. 2: 166. 1844 Atocion
asterias (Griseb.) Tzvelev in Novosti Sist. Vyssh.
Rast. 33: 97. 2001 (published on 30 March 2001,
and pre-dates the combination by M. Lidén in
Nordic J. Bot. 20: 517, published on 20 May 2001)
– Type: Greece, W Macedonia, ‘juxta rivulos in flor.
Alp. Nidgé (Mount Voras) social’ – according to
Strid (1986) this is Mount Piperitsa south-west of
Kajmakcˇalan, Grisebach 738 (holotype, GOET!).
=Silene asterias Griseb. subsp. radikae Trinajstic´
& Zi.Pavletic´ in Suppl. Fl. Anal. Jugosl. 6: 14. 1979
REVISION OF ATOCION AND VISCARIA 205
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S. asterias Griseb. f. radikae (Trinajstic´ & Zi.Pavletic´)
Micevski in Prilozi Oddel. Biol. Med. Nauki, Make-
donska Akad. Nauk. Umet. 9: 45. 1988, publ. 1991 –
Type: Macedonia, ‘Fluvius Radika in Makedonija’
(holotype). Note: the holotype is according to the
protologue deposited in ‘Hb. I Trinajstic´, ZA 11934’,
but it is not there (M. Plazibat, pers. comm.). It is
probably in the private collection of I. Trinajstic´.
Description: Plant (35–)60–90(−100) cm high peren-
nial. Rhizome (3–)5–10(−13) mm wide. Stem erect,
green, (2–)3–7(−9) mm wide, rarely branching in lower
portion, sometimes in upper portion, glabrous, upper
portion of the upper internodes glutinous. Distinct
internodes (three–)four to six(–seven), lowermost 1–4
cm long, uppermost 5–25 cm long. Basal leaves green,
glabrous, lower portion usually ciliate, oblanceolate–
spathulate, (5–)6–18(−23) cm long, 15–25(−30) mm
wide, obtuse or acute, base tapering or slightly
connate–perfoliate, sessile. Stem leaves green, gla-
brous, lower portion usually ciliate, oblanceolate to
ovate–lanceolate, 4–10(−15) cm long, 10–30(−40) mm
wide, obtuse or acute, base connate–perfoliate, sessile.
Inflorescence approximately capitate, densely con-
gested. First flower pedicel 0.05–0.20(−0.30) cm long.
Lower-portion bracts opaque, ovate or ovate–
lanceolate, 7–20(−30) mm long, 5–10 mm wide, acute,
base rounded. Upper-portion bracts scarious, ovate–
lanceolate, 5–8(−10) mm long, 1–2 mm wide, acute,
base rounded. Flowers diurnal. Anthophore glabrous,
(5.0–)6.0–8.5(−9.0) mm long. Calyx mostly purple–
green, glabrous, cylindrical–clavate, (10–)12–17(−18)
mm long, 2–4 mm wide. Calyx primary veins ten,
slightly raised and slightly anastomosing. Calyx teeth
mostly homogeneous, triangulate–lanceolate, (1.2–)
2.0–4.0(−5.0) mm long, 1.3–2.5 mm wide, acuminate,
margin hairs absent. Corolla upper and lower surface
carmine. Petal claw glabrous, 5.5–7.5 mm long, 0.7–
1.6 mm wide. Petal limb 3.8–6.2 mm long, 1.0–2.5 mm
wide, entire. Coronal scales none or two, 0–0.3 mm
long. Stamen filaments 8–12 mm long, glabrous.
Anthers exserted, violet–brown. Gynoecium styles
three, exserted. Fruit stalks erect, glabrous, first fruit
stalk 0.05–0.20(−0.30) cm long. Fruits capsules,
robust, opaque, ellipsoid or obovoid, (5–)6–8 mm long,
2–4 mm wide. Seeds reniform, 0.6–0.8 mm wide, 0.50–
0.65 mm high. Hilum slightly sunken, side flat, groove
absent. Testa strongly papillate. 2n= 24.
Distribution: Viscaria asterias is endemic to the
central part of the Balkan Peninsula (Albania,
Kosovo, Montenegro, Serbia, Macedonia, Greece, Bul-
garia; Jalas & Suominen, 1986).
Habitat: Along streams and in wet meadows on sili-
ceous substrates.
Taxonomic remarks: Viscaria asterias has always
been considered to be closely related to Atocion taxa,
but a phylogenetic study (Frajman et al., 2009b) has
shown its unambiguous position in Viscaria. Boissier
(1867) classified the species in a monotypic series
Armeroideae (Table 1). Silene asterias subsp. radikae
is said to have calyx approximately 8 mm long and
long–mucronate calyx teeth (Trinajstic´ & Pavletic´,
1979). The calyx is short compared with the variation
range in V. asterias, whereas the long–mucronate
calyx teeth can be found also in plants with longer
calyces from other parts of the species range.
3. VISCARIA VULGARIS Bernh., Syst. Verz.: 200. 1800
Lychnis viscaria L., Sp. Pl.: 436. 1753 Silene
viscaria (L.) Borkh., Rhein. Mag. Enweit. Naturk.
1: 520. 1793. Note: ‘Silene viscaria (L.) Jess., Deut.
Excurs.-Fl.: 280. 1879’ is often incorrectly listed as
the first published combination in Silene!Viscago
vulgaris Röhl., Deut. Fl. 2: 275. 1796, nom. illeg.
Lychnis angustifolia Gray, Nat. Arr. Brit. Pl. 2:
648. 1821, nom. illeg. Steris viscaria (L.) Raf.,
Autik. Bot. 1: 27. 1840 Viscaria viscosa Asch., Fl.
Brandenburg 1: 85. 1860, nom. illeg. – Type: Herb.
Linn. no. 602.2 (LINN!), lectotype designated by
Ikonnikov in Novosti Sist. Vyssh. Rast. 24: 80.
1987. Jonsell & Jarvis in Nordic J. Bot. 14: 157.
1994, designated the same specimen as lectotype,
but their typification is pre-dated by that of
Ikonnikov.
Note: Ascherson & Graebner (1929) used the name
Viscaria viscaria (L.) Voss, in Vilm. Blumengärtn., ed.
3, 1: 100. 1894, nom. inval., for this species. In syn-
onymy they also list Lychnis viscosa Scop., Fl.
Carniol. 1, ed. 2: 306. 1772, but L. viscosa Scop. is
clearly based on Cucubalus viscosus L., Sp. Pl. 1: 414.
1753, nowadays known as Silene viscosa (L.) Pers. As
the latter does not grow in Carniolia (today part of
Slovenia), Scopoli must have misapplied the name for
some other species, possibly V. vulgaris.
=Viscaria neglecta G.Don, Gen. Hist. 1: 415. 1831
Lychnis neglecta G.Don ex Loudon, Hort. Brit.: 186.
1830, nom. nud. Viscaria alba Hort. ex Loudon,
Hort. Brit.: 186. 1830, pro syn., nom. inval. Viscaria
albiflora Sweet, Hort. Brit., ed. 2: 54. 1830, nom. nud.
– Type not designated (specimen not at BM!).
=Viscaria purpurea Wimm., Fl. Schles., 67. 1841,
nom nud.
=Viscaria atropurpurea Griseb., Spic. Fl. Rumel. 1:
166. 1843 Lychnis atropurpurea (Griseb.) Nyman,
Syll. Fl. Eur.: 229. 1855 Viscaria vulgaris subsp.
atropurpurea (Griseb.) Stoj. in Annu. Univ. Sofia Phys.-
Math. 3 (Sci. Nat.) 35: 326. 1939 Steris atropurpurea
(Griseb.) Holub in Folia Geobot. Phytotax. 12: 429.
1977 Silene atropurpurea (Griseb.) Greuter & Burdet
in Willdenowia 12: 189. 1982 – Type: Greece, west
206 B. FRAJMAN ET AL.
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210
Macedonia, ‘in form. humidis fl. alp. Nidgé (Mount
Voras) raro’, Griesebach 752 pro parte, right-hand
plant only (GOET!), lectotype designated by Greuter in
Strid & Tan (eds.), Fl. Hellenica 1: 248. 1997.
=Viscaria sartorii Boiss. in Diagn. Pl. Orient., ser
2, 1: 79. 1854 Lychnis sartorii (Boiss.) Hayek in
Feddes Repert. Beih. 30: 287. 1924 Steris sartorii
(Boiss.) Ikonn. in Novosti Sist. Vyssh. Rast. 24: 81.
1987 Lychnis atropurpurea subsp. sartorii (Boiss.)
Micevski in Prilozi Oddel. Biol. Med. Nauki Maked.
Akad. Nauk. Umet., 9: 48. 1988, publ. 1991 – Type:
Greece, Peloponnisos, Mount Parnonas: ‘in castaneto
prope Hag. Petrum Peloponnesi’, leg. Sartori,
20.5.1837, Sartori 1824 (holotype, G-BOIS!). Note:
Strid, Mount Fl. Greece 1: 135. 1986, designated
another specimen (‘Mt. Parnassos’, leg. Samartini &
Guicciardi, 23.6.1857, G-BOIS) as lectotype, but it
could only be a neotype as it was collected after the
publication of V. sartorii. The specimen cited by Bois-
sier in the protologue is certainly the original
element, and thus the holotype, which takes prec-
edence over a neotype (McNeill et al., 2007).
=Lychnis splendens Hort. ex Vilm., Fl. Pleine Terre,
ed 2: 524. 1866 – Type not designated (synonym
according to Ascherson & Graebner, 1929).
=Viscaria vulgaris Bernh. subsp. valesiaca Thell.
in Ber. Schweiz. Bot. Ges. 22: 122. 1913 Viscaria
vulgaris ssp. helvetica Rouy et Fouc., Fl. France
3:100. 1896, nom. illeg., non Don Viscaria vallesiaca
(Thell.) Bergmans, Vaste Pl. Rotsheesters: 594. 1924 –
Type not designated. Note: According to Thellung (l.c.)
this plant is a probably a hybrid between V. vulgaris
and V. alpina.
=Lychnis viscaria subsp. subnivalis Panov in
Fitologija 48: 8–9. 1996 – Type: ‘Bulgaria occidentalis.
Mons Rila. In saxosis et lapidosis circum nives
tabescentesin loco “Gradinski esera” supra urbem
Rila, c. 2200–2300 m s.m.’. Leg. P. Panov, 16.7.1953,
(holotype, SOM 147882!).
Description: Plant (15–)25–80(−100) cm high peren-
nial. Rhizome 3–8 mm wide. Stem erect, green, 1.5–
5.0(−6.0) mm wide, not branching in lower portion,
rarely in upper portion, glabrous, upper portion of the
upper internodes glutinous. Distinct internodes two to
five, lowermost 2–10(−17) cm long, uppermost (2–)3–
18(−20) cm long. Basal leaves green, glabrous, lower
portion usually ciliate, linear–lanceolate to oblanceo-
late, (2.5–)3.0–15.0(−20.0) cm long, (2-)3–10(−15) mm
wide, acute, base slightly connate–perfoliate, sessile.
Stem leaves green, glabrous, lower–portion usually
ciliate, oblanceolate to ovate–lanceolate, (1.5–)3.0–
12.0(−18.0) cm long, 2–8(−12) mm wide, acute, base
connate–perfoliate, sessile. Inflorescence cymose–
paniculate, lax or slightly congested. First flower
pedicel (0.1–)0.2–0.4(−0.6) cm long. Lower-portion
bracts opaque, ovate–lanceolate, (8–)12–50(−80) mm
long, (1–)2–4(−6) mm wide, acute or obtuse, base
rounded. Upper portion bracts slightly scarious, ovate–
lanceolate, 2.5–7.0(−9.0) mm long, (0.5–)1.0–2.0(−2.5)
mm wide, acute, base rounded. Flowers diurnal.
Anthophore glabrous, 1.0–5.0(−5.5) mm long. Calyx
purple–violet to green, glabrous or sometimes pubes-
cent or glandular–pubescent, clavate or campanulate,
(6–)7–15(−16) mm long, (3.0–)3.5–6.0(−7.5) mm wide.
Calyx primary veins ten, slightly raised and slightly
anastomosing. Calyx teeth mostly homogeneous, tri-
angulate, (1.0–)1.2–2.5(−3.0) mm long, (1.2–)1.4–
2.5(−2.8) mm wide, acute or obtuse, margin hairs
absent. Corolla upper and lower surface purple to pink
or rarely white. Petal claw glabrous, 5.5–12.0 mm long,
1–3 mm wide. Petal limb (3.3–)4.0–10.0(−11.0) mm
long, (1.8–)2.0–8.0 mm wide, slightly emarginate or
entire, lobes 0–1 mm long. Coronal scales two, usually
linear, (0.5–)1.0–3.5 mm long, apex rounded to obtuse.
Stamen filaments 5.5–14.0 mm long, glabrous.
Anthers exserted, pink to lilac. Gynoecium styles five,
exserted. Fruit stalks erect, glabrous, first fruit stalk
(0.1–)0.2–0.4(−0.6) cm long. Fruits capsules, robust,
opaque, ellipsoid or ovoid, (6-)7–9(−10) mm long,
3–5 mm wide. Seeds reniform, 0.50–0.75 mm wide,
0.40–0.65 mm high. Hilum slightly sunken, side flat,
groove present. Testa papillate. 2n= 24.
Distribution: Viscaria vulgaris is distributed in most
parts of Europe (excluding the Iberian Peninsula,
northern Scandinavia, northern Russia, most of Italy
south of the Po river and southern Greece; Jalas &
Suominen, 1986). It is an occasional and adventive
garden escape in eastern North America (Morton,
2005).
Habitat: Dry, neutral to acidic grounds on siliceous or
ophiolitic substrate, rarely on limestone. Often on
sandy, gravely or rocky grounds, in open forests and
forest clearings, rocky outcrops and ledges and dry
grasslands (Meusel & Mühlberg, 1978–1979; Greuter,
1997; Kurtto & Wesenberg, 2001).
Taxonomic remarks: Viscaria atropurpurea has often
been considered as a species [e.g. Boissier, 1867;
Rohrbach, 1869–70; Bornmüller, 1937 as V. sartorii
(V. atropurpurea pro parte)], which differs from V. vul-
garis by its shorter anthophore [‘carpophore’ was used
by most botanists until Greuter, (1995) introduced the
more appropriate term ‘anthophore’]. Bornmüller
(1937) reported rather extensive variation in V. vul-
garis s.s., but recognized one species with a long
(V. vulgaris) and one with a relatively short antho-
phore (as V. sartorii). Stojanoff (1939), in his morpho-
logical study of V. atropurpurea and V. vulgaris (with
limited herbarium material of the latter), argued that
REVISION OF ATOCION AND VISCARIA 207
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210
the morphological variability in this species group is
more complex than merely the difference in antho-
phore length and is mostly ecologically correlated
(Stojanoff, 1939: table 1). He argued that the variation
in anthophore length is not discretely distributed, but
rather varies from one extreme to the other (short in
Greece and gradually longer towards the north and
north-west). Chater (1964) followed Stojanoff’s
approach and recognized two subspecies, differing in
anthophore length, and added: ‘none of the other
variable characters is correlated with the carpophore
(–) capsule length ratio to a significant degree’. Greuter
(1997) maintained specific rank for V. atropurpurea,
arguing that the vicarious taxa V. atropurpurea and
V. vulgaris appear to differ consistently in the antho-
phore length (at least 3 mm long in V. vulgaris and
shorter than 2 mm in V. atropurpurea), and in the
usually paler petals in V. vulgaris. A certain cline in
anthophore length (and consequently calyx shape) is
observed from south-east towards north and west, but
seems not to be correlated with other characters.
Moreover, our morphological study shows that antho-
phores shorter than 3 mm also occur in V. vulgaris s.s.
The intensity of petal colour is also rather variable
across the range of the species (B. Frajman, pers.
observ.). Moreover, patterns revealed by a phylogenetic
study based on different DNA regions (Frajman et al.,
2009b) do not correlate with differences in anthophore
length, and V. vulgaris and V. atropurpurea are best
treated as synonyms.
Hybrids between V. alpina and V. vulgaris
(V. alpina xvulgaris) have been reported; they are
only known from Scandinavia from the places where
both parental taxa grow in vicinity (Meusel &
Mühlberg, 1978–1979; Kurtto & Wesenberg, 2001).
The name V. media Fr. ex Hartman, Handb. Skand.
Fl., ed. 11: 232. 1879 has been used for the hybrids. –
Type: ‘Exs. H.N.f. 15, n38; Viscaria purpureo-alpina s.
media. Blekingia. Vedeby.’ Leg. J. A. Ankarcrona (UPS
200490!), lectotype, designated here. Several dupli-
cates (isolectotypes) from Fries’s Herbarium normale
are according to Stafleu & Cowan (1976–83) deposited
in the following herbaria (abbreviations according to
Thiers, 2013): B, BM, BP, C, CGE, K, L, LD, LE, LZ,
P, S, W, WAG.
Recently, a new species Silene hamzaoglui Budak
(Budak & Koç, 2011) has been published from Turkey.
It resembles V. vulgaris, but its identity and taxo-
nomic position still need to be examined.
ACKNOWLEDGEMENTS
The Swedish Research Council for Environment, Agri-
cultural Sciences and Spatial Planning and the
Swedish Research Council kindly provided financial
support for this study. The fieldwork and analyses
were partially supported by the scholarships of Helge
Ax:son Johnsons Stiftelse to B.F. We are most grateful
to the curators from the herbaria BM, BRNM, C, CGE,
CL, DWC, E, G-BOIS, GH, GOET, K, KRAM, LE,
LINN, LJM, LJU, LW, M, O, P, PH, PR, PRC, REG, S,
SAV, SKO, SOM, UPS, W, WIS, WU and ZA for
providing us with specimens, and to A. Strid for
providing us with the distribution data of some species
from Greece, D. R. Letz (Bratislava) for providing the
photographs of the type of S. vandasii and M. Ronikier
(Krakow) for help with the literature and the types.
Many thanks are due to M. Thulin for help with
nomenclatural issues and comments on a previous
version of the manuscript and to P. Schönswetter for
the comments on a previous version of the manuscript.
The comments of the four anonymous reviewers and
the editor Mike Fay greatly improved the manuscript.
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© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194–210
... (bas. A. compactum (Fisch.) Tzvel.), поширена на Кавказі, Балканах та частково в Передній Азії (Шишкин 1936;Frajman et al. 2013). Пізніше М.М. ...
... A. hypanicum та A. сompactum є досить близькими за морфологією, тому в західних публікаціях (Chater et al. 1993;Frajman et al. 2009Frajman et al. , 2013 ...
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Silene hamzaoglui Budak sp. nova is described as a new species from Çekerek (Yozgat, Turkey). Th e specimen was collected from calcareous rocky places, at an altitude between 1140 and 1180 m. Th e new species is similar to Silene duralii Y.Bağcı and Silene capillipes Boiss. & Heldr. Th e diff erences among these 3 species are discussed. Çekerek'ten (Yozgat/Türkiye) yeni bir tür Silene hamzaoglui (Caryophyllaceae) Özet: Silene hamzaoglui Budak sp. nova Çekerek'ten yeni bir tür olarak tanımlandı (Yozgat/Türkiye). Bitki kalkerli kayalık yerlerden, 1140-1180 metreler arasından toplandı. Yeni tür Silene duralii Y.Bağcı ve Silene capillipes Boiss. & Heldr. türlerine yakındır. Türler arasındaki farklar tartışıldı.