TAXON 67 (1) • February 2018: 83–112
Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
Untangling phylogenetic patterns and taxonomic confusion in tribe
Caryophylleae (Caryophyllaceae) with special focus on generic
boundaries
Hossein Madhani,1 Richard Rabeler,2 Atefeh Pirani,3 Bengt Oxelman,4 Guenther Heubl5 & Shahin Zarre1
1 Department of Plant Science, Center of Excellence in Phylogeny of Living Organisms, School of Biology, College of Science,
University of Tehran, P.O. Box 14155-6455, Tehran, Iran
2 University of Michigan Herbarium-EEB, 3600 Varsity Drive, Ann Arbor, Michigan 48108-2228, U.S.A.
3 Department of Biology, Faculty of Sciences, Ferdowsi University of Mashhad, P.O. Box 91775-1436, Mashhad, Iran
4 Department of Biological and Environmental Sciences, University of Gothenburg, Box 461, 40530 Göteborg, Sweden
5 Biodiversity Research – Systematic Botany, Department of Biology I, Ludwig-Maximilians-Universität München, Menzinger Str. 67,
80638 München, Germany; and GeoBio Center LMU
Author for correspondence: Shahin Zarre, zarre@khayam.ut.ac.ir
DOI https://doi.org/10.12705/671.6
Abstract Assigning correct names to taxa is a challenging goal in the taxonomy of many groups within the Caryophyllaceae. This
challenge is most serious in tribe Caryophylleae since the supposed genera seem to be highly artificial, and the available morphological
evidence cannot effectively be used for delimitation and exact determination of taxa. The main goal of the present study was to re-assess
the monophyly of the genera currently recognized in this tribe using molecular phylogenetic data. We used the sequences of nuclear
ribosomal internal transcribed spacer (ITS) and the chloroplast gene rps16 for 135 and 94 accessions, respectively, representing all 16
genera currently recognized in the tribe Caryophylleae, with a rich sampling of Gypsophila as one of the most heterogeneous groups
in the tribe. Phylogenetic trees were reconstructed using maximum parsimony and Bayesian inference methods. The results show
that most of the large genera of Caryophylleae are not monophyletic. As a result, we propose a new classification system matching
both molecular phylogenetic and morphological evidence. The main taxonomic conclusions include: (1) the description of three new
genera, (2) treating five small genera as synonyms, (3) resurrecting the genus Heterochroa with six species, and (4) proposing 23
new combinations plus 2 replacement names at the specific level. As a result, we recognize 14 genera in Caryophylleae. A diagnostic
key to all genera of Caryophylleae is provided.
Keywords carnation; Dianthus; Gypsophila; Saponaria; systematics; taxonomic revision
Supplementary Material DNA sequence alignments are available from https://doi.org/10.12705/671.6.S
INTRODUCTION
Assigning correct names to plant taxa as the basic elements of ecosystems is a critical first step for any biodiversity
inventory or monitoring program. Such inventory programs
are very important for resource management and conservation planning. In the flora of SW Asia, Caryophyllaceae are
abundant in various natural ecosystems, but the delimitation of taxa in the family is problematic with issues such
as clinal trends in morphological traits blurring the borders
between taxa, frequent hybridization in some genera (such as
Dianthus L., see Vítová & al., 2015), and seemingly artificial
taxonomic borders appearing at various ranks.
The focal group of the present study is tribe Caryophylleae
Lam. & DC., including about 630 species assigned currently to 16 genera (Hernández-Ledesma & al., 2015). It is,
after tribe Sileneae DC., the second-largest tribe in family Caryophyllaceae (Bittrich, 1993; Harbaugh & al., 2010;
Greenberg & Donoghue, 2011; Pirani & al., 2014; HernándezLedesma & al., 2015; R. Rabeler, personal data). The members
of the tribe are primarily Holarctic, with their diversity centered in Mediterranean and Irano-Turanian regions and few
taxa extending to Africa. Like many other members of the large
clade Plurcaryophyllaceae (sensu Greenberg & Donoghue,
2011), they are perennial or annual herbs, sometimes woody
at base or even spiny cushions (e.g., Acanthophyllum C.A.Mey.
spp.) characterized by five fused sepals, five more or less
clawed petals, ten free stamens, and capsules opening with
four, or rarely six, teeth (Bittrich, 1993). Some representative
images of these plants are presented in Fig. 1. Contrary to
the members of closely related tribe Sileneae, the commissural veins are absent or scarcely evident on the calyx tube in
members of Caryophylleae. Many species are well-known ornamental plants (e.g., carnation: Dianthus spp., baby’s-breath:
Gypsophila L. spp.), used as source of triterpene saponins
(e.g., Acanthophyllum spp.), or known as important weeds
(e.g., Gypsophila spp.). Some species are important components of mountainous steppes (e.g., Acanthophyllum spp.) or
saxicolous vegetation (e.g., Dianthus spp., Gypsophila spp.)
in Europe, Central and Southwest (SW) Asia. The largest
Article history: Received: 12 Feb 2017 | returned for (first) revision: 6 Apr 2017 | (last) revision received: 29 Oct 2017 | accepted: 20 Nov 02017 |
published: online fast track, n/a; in print and online issues, 6 Mar 2018 || Associate Editor: Sigrid Liede-Schumann || © International Association
for Plant Taxonomy (IAPT) 2018, all rights reserved
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83
Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
genera are Dianthus (ca. 300 spp.), Gypsophila (ca. 150 spp.),
Acanthophyllum (ca. 90–100 spp.), Petrorhagia (Ser.) Link
(ca. 33 spp.), and Saponaria L. (ca. 30 spp.).
The monophyly of tribe Caryophylleae and some of its large
genera such as Dianthus and Acanthophyllum are confirmed
TAXON 67 (1) • February 2018: 83–112
by both morphological and molecular evidence (Fior & al.,
2004; Harbaugh & al., 2010; Greenberg & Donoghue, 2011;
Pirani & al., 2014), although some transfers and synonymy are
still necessary to make these genera entirely monophyletic.
Dianthus (including Velezia L.) as the largest genus of the tribe
Fig. 1. (in two parts) Selected members of Caryophylleae in their natural habitat. A & B, Gypsophila leioclada (by H. Madhani); C, Psammophiliella
muralis (by Natalia Gamow); D, Gypsophila acantholimoides (by H. Madhani); E, G. pilosa (by H. Madhani); F, G. desertorum (by Martin
Schnittler in http://floragreif.uni-greifswald.de); G, G. violacea (by I. Khan in http://www.plantarium.ru); (continued on next page)
84
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TAXON 67 (1) • February 2018: 83–112
Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
is monophyletic (Harbaugh & al., 2010; Valente & al. 2010;
Greenberg & Donoghue, 2011) and is characterized by the presence of an epicalyx consisting of discrete bracteoles subtending
the calyx (except Velezia), numerous fine veins on the calyx
tube, and dentate or fimbriate petal apices in most species.
Acanthophyllum, with a predominantly cushion habit and spiny
leaves, should include taxa formerly assigned to Allochrusa
Bunge, Ochotonophila Gilli, Scleranthopsis Rech.f. and part
of Diaphanoptera Rech.f (Pirani & al., 2014). However, the
monophyly of other genera such as Bolanthus (Ser.) Rchb.,
Fig. 1. (continued from previous page) H, Psammosilene tunicoides (by J.T. Johansson in http://angio.bergianska.se); I, Bolanthus fruticulosus
(by Armin Jagel); J, G. aretioides (by H. Madhani); K, Saponaria viscosa (by S. Banquet in http://www.plantarium.ru); L, G. saponarioides
(by H. Madhani); M, Gypsophila montserratii (by A. Gutiérrez & S. Fajarnés); N, Gypsophila bazorganica (by Navid Madhani); O, Petrorhagia
saxifraga (by D. Oreshkin in http://www.plantarium.ru).
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85
Candolle
(1824)
Fenzl
(1840)
Bentham & Hooker
(1862)
Diantheae
Dianthus
subg. Tunica
Velezia
Version of Record
Lychnideae
Cucubalus
Gypsophila
sect. Heterochroa
Lychnis
Saponaria
Silene
Vaccaria
Viscaria
Drypideae
Drypis
Acanthophyllum
Sileneae
Acanthophyllum
Cucubalus
Dianthus
Drypis
Gypsophila
sect. Ankyropetalum
Lychnis
sect. Viscaria
Saponaria
sect. Vaccaria
sect. Agrostema
Silene
Tunica
Uebelinia
Velezia
Pax & Hoffmann
(1934)
Schischkin
(1936)
Šourková
(1978)
Bittrich
(1993)
Harbaugh & al.
(2010)
Diantheae
Dianthus
Tunica
Velezia
Diantheae
Acanthophyllum
Dianthus
Gypsophila
subg. Ankyropetalum
subg. Bolanthus
Phrynella
Saponaria
Tunica
sect. Kohlrauschia
Vaccaria
Velezia
Diantheae
Acanthophyllum
subg. Allochrusa
Dianthus
Gypsophila
Kohlrauschia
Saponaria
Tunica
Vaccaria
Velezia
Diantheae
Dianthinae
Dianthus
Petrorhagia
Velezia
Gypsophilinae
Gypsophila
Phrynella
Saponaria
Vaccaria
Caryophylleae
Acanthophyllum
Allochrusa
Ankyropetalum
Bolanthus
Cyathophylla
Dianthus
Diaphanoptera
Gypsophila
Kohlrauschia
Ochotonophila
Petrorhagia
Phrynella
Pleioneura
Saponaria
Scleranthopsis
Vaccaria
Velezia
Caryophylleae
Acanthophyllum
Allochrusa
Dianthus
Gypsophila
Petrorhagia
Saponaria
Vaccaria
Velezia
Lychnideae
Acanthophyllum
Allochrusa
Ankyropetalum
Cucubalus
Drypis
Githago
Gypsophila
Lychnis
Melandrium
Saponaria
Silene
Tunica
Lychnideae
Cucubalinae
Cucubalus
Drypidinae
Drypis
Sileninae
Heliosperma
Lychnis
Melandrium
Pertrocoptis
Silene
Uebelinia
Viscaria
Lychnideae
Agrostemma
Coronaria
Cucubalus
Lychnis
Melandrium
Petrocoma
Silene
Viscaria
Lychnideae
Agrostemma
Behenantha
Coronaria
Cucubalus
Lychnis
Melandrium
Petrocoma
Petrocoptis
Pleconax
Polyschemone
Schischkiniella
Steris (Viscaria)
Silene
Uebelinia
Sileneae
Agrostemma
Cucubalus
Lychnis
Petrocoptis
Silene
Uebelinia
Sileneae
Agrostemma
Lychnis
Petrocoptis
Silene
Drypideae
Acanthophyllinae
Acanthophyllum
Drypidinae
Drypis
Drypideae
Drypis
Insufficiently
known genus
Psammosilene
TAXON 67 (1) • February 2018: 83–112
Sileneae
Banffya
Cucubalus
Dianthus
Drypis
Gypsophila
sect. Petrorhagia
Lychnis
sect. Viscaria
sect. Agrostema
Saponaria
sect. Bolanthus
sect. Vaccaria
Silene
Velezia
Boissier
(1867)
Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
86
Table 1. Classification history of Caryophylleae (tribes in bold, subtribes underlined) and its allied genera in Sileneae.
TAXON 67 (1) • February 2018: 83–112
Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
Gypsophila, Petrorhagia, Saponaria, and Vaccaria Wolf has
not been adequately addressed (Kadereit & al., 2016) in previous molecular phylogenetic studies (Fior & al., 2006; Harbaugh
& al., 2010; Greenberg & Donoghue, 2011).
The taxonomy of this group has been subject of several
treatments (for a review, see Table 1). The confusion began as
Haller (1742) described Tunica with the names Dianthus and
Caryophyllus below it (probably as synonyms or equivalents).
Without naming any species of Tunica, Ludwig (1757) applied
this name beside Dianthus, making the former a synonym of
the latter. Using the generic name Tunica, replacing Dianthus,
was followed by Scopoli (1771) when he replaced even the
type of Dianthus with a Tunica (as T. caryophyllus (L.) Scop.).
Tunica was later used by several authors (e.g., Boissier, 1867)
as a genus distinct from Dianthus; the genus in this usage is
now treated as a synonym of Petrorhagia (for a detailed bibliography see Ball & Heywood, 1964). In addition, Linnaeus
(1753) could not clearly delimit Gypsophila, Saponaria, and
Tunica; some of the species that he described as members of
Gypsophila are considered today as members of Saponaria,
Petrorhagia, and Arenaria L. There are only four genera of
Caryophylleae (Dianthus, Gypsophila, Saponaria, Velezia) in
its modern definition (sensu Harbaugh & al., 2010) that were
first described by Linnaeus (1753). These genera plus a few
more were assigned to tribe Sileneae by Candolle (1824).
Among the genera currently assigned to Caryophylleae,
six genera were recognized by Fenzl (1840) who classified
them in three different tribes: Diantheae, Lychnideae Fenzl
and Drypideae Fenzl. Bentham (1862) put all these genera
in tribe Sileneae. In Flora Orientalis, Boissier (1867) adopted a system almost similar to Fenzl (1840) but added
the genus Tunica (in this usage, a synonym of Petrorhagia,
see Ball & Heywood, 1964; Rabeler, 1984) to this list and
divided these seven genera between two tribes: Diantheae
and Lychnideae. Pax & Hoffmann (1934) also accepted the
tribe name Diantheae and added the genus Phrynella Pax &
K.Hoffm. to it in Die natürlichen Pflanzenfamilien. In Flora
U.R.S.S. (Schischkin, 1936), the tribe Diantheae encompasses
eight genera also including Acanthophyllum (divided into
two subgenera: subg. Acanthophyllum and subg. Allochrusa
(Bunge) Schischk.) and Kohlrauschia Kunth as well as Tunica
in the sense of Petrorhagia. In other floristic works such as
Flora Europaea (Tutin & al., 1964), Flora of Turkey (Reeve
& al., 1967) and Flora Iranica (Rechinger, 1988), the traditional subfamilial classification system (including three
subfamilies: Illecebroideae Arn. [= Paronychioideae A.St.Hil
ex Fenzl], Minuartioideae DC. [= Alsinoideae Beilschm.],
Caryophylloideae Arn. [= Silenoideae Arn.]) has been applied to these genera, and Caryophylleae is assigned to subfamily Caryophylloideae.
There are relatively few monographs of genera of
Caryophylleae. Bittrich (1993) provides the most recent
comprehensive synopsis of the Caryophylleae, dividing the
tribe into 17 genera. The monographic work on Gypsophila
and its related genera (Barkoudah, 1962) has provided the
primary source for determination of species in Gypsophila
and Bolanthus. The genus Petrorhagia was revised by Ball &
Heywood (1964), with Rabeler (1984) providing additional
nomenclatural clarification. No recent monograph exists for
Dianthus, with the most recent infrageneric classification
appearing in Pax & Hoffman (1934), or for Saponaria, with
the only monograph published by Simmler (1910) and Shults
(1989) providing additional information for taxa occurring
in Russia.
Recent molecular phylogenetic studies on Caryophyllaceae
have indicated the necessity for major changes in the classification of the family and rejected the subfamilial system, which
has commonly been in use (Fior & al., 2006; Harbaugh &
al., 2010; Greenberg & Donoghue, 2011). Today, the family is
divided into 11 tribes including Caryophylleae with Dianthus
caryophyllus L. as type (Harbaugh & al., 2010). Until now, 12
of the Caryophylleae genera accepted by Hernández-Ledesma
& al. (2015) have been included in molecular analyses, but
genera such as Gypsophila, Petrorhagia and Saponaria are
poorly represented.
Gypsophila is one of the most heterogeneous and largest
groups. The genus comprises approximately 150 species of
annual or perennial herbaceous, creeping or cushion-forming
plants, inhabiting primarily the mountainous steppes in the
north temperate part of the Old World with a diversification hotspot in the Irano-Turanian region (Barkoudah, 1962;
Amini & al., 2011). These species also show major variation in inflorescence type ranging from many-flowered lax
thyrses or panicles (e.g., G. elegans M.Bieb., G. paniculata L.,
G. pilosa Huds.) to compact head-like cymes (G. capitata
M.Bieb., G. capituliflora Rupr., G. caricifolia Boiss.), and
few-(uni-)flowered raceme-like monochasia (e.g., G. bazorganica Rech.f., G. saponarioides Bornm. & Gauba). The
morphological differences between Ankyropetalum Fenzl,
Bolanthus, Gypsophila and Phrynella are minor (Table 2) and
their delimitations appear artificial.
The main focus of the present study is to clarify the
generic delimitation in Caryophylleae. The detailed aims
of the study are: (1) to elucidate phylogenetic relationships
and natural groupings in Caryophylleae, (2) to clarify the
limits of the genera in the tribe, (3) to test the monophyly of
Gypsophila and its closely related genera, and (4) to assess
evolutionary progressions for morphological traits applied
in former generic classifications of the tribe Caryophylleae.
MATERIALS AND METHODS
Taxon sampling. — This investigation is based primarily upon specimens deposited in the herbaria: B, G, LE, M,
MSB and TUH. Over 2000 herbarium sheets of the representatives of Caryophylleae have been determined/revised
and studied.
The present study is the largest phylogenetic study on the
tribe in terms of both numbers of genera and species. Since
the tribe itself was proven to be monophyletic in previous
analyses with tribe Eremogoneae Rabeler & W.L.Wagner or
Sileneae as outgroup (Harbaugh & al., 2010; Greenberg &
Donoghue, 2011), we included 12 representatives of the most
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Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
Table 2. Characteristics of genera of Caryophylleae according to Bittrich (1993) and revisions proposed by Pirani & al. (2014).
Gypsophila
Saponaria
Type
G. repens L.
S. officinalis L. A. gypsophiloides Fenzl
Pl. griffithiana V. pyramidata
(Boiss.) Rech.f. Medik.
A. mucronatum
C.A.Mey.
Ph. ortegioides
Pax & K.Hoffm.
Chromosome
number
2n = 24, 26, 28,
30, 34, 36, 48,
51, 60, 68
2n = 28, 56
–
2n = 24, 30
2n = 26, 30,
60, 90
–
No. of species
ca. 150
ca. 30
4
1
1
ca. 90–100
Habit
Annual and
perennial, sometimes densely
caespitose
Perennial,
rarely annual
herbs
Perennial,
densely caespitose to densely
pulvinate
Perennial
herbs with
thick rhizome
(geophyte)
Annual glabrous Small, shrubby,
herbs
cushion-forming
perennials with
spiny leaves, or
rarely perennial
herbs
Perennial herb
with woody
caudex, rigid
stem,
Seed shape
Reniform
GlobularReniform to
Reniformreniform,
almost globular globose, with
marginal hilum, compressed
with granular
wrinkles
Reniform or
globose
Obovoidreniform
Oblong, commashaped, with a
marginal hilum
Inflorescence
type/position
Paniculate or
head-like cyme
Lax cymes
Lax or dense
paniculate or
capitate cymes,
rarely solitary
Panicle cymes
Inflorescence
terminal, regularly dichasial,
lax, richly
branched
Dichasial cymes
arranged in
terminal heads
and/or axillary
verticillasters,
sometimes
paniculate or
corymbose
Dichasial inflorescence very
often reduced to
solitary sessile
flowers
Calyx
Hemispherical,
Cylindrical,
campanulate, tur- obscurely
15–25-veined
binate, or rarely
tubular, 5-nerved
with broad scarious commissures,
5-toothed
Tubular,
15-nerved,
5-toothed
Ovoid with
5 prominent
angles
Tubularturbinate,
5–15-nerved,
5-toothed
Long-campanulate, up to about
1/3 divided in
teeth, 5-costate,
the ribs alternating with hyaline
intervals
Embryo
Peripheral;
(Curved) pe(Circular) peripheral, central endosperm
ripheral, central
central
endosperm, with endosperm
prominent radicle
Curved
Curved
Unciform or
strongly curved
Hook-shaped,
with a straight
prominent radicle
Petals
Entire or slightly
emarginate
Mostly disClawed, limb
tinctly clawed, 3(–5)-partite
coronal scales
usually present
Pink, with narrowly winged
claw, limb
entire, coronal
scales present
5, pink or
purple, clawed,
limb entire, dentate or notched,
coronal scales
absent
Limb entire,
Cuneate, with
sometimes bifid, a bare claw and
gradually taper- without corona
ing into claw
Capsule
Always exceeding the calyx,
dehiscence
Opening by
4(6) teeth
Dehiscing from
base by irregular longitudinal
slits
Capsule opening by 4 teeth
1- or 2-seeded,
With papery
exocarp opening mode of dehisby 4 teeth, encence various
docarp dehiscing irregularly
Long-ovoid,
opening with 4
fissures extending
to the middle,
with 1–3 seeds
Ovules number
4–36
2–14
8–10
12
Numerous
4(–21)
8
Special features
–
Seeds usually
smooth and
shiny
Gynophore
absent, ovary
more or less
sessile
Coronal scales
present
Seeds shiny
Bracts and
leaves mostly
spiny
Calyx with one
to four pairs of
bracteoles at base
(epicalyx)
Distribution
Temperate Eurasia, E Mediterranean, and
Irano-Turanian
region, Egypt,
Arabia, Somalia,
Australia
Temperate
Eurasia,
chiefly in
Mediterranean
and IranoTuranian
region
Turkey, Iran,
Iraq, Lebanon,
Syria, Palestine
From C Asia to C, E & S
Europe; Asia
W Himalaya;
Afghanistan
C, W & SW
Asia, Siberia
S & C Turkey
88
Ankyropetalum
2n = 26
Campanulatetubular,
5-nerved with
5 small teeth
Pleioneura
Version of Record
Vaccaria
Acanthophyllum Phrynella
1
TAXON 67 (1) • February 2018: 83–112
Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
Tabel 2. Continued.
Bolanthus
Petrorhagia
Velezia
Dianthus
Cyathophylla
Type
B. hirsutus
(Labill.)
Barkoudah
P. saxifraga
(L.) Link
V. rigida L.
D. caryophyllus L.
P. tunicoides
C. chlorifolia
(Poir.) Bocquet W.C.Wu &
C.Y.Wu
& Strid
D. khorasanica
Rech.f.
Chromosome
number
2n = 20, 30
2n = 26, 28,
30, 60
2n = 28
2n = 30, 60, 90
2n = 30
–
No. of species
ca. 18
ca. 33
2
ca. 300
Habit
Perennial, puAnnual and
berulent to hirsute, perennial
herbs
grey-green to
velvety herbs, with
thin short stems
and small leaves
Seed shape
Comma-shaped,
compressed on
both sides, with
flat or channelled
back; testa with
small tubercles;
hilum marginal
Peltate with
facial hilum
Psammosilene Diaphanoptera
2n = 28
1
1
6
Annual herbs
Perennial
puberulent
herbs
Perennial and
tufted herbs,
woody at base
Peltate with
facial hilum
Roundish,
reddish brown
to blackish,
Peltate
Reniform
Solitary or
capitate terminal
cymes often subtended by bracts
Terminal heads Terminal
cymes
subtended by
bowl-shaped,
entire involucre
Lax few-flowered cymes
Tubular, usually
(5–)15-nerved,
without scarious
commissures,
1–3 bracts in
form of epicalyx
present
Tubular with
many parallel
veins, without
membranous
commissures,
5-toothed
Cylindrical
with 5 short
teeth, without
scarious commissures and
commissural
veins
Tubular,
15-nerved,
5-toothed
Turbinate or
vase-shaped
Straight
Straight
Curved
Straight
Curved
Annual or perenAnnual, rigid,
dichotomously nial herbs, rarely
branching herbs subshrubs
Peltate with
facial hilum
Inflorescence type/ Contracted dichaposition
sial cymes
Solitary or
Panicles or
capitate cymes monochasial
cymes
Calyx
Tubiform, pentagonal, turbinate
at base, with small
teeth and with 5
projecting ribs
alternating with
comparatively
broad hyaline
bands
Cylindrical or
campanulate,
5-toothed,
5–15-nerved,
with scarious
commissures
Embryo
Hook-shaped, with Straight
long prominent
radicle, peripheral
Petal
Entire, cuneate,
with small limb
and winged claw
5, entire or
bifid, clawed
or not, coronal
scales absent
5(10), longclawed
with small
2–4-toothed or
emarginated
limb, coronal
scales absent
Linear oblong,
White, pink
inconspicuous
or red, rarely
yellow, limb
entire, toothed or
fimbriate, without
coronal scales,
claw long, with
two longitudinal
ridges
5, purpleviolet, inconspicuously
clawed, entire
5, rose or
violet, entire or
emarginate
Capsule
Longer than sepals, with 4 teeth
Opening by
4 teeth
4 teeth splitting
into 2 valves
Opening by
4 teeth
Membranous,
probably
indehiscent,
1-seeded
4-valved
Ovules number
8–20
Numerous
Few
Numerous
Unknown
2
(6–)8–19
Special features
Seeds smooth and
shiny
Leaves narSepal hardened
row, grass-like at the apex and
margin
Epicalyx scales
two to many
Bowl-shaped
involucre
subtending
inflorescence
Nearly nonsplitting,
1-seeded and
membranous
in texture
Inflated calyx,
winged at ribs,
caespitose
woody cushions
Distribution
Greece, S Turkey,
Syria, Lebanon,
Palestine
Europe, from
Mediterranean
to Kashmir,
and on Canary
Islands
Sino-Himalaya,
and from the
Mediterranean
east to
Afghanistan
Europe, Asia,
especially
Mediterranean,
Africa
Mountains of
Greece and
Turkey
China, prov.
Yunnan
High altitude
of NE Iran,
Turkmenistan,
and Afghanistan
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Opening by
4 teeth
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Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
closely related tribes to Caryophylleae (i.e., Alsineae Lam.
& DC., Arenarieae Kitt., Eremogoneae and Sileneae) in our
analyses in order to keep the effect of sequence homoplasy
on tree reconstruction at minimum. In general, six species
of different clades known in Silene L. (Sileneae), one species
each of Cerastium L. and Stellaria L. (Alsineae), two species of Eremogone Fenzl (Eremogoneae), one species each
of Arenaria and Moehringia L. (Arenarieae) were selected
as outgroups according to the results of previous molecular
phylogenetic studies (Harbaugh & al., 2010; Greenberg &
Donoghue, 2011). Within the tribe, we tried to perform a
balanced sampling representing almost all major morphological lineages known to us based on personal experience
and examining pertinent literature. Therefore, the present
study allows us to define the genera in Caryophylleae more
appropriately, matching morphological, geographical and
molecular phylogenetic evidence.
We produced a dataset of nrDNA ITS with 136 (59 new
sequences and 77 obtained from GenBank) accessions representing 112 species, and a dataset of cpDNA rps16 with
94 (58 new sequences and 36 obtained from GenBank) accessions representing 85 species. Dissatified with the poor
sampling of Gypsophila in previous molecular phylogenetic
studies, we focused more intensively on this genus, and
generated sequences for 44 and 34 species of this genus for
nrDNA ITS and cpDNA rps16, respectively. For this purpose
we attempted to sample all subgenera and sections recognized in Barkoudah (1962). Representatives of four formerly
recognized genera of Caryophylleae, i.e., Ankyropetalum,
Bolanthus, Cyathophylla Bocquet & Strid, and Phrynella as
well as the new genera described in this paper, i.e. Balkana
gen. nov., Graecobolanthus gen. nov., were sequenced in
this study for the first time. The voucher specimens for the
sequences generated in this study (Appendix 1) are deposited in B, M, MSB and/or TUH.
DNA extraction, amplification and sequencing. — The
present study was performed based on two molecular markers, one nuclear: the internal trascribed spacer (ITS) region
of the ribosomal cistron (consisting of ITS1, the intervening
5.8S gene, and ITS2) and one plastid intron: rps16. DNA was
extracted from dried leaf material using a NucleoSpin Plant
DNA extraction kit (Macherey-Nagel, Düren, Germany) according to the manufacturers’ protocol. Amplification of the
ITS region was performed using the primer pair Leu1 (Vargas
& al., 1998) and ITS4 (White & al., 1990). In some difficult
cases ITS2 and ITS3 were used, as described by White & al.
(1990). For the plastid region (complete intron rps16), we used
the primers rpsF and rpsR2R or rpsF and rpsR3R (Oxelman &
al., 1997; Petri & Oxelman, 2011; Kool & al., 2012). All PCR
amplifications were performed in a Thermocycler T-Personal
48 (Biometra, Göttingen, Germany), Primus 96 plus (MWG:
Biotech, Ebersberg, Germany), or 2720 (Applied Biosystems,
Carlsbad, California, U.S.A.).
Cycle sequencing was done using BigDye Terminator
v.3.1, Cycle Sequencing Kit (Applied Biosystems). DNA
samples were sequenced with ABI3730 DNA Analyser 48well capillary sequencer (Applied Biosystems).
90
TAXON 67 (1) • February 2018: 83–112
Alignment and tree reconstruction. — We edited the
sequences with Geneious v.8.0.5 (Kearse & al., 2012), and
conducted the multiple alignment using MAFFT v.7 with
default parameters (Katoh & Standley, 2013). The alignments
were manually corrected using Mesquite v.3.02 (Maddison
& Maddison, 2011).
Alignments of the present phylogenetic datasets are available
as supplementary material (https://doi.org/10.12705/671.6.S).
The beginning and end of the alignments, where the sequences
of many individuals lack sharp electropherogram peaks, were
trimmed prior to analysis. The ITS and rps16 datasets were
analyzed separately as their taxon composition differed. Two
kinds of analyses were run on each dataset separately: Bayesian
inference (BI) and maximum parsimony (MP). Before running BI, the optimal substitution models were estimated using
the Akaike information criterion (AIC) in jModelTest v.0.1.1
(Posada, 2008). General time reversible model with gammashaped rate variation and a proportion of invariable sites
(GTR + I + Γ) was estimated as the best-fit model for both ITS
and rps16 markers. For BI we used MrBayes v.3.2.6 (Ronquist
& Huelsenbeck, 2003) under CIPRES server (Miller & al.,
2010) with the number of MCMC generations for ITS and rps16
datasets set to 40 million. Trees were sampled every 1000
generations with the default of three “heated” and one “cold”
chain, and pre-stationarity MCMC samples were discarded as
burn-in (2500 samples as calculated by Tracer v.1.6 software,
Rambault & al., 2014). The remaining trees were summarized
in a 50% majority-rule consensus tree for each dataset. MP
analyses were performed using PAUP* v.4.0b10 (Swofford,
2003) with the following parameters: all characters unordered
and equally weighted, heuristic search with random sequence
addition, tree-bisection-reconnection branch swapping, 50
random-addition-sequence replicates, and MAXTREES option
set to 10,000. The obtained trees were summarized in a strict
consensus tree. Bootstrapping was done using maximum likelihood method as implemented in RAxML-HPC2 on XSEDE v.8
(Stamatakis, 2014) the following settings: model = GTRCAT,
bootstrap nreps = 1000 (summarized in a 50% majority-rule
consensus tree as a cladogram).
Morphological character mapping. — A data matrix of
four morphological characters for all taxa included in the
phylogenetic analysis of ITS was prepared. The selected characters were: (1) membranous commissures of calyx (present/
absent), (2) bracteoles (present/absent), (3) seed shape (reniform or pyriform/reniform-oblong/peltate/comma shaped),
and (4) embryo shape (curved/straight/hook-shaped). The evolutionary pathways were reconstructed using Mesquite v.3.02
(Maddison & Maddison, 2011). We employed the Markov
k-state 1 (Mk1) parameter model of evolution for the ML
reconstructions, with equal probability for any particular
character change.
Typification information. — Details about type specimens
of the basionyms of the new combinations and resurrected
names that we have included are based on examining protologues and searching major indices (Tropicos, http://www.
tropicos.org/; JSTOR Global Plants, https://plants.jstor.org), as
well as websites of several individual herbaria (BM, BR, E,
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TAXON 67 (1) • February 2018: 83–112
Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
G, GH, K, KEW, L, LINN, OS, P, US, WU) for extant specimens. We have examined a digital image from one (or more)
of these sources for any specimen where we cite a barcode
in the type citations.
RESULTS
The alignment of the ITS dataset needed introduction
of ten large indels (≥ 5 bp). The final ITS matrix for 135
terminals comprised a total of 700 characters, whereas the
rps16 dataset with 94 terminals was finally 1034 bp long and
included 29 large indels. A list of alignment characteristics
and parsimony statistics is presented in Table 3. The results
of MP and BI for both datasets were congruent. Therefore, we
present and discuss only the 50% majority-rule trees obtained
from BI of the ITS (Fig. 2) and rps16 (Fig. 3). For a better
comparison of the obtained trees and checking for congruency between the two datasets, the mirror image of obtained
trees is shown in Fig. 4.
All trees are congruent in showing tribe Caryophylleae
as monophyletic with Silene (in rps16 trees) or Eremogone (in
ITS trees) as sister group. We recognize three main clades in
both trees which are called Gypsophilinae, Saponaria s.str.
and Caryophyllinae. In the ITS tree (Fig. 2), Psammosilene
W.C.Wu & C.Y.Wu, missing in the rps16 tree, is sister to these
three. The largest clade in the tribe in terms of species number
is Caryophyllineae and can be subdivided into four clades:
(1) Acanthophyllum s.l. (incl. Allochrusa, Diaphanoptera);
(2) Cyathophylla (including Cyathophylla spp., Saponaria
viscosa C.A.Mey. and a few species placed in Gypsophila by
Barkoudah, 1962); (3) Dianthus s.l. (including Bolanthus spp.,
Phrynella ortegioides (Fisch. & C.A.Mey.) Pax & K.Hoffm.,
Gypsophila confertifolia, Petrorhagia spp., Psammophiliella
spp., G. spergulifolia Griseb., Dianthus spp., and Velezia
rigida); and (4) Petroana (including Gypsophila montana and
G. montserratii). Gypsophilinae and Saponaria s.str. show a
sister relationship in the rps16 tree, and their clade is in turn
sister to Caryophyllinae, while Saponaria s.str. forms the
sister to the Caryophyllinae in the ITS topology (Fig. 2). The
Gypsophilinae clade embraces most species of Gypsophila
including the type of the genus (G. repens L.) along with
the representatives of the genera Vaccaria, Ankyropetalum,
and Bolbosaponaria Bondarenko as well as one species of
Diaphanoptera (D. afghanica Podlech).
some smaller genera and little knowledge of the morphology
of many crucial taxa are the main reasons for this uncertainty.
Since many ornamental and medicinal plants are representatives of this tribe, and due to frequent occurrence of the
members of this tribe in natural ecosystems in SW Asia,
providing the correct taxonomic name to these taxa is very
important, though challenging.
The general topology of the trees obtained here supports
the presence of three major clades in Caryophylleae, fitting partly with the subtribal system proposed by Šourková
(1978). Our phylogenies suggest adding two subtribes to
subtribes Dianthinae Šourková (homotypic synonym of
Caryophyl linae (Juss.) Rabeler & Bittrich, see Rabeler
& Bittrich, 1993) and Gypsophilinae Šourková, namely
subtribes Saponariinae and Psammosileninae. However,
as three of the four subtribes would each include only one
genus, this system seems unnecessary. Therefore, we prefer
to name not all clades formally as shown on the trees (Figs.
2–4). According to the results we obtained, we provide
below a detailed survey on the delimitation of genera in
tribe Caryophylleae.
Gypsophila. — As circumscribed currently, this is a
heterogeneous group morphologically (Bittrich, 1993). Our
study shows that a major revision is necessary to make this
genus monophyletic. All analyses conducted here show
clearly that the generic names Ankyropetalum, Bolbosaponaria (already included in the genus, see Bittrich, 1993),
Dichoglottis Fisch. & C.A.Mey. (already included in the
genus, see Barkoudah, 1962), Vaccaria and one species of
Diaphanoptera (D. afghanica) are nested within Gypsophila.
In each of these cases, the morphological diagnostic features
are shared by certain species assigned to Gypsophila. In the
case of Ankyropetalum the deeply incised petals as well as
exserted stamens (in addition to other features mentioned
by Barkoudah, 1962) might represent autapomorphies. The
membranous commissures and the winged veins of the calyx
as well as its inflated form in Vaccaria are also evident in
Diaphanoptera khorasanica Rech.f. and partly in D. afghanica as well as Bolbosaponaria bucharica (B.Fedtsch.)
Bondarenko (also nested within Gypsophila in our analyses). The possible inclusion of Vaccaria in Gypsophila has
been suggested before (Kadereit & al., 2016). The genus
Diaphanoptera is clearly polyphyletic with three species
Table 3. Alignment characteristics and statistics of phylogenetic analyses of rps16 and ITS datasets in Caryophylleae.
DISCUSSION
rps16
Despite many molecular phylogenetic studies in Caryophyllaceae during recent years (Fior & al., 2006; Harbaugh
& al., 2010; Greenberg & Donoghue, 2011; Pirani & al., 2014;
Dillenberger & Kadereit, 2014; Sadeghian & al., 2015) the
generic boundaries in the tribe Caryophylleae are still blurred
(Pirani & al., 2014; Hernández-Ledesma & al., 2015). The
scanty sampling of some large genera in the tribe, such as
Gypsophila and Saponaria, as well as missing sequences of
ITS
Number of terminals
94
135
Sequence length [bp]
414–810
242–648
Aligned length [bp]
1034
700
Constant characters [bp]
541
261
Parsimony-uninformative characters [bp]
158
82
Parsimony-informative characters [bp]
335
357
Parsimony-informative characters [%]
32.4
51
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91
A
Cyathophylla clade
Dianthus s.l.
Petroana clade
Saponaria s.str.
1
96/100
Petrorhagia
Velezia
1
100/100
Dianthus
Dianthus
Petrorhagia
Petrorhagia
Graecobolanthus
Balkana
Psammophiliella
Bolanthus
Gypsophila
Psammophiliella
Phrynella
Bolanthus
Bolanthus
Gypsophila
Petroana
Gypsophila
Cyathophylla
Saponaria
Heterochroa
Gypsophila
Diaphanoptera
Gypsophila
Gypsophila
Acanthophyllum
Acanthophyllum
Diaphanoptera
Acanthophyllum s.l.
Caryophyllinae
1
100/100
1
90/97
Allochrusa
Saponaria
Saponaria
Pleioneura
TAXON 67 (1) • February 2018: 83–112
Petrorhagia armerioides
Velezia rigida 1
V. rigida 2
Dianthus cyri
D. deltoides
1
1
D. armeria
96/96
100/100
1 0.96 D. chinensis
D. carthusianorum
0.95
D. andrzejowskianus
Petrorhagia dubia
1
0.98 100/100
P. prolifera
1
P. saxifraga
0.85
P. thessala
Bolanthus graecus
Gypsophila spergulifolia
Psammophiliella muralis 1
1
1
P. muralis 2
100/100
Phrynella ortegioides
100/100
Bolanthus minuartioides
B. cherlerioides
1
1
B. huber-morathii
84/89
100/99
Gypsophila confertifolia
Gypsophila montana
1
G. montserratii 2
1
99/100
G. montserratii 1
89/Saponaria viscosa
1
Gypsophila desertorum 2
1
1
98/100
100/100 G. desertorum 1
100/100
G. violacea
Diaphanoptera ekbergii
0.99
G. herniarioides
-/77
1
G. cerastioides 2
-/75
1
100/100 G. cerastioides 1
Acanthophyllum allochrusoides
A. grandiflorum
G. myriantha
1
G. honigbergeri
-/79
A.
scapiflorum
1
A. laxiflorum
-/76
A. aphananthum
A. spinosum
Diaphanoptera stenocalycina
1
1
-/79 90/90 D. lindbergii
1
A. kandaharicum
100/100
1
A. stocksianum
-/77 1
A. kabulicum
99/99 -/96
A. borsczowii
Allochrusa bungei 1
1
Al. bungei 2
100/100
Al. versicolor
0.96
79/96
A. sordidum
1
A. oppositiflorum
84/92 1
A. mucronatum
92/88
Saponaria ocymoides 1
1
1 100/100
S. ocymoides 2
0.99 84/99 S. pumila
-/89
S.
officinalis
1
1
0.87
100/100 S. officinalis 2
1
Pleioneura griffithiana
100/100
S. prostrata
1
S. glutinosa
81/96
0.77
Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
Version of Record
Fig. 2. Majority-rule consensus tree inferred from Bayesian analysis of ITS data in tribe Caryophylleae. Numbers above the branches indicate
posterior probability values; those below branches are MP/ML bootstrap values. The generic names inside the grey boxes are those accepted
in the present study. Species names are according to www.ipni.org and do not follow the taxonomic treatments suggested in the present study.
The generic names in the column right to the grey area are those accepted by Bittrich (1993), Pirani & al. (2014), and Hernández-Ledesma
& al. (2015). Values below 0.5 for posterior probability and below 50% for ML/MP bootstrap are not shown.
92
Accepted genera based on
Accepted genera
Bittrich (1993); Pirani & al. (2014);
based on present study Hernández-Ledesma & al. (2015)
Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
Psammosilene
Ankyropetalum
Vaccaria
Gypsophilinae
0.93
1
93/98
A
outgroup
0.99
-/70
0.8
G. elegans 1
0.97
G. aretioides
0.79
G. antari
G. nabelekii
G. linearifolia
0.98
G. pilosa 1
1
0.97
G. pilosa 2
100/100
1
G. capillaris 2
G. capillaris 1
99/99
G. petraea
G. fastigiata 2
0.94
G. fastigiata 1
0.99
G. cephalotes 1
G. cephalotes2
0.75
G.
repens
1
0.97
G. repens 2
G. patrinii 2
G. pacifica
0.84
G. capituliflora 2
G. capituliflora 1
G. patrinii 1
0.94
G. acutifolia
G. scorzonerifolia
G. stevenii
Gypsophila
G.
globulosa
0.91
G. capitata
G. bermejoi
G. glomerata
G. uralensis
G. leioclada
G. aucheri 2
0.96
1
G. aucheri 1
-/77
G. curvifolia 2
1
G. curvifolia 1
1
G. virgata
100/100
G. laricina
1 G. pinifolia 2
1
1 G. pinifolia 1
82/94
G. pilulifera
G. venusta
Diaphanoptera afghanica
1
100/100 1 G. bucharica 2
1
99/100 G. bucharica 1
G. tomentosa
-/92 1
98/99 1 G. perfoliata
95/98 G. oblanceolata
G. arrostii
1
G. paniculata
94/98 1
G. bicolor
83/93
0.99 Vaccaria hispanica 3
1
94/88
V. hispanica 1
100/100
V. hispanica 2
1
G. heteropoda 2
92/93
G. heteropoda 1
78/1
0.94
Ankyropetalum gypsophiloides
100/100
G. sp.
G. viscosa
Psammosilene tunicoides
Psammosilene
Gypsophila elegans 3
0.96
G. arabica
1
G. silenoides
97/86 G. acantholimoides
1
100/100
Bolbosaponaria
aberrant taxa. Such a treatment has already been applied
to G. muralis L. and three related taxa now considered as
Psammophiliella Ikonn. (Hernández-Ledesma & al., 2015:
based on results obtained by Greenberg & Donoghue, 2011,
and Pirani & al., 2014). Our results show that Gypsophila violacea (Ledeb.) Fenzl, G. desertorum (Bunge) Fenzl, G. montserratii Fern.Casas, G. montana Balf.f., G. spergulifolia,
G. confertifolia Hub.-Mor., G. cerastioides D.Don, G. herniarioides Boiss., G. honigbergeri (Fenzl) Boiss. and G. myriantha
Rech.f. should also be transferred to other genera.
The last four species listed above are placed in the
Acanthophyllum s.l. clade. Gypsophila cerastioides is neither spiny nor shows the typical many-flowered rich axillary
inflorescences known in most species of Acanthophyllum.
Gypsophila
that are woody at base nested within Acanthophyllum (see
also Pirani & al., 2014) and one species in Gypsophila. In
line with previous analyses, the genus Diaphanoptera cannot be maintained and should be reduced to a synonym
under Acanthophyllum (Pirani & al., unpub. data). It seems
that the unique tuberous roots and long tubular calyces
with membranous commissures between the calyx veins
in Bolbosaponaria are not applicable at generic rank, but
may be useful for separating lower ranks such as sections.
However, the subgeneric classification of Gypsophila needs
a richer sampling and is not addressed here, but it is included
in the scope of our future investigations.
Another important finding of our study relating to
Gypsophila is the necessity to exclude some morphologically
Diaphanoptera
TAXON 67 (1) • February 2018: 83–112
Caryophylleae
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Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
The broad leaves, many-ovulate capsules and large petals in
this species indicate rather its unique position that led some
earlier taxonomists to assign it to the genus Acosmia Benth.
(or Timaeosia Klotzsch). In the ITS tree presented here, this
species is closely related to Diaphanoptera ekbergii Hedge
& Wendelbo, and G. herniarioides.
In line with our molecular phylogenetic results, the
woody cushion-forming habit, the well-developed short lateral branches in G. herniarioides resulting in a false verticillate phyllotaxy as well as dense and many-flowered corymbose inflorescences, and 4- to 12-ovulate ovary fit well
with Acanthophyllum, but are absent or rarely observed in
Gypsophila. However, the Acanthophyllum s.l. clade includes
some other unarmed plants such as members of Allochrusa,
and therefore, recognition of neither G. herniarioides nor
Allochrusa spp. under separate genera are supported by molecular data (see also Pirani & al., 2014). Our extended taxon
sampling (especially regarding the number of genera sampled)
for both markers, does not allow Allochrusa to be separated
from Acanthophyllum. The species of the genus Allochrusa
were considered once as members of Acanthophyllum subg.
Allochrusa (Schischkin, 1936) and our molecular phylogenetic
studies corroborate the taxonomic treatment performed by
Pirani & al. (2014) and contradict the treatment by HernándezLedesma & al. (2015) where it was recognized provisionally
at the generic level. According to this concept, it is necessary
to resurrect the generic name Acanthophyllum for some taxa
treated as Allochrusa in recent taxonomic surveys (see under
Taxonomic implications).
Gypsophila honigbergeri is spiny (at least at leaf apex)
and shows characteristic capsules of Acanthophyllum (low
ovule number, one-seeded), and was suggested previously to
be a member of this genus (Barkoudah, 1962). Aggregation
of some characters, i.e., cylindrical calyx, shortly exserted
petals and long stamens, which are unusual characters in
Gypsophila, associated with a cushion-forming habit resembling Acanthophyllum drove Rechinger (1988) to describe the
distinct subgenus Kabulianthe under Gypsophila. Ikonnikov
(2004) even elevated this taxon to generic rank and introduced
the combination Kabulianthe honigbergeri (Fenzl) Ikonn.
Gypsophila myriantha is also spiny and similar to Acanthophyllum in general habit except for the open paniculate inflorescence that is similar to many species of Gypsophila.
The capsules in this species have not been seen, but the
number of ovules is lower than 12, associating it again with
Acanthophyllum. Our molecular phylogenetic investigations
clearly place this species in Acanthophyllum.
Gypsophila desertorum and G. violacea form a distinct
clade sister to Acanthophyllum s.l. + Dianthus s.l. clades. Both
species are representatives of Gypsophila sect. Heterochroa
TAXON 67 (1) • February 2018: 83–112
(Bunge) A.Braun (for notes on correct name and typification
of this taxon see Rabeler, 1993) and show a south to east
Siberian distribution. Among the species Barkoudah originally assigned to G. sect. Heterochroa, G. cerastioides and
G. hernarioides, should be transferred to Acanthophyllum and
are restricted to Afghanistan and Pakistan (see above), while
other species show a more northern distribution reaching Far
East and Central Asia. This latter group including G. antoninae Schischk., G. desertorum, G. microphylla (Schrenk)
Fenzl, G. sericea (Ser.) Krylov (≡ Heterochroa petrea Bunge:
type of the genus Heterochroa), G. turkestanica Schischk.
and G. violacea are low herbaceous plants with short internodes, small leaves, short pedicels and short campanulate
calyces with calyx teeth mostly scarious at margins, and lack
calcium oxalate crystals (which are present in most species
of Gypsophila: Barkoudah, 1962). Our phylogenetic results,
in accordance with geographical distribution patterns and
morphology, suggest resurrection of the genus Heterochroa
Bunge similar to Gypsophila sect. Heterochroa as defined
by Barkoudah (1962) but excluding G. cerastioides, G. hernarioides and G. honigbergeri that should be transferred to
Acanthophyllum. No material of G. glandulosa (Boiss.) Walp.
was available to us; a species distributed in Turkey showing
intermediate morphological characters between Heterochroa,
Gypsophila and Acanthophyllum.
Gypsophila montserratii and G. montana (≡ Saponaria
montana (Balf.f.) Barkoudah) are characterized by a very
small calyx and petals. The haploid base chromosome number
is known only for the former (x = 13: Löve, 1973), which is
neither congruent with Gypsophila (mostly x = 17) nor with
Dianthus (x = 15). They form a highly supported clade sister
to the Dianthus s.l. clade in the ITS and rps16 trees (Figs.
2, 3). Like Miller & Cope (1996), who treated G. montana
within Gypsophila, we were not able to trace any important
morphological character that supported Barkoudah (1962)
transferring G. montana to Saponaria, but the morphological
resemblance of this taxon to G. montserratii is strong enough
to correlate it with the latter. These two species are geographically remote from each other: G. montserratii is known from
the Iberian Peninsula, and G. montana is distributed in mountains and wadi-beds of Socotra (an archipelago in Arabian
Sea, territory of Yemen) (Miller & Cope, 1996). In line with
our molecular phylogenetic analyses and morphological evidence, we suggest describing a new genus, Petroana, for this
group (see below under Taxonomic implications).
Saponaria. — Seven species of Saponaria are included in
our analyses, most of which form a clade in both analyses that
includes Pleioneura griffithiana (Boiss.) Rech.f. Although the
Saponaria clade is fully supported in the analyses of both markers, its placement varies between the markers used (see under
Fig. 3. Majority-rule consensus tree inferred from Bayesian analysis of rps16 data in tribe Caryophylleae. Numbers above the branches
indicate posterior probability values; those below branches are MP/ML bootstrap values. Species names are according to www.ipni.org
and do not follow the taxonomic treatments suggested in the present study. The generic names inside the grey boxes are those accepted
in the present study. The generic names in the column right to the grey area are those accepted by Bittrich (1993), Pirani & al. (2014), and
Hernández-Ledesma & al. (2015). Values below 0.5 for posterior probability and below 50 for ML/MP bootstrap are not shown.
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Cyathophylla clade
Petroana clade
Dianthus s.l.
Petrorhagia alpina
1
1
74/82
P. candica
0.98 91/94
P. armerioides
1
Dianthus
Dianthus carthusianorum
D. armeria
1 89/93
76/91 0.82
P. saxifraga
Petrorhagia
0.99
Bolanthus fruticulosus
1
Graecobolanthus
-/76
B.
graecus
100/100
Psammophiliella muralis
Psammophiliella
1
Phrynella ortegioides
91/94
1
B.
cherlerioides
Bolanthus
0.85 100/100
0.86
B. huber-morathii
-/92
-/75
Balkana
Gypsophila spergulifolia
1
Gypsophila montserratii
Petroana
G. montana
100/100
1
Cyathophylla chlorifolia Cyathophylla
1 86/91
Saponaria viscosa
88/94
Gypsophila desertorum
Heterochroa
Diaphanoptera ekbergii 2
1
D. ekbergii1
0.99
Acanthophyllum laxiflorum
0.94
A. scapiflorum
G. honigbergeri
Diaphanoptera lindbergii
1
D. stenocalycina 2
99/100
D. stenocalycina 1
A. allochrusoides 2
A. allochrusoides 1
A. spinosum
Acanthophyllum
1
A. kabulicum
90 /91
A. borsczowii
A. stocksianum
A. kandaharicum
A. grandiflorum
1
A. aphananthum 2
99/100
A. aphananthum 1
Gypsophila cerastioides
A. oppositiflorum
Allochrusa bungei 2
1
Al. bungei 1
99/99
Al. versicolor
A. mucronatum 2
A. mucronatum 1
A. sordidum
Gypsophila antari
G. capillaris
G. linearifolia
G. paniculata
G. aucheri
G. patrinii
G. capituliflora
G. cephalotes
G. fastigiata
G. repens
G. pinifolia
G. petraea
G. arrostii
G. acutifolia
G. curvifolia
Gypsophila
G. oblanceolata
G. capitata
G. bucharica
1
G. viscosa
100/100
Ankyropetalum gypsophiloides
G. perfoliata
G. tomentosa
G. elegans 2
G. elegans 3
G. venusta
0.95
G. laricina
1
G. leioclada
88/98
G. bicolor
G. acantholimoides
G. nabelekii
G. pilosa
Saponaria sicula
S. ocymoides 1
1
S. ocymoides 2
Saponaria
100/100
S. officinalis 1
S. officinalis 2
S. prostrata
Pleioneura griffithiana
outgroup
Petrorhagia
Dianthus
Petrorhagia
Bolanthus
Psammophiliella
Phrynella
Bolanthus
Gypsophila
Gypsophila
Cyathophylla
Gypsophila
Diaphanoptera
Diaphanoptera
Acanthophyllum
Gypsophila
Allochrusa
Gypsophila
Bolbosaponaria
Ankyropetalum
Saponaria s.str.
1
94/100
Gypsophilinae
Caryophylleae
Acanthophyllum s.l.
Caryophyllinae
Accepted genera based on
Accepted genera
Bittrich (1993); Pirani & al. (2014);
based on present study Hernández-Ledesma & al. (2015)
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Pleioneura
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Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
Results). Our results show clearly that Pleioneura is nested
within Saponaria in spite of the unique stigma (positioned
just at the end of the style) and the membranous commissures
between the calyx veins. In contrast to Bittrich (1993) and
Hernández-Ledesma & al. (2015), who recognized Pleioneura,
we suggest its synonymy within Saponaria.
Among the species currently assigned to Saponaria,
S. viscosa is nested within the Cyathophylla clade (Fig. 3).
Unfortunately, the ITS sequence for Cyathophylla chlorifolia Bocquet & Strid could not be generated, but the rps16 sequences put C. chlorifolia and S. viscosa together clearly with
full support. The perfoliate leaves and congested inflorescence
in Cyathophylla are the most important morphological features
separating it from Saponaria. However, the pedicels in S. viscosa are relatively short, so that the young inflorescences do
show a congested form. In general, only the ovate to rounded
leaves in C. chlorifolia, which look perfoliate at the base, can
be considered as important characters separating these taxa.
Furthermore, C. chlorifolia is found in Greece and Turkey,
whereas S. viscosa is distributed in eastern Turkey, Azerbaijan,
Iraq, Iran and Turkmenistan. The distribution of these species
overlaps somewhat in Ankara Province in Turkey.
Dianthus s.l. — Bolanthus, Dianthus, Petrorhagia, Phrynella, Psammophiliella, and Velezia are the main representatives of the clade Dianthus s.l.; our study is the first molecular investigation including genera Bolanthus and Phrynella.
Furthermore, we extended the sampling of Petrorhagia, supporting its paraphyly as previously indicated but based on a
different grouping of sampled taxa (Greenberg & Donoghue,
2011). The relatively rich sampling of Dianthus in previous
analyses (Valente & al., 2010; Greenberg & Donoghue, 2011)
indicated that Dianthus was monophyletic with Velezia nested
within (Harbaugh & al., 2010; Kemler & al., 2013). Despite
low sampling here, our analyses also suggest inclusion of
Velezia as well as a few species of Petrorhagia (excluding its
type: P. saxifraga (L.) Link) in Dianthus.
Petrorhagia. — Morphologically, species of Petrorhagia
either have conspicuous bracts encircling the calyx (including the type) or such bracts are missing. Our analyses clearly
suggest including this latter group in Dianthus; most of these
species have in the past been placed within Gypsophila
(Grisebach, 1843).
In both trees (Figs. 2, 3) the Dianthus s.l. clade is divided into five main lineages with unresolved or moderately
to low-supported relationships: (1) Bolanthus core group
including some species of this genus as well as Gypsophila
confertifolia and Phrynella; (2) Psammophiliella muralis; (3)
G. spergulifolia; (4) the core group of Petrorhagia (including
its type) along with the second group of Bolanthus (B. graecus, B. fruticulosus); and (5) Dianthus, Velezia and the second
TAXON 67 (1) • February 2018: 83–112
group of Petrorhagia (including P. armerioides, P. alpina and
P. candica).
The findings of our analyses indicate that a group of species currently assigned to Petrorhagia sect. Pseudotunica
(Fenzl) Post & Kuntze and sect. Pseudogypsophila (A.Braun)
P.Ball & Heywood, which are characterized by the absence of
conspicuous epicalyx bracts, are also associated with Dianthus
and form a common clade with Velezia. Since Dianthus sect.
Armeriastrum Ser. forms the most basally branching clade of
the genus (as currently circumscribed, see Valente & al., 2010)
and the clade composed of Velezia rigida and part of P. sect.
Pseudotunica are sister to the Dianthus crown group, it seems
that it is inevitable to extend the formerly suggested lumping
approach to place P. sect. Pseudotunica in Dianthus. However,
as the type of this section was not available to us, the formal
synonymy of this section should wait for further investigations,
but the few species of this section analyzed here are transferred
to Dianthus (see under Taxonomic implications).
Bolanthus. — Bolanthus fruticulosus and B. graecus are
both distributed in Greece (Strid, 1986) slightly disjunct from
most species of the genus, which is known from Turkey and
Middle East. The representatives of the genus from both areas
are mostly saxicolous plants with showy petals rounded at the
apex as well as conspicuous calyx veins and narrow commissural membranes. However, the petals in the Greek species turn
abruptly downwards and become clearly deflexed shortly after
anthesis, while in species from Turkey and the Middle East the
petals might be recurved, but not abruptly deflexed. Our results
agree with this morphological characteristic and geographical
data, suggesting a new genus for the Greek representatives
formerly assigned to Bolanthus is needed (see Graecobolanthus
under Taxonomic implications).
Gypsophila confertifolia, which shows an overlapping
distribution pattern with the main group of Bolanthus centered in the East-Mediterranean phytogeographic subregion
(sensu Eig, 1931), is characterized by a tubular calyx and
short pedicels, which give rise to capitate inflorescences, resembling some species of Bolanthus such as B. minuartioides
(Jaub. & Spach) Hub.-Mor. The main difference between this
species and members of Bolanthus is its annual habit. Our
ITS trees (Fig. 2) suggest that this species is more closely
related to Bolanthus rather than Gypsophila or, as suggested by
Ikonnikov (1976), Psammophiliella. This assumption is also
supported by geographical and morphological data. Based
on available data, G. confertifolia should be transferred to
Bolanthus.
A similar situation involves the obligate serpentinophyte
taxon (Marin & Tatić, 2001; Jakovljević & al., 2011) G. spergulifolia. Morphologically this species is characterized by a basal
aggregate of linear leaves, which are triangular in cross-section,
Fig. 4. Tanglegram of Dendroscope program package (Huson & Scornavacca, 2012) comparing the phylogenies of ITS (left side) and rps16
datasets (right side) based on the 50% majority-rule consensus trees obtained from Bayesian phylogenetic analyses. Numbers above the
branches indicate posterior probability values; those below branches are MP/ML bootstrap values. Values below 0.5 for posterior probability and below 50% for ML/MP bootstrap are not shown. Grey boxes represent same genera of Caryophylleae as mentioned in the key
at the bottom right of the figure.
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1
100/100
Caryophyllinae
1
100/100
1
84/89
1
100/99
1
99/100
1
98/100
Cyathophylla clade
Acanthophyllum s.l.
1
79
1
100/100
0.96
79/96
1
100/100
Gypsophilinae
1
93/98
Caryophyllinae
0.85
Dianthus s.l.
0.95
Acanthophyllum s.l.
Dianthus s.l.
1
96/96
1
96/100
Petrorhagia armerioides
Petrorhagia candica
Velezia rigida
P. alpina
V. rigida
Dianthus cyri
P. armerioides
1
D. deltoides
Dianthus carthusianorum
D. chinensis
89/93
D. armeria
D. armeria
D. carthusianorum
Petrorhagia saxifraga
0.82
D. andrzejowskianus
0.99
Bolanthus fruticulosus
Petrorhagia dubia
-/76
B. graecus
P. prolifera
1
P. saxifraga
Psammophiliella muralis
P. thessala
91/94
Phrynella ortegioides
Bolanthus graecus
Bolanthus
cherlerioides
Gypsophila spergulifolia
0.85
0.86
Psammophiliella muralis
B. huber-morathii
-/92
-/75
P. muralis
Gypsophila
spergulifolia
Phrynella ortegioides
1
G. montserratii
Bolanthus minuartioides
B. cherlerioides
G. montana
100/100
B. huber-morathii
Cyathophylla chlorifolia
Gypsophila confertifolia
1
Saponaria viscosa
G. montana
88/94
G. montserratii
G. desertorum
G. montserratii
Cyathophylla clade
Diaphanoptera ekbergii
Saponaria viscosa
D. ekbergii
Gypsophila desertorum
G. desertorum
Acanthophyllum laxiflorum
G. violacea
A. scapiflorum
Diaphanoptera ekbergii
A. honigbergeri
G. herniarioides
G. cerastioides
Diaphanoptera lindbergii
1
G. cerastioides
Di. stenocalycina
Acanthophyllum allochrusoides
99/100
Di.
stenocalycina
A. grandiflorum
A. allochrusoides
G. myriantha
A. honigbergeri
A. allochrusoides
A. scapiflorum
A. spinosum
A. laxiflorum
1
A. kabulicum
A. aphananthum
A. spinosum
90/91
A. borsczowii
Diaphanoptera stenocalycina
A. stocksianum
D. lindbergii
A. kandaharicum
A. kandaharicum
A. stocksianum
A. grandiflorum
A. kabulicum
1
A. aphananthum
A. borsczowii
99/100
A. aphananthum
Allochrusa bungei
Al. versicolor
G. cerastioides
Al. bungei
A. oppositiflorum
A. sordidum
Allochrusa
bungei
A. oppositiflorum
1
A. mucronatum
Al. bungei
Saponaria ocymoides
99/99
Al.
versicolor
S. ocymoides
A. mucronatum
S. pumila
S. officinalis
A. mucronatum
S. officinalis
A. sordidum
Pleioneura griffithiana
G. antari
S. prostrata
S. glutinosa
G. capillaris
Gypsophila elegans
G. linearifolia
G. arabica
G. paniculata
G. silenoides
G. acantholimoides
G. aucheri
G. elegans
G. patrinii
G. aretioides
G. capituliflora
G. antari
G. nabelekii
G. cephalotes
G. linearifolia
G. fastigiata
G. pilosa
G. repens
G. pilosa
G. capillaris
G. pinifolia
G. capillaris
G. petraea
G. petraea
G. arrostii
G. fastigiata
G. acutifolia
G. fastigiata
G. cephalotes
G. curvifolia
G. cephalotes
G. oblanceolata
G. repens
G. capitata
G. repens
G. patrinii
G. bucharica
G. pacifica
1
G. viscosa
G. capituliflora
100/100
Ankyropetalum gypsophiloides
G. capituliflora
G. patrinii
G. perfoliata
G. acutifolia
G. tomentosa
G. scorzonerifolia
G. elegans
G. capitata
G. stevenii
G. elegans
G. globulosa
G. venusta
G. bermejoi
G. laricina
G. glomerata
1
G. uralensis
G. leioclada
88/98
G. leioclada
G.
bicolor
G. aucheri
G. acantholimoides
G. aucheri
G. curvifolia
G. nabelekii
G. curvifolia
G. pilosa
G. virgata
Saponaria
sicula
G. laricina
G. pinifolia
S. ocymoides
1
G. pinifolia
S. ocymoides
G. pilulifera
100/100
S. officinalis
G. venusta
Diaphanoptera afghanica
S. officinalis
G. bucharica
S. prostrata
G. bucharica
Pleioneura griffithiana
G. tomentosa
G. perfoliata
G. oblanceolata
Graecobolanthus
Psammophiliella
Acanthophyllum
G. arrostii
G. paniculata
Gypsophila
Psammosilene
Balkana
G. bicolor
Vaccaria hispanica
V. hispanica
Bolanthus
Heterochroa
Saponaria
V. hispanica
G. heteropoda
Cyathophylla
Petroana
G. heteropoda
Ankyropetalum gypsophiloides
G. sp.
Dianthus
Petrorhagia
G. viscosa
Psammosilene tunicoides
rps16
ITS
1
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Gypsophilinae
0.75
1
100/100
0.94
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relatively long pedicels and a short campanulate-turbinate
calyx. Furthermore, the petals are uniquely bicolored, red on the
outer surface, but with a white or pink inner surface. It shows
also an isolated geographic distribution, endemic to the Balkan
region (in Albania and Serbia). Our results suggest recognizing
this taxon as distinct from both Bolanthus and Gypsophila (see
Balkana under Taxonomic implications), although the commissural membranes of the calyx known in both genera occur also
A
TAXON 67 (1) • February 2018: 83–112
in this taxon. In our phylogenetic trees this species is placed
more closely to Bolanthus rather than to Gypsophila.
Phylogenetic analyses of both markers clearly place
Phrynella within Bolanthus. Morphological and geographical data also support this relationship. The monotypic genus
Phrynella is known only from Turkey, where several species of Bolanthus are also found, sometimes in similar regions (Reeve & al., 1967). Short internodes, 1- to 3-flowered
B
e
s
mo
am
m
am
oup
Ps
Dian
Dian
Bracteole
absent
present
thus
Membranous
Commissures
absent
present
thus
outgr
o
outgr
up
Ps
ne
ile
en
il
os
Pet
rorh
agi
Ba Gr
a
lkaaec
Ps n ob
a
am
ola
nth
m
op
us
hi
lie
lla
Pet
rorh
C
a
an
tro
lla
Pe
phy
tho
Cya
hroa
eroc
Het
a
an
tro
lla
Pe
phy
tho
Cya
hroa
eroc
Het
agi
Ba Gr
a
l a
Ps kan ecob
a
am
ola
nth
m
op
us
hi
lie
lla
D
a
an
tro
lla
Pe
phy
tho
Cya
hroa
eroc
Het
a
an
tro
lla
Pe
phy
tho
Cya
hroa
eroc
Het
Pet
rorh
agi
Ba Gr
a
a
lka ec
Ps n ob
am a ol
an
m
thu
op
s
hi
lie
lla
Dian
Embryo shape
curved
straight
hook-shaped
thus
outgr
thus
Dian
Seed shape
reniform
reniform-oblong
peltate
comma-shaped
oup
a
Ps
outgr
P
o
mm
e
en
sil
oup
sa
e
en
sil
o
mm
Pet
rorh
agi
Ba Gr
a
lka aec
Ps n ob
am a ol
an
m
thu
op
s
hi
lie
lla
Fig. 5A–D. Evolutionary histories of four selected morphological characters mapped on Bayesian semi-strict consensus tree of nrITS sequences
in Caryophylleae. The characters and their states are mentioned in the center of each circular tree.
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axillary inflorescences, sessile flowers, and petals slightly
overtopping the calyx are shared by Phrynella ortegioides
and several species of Bolanthus. Our results are partly in
accordance with Barkoudah’s (1962) morphological concept,
associating Phrynella with B. minuartioides. However, he
transferred the latter to Acanthophyllum mainly due to its rigid
habit, linear parallel-nerved leaves, sessile axillary and terminal flowers and unequal stamens (Barkoudah, 1962). The
analyses presented here, despite showing a close relationship
between Ph. ortegioides and B. minuartioides, rejects their
affinity with Acanthophyllum.
Evolution of selected morphological characters. — Mapping of morphological characters on the consensus ITS tree
shows that most of the diagnostic traits used formerly in tribe
Caryophylleae are homoplasious and not useful for defining
the boundaries between the genera (Fig. 5A–D). Therefore,
from a taxonomic point of view, it is inevitable that we apply
a combination of characters for defining the genera in this
group, noting that only a few genera might show no overlap
in these characters. It appears as if membranous commissures between the sepals evolved in basal branches of the ITS
tree (Fig. 5A), suggesting a reversal of this character in some
species or species groups in Acanthophyllum and Dianthus.
However, in Psammosilene, which occupies a basal position
in Caryophylleae, commissural membranes are absent, and
the membranous parts of the calyx are confined to tooth margins. Seed shape (Fig. 5C) in the members of Caryophylleae
is quite variable and, therefore, we determined four states for
this character. Peltate, dorsiventrally compressed seeds, which
have been considered as a major synapomorphy for Dianthus,
are shared by Petrorhagia, Psammosilene and Velezia, among
which Psammosilene is not closely related to Dianthus. Within
the Dianthus s.l. clade the peltate shape is the most common
state, but the reniform type (as in Gypsophila spergulifolia =
Balkana spergulifolia (Griseb.) Madhani & Zarre: see below
under Taxonomic implications) and the comma-shaped type (as
in Bolanthus spp.) represent some cases of homoplasy regarding this character state. Ball & Heywood (1964) considered
the peltate seeds as a feature separating the genera Dianthus,
Petrorhagia and Velezia from all other members of Silenoideae,
but recent studies reported this seed type also in Psammosilene
(Bittrich, 1993). In Acanthophyllum s.l. and Heterochroa, the
common seed shape is the reniform-oblong, which can be considered as intermediate between reniform and peltate types. A
detailed seed micromorphological investigation in the Dianthus
s.l. clade, like those already conducted on Gypsophila (Amini &
al., 2011), Velezia (Poyraz & Ataşlar, 2010) and Acanthophyllum
(Pirani & al., in prep.), will shed light on the evolutionary pathways of this character.
The curved embryo is the most common type in the basally
branching clades of Caryophylleae except Psammosilene, which
has a straight embryo, but the straight and hook-shaped ones are
most common in members of the Dianthus s.l. clade suggesting a derived position for this state (Fig. 5D). Embryo shape is
otherwise correlated with seed shape, so that reniform/reniformoblong, peltate, and comma-shaped seeds show curved, straight,
and hook-shaped embryos, respectively.
Another important morphological character addressed in
the present study is the presence/absences of bracteoles (Fig.
5B). Presence of bracteoles in Allochrusa, Diaphanoptera,
Ochotonophila and Scleranthopsis, reflects their phylogenetic position nested within Acanthophyllum. However, absence of bracteoles in other taxa of Acanthophyllum s.l. clade
(Gypsophila cerastioides, G. hernarioides, G. honigbergeri,
G. myriantha), suggests at least two reversals for this character in this clade.
TAXONOMIC IMPLICATIONS
The molecular phylogenetic analyses performed here, together with morphological and geographic evidence, suggest a
new generic concept in tribe Caryophylleae. It is necessary to
transfer some species to other genera or describe new genera
in order to make them natural and monophyletic. Based on this
new concept, we provide a diagnostic key to all genera we currently recognize in the tribe and then brief descriptions of each
genus (most based on Bittrich, 1993 and modified as necessary).
New combinations are made only when material of the taxa was
available to us for analysis. Therefore, for many taxa we did not
propose any taxonomic change due to lack of specimens for
detailed herbarium or molecular examination.
Diagnostic key to genera of Caryophylleae. — In many
genera there are a few species showing some deviation from
the typical variation exhibited within the genus. In most cases,
it is necessary to use a combination of characters in order to
determine the genus with certainty.
1. Seeds peltate, with central (facial) hilum; embryo
straight ...................................................... 2
1. Seeds reniform, reniform-oblong or comma-shaped, with
lateral hilum; embryo curved or hook-shaped ........ 5
2. Leaves with short petiole, ovate; stamens 5; capsules
membranous, nearly indehiscent ........ Psammosilene
2. Leaves sessile, linear, subulate, grass-like; stamens (5)10;
capsules papery, dehiscent ............................... 3
3. Calyx without membranous commissures, with 35 or
more veins, rarely 5- to 15-nerved (cf. Velezia); calyx tube
long tubular, teeth straight ...... Dianthus (incl. Velezia)
3. Calyx with membranous commissures, with 5‒15 veins;
calyx tube variously shaped, if tubular the teeth recurved
to deflexed ................................................... 4
4. Seeds > 1.5 mm, with thin margin, smooth on surface
.......................... Dianthus (incl. Petrorhagia p.p.)
4. Seeds < 1.5 mm, with thickened margin, reticulate on
surface .......................................... Petrorhagia
5. Seeds comma-shaped (or oblong), with hook-shaped
embryo ........................................................ 6
5. Seeds reniform to reniform-oblong, embryo curved .. 7
6. Petals turning abruptly downward and becoming clearly
deflexed (Greece) ....................... Graecobolanthus
6. Petals recurved gradually (Turkey to the coastal mountains of Syria, Lebanon and Palestine) ...................
................................. Bolanthus (incl. Phrynella)
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Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
7.
7.
8.
8.
9.
9.
10.
10.
11.
11.
12.
12.
13.
13.
14.
14.
15.
15.
16.
16.
17.
17.
18.
100
Calyx bladdery inflated, or turbinate, constricted at teeth,
commissural regions membranous hyaline, sometimes
wing-like ................................... Diaphanoptera
Calyx campanulate to tubular, if inflated, commissural regions papery or leafy and main veins with leafy
wings ......................................................... 8
Bracteoles present, leafy, papery or rarely membranous;
calyx papery in texture or only membranous at intervals
.............................................. Acanthophyllum
(incl. Allochrusa, Ochotonophila, Scleranthopsis)
Bracteoles absent ........................................... 9
Calyx bladdery inflated, nerves prominent and thick,
costate, or winged, midveins 5; bracteoles membranous
hyaline .......................... Gypsophila (cf. Vaccaria)
Calyx tubular, campanulate, or obconical, not much inflated, lateral nerves obscure, not prominent and thick,
midveins 5 or more; bracteoles absent ................ 10
Calyx obscurely nerved or with 15–25 nerves, commissures absent or present; petals inconspicuous, or clawed,
mostly with appendages .................................. 11
Calyx 5-nerved, with membranous commissures; petals
not or only indistinctly clawed, without appendages ..12
Plants annual; inflorescences congested; capsule slightly
longer than the calyx; coronal scales absent ............
.................................................. Cyathophylla
Plants annual, biennial or perennial; inflorescences usually lax; capsule mostly shorter than the calyx; coronal
scales mostly present ............................ Saponaria
Leaves fleshy, spathulate; flowers very small: calyx < 4mm,
corolla < 5 mm; seed testa with swollen cells tuberculate
on periclinal wall, testa cells polygonal-oblong, moderately elongated (Iberian Peninsula, Socotra) .. Petroana
Leaves not fleshy or subfleshy, linear to ovate; flowers small or large; seed testa variously shaped, with or
without tubercles ......................................... 13
Petals bicolored, red on the outer surface, white or pink
on the inner surface; leaves triquetrous, mostly 3 or 4 at
each node (Albania, Serbia, Bosnia) ........... Balkana
Petals always concolored, variously colored; leaves slender, in few species triquetrous, then the plants mostly
caespitose, paired at nodes .............................. 14
The stigmatic surface terminal; ovules less than 24 .. 15
The stigmatic surface extending along the inner side of
styles; ovules 24‒36 .......................................20
Stem nodes with small lateral shoots in leaf axils giving a verticillate appearance; leaves acerose, spiny, or
terminating to a spine .................. Acanthophyllum
Lateral shoots in leaf axils absent; leaves not spiny except
in Gypsophila acantholimoides and G. pinifolia ....... 16
Capsules shorter than the calyx ......................... 17
Capsules exceeding the calyx ........................... 18
Plants annual, shorter than 10 cm, covered by long glandular hairs ...................... Bolanthus confertifolius
Plants perennial, if annual then taller than 10 cm and
glandular hairs absent or short ................ Gypsophila
Plants perennial; capsules ± indehiscent .................
...................... Acanthophyllum (cf. A. cerastioides)
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18. Plants annual or perennial; capsules dehiscent ....... 19
19. Calyx without membranous commissural intervals or
with very narrow ones, calcium oxalate crystals absent;
stamens shorter than the petals ............. Heterochroa
19. Calyx with membranous commissural intervals encompassing calcium oxalate crystals; stamens longer (or
sometimes shorter) than petals ......................... 20
20. Annual plants with fibrous roots, puberulent below and
glabrous in inflorescence (subcosmopolitan, absent in
Australia and New Zealand) .......... Psammophiliella
20. Annual or perennial plants with tap root, variously
hairy ............................................. Gypsophila
Accepted genera and taxonomic changes
1. Acanthophyllum C.A.Mey., Verz. Pfl. Casp. Meer.: 210.
1831 – Type (designated by Schiman-Czeika in Rechinger,
Fl. Iranica 163: 274. 1988): Acanthophyllum mucronatum
C.A.Mey.
= Timaeosia Klotzsch in Klotzsch & Garcke, Bot. Ergebn.
Reise Waldemar: 138, t. 33. 1862 – Type: T. cerastioides
(D.Don) Klotzsch (≡ Acanthophyllum cerastioides (D.Don.)
Madhani & Zarre).
= Allochrusa Bunge ex Boiss., Fl. Orient. 1: 559. 1867 ≡
Acanthophyllum subg. Allochrusa (Bunge) Schischk., Fl.
URSS 6: 608. 1936 ≡ Acanthophyllum sect. Allochrusa
(Bunge ex Boiss.) Pirani & Rabeler in Phytotaxa 303(2):
198. 2017 – Type (designated by J.J. Swart, ING Card
13030, 1 Apr 1961): Allochrusa versicolor (Fisch. &
C.A.Mey.) Boiss. (≡ Acanthophyllum versicolor Fisch.
& C.A.Mey.).
= Ochotonophila Gilli in Feddes Repert. Spec. Nov. Regni Veg.
59: 169. 1956 ≡ Acanthophyllum sect. Ochotonophila (Gilli)
Pirani in Taxon 63(3): 604. 2014 – Type: O. allochrusoides
Gilli (≡ Acanthophyllum alluchrusoides (Gilli) Pirani).
= Kuhitangia Ovcz. in Dokl. Akad. Nauk Tadzhiksk. S.S.R.
10: 50. 1967 – Type: K. popovii (Preobr.) Ovcz. (≡
Acanthophyllum popovii (Preobr.) Barkoudah).
= Scleranthopsis Rech.f. in Ann. Naturhist. Mus. Wien 70: 37.
1967 – Type: S. aphanantha (Rech.f.) Rech.f. (≡ Acanthophyllum aphananthum Rech.f.).
= Kabulianthe (Rech.f.) Ikonn. in Bot. Žhurn. (Moscow
& Leningrad) 89(1): 114. 2004 ≡ Gypsophila subg.
Kabulianthe Rech.f., Fl. Iranica 163: 244. 1988 – Type:
K. honigbergeri (Fenzl) Ikonn. (≡ Acanthophyllum honigbergeri (Fenzl) Barkoudah).
Diagnosis. – Most of the species are spinose and cushionlike, often bracteate and bracteolate, and oblong- reniform seeds.
Description. – Small shrubby, tufted perennial plants,
rarely perennial herbs (e.g., Acanthophyllum cerastioides, see
below); leaves subulate, acerose, spiny, spring leaves herbaceous, or rarely thinly herbaceous with mostly spinulose apex;
flowers often in fragile, ± dense, globose heads, or lax panicles
(sect. Allochrusa) and rarely solitary; calyx tubular-turbinate,
or rarely campanulate to campanulate-tubular, 5- to 15-nerved,
5-toothed, sometimes with narrow membranous commissures;
petals 5, white, pink, rose or lilac, limbs entire, rarely bifid;
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stamens 10; styles 2; ovary with 4–12(–20) ovules, on a short
gynophore; capsule often with 1–2 seeds and opening in various ways; seeds oblong-reniform, or rarely reniform; embryo
curved (2n = 26, 30, 60, 90; species: ca. 90–100.
Distribution. – SW and Central Asia, one species in China.
Habitat. – Acanthophyllum grows in subalpine steppes,
on gravelly, sandy or stony hills and rocky slopes.
Resurrected names
Acanthophyllum bungei (Boiss.) Trautv. in Trudy Imp. S.Peterburgsk. Bot. Sada 2: 511. 1873 ≡ Allochrusa bungei
Boiss., Fl. Orient. 1: 560. 1867 – Holotype: Iran, East
Azerbaijan, inter Marand to Jolfa, 1859, A. Bunge s.n. (G
barcode G00150411!; isotypes: K barcode K000725639!,
W No. W 1988-0009970!).
Acanthophyllum gypsophiloides Regel, Descr. Pl. Nov. Rar.
Fedtsch.: 15. 1882 ≡ Allochrusa gypsophiloides (Regel)
Schischk. in Trudy Bot. Inst. Akad. Nauk S.S.S.R., Ser. 1,
Fl. Sist. Vyssh. Rast. 4: 306. 1937 – Type: [Central Asia,
Kazakhstan], Kara-Tau, A. Fedtschenko s.n. (LE).
Acanthophyllum honigbergeri (Fenzl) Barkoudah in Wentia
9: 182. 1962 ≡ Silene honigbergeri Fenzl in Endlicher &
Fenzl, Sert. Cabul.: 3. 1836 ≡ Gypsophila honigbergeri
(Fenzl) Boiss., Fl. Orient. 1: 558. 1867 ≡ Kabulianthe
honigbergeri (Fenzl) Ikonn. in Bot. Zhurn. (Moscow &
Leningrad) 89(1): 114. 2004 – Holotype: [Afghanistan]
E montibus prope Kabul, J.M. Honigberger s.n. (W No.
W 0048192!; possible isotypes: W No. W 0048249 [not
imaged], W No. W 1914-0006602 [not imaged]).
= Gypsophila galiifolia Gilli in Feddes Repert. Spec. Nov. Regni
Veg. 59: 165. 1956 – Lectotype (designated by Rechinger,
Fl. Iranica 163: 245. 1988): [Afghanistan] near Kabul, A.
Gilli 1264 (W No. W 1969-0001338!).
Acanthophyllum paniculatum Regel & Herder in Bull. Soc.
Imp. Naturalistes Moscou 39(1): 539. 1866 ≡ Allochrusa
paniculata (Regel & Herder) Ovcz. & Czukav. in Ovczinnikov, Fl. Tadzhiksk. S.S.R. 3: 611. 1968 – Holotype:
Central Asia, [Kazakhstan, Dzhungarian Alatau] die Hügel
in der Nähe des Piquet Sary-bulka am Füsse des Alatau
der sieben Flüsse, 2500 ft., ?.1857, C.L. Semenov s.n. (LE).
Acanthophyllum tadshikistanicum (Schischk.) Schischk.,
Fl. URSS 6: 801. 1936 ≡ Allochrusa tadshikistanica
Schischk. in Trudy Bot. Muz. 24: 40. 1932 – Type:
Tadjikistan, (Buchara olim), in itinere Kizil-su et Sarai, in
declivitatibus montanis ad ripam dextram flum. Kizil-su,
D. Divnogorskaja s.n. (LE).
Acanthophyllum transhyrcanum Preobr. in Bot. Mater. Gerb.
Glavn. Bot. Sada R.S.F.S.R. 1(3): 1. 1920 ≡ Allochrusa
transhyrcana (Preobr.) Czerep., Sosud. Rast. S.S.S.R.: 154.
1981 ≡ Diaphanoptera transhyrcana (Preobr.) Rech.f. &
Schiman-Czeika in Rechinger, Fl. Iranica 163: 335. 1988
– Holotype: Turkmenistan: Mulla-Kara, in deserto prope
Balchan, 1889, A. Antonow s.n. (LE; isotype: W No.
W 1986-0005948!).
Acanthophyllum versicolor Fisch. & C.A.Mey. in Index
Seminum (St. Petersburg [Petropolitanus]) 4: 31. 1838
≡ Allochrusa versicolor (Fisch. & C.A.Mey.) Boiss., Fl.
Orient. 1: 559. 1867 – Holotype: [Azerbaidzhan, Armenia],
in locis lapidosis aridissimis desertisque salsis provinciae
Nakitschiwan, J.N. Szovits s.n. (LE; possible isotype: US
barcode 00289322!).
New combinations
Acanthophyllum cerastioides (D.Don) Madhani & Zarre, comb.
nov. ≡ Gypsophila cerastioides D.Don, Prodr. Fl. Nepal.:
213. 1825 ≡ Timaeosia cerastioides (D.Don) Klotzsch in
Klotzsch & Garcke, Bot. Ergebn. Reise Waldemar: 138.
1862 – Holotype: [Nepal], Gosaingsthan, 1980–3900 m,
1829, M. Wallich Cat. 644 (K barcode K000725774!; isotypes: E barcode E00301689, G barcode G00226904!).
Acanthophyllum herniarioides (Boiss.) Madhani & Zarre,
comb. nov. ≡ Gypsophila herniarioides Boiss., Fl. Orient.,
Suppl.: 84. 1888 – Holotype: Afghanistan, Kurrum valley,
Sikaram, 7 Aug 1879, J.E. Aitchison 961 (G; isotype: K
barcode K000725698!).
Acanthophyllum myrianthum (Rech.f.) Madhani & A.Pirani,
comb. nov. ≡ Gypsophila myriantha Rech.f. in Anz.
Österr. Akad. Wiss., Math.-Naturwiss. Kl. 105: 11. 1969
– Holotype: Afghanistan, Deh Kundi, in declivibus saxosis aridis (Tonschiefer) 10 km W Shahrestan, 33°40′ N,
66°35′ E, versus Deh Kundi, 34°10′ N, 66°07′ E, 2200 m,
2 Jul 1967, K.H. Rechinger 36812 (W No. W 1969-0013845!;
isotypes; B barcode B 10 0365629!, E barcode E00301855!,
G barcode G00226867!, K barcode K000725739!, LE barcode LE 00012091!, MO barcode MO-176943!, S No. S-G8666!, US barcode 00103460!).
Acanthophyllum persicum (Boiss.) A.Pirani & Rabeler, comb.
nov. ≡ Saponaria persica Boiss., Diagn. Pl. Orient., ser. 1,
1: 18. 1843 ≡ Allochrusa persica (Boiss.) Boiss., Fl. Orient.
1: 560. 1867 – Lectotype (designated here) [following
annotation on herbarium sheet by Schiman-Czeika, 1982]:
[Iran] in lapidosis circa Tabriz, P.M.R. Aucher-Eloy 4242
(G [herb. Boissier] barcode G00226480!; isolectotypes: G
barcode G00226470!, K barcode K000725640!).
Acanthophyllum sedifolium (Kurz) Madhani & Zarre, comb.
nov. ≡ Gypsophila sedifolia Kurz in Flora 55: 285. 1872 –
Holotype: [India] Kashmir; Zanskar, 12–13,000 ft., 2 Jul
1848, T. Thomson s.n. (B, destroyed) – Lectotype (designated here): (K barcode K000725777!).
Acanthophyllum takhtajanii (Gabrieljan & Dittrich) A.Pirani &
Rabeler, comb. nov. ≡ Allochrusa takhtajanii Gabrieljan &
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Dittrich in Biol. Žhurn. Armenii 43: 184. 1990 – Holotype:
Armenia, Ararat Distr., lower part of Uztz mountains near
Surenavan, 800 m, 20 Jun 1986, E.T. Gabrieljan & K.G.
Tamanian s.n. (ERE barcode ERE0000046!; isotype: G
barcode G00226486!).
2. Balkana Madhani & Zarre, gen. nov. – Type: Balkana spergulifolia (Griseb.) Madhani & Zarre.
Diagnosis. – This monotypic genus differs from all other
genera of Caryophylleae by its verticillate phyllotaxy at least
at some nodes, as well as leaves distinctly swollen and connate at base.
Description. – Perennial herbs; leaves linear, triquetrous,
(2–)3‒4 at each node; flowers in terminal panicles composed of
dichasial partial inflorescences; pedicel longer than the calyx;
calyx campanulate-turbinate with 5 veins and membranous
commissures; petals 5, bicolored, outer surface red, inner surface white or pink; stamens 10; styles 2; ovules ca. 16; capsule
opening by 4 teeth; seeds reniform with small flat tubercles;
embryo curved; monotypic.
Etymology. – Balkana is named after the general distribution of the plants which is centered in Balkan Peninsula.
Distribution. – The genus is a Mediterranean element distributed in the inner and western part of the Balkan Peninsula:
Albania, Serbia, Bosnia-Herzegovina.
Habitat. – These plants are thermophilous, growing on
limestone or serpentine soils.
Included species
Balkana spergulifolia (Griseb.) Madhani & Zarre, comb. nov.
≡ Gypsophila spergulifolia Griseb., Spic. Fl. Rumel. 1:
183. 1843, non G. spergulifolia (Jaub. & Spach) Boiss.,
Fl. Orient. 1: 559. 1867 – Holotype: W Albania, Mt. Puka
near Alessia, A.H.R. Griesbach s.n. (GOET barcode
GOET005978!; isotype: K; possible isotype: M barcode
M-0242533!).
3. Bolanthus (Ser.) Rchb., Deut. Bot. Herb.-Buch: 205. 1841 ≡
Saponaria sect. Bolanthus Ser. in Candolle, Prodr. 1: 366.
1824 ≡ Gypsophila sect. Bolanthus (Ser.) Boiss., Fl. Orient.
1: 537. 1867 – Type (designated by Barkoudah in Wentia 9:
168. 1962): B. hirsutus (Labill.) Barkoudah.
= Phrynella Pax & K.Hoffm. in Engler & Prantl, Nat. Pflanzenfam., ed. 2, 16c: 364. 1934 ≡ Phryna (Boiss.) Pax & K.Hoffm.
in Engler & Prantl, Nat. Pflanzenfam., ed. 2, 16c: 351. 1934,
non Phryna Bubani 1901, syn. nov. – Type: P. ortegioides
(Fisch. & C.A.Mey.) Pax & K.Hoffm. (≡ Bolanthus ortegioides (Fisch. & C.A.Mey.) Madhani & Rabeler).
Diagnosis. – Seeds in the members of this genus (except
B. confertifolius) are comma-shaped and the embryo is hookshaped. The projecting veins on the tubular calyx give a pentagonal shape to the calyx in these plants. Unlike its related
genus Graecobolanthus, the petals in members of Bolanthus
are recurved gradually, not abruptly deflexed.
Description. – Perennials, low prostrate or cushion-forming, or rarely annual (B. confertifolius), plants hairy; leaves
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small and linear; flowers small, in paniclulate to subcapitate
dichasia or solitary; calyx tubular-turbinate with 5 projecting veins and membranous commissures; petals 5, recurved
gradually, white or pink with purple veins; stamens 10; styles 2;
ovary on a short gynophore; capsule opening by 4 teeth; seeds
comma-shaped with prominent radicle; embryo hook-shaped;
species ca. 10.
Etymology. – From the Greek bolosi: lump + nugget +
anthos: flower.
Distribution. – The genus includes East Mediterranean
elements, reaching higher elevations in western parts of the
Mediterranean from Turkey to Syria, Lebanon and Palestine.
Habitat. – These plants mostly grow in crevices of calcareous rock and on stony slopes.
New combinations
Bolanthus confertifolius (Hub.-Mor.) Madhani & Heubl,
comb. nov. ≡ Gypsophila confertifolia Hub.-Mor. in Feddes
Repert. Spec. Nov. Regni Veg. 52: 42. 1943 – Holotype:
[Turkey, C2] Muğla: Muğla-Fethiye, 141 km SE of Muğla,
7 Jun 1938, J. Reese s.n. (G barcode G00006010! [ex Hb.
Huber-Morath]; isotype: BASBG).
Bolanthus ortegioides (Fisch. & C.A.Mey.) Madhani &
Rabeler, comb. nov. ≡ Tunica ortegioides Fisch. & C.A.
Mey. in Ann. Sci. Nat., Bot., sér. 4, 1: 36. 1854 (“artegioides”) ≡ Saponaria ortegioides (Fisch. & C.A.Mey.)
Boiss. & Balansa in Boissier, Diagn. Pl. Orient., ser. 2, 6:
25. 1859 ≡ Gypsophila ortegioides (Fisch. & C.A.Mey.)
Boiss., Fl. Orient. 1: 552. 1867 ≡ Phryna ortegioides (Fisch.
& C.A.Mey.) Pax & K.Hoffm. in Engler & Prantl, Nat.
Pflanzenfam., ed. 2, 16c: 351. 1934 ≡ Phrynella ortegioides (Fisch. & C.A.Mey.) Pax & K.Hoffm. in Engler &
Prantl, Nat. Pflanzenfam., ed. 2, 16c: 364. 1934 – Holotype:
[Turkey, B5], Ali Dag, near Kayseri, M. de Tchihatcheff
601 (P barcode P01903203!).
= Tunica xylorrhiza Boiss. in Ann. Sci. Nat., Bot., sér. 4, 2: 246.
1854 – Type: [Turkey, Tokat] In locis montosis sylvaticis
Ponti meridionalis inter pagum Almus et urbem Niksar,
M. de Tchihatcheff s.n. (G [herb. Boissier]).
Note. – The type specimen of Tunica ortegioides is probably also the type of T. xylorrhiza; further study is required to
clarify whether these names are based on same collection/plant.
4. Cyathophylla Bocquet & Strid in Strid, Mount. Fl. Greece
1: 175. 1986 – Type: C. chlorifolia (Poir.) Bocquet & Strid
(≡ Cucubalus chlorifolius Poir.).
Diagnosis. – Capsules exceeding the calyx, congested inflorescences in both species of this genus, perfoliate leaves
in the type (C. chlorifolia), and dense glandular indumentum
covering the entire plant in C. viscosa (C.A.Mey.) Madhani &
Rabeler, are the characteristics separating Cyathophylla from
Saponaria.
Description. – Annual herbs, glabrous or with glandular
hairs covering entire plant; leaves linear-lanceolate or ovate to
rounded, perfoliate or shortly petiolate; flowers in a capitate
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inflorescence; calyx tubular-cylindric or tubular-ovoid, 5–15
veins lacking membranous commissures; petals 5, rose or pink,
linear to linear-oblong; stamens 10; styles 2; capsule oblongovate, opening by 4 teeth; seeds reniform with flat tubercules;
embryo hookshaped; 2 species.
Etymology. – From the Greek cyath: a cup + phylla: leaves.
Distribution. – Greece, Turkey, Iraq, Azerbaijan, Iran,
and Turkmenistan.
Habitat. – In rocky slopes, mostly on limestone.
New combination
Cyathophylla viscosa (C.A.Mey.) Madhani & Rabeler, comb.
nov. ≡ Saponaria viscosa C.A.Mey., Verz. Pfl. Casp.
Meer.: 212. 1831 – Holotype: [Azerbaijan], Talysh, Swant,
C.A. von Meyer 145 (LE; isotype: E).
5. Dianthus L., Sp. Pl.: 409. 1753 – Type (designated by
Hitchcock & Green, in Sprague, Nom. Prop. Brit. Bot.:
155. 1929): D. caryophyllus L.
= Velezia L., Sp. Pl.: 332. 1753 – Type: V. rigida L.
Diagnosis. – Peltate seeds and straight embryos are characteristic features of the members of this genus. An epicalyx
is often present and membranous commissures are usually
absent (except: Dianthus candicus (P.W.Ball & Heywood)
Madhani & Heubl, D. recticaulis Ledeb., D. nudiflorus Griff.
and D. tunicoides (Ser. ex DC.) Madhani & Heubl, see below).
Description. – Annual or perennial herbs, rarely subshrubs;
leaves grass-like, often linear to oblong (ovate); flowers mostly
hermaphrodite or rarely unisexual, solitary or in capitate (rarely
paniculate or monochasial) cymes, often subtended by bracts,
and calyx mostly subtended by two or many epicalyx scales;
calyx tubular with 20–60 parallel veins, without membranous
commissures or rarely 5- to 15-veined (former Petrorhagia
spp. and Velezia rigida) and sometimes with membranous commissures (former Petrorhagia spp.), 5-toothed; petals white,
pink or red, rarely yellow, limb entire, toothed or fimbriate,
without coronal scales; stamens 10; styles 2; ovary on a short
gynophore; capsule opening by 4 teeth; seeds many, peltate;
embryo straight; 2n = 26 (Dianthus tunicoides), 28, 30, 60, 90;
species more than 300.
Etymology. – Dianthus is the contracted form of Diosanthos; from Greek Dios-: of Zeus + anthos: flower.
Distribution. – Europe, Asia, and Africa, especially in
Mediterranean regions; introduced in North and South America,
Hawaii, and Australia.
Habitat. – Mostly in hillsides, mountain slopes, dry meadows, and rocky hills.
Resurrected name
Dianthus nudiflorus Griff., Not. Pl. Asiat. 4: 466. 1854 –
Holotype: Afghanistan, barren rocky mountains around
Otipore, 7 Apr 1839, W. Griffith 132 (K).
= Velezia rigida L., Sp. Pl.: 332. 1753 – Lectotype (designated
by Strid in Taxon 53: 1053. 2004): Loefling 307, Herb. Linn.
No. 326.1 (LINN).
Note. – Since the name Dianthus rigidus had already been
used by Marschall von Bieberstein (Fl. Taur.-Caucas. 1: 325.
1808), this name was not available for a new combination.
Therefore, one of the oldest synonyms of this species under
Dianthus (i.e., D. nudiflorus Griff. 1854) is resurrected here.
New combinations
Dianthus candicus (P.W.Ball & Heywood) Madhani & Heubl,
comb. nov. ≡ Petrorhagia candica P.W.Ball & Heywood,
Bull. Brit. Mus. (Nat. Hist.), Bot. 3: 141. 1964 ≡ Fiedleria
candica (P.W.Ball & Heywood) Ovcz., Fl. Tadzhikskoi
S.S.R. 3: 608. 1968 – Holotype: [Greece], Crete, Sitia, May
1846, T.H.H. Heldreich s.n. (BM; isotypes: CGE, K barcodes K000725564! & K000725566!).
Dianthus strictiformis Madhani & Zarre, nom. nov., non
Dianthus strictus Banks ex Sol. in Russel, Nat. Hist.
Aleppo, ed. 2, 2: 252. 1794 ≡ Gypsophila stricta Bunge in
Ledeb., Fl. Altaic. 2: 129. 1830 ≡ Tunica stricta (Bunge)
Fisch. & C.A.Mey. in Index Seminum (St.Petersburg
[Petropolitanus]) 4: 50. 1837 ≡ Dianthus recticaulis Ledeb.,
Fl. Ross. 1(2): 287. 1842, nom. superfl. [citing G. stricta and
Tunica stricta in synonymy] – Holotype: Russia, in siccis
prope Buchtorminsk et Alexandrowsk, in rupestribus prope
pagum Krasnojarsk ad fl. Irtysch, C.F. Ledebour s.n. (LE).
= Petrorhagia alpina (Hablitz) P.W.Ball & Heywood in Bull.
Brit. Mus. (Nat. Hist.), Bot. 3: 145. 1964 ≡ Gypsophila alpina Hablitz in Neueste Nord. Beytr. Phys. Geogr. ErdVölkerbeschreib. 4: 57. 1783 ≡ Tunica alpina (Hablitz)
Bobrov in Bot. Zhourn. S.S.S.R. 43: 1546. 1958 ≡ Fiedleria
alpina (Hablitz) Ovcz. in Dokl. Akad. Nauk Tadzhiksk.
S.S.R. 7: 52. 1967 – Holotype: [Russia] Siberia, Irtysh River,
gravelly banks, 2000–3000 m, C.L. von Hablitz s.n. (LE).
Dianthus tunicoides Madhani & Heubl, nom. nov., non
D. armerioides Raf. in J. Bot. (Paris) 4: 269. 1814 ≡
Gypsophila armerioides Ser. ex DC., Prodr. 1: 353. 1824
≡ Tunica sibthorpii Boiss. in Diagn. Pl. Orient., ser. 1, 8:
61. 1849, nom. illeg., ≡ Tunica armerioides (Ser. ex DC.)
Halácsy, Consp. Fl. Graec. 1: 194. 1900 ≡ Petrorhagia
armerioides (Ser. ex DC.) P.W.Ball & Heywood in Bull.
Brit. Mus. (Nat. Hist.), Bot. 3: 139. 1964 ≡ Fiedleria armerioides (Ser. ex DC.) Ovcz., Fl. Tadzhikskoi S.S.R. 3: 608.
1968 – Holotype: Turquie [Turkey], Troade, G.A. Olivier
s.n. (G-DC barcode G00214254!; isotype: MW).
6. Diaphanoptera Rech.f. in Repert. Spec. Nov. Regni Veg. 48:
41. 1940 – Type: D. khorasanica Rech.f.
Diagnosis. – The members of this genus are characterized
by their membranous winged-vein calyces.
Description. – Perennial tufted plants, woody at base and
sometimes glandular pubescent; leaves slightly succulent; flowers in lax few-flowered cymes; bracts and bracteoles often present; calyx turbinate, membranous, with 5 prominent or winged
veins, 5-toothed; petals 5, rose or violet, entire or emarginate;
stamens 10; styles 2; ovary obovate, on a gynophore, ovules
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Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
(6–)8–19; capsule opening by 4 teeth; seeds reniform; embryo
curved; species 6.
Etymology. – From Ancient Greek diaphanḗs: transparent
+ ptron: wing.
Distribution. – Mountainous areas up to 3800 m in NE
Iran, Turkmenistan, and Afghanistan.
Habitat. – High altitudes of mountain peaks, on serpentine,
calcareous or rocky substrates.
7. Graecobolanthus Madhani & Rabeler, gen. nov. – Type:
G. graecus (Schreb.) Madhani & Rabeler (≡ Saponaria
graeca Schreb.).
Diagnosis. – This new genus differs from Bolanthus
mainly by its abruptly deflexed petals.
Description. – Perennial herbs; caudex woody and thick,
leaves paired at nodes, small, linear, linear-lanceolate, lanceolate or spathulate; flowers in lax dichasial or capitate inflorescences; calyx tubular-turbinate with 5 winged veins and
membranous commissures; petals 5, abruptly deflexed, white
or purple; stamens 10; styles 2, stigmatic surface all along the
inner side; ovary on a short gynophore; ovules 8–28; capsule
opening by 4 teeth; seeds comma-shaped with small tubercles
on testa and with a prominent radicle; embryo hook-shaped;
2n = 20; species 8.
Etymology. – From Latin Graeco: Greek + Bolanthus.
Distribution. – The members of this genus are Mediterranean elements restricted to Greece, in particular to Pelopon nese Peninsula.
Habitat. – The members of this genus are found in mountainous areas and mainly inhabit rocky and stony slopes.
Included species
Graecobolanthus chelmicus (Phitos) Rabeler & Madhani,
comb. nov. ≡ Bolanthus chelmicus Phitos in Bot. Chron.
(Patras) 1(1): 40. 1981 – Holotype: Greece, prov. Achaia:
mons Chelmos, supra pagum Peristera, in declivibus orientalibus, 1100–1200 m, Georgiadis 1783 (UPA).
Graecobolanthus creutzburgii (Greuter) Rabeler & Madhani,
comb. nov. ≡ Bolanthus creutzburgii Greuter in Candollea
20: 210. 1965 – Holotype: Greece, Creta, prov. Pirjotísi,
NW-Hang des Berges Mavri, ob. der Quelle Skaronero,
Tripolitaa-Kalk, 1800–1900m, 30 Jun 1961, W. Greuter 3733
(PAL [herb. Greuter]; isotypes: G barcode G00226560!, W
No. W 1966-0016564!, Z).
Graecobolanthus fruticulosus (Bory & Chaub.) Madhani
& Zarre, comb. nov. ≡ Saponaria fruticulosa Bory &
Chaub. in Bory & al., Exp. Sci. Morée, Bot.: 118. 1832 ≡
Gypsophila fruticulosa (Bory & Chaub.) Boiss., Fl. Orient.
1: 556. 1867 ≡ Bolanthus fruticulosus (Bory & Chaub.)
Barkoudah in Wentia 9: 164. 1962 – Holotype: [Greece,
Peloponnese] Coteaux de Laconie, 1829, J.B.G.M. Bory
s.n. (P barcode P04982600!).
Note. – Phitos (1981) designated a lectotype (P04982599)
that he notes was collected in 1833. Since this specimen was
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collected after the species protologue, this lectotypification
cannot be considered as effective (McNeill & al., 2012: Art. 9.3).
Graecobolanthus graecus (Schreb.) Madhani & Rabeler,
comb. nov. ≡ Saponaria graeca Schreb. in Nova Acta
Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 4: 138.
1770 ≡ Gypsophila graeca Britten in J. Bot. 44: 345. 1906
≡ Bolanthus graecus (Schreb.) Barkoudah in Wentia 9: 164.
1962 – Lectotype (designated by Phitos in Strid & Tan,
Fl. Hellenica 1: 327. 1997): “Lychnis pumila, umbellifera,
Polygoni folio, flore albo, cum circulo atro-purpureo”,
Tournefort 3032 (P-TRF).
= Cucubalus polygonoides Willd., Sp. Pl. 2: 690. 1799 ≡
Silene polygonoides (Willd.) Pers., Syn. Pl. 1: 500. 1805
≡ Saponaria polygonoides (Willd.) Jaub. & Spach, Ill.
Pl. Orient. 5: 2, t. 402. 1853 ≡ Gypsophila polygonoides (Willd.) Halácsy in Denkschr. Akad. Wiss. Wien,
Math.-Naturwiss. Kl. 61: 473. 1894 – Holotype: Greece,
Náxos, [Gundelsheimer ex] W. Wierweg 1 (B-W barcode
B -W 08601 -01 0!).
= Gypsophila ocellata Sm. in Sibthorp & Smith, Fl. Graec.
Prodr. 1: 281. 1809 ≡ Gypsophila hirsuta var. ocellata (Sm.)
Boiss., Fl. Orient. 1: 556. 1867 ≡ Gypsophila polygonoides
subsp. ocellata (Sm.) Hayek in Repert. Spec. Nov. Regni
Veg. Beih. 30(1): 220. 1924 – Lectotype (designated by
Phitos in Strid & Tan, Fl. Hellenica 1: 327. 1997): [Greece,
Evonia] in Delphi monte Euboeae, Sibthorp s.n. (OXF [IDC
photo 43: A4]).
Graecobolanthus intermedius (Phitos) Rabeler & Madhani,
comb. nov. ≡ Bolanthus intermedius Phitos in Bot. Chron.
(Patras) 1(1): 39. 1981 – Holotype: Greece, Ins. Euboea,
in saxosis serpentinicis et magesiticis litorcis ad pagu
Mantudi [rocky shore, Mantudi], Georgiadis 1657 (UPA).
Graecobolanthus laconicus (Boiss. & Heldr. ex Boiss.)
Madhani & Zarre, comb. nov. ≡ Gypsophila fasciculata
var. laconica Boiss. & Heldr. ex Boiss., Fl. Orient. 1: 556.
1867 ≡ Gypsophila laconica Boiss. & Heldr. ex Boiss., Fl.
Orient., Suppl.: 88. 1888 ≡ Bolanthus laconicus (Boiss.
& Heldr. ex Boiss.) Barkoudah in Wentia 9: 163. 1962 –
Lectotype (designated by Phitos in Bot. Chron. (Patras) 1:
36. 1981): Greece, Peloponnesus, in regione media montis
Malevo (Napvov) prope Vromopigadon, 2000 ft, 7–19 Jul
1850, G.H. Orphanides, Fl. Graeca Exs. 1 (ATHU; isolectotypes: BR barcode 000006970062!, FI, G [herb. Boissier],
JE barcodes JE00015406! & JE00015407!, K barcodes
K000725783! & K000725784!, L, LD barcode 1006486!,
US barcode 00589410!, WU No. 0073639!).
Graecobolanthus thessalus (Jaub. & Spach) Madhani & Zarre,
comb. nov. ≡ Saponaria thessala Jaub. & Spach, Ill. Pl.
Orient. 5: 2. 1853 ≡ Gypsophila thessala (Jaub. & Spach)
Halácsy in Consp. Fl. Graec. 1: 191. 1900 ≡ Gypsophila
polygonoides subsp. thessala (Jaub. & Spach) Hayek in
Repert. Spec. Nov. Regni Veg. Beih. 30(1): 221. 1924 –
Holotype: Greece, Thessaliae, prope Vólos, 1837, P.M.R.
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Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
Aucher-Eloy 566 (P barcode P05075275!, isotypes: BM
barcodes BM000572757! & BM000810718!, FI, K barcodes
K000725780! & K000725781!, P).
Graecobolanthus thymifolius (Sm.) Rabeler & Madhani,
comb. nov. ≡ Gypsophila thymifolia Sm. in Sibthorp &
Smith, Fl. Graec. Prodr. 1: 282. 1809 ≡ Saponaria thymifolia Boiss., Diagn. Pl. Orient, ser. 1, 1: 17. 1843 ≡ Gypsophila
hirsuta var. thymifolia Boiss., Fl. Orient. 1: 556. 1867 ≡
Bolanthus thymifolius (Sm.) Phitos in Bot. Chron. (Patras)
1: 39. 1981 – Lectotype (designated by Phitos in Strid &
Tan, Fl. Hellenica 1: 328. 1997): [Greece, Sterea Ellas] in
monte Parnasso, Sibthorp s.n. (OXF [IDC photo 43: A5]).
8. Gypsophila L., Sp. Pl.: 406. 1753 – Type (designated by
Hitchcock & Green in Sprague, Nom. Prop. Brit. Bot. 154.
1929): G. repens L.
= Rokejeka Forssk., Fl. Aegypt.-Arab.: 90. 1775 – Type: R. capillaris Forssk. (≡ Gypsophila capillaris (Forssk.) C.Chr.).
= Vaccaria Wolf, Gen. Pl.: III. 1776. – Lectotype (designated
by Phillips, Gen. S. African Fl. Pl., ed. 2: 330. 1951):
V. pyramidata Medik. (≡ Gypsophila hispanica Mill.).
= Hagenia Moench, Methodus: 61. 1794, non J.F.Gmel. 1791 –
Type: H. filiformis Moench (= Gypsophila pilosa Huds.).
= Arrostia Raf., Caratt. Nouv. Gen.: 75. 1810 – Type: A. dichotoma Raf. (≡ Gypsophila arrostii (Raf.) Guss.).
= Dichoglottis Fisch. & C.A.Mey. in Index Seminum (St.
Petersburg [Petropolitanus]) 1: 25. 1835 – Type: D. linearifolia Fisch. & C.A.Mey. (≡ Gypsophila linearifolia
(Fisch. & C.A.Mey.) Boiss.).
= Ankyropetalum Fenzl in Bot. Zeitung (Berlin) 1: 393. 1843
– Type: A. gypsophiloides Fenzl (≡ Gypsophila gypsophiloides (Fenzl) Blakelock).
= Bolbosaponaria Bondarenko in Kovalevskaja, Opred. Rast.
Sred. Azii 2: 327. 1971 – Type: B. sewerzowii (Regel &
Schmalh.) Bondarenko (“severtzowii”) (≡ Saponaria
sewerzowii Regel & Schmalh (“sewerzowi”).
= Pseudosaponaria (F.N.Williams) Ikonn. in Novosti Sist.
Vyssh. Rast. 15: 144. 1979 – Type: P. pilosa (Huds.) Ikonn.
(≡ Gypsophila pilosa Huds.).
Diagnosis. – The members of this genus are separated from
Acanthophyllum, Heterochroa and Saponaria by the presence of
distinct membranous commissures and calcium oxalate druses
in the mesophyll of the calyx. They differ from Acanthophyllum,
Bolanthus and Petrorhagia by reniform or reniform-globular
seeds. The concolored petals and opposite phyllotaxy can discriminate these species from Balkana (see above).
Description. – Annual or perennial herbs to tufted caespitose or pulvinate subshrubs; leaves linear or even spiny to
lanceolate and ovate, often somewhat fleshy; flowers hermaphrodite or sometimes unisexual, in many-flowered lax thyrses or
panicles, or compact head-like or few-(uni-)flowered racemelike monochasia; bracts present; pedicel longer than the calyx;
calyx campanulate-turbinate or tubular, mostly with calcium
oxalate druses, with 5 veins and membranous commissures, or
inflated with 5 winged veins and lacking membranous commissures; petals 5, white, pink, or purple, often concolorous
(upper and lower side) with purple veins; stamens 10, rarely
5; styles 2(–3), stigmatic surface extending all along the style;
ovules 4–36; capsule opening by 4 valves; seeds reniform;
embryo curved; 2n = 12, 24, 26, 28, 30, 34, 36, 48, 51, 60, 68;
species ca. 150.
Etymology. – From Greek gypsos: chalk, gypsum + philos:
loving.
Distribution. – Temperate regions of Eurasia, Africa,
Pacific Islands, with one species extending to Australia; introduced in North and South America.
Habitat. – Mostly in steppes on calcareous hills, dry or
rocky slopes, and sandy soils, sometimes weeds on farms, some
species ruderals growing along roadsides.
Resurrected names
Gypsophila arsusiana (Kotschy ex Boiss.) F.N.Williams in J.
Bot. 27: 322. 1889 ≡ Ankyropetalum arsusianum Kotschy
ex Boiss., Fl. Orient. 1: 533. 1867 – Syntypes: [Turkey
C5 Hatay], Mount Amanus, supra Arsus, 2 Jul 1862, Th.
Kotschy 117 (G [herb. Boissier]; isosyntypes: JE barcodes
JE00015413! & JE00015412!, K barcode K000725802!, L
barcode L 0038665!, P barcodes P01903163!, P01903164!
& P01903165!); Mount Akkerdagh prope Marasch,
Haussknecht s.n. (G [herb. Boissier], JE).
Gypsophila bucharica B.Fedtsch. in Trudy Imp. S.-Peterburgsk. Bot. Sada 32: 7. 1911 ≡ Saponaria bucharica
(B.Fedtsch.) Preobr. ex Popov in Trudy Turkestansk.
Gosud. Univ. 4: 24 1922 ≡ Bolbosaponaria bucharica
(B.Fedtsch.) Bondarenko in Opred. Rast. Sred. Azii 2:
292. 1971 – Holotype: [Tajikistan], Viloyati Khatlon
(Qurghonteppa), Chanatus Buchara, Prov. Baldschuan, in
montibus ad pagum Tutkaul in valle fluvii Wachsch, 8 May
1906, G.G. Morren s.n, Anonymous, Ed. Horti Bot. Imp.
10 (LE; isotype: FR barcode FR-0030878!).
Gypsophila gypsophiloides (Fenzl) Blakelock in Kew Bull.
12(2): 193. 1957 ≡ Ankyropetalum gypsophiloides Fenzl
in Bot. Zeitung (Berlin) 1: 393. 1843 – Syntypes: [Turkey
C8 Mardin], zwischen Mardin, Assuauer und Tichalaga,
Th. Kotschy 356 (W, destroyed; isosyntypes: E barcode
E00301841!, K barcodes K000725796! & K000725797!);
[Iraq, Kurdistan:] prope Gara, Jul 1841, Th. Kotschy,
Pl. Alepp. Kurd. Moss. 406 (W, destroyed; isosyntypes:
BM barcode BM000572761!, K barcode K000725795!,
HAL barcode HAL0117954!, P barcodes P04982620!,
P04982621!, P04982623! & P04982624!).
Gypsophila reuteri (Boiss. & Hausskn.) F.N.Williams in J. Bot.
27: 322. 1889 ≡ Ankyropetalum reuteri Boiss. & Hausskn.
in Boissier, Fl. Orient. 1: 533. 1867 – Holotype: [Turkey],
Maras, Akkerdagh, 15 Jul 1861, H.C. Haussknecht s.n. (G
[herb. Boissier]; isotype: JE barcode JE00015414!).
Gypsophila vaccaria (L.) Sm. in Sibthorp & Smith, Fl. Graec.
Prodr. 1: 279. 1809 ≡ Saponaria vaccaria L., Sp. Pl.: 409.
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Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
1753 ≡ Lychnis vaccaria (L.) Scop., Fl. Carniol., ed. 2, 1:
303. 1771 ≡ Vaccaria vulgaris Host, Fl. Austriac. 1: 518.
1827 ≡ Silene vaccaria (L.) E.H.L.Krause, Deutschl. Fl.,
ed. 2, 5: 120. 1901 – Lectotype (designated by Sell, 1980, on
the sheet): Cultivated material from the garden of George
Clifford III: Hartekamp Garden, Holland, Hort. Cliff. 166
(BM barcode BM000628472!).
= Saponaria hispanica Mill., Gard. Dict., ed. 8, in Errata. 1768
≡ Vaccaria hispanica (Mill.) Rauschert in Wiss. Z. MartinLuther-Univ. Halle-Wittenberg, Math.-Naturwiss. Reihe
14: 496. 1965 – Type: not specified (indicated as “… grows
naturally in Spain”).
= Saponaria segetalis Neck., Delic. Gallo-Belg. 1: 194.
1768 ≡ Vaccaria segetalis (Neck.) Garcke ex Asch., Fl.
Brandenburg 1: 84. 1860, nom. illeg. (cited S. vaccaria L.
in synonymy).
= Saponaria rubra Lam., Fl. Franç. 2: 541. 1779, nom. illeg.
(cited S. vaccaria L. in synonymy).
= Vaccaria pyramidata Medik. in Philos. Bot. 1: 96. 1789 –
Syntypes (?): East India, W. Roxburgh s.n. (K barcodes
K000725844! & K000725845!).
= Vaccaria parviflora Moench, Methodus: 63. 1794, nom. illeg.
(cited Saponaria vaccaria L. in synonymy).
= Saponaria perfoliata Roxb. ex Willd., Enum. Hort. Berol.:
464. 1809 ≡ Vaccaria perfoliata (Roxb. ex Willd.)
Sweet, Hort. Brit., ed. 2: 51. 1830 – Holotype (?): India ?,
W. Roxburgh s.n. (B barcode B -W 08501 -01 0!; isotype:
BR barcode 00006981341!).
= Saponaria vaccaria var. grandiflora Fisch. ex DC., Prodr. 1:
365. 1824 ≡ Vaccaria grandiflora (Fisch. & DC.) Jaub. &
Spach, Ill. Pl. Orient. 3: 40, t. 231. 1847 ≡ Vaccaria perfoliata var. grandiflora (Fisch. ex Seringe) Halacsy, Consp.
Fl. Graec. 1: 190. 1900 ≡ Vaccaria hispanica subsp. grandiflora (Fisch. ex DC.) Holub in Folia Geobot. Phytotax. 11:
83 .1976 ≡ Vaccaria hispanica var. grandiflora (Fisch. ex
DC.) J.Léonard in Bull. Jard. Bot. Natl. Belg. 55: 298. 1985
– Holotype: [Georgia] Iberia, Tiflis, 1819, F.E.L. Fischer
s.n. (G-DC barcode G00211736!).
= Vaccaria arvensis Link, Handbuch 2: 240. 1829, nom. illeg.
(cited V. pyramidata Medik. in synonymy).
= Vaccaria sessilifolia Sweet, Hort. Brit., ed. 2.: 51. 1830, nom.
illeg. (cited V. pyramidata Medik. in synonymy).
= Vaccaria oxyodonta Boiss., Diagn. Pl. Orient., ser. 2, 1: 68.
1854 ≡ Saponaria oxyodonta (Boiss.) Boiss., Fl. Orient. 1:
525. 1867 ≡ Vaccaria pyramidata var. oxyodonta (Boiss.)
Zohary, Fl. Palaestina 1: 104. 1966 ≡ Vaccaria hispanica
subsp. oxyodonta (Boiss.) Greuter & Burdet in Willdenowia
12: 191. 1982 – Lectotype (designated by Rechinger, F.
Iranica 163: 339. 1988): Afghanistan, W. Griffith Herb.
Late East Ind. Comp. 309 (G; isotype: C).
= Saponaria liniflora Boiss. & Hausskn. in Boissier, Fl. Orient.
1: 525. 1867 ≡ Vaccaria liniflora (Boiss. & Hausskn.)
Bornm. in Notizbl. Bot. Gart. Berlin-Dahlem 7: 142. 1917
≡ Vaccaria pyramidata var. liniflora (Boiss. & Hausskn.)
Cullen in Notes Roy. Bot. Gard. Edinburgh 27: 214. 1967
≡ Vaccaria hispanica subsp. liniflora (Boiss. & Hausskn.)
Greuter & Burdet in Willdenowia 12: 191. 1982 ≡ Vaccaria
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TAXON 67 (1) • February 2018: 83–112
hispanica var. liniflora (Boiss. & Hausskn.) J.Léonard
in Bull. Jard. Bot. Natl. Belg. 55: 298. 1985 – Syntypes:
[Turkey, C7] Inter segetes, inter Orfa et Karan, 1865, H.K.
Haussknecht, Iter syriaco-armeniacum s.n. (or 542) (JE
barcodes JE00015333!, JE00015334!, JE00015335! &
JE00015336!; isosyntype: K barcode K000725843!).
= Vaccaria perfoliata Halacsy, Consp. Fl. Graec. 1: 189. 1900,
nom. illeg., non (Roxb. ex Willd.) Sweet 1830.
= Vaccaria brachycalyx Pau in Trab. Mus. Ci. Nat., Ser. Bot.
14: 10. 1918 – Type: [Iran, Prov. Khuzestan] Gotevend
y Valle de Bazouft, bajo Karum, 400 m, Martinez de la
Escalera s.n. (MA).
9. Heterochroa Bunge in Ledeb., Fl. Altaic. 2: 131. 1830 ≡
Gypsophila sect. Heterochroa (Bunge) A.Braun in Flora
26: 383. 1843. – Type: H. petraea Bunge.
Diagnosis. – Reniform-oblong seeds and narrow membranous commissures are the main characteristic features of
this genus.
Description. – Perennial low herbs, or dense caespitose
plants, mostly with thick roots, often glandular hairy; leaves
very small, linear, linear-subulate, linear-lanceolate, lanceolate,
or ovate; inflorescence dichasial, lax-dichasial, or solitary;
calyx campanulate or widely campanulate, either with narrow
or without membranous commissures and without calcium
oxalate druses; petals 5, white to purple; stamens 10, often
shorter than petals; styles 2; ovules 8–24; capsule opening by
4 valves; seeds reniform, slightly oblong; embryo curved; 2n
= 34, 36; species 6.
Etymology. – From Greek hetero: different + chroa: skin,
color of skin.
Distribution. – Kazakhstan (Turkestan), Russia (West
Siberia; Altai; Far East, Kamchatka peninsula), Mongolia and
N China.
Habitat. – On stony or rubble hills and alpine zones, or on
stony semi-desert soils.
Resurrected names
Heterochroa desertorum Bunge in Mém. Acad. Imp. Sci.
St.-Pétersbourg Divers Savans 2: 543. 1835 ≡ Gypsophila
desertorum (Bunge) Fenzl in Ledebour, Fl. Ross. 1: 292.
1842 – Holotype: [Russia, West Siberia], Altai, along the
river Tshuya, 1832, A. Bunge s.n. (LE; isotype: P barcode
P04980638!; possible isotypes: CAS barcode 00123503!, E
barcode E00301730!, K barcode K000725758!, L barcode
L 0038678!).
Heterochroa microphylla Schrenk in Fischer & Meyer, Enum.
Pl. Nov. 1: 92 1841 – Holotype: [Kazakhstan] Alatau,
Tarbagatai Mts., 1840, A.G. Schrenk s.n. (LE; isotypes:
BR, P barcode P04980989!).
Heterochroa petraea Bunge in Ledebour, Fl. Altaic. 2: 131.
1830 ≡ Gypsophila petraea (Bunge) Fenzl in Ledebour,
Fl. Ross. 1: 291. 1842, nom. illeg., non (Baumg.) Rchb.
1830 – Syntypes: [Russia: West Siberia], Altai, Baschkaus,
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Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
A. Bunge s.n. (LE; possible isosyntypes: BR, E barcode
E00301729!, G barcode G00226863!, HAL barcode
HAL0117888!, K barcodes K000725762!, K000725763! &
K000725764!, M, P barcodes P04981532! & P04981533!);
prope Riddersk, collector? (LE).
Heterochroa violacea (Ledeb.) Walp., Repert. Bot. Syst. 5: 81.
1845 ≡ Gypsophila violacea (Ledeb.) Fenzl in Ledebour,
Fl. Ross. 1: 291. 1842 ≡ Arenaria violacea Ledeb. in Mém.
Acad. Imp. Sci. St. Pétersbourg Hist. Acad. 5: 533–534.
1815 – Holotype: [Russia, Far East], Okhotsk, Yablonov
Mts, near the city Okhotsk, D. Redowsky s.n. (LE).
New combinations
Heterochroa antoninae (Schischk.) Madhani & Zarre, comb.
nov. ≡ Gypsophila antoninae Schischk., Fl. URSS 6:
744. 1936; and in Trudy Bot. Inst. Akad. Nauk S.S.S.R.,
Ser. 1, Fl. Sist. Vyssh. Rast. 3: 180. 1937 – Holotype:
Turkmenistan, Kopet Dag, Kyzyl Chasar, 27 Jun 1934, A.
Borissova s.n. (LE).
= Gypsophila porphyrantha Rech.f. & Aellen in Bot. Jahrb.
Syst. 75: 356. 1951 – Holotype: [Iran], Khorasan, Montes
Hezar Masjed, inter Gash et Talqur, 1600–1800 m, 7–10
Jun 1948, K.H. Rechinger & F. Rechinger, Iter Iranicum
II, 5173 (W No. W 1960-0001226!; isotypes: G barcodes
G00006044! & G00006045!).
Heterochroa turkestanica (Schischk.) Madhani & Zarre,
comb. nov. ≡ Gypsophila turkestanica Schischk. in Trudy
Bot. Muz. 24: 38. 1932 – Holotype: [Russia], Middle Asia,
Tien Shan, Alexandrov Mts., between the rivers Tshatshke
and Terek, 20 Jul 1930, M. IIjin s.n. (LE).
10. Petroana Madhani & Zarre, gen. nov. – Type: P. montserratii (Fern. Casas.) Madhani & Zarre (≡ Gypsophila
montserratii Fern.Casas.).
Diagnosis. – It is similar to Gypsophila but differs in having spathulate and fleshy leaves as well as seeds with testa cells
moderately polygonal (not elongated as in Gypsophila), swollen
and tuberculate on periclinal walls.
Description. – Perennial herbs; leaves spathulate and fleshy,
paired at each node, and sometimes condensed; flowers small,
arranged in dichasial lax inflorescences; pedicel 4–6 mm long;
calyx campanulate with membranous commissures; petals 5,
concolorous, white to pinkish; stamens 10; ovary on a very short
gynophore; ovules ca. 16; styles 2, stigma terminal; capsule
opening by 4 valves; seeds subreniform, testa cells polygonal,
swollen and tuberculate; embryo curved; 2n = 26; species 2.
Etymology. – Petro: rock + ana: pertaining.
Distribution. – The two members of this genus show a
disjunct distribution pattern; P. montserratii is a mountainous
element in the Iberian peninsula and P. montana is found in
Yemen (N & S), Socotra, Oman and Somalia.
Habitat. – Open rocky slopes and gravelly wadi-bed (for
P. montana) and cracks in limestone, overhangs, and vertical
walls (for P. montserratii).
Included species
Petroana montana (Balf.f.) Madhani & Zarre, comb. nov. ≡
Gypsophila montana Balf.f. in Proc. Roy. Soc. Edinburgh
11: 501. 1882 ≡ Saponaria montana (Balf.f.) Barkoudah
in Wentia 9: 183. 1962 – Lectotype (designated here):
Socotra, Feb–Mar 1880, Balfour, Cockburn & Scott
442 (E barcode E00239367!; isolectotype: P barcode
P05018082!).
Petroana montserratii (Fern.Casas) Madhani & Zarre, comb.
nov. ≡ Gypsophila montserratii Fern.Casas in Publ. Inst.
Biol. Aplicada 52: 121. 1972 – Holotype: Spain, Fuensanta
nomen auclit, inter oppidula El-che de la Sierra et Yeste
(Albacete), ubi lege1’Unt, 14 Jul 1971, J. Molero &
J. Fernandez Casas s.n. (GDA; isotypes: BC, BCC, BCF,
JACA, MA barcode MA 327148!, SEV barcode SEV 9303!).
11. Petrorhagia (Ser.) Link, Handbuch 2: 235. 1831 ≡ Gypsophila
sect. Petrorhagia Ser. in DC., Prodr. 1: 354. 1824 – Type
(designated by Britton & Brown, Ill. Fl. N.U.S., ed. 2, 2:
72. 1913): P. saxifraga (L.) Link.
= Imperatia Moench, Methodus: 60. 1794 – Type: I. filiformis
Moench (≡ Petrorhagia saxifraga (L.) Link).
= Tunica Ludw., Inst. Regn. Veg., ed. 2: 129. 1757 – Type:
T. saxifraga Scop. (≡ Petrorhagia saxifraga (L.) Link).
= Kohlrauschia Kunth, Fl. Berol. 1: 108. 1838 ≡ Dianthus sect.
Kohlrauschia (Kunth) Fenzl in Endlicher, Gen. Pl.: 971.
1840 ≡ Petrorhagia sect. Kohlrauschia (Kunth) Ball &
Heywood in Bull. Brit. Mus. (Nat. Hist.), Bot. 3. 1964 –
Type: K. prolifera (L.) Kunth (≡ Petrorhagia prolifera (L.)
P.W.Ball & Heywood).
= Fiedleria Rchb., Deut. Bot. Herb.-Buch: 206. 1841 ≡ Tunica
sect. Fiedleria (Rchb.) Graebn. in Ascherson & Graebner,
Syn. Mitteleur. Fl. 5(2): 272. 1921 – Type: F. illyrica (Sm.)
Rchb. (≡ Petrorhagia illyrica (Ard.) P.W.Ball & Heywood).
Diagnosis. – Peltate seeds and a straight embryo along with
membranous calyx commissures as well as the presence of an
epicalyx in most members of Petrorhagia, separate it from
Bolanthus, Dianthus, Gypsophila and Saponaria.
Description. – Annual, biennial, or perennial herbs,
sometimes woody at base; leaves often narrow and grass-like,
linear, subulate to oblong; flowers hermaphrodite or sometimes unisexual in panicles, capitate or fasciculate cymes, or
solitary; bracts and bracteoles absent or present, when present
often surrounding the calyx as an epicalyx; calyx campanulate, cylindrical or tubular, 5- to 15-veined, 5-toothed with
broad membranous commissures; petals 5, white to pink or
reddish-lilac; stamens 10; styles 2; capsule opening by 4 teeth;
seeds numerous, peltate, dorsiventrally compressed, with facial
hilum; embryo straight; 2n = 26, 28, 30, or 60; species ca. 30.
Etymology. – From the Greek petra: rock + rhagas: a chink
or break.
Distribution. – Europe (Mediterranean region), C and SW
Asia, Africa (Mediterranean region); introduced in North (with
only four species being native) and South America, Africa
(Republic of South Africa), Hawaii, Australia.
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Habitat. – These plants mostly grow in rocky cliffs of
mountainous areas, calcareous or rocky substrates.
12. Psammophiliella Ikonn. in Novosti Sist. Vyssh. Rast. 11:
116. 1976 ≡ Psammophila Fourr. ex Ikonn., Novosti Sist.
Vyssh. Rast. 8: 273. 1971, nom. illeg., non Schult. 1822 –
Type: P. muralis (L.) Ikonn. (≡ Gypsophila muralis L.).
– “Psammophila” Fourr. in Ann. Soc. Linn. Lyon, sér. 2, 16:
345. 1868, nom. nud., not validly published.
Diagnosis. – Roots more or less fibrillary (P. muralis), and
the stigmatic surface extending along the inner side of the style
are the diagnostic features of these species.
Description. – Annual herbs; leaves linear to lanceolate;
flowers in lax dichasial cymes; bracts present; calyx campanulate or turbinate-tubular, with membranous or inconspicuously
membranous commissures; petals 5, white or pink; stamens
10, the stigmatic surface extending all along the style; styles 2;
capsule opening by 4 teeth, oblong; seeds reniform, with flat
tubercles; embryo curved; 2n = (30), 34; species 4.
Etymology. – From Greek psammos: sand + philia: loving.
Distribution. – Central Asia; P. muralis is native to Europe
and introduced to North America.
Habitat. – Sandy and calcareous hills, and sometimes ruderal environments (P. muralis).
= Pleioneura Rech.f. in Bot. Jahrb. Syst. 75(3): 357. 1951 – Type:
P. griffithiana (Boiss.) Rech.f. (≡ Saponaria griffithiana
Boiss.).
Diagnosis. – Distinctly clawed petals with coronal appendages separate the members of this genus from other related
genera, i.e., Gypsophila.
Description. – Perennial or rarely annual herbs; leaves
lanceolate to ovate or linear; inflorescence formed of lax or
densely paniculate or capitate cymes, rarely solitary; bracts
herbaceous, bracteoles absent; calyx tubular, obscurely 15- to
25-veined, without membranous commissures or with very
narrow ones; petals 5, mostly distinctly clawed, coronal scales
usually present; stamens 10; styles 2, rarely 3; ovary on a very
short gynophore; ovules ca. 16; capsules opening by 4, rarely
6, teeth; seeds reniform-globular with a distinct hilum; embryo
curved; 2n = 28, 56; species ca. 30.
Etymology. – From the Latin sapo: soap.
Distribution. – Temperate Eurasia, mainly in the
Mediterranean and Irano-Turanian regions, Mediterranean
Africa; introduced to North and South America, India, and
Australia .
Habitat. – Various habitats, sometimes on serpentine and
calcareous soils.
Resurrected name
13. Psammosilene W.C.Wu & C.Y.Wu in King & al., Icon. Pl.
Medic. Libro Tien-Nan-Pen-Tsao Lanmaoano 1: [s.n.], t. 1.
1945 – Type: P. tunicoides W.C.Wu & C.Y.Wu.
Diagnosis. – The membranous capsules in this genus are
unique among the members of tribe Caryophylleae.
Description. – Perennial herbs, puberulous; leaves ovate,
subsessile; flowers in terminal cymes; bracts leafy; calyx tubular, 15-veined and 5-toothed, densely glandular pubescent,
veins green; petals 5 purple-violet, inconspicuously clawed;
stamens 5; styles 2; ovary sessile, membranous, narrowly obovoid; ovules 2; capsule membranous, probably indehiscent,
1-seeded, enclosed by the persistent sepals; seeds peltate, embryo straight; species 1.
Etymology. – From Greek psammos: sand + Silene.
Distribution. – China (prov. Yunnan).
Habitat. – Rocky mountain slopes, dry pastures, calcareous
rock crevices, forests.
14. Saponaria L., Sp. Pl.: 408. 1753 – Type (designated by
Hitchcock & Green in Sprague, Nom. Prop. Brit. Bot. 155.
1929): S. officinalis L.
= Bootia Neck., Delic. Gallo-Belg.: 193. 1768, nom. illeg., non
Adans. 1763 ≡ Saponaria sect. Bootia (Neck.) DC., Prodr.
1: 365. 1824 – Type: B. vulgaris Neck. (≡ Saponaria officinalis L.).
= Saponaria sect. Proteinia Ser. ex DC., Prodr. 1: 366. 1824
≡ Proteinia (Ser. ex DC.) Rchb., Deut. Bot. Herb.-Buch:
205. 1841 – Type (designated by Schulz in Bot. Zhurn.
(Moscow & Leningrad) 69: 1479. 1984): Saponaria orientalis L.
= Spanizium Griseb., Spic. Fl. Rumel. 1: 180. 1843 – Type:
S. ocymoides (L.) Griseb. (≡ Saponaria ocymoides L.).
108
Saponaria griffithiana Boiss., Diagn. Pl. Orient., ser. 2, 1: 70.
1854 ≡ Pleioneura griffithiana (Boiss.) Rech.f. in Bot. Jahrb.
Syst. 75: 357. 1951 – Holotype: [Afghanistan], Afghania,
2100–3800 m, W. Griffith 1642 (G [herb. Boissier]; isotypes: K barcodes K000725812! & K000725813!; possible isotypes [Griffith s.n. in Herb East India Co. 308]:
C barcode C10009153!, GH barcode 00096746!, K barcode
K000725815!, P barcode P05017981!, S No. S08-18!, W
No. W 0047801!).
AUTHOR CONTRIBUTIONS
HM: Specimen study, plant collection, laboratory procedures, molecular
analyses, manuscript preparation. RR: Nomenclatural research, manuscript revision, providing some references, comments on consensus
trees, some taxonomic novelties in the study group. AP: Providing some
sequences, commenting the manuscript. BO: Manuscript revision, providing some sequences. GH: Providing laboratory and technical facilities,
providing some plant materials, manuscript revision. SZ: Supervising the
study, manuscript revision. — HM, https://orcid.org/0000-0003-03609527, hossein.madhani@ut.ac.ir; RR, https://orcid.org/0000-0002-67650353, rabeler@umich.edu); AP, https://orcid.org/0000-0003-4937-5711,
atefeh.pirani@gmai.com; BO, https://orcid.org/0000-0002-6104-4264,
bengt.oxelman@bioenv.gu.se; GH, heubl@lrz.uni-muenchen.de; SZ,
https://orcid.org/0000-0001-9159-1800, zarre@khayam.ut.ac.ir)
ACKNOWLEDGEMENTS
This paper presents some results of the project, entitled: Molecular
phylogeny of Caryophylleae (Caryophyllaceae), supported partly
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TAXON 67 (1) • February 2018: 83–112
Madhani & al. • Phylogeny and taxonomy of Caryophylleae (Caryophyllaceae)
by the Iran National Science Foundation (INSF, grant number
96003422). HM is grateful to the University of Tehran for supporting his master thesis on Caryophylleae. We are grateful to the
curators of the herbaria B, G, LE, M, MSB, TUH, W and WU for
their generous assistance, giving us the opportunity to study sheets
of this plant group and remove samples for DNA extraction. SZ also
appreciates financial supports provided by Alexander Humboldt
Stiftung (Germany) as well as University of Tehran. Nicholas Turland
(B) and Kanchi Gandhi (GH) provided nomenclatural assistance.
James Solomon (MO) kindly provided a copy of Phitos (1981). We
are grateful to the editors and reviewers handling this manuscript
who helped us to improve the methodology, taxonomic treatments
as well as linguistic errors.
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Appendix 1. Voucher information: species name, geographical origin, collector(s), voucher (herbarium), GenBank accession numbers for ITS and rps16,
respectively (* indicates sequences new for this study). Species names follow the taxonomic treatment suggested in the present study. Voucher data are
given only for accessions used in generating new sequences or those generated previously by our team (Pirani & al., 2014).
Acanthophyllum aphananthum Rech.f. 1, Afghanistan, Kabul, Rechinger 31265 (M), KF924626, KF924681; Acanthophyllum aphananthum 2, Afghanistan,
Prov. Kabul, Kabul, Band-I Kharghak, 34°32′ N 69°06′ E, 2050 m, 24.6.1965, Rechinger 31265 (MSB), –, *MF401175; Acanthophyllum allochrusoides
(Gilli) A.Pirani 1, Afghanistan, Bamian, Wendelbo & Ekberg W9801 (GB), KF924627, KF924682; Acanthophyllum allochrusoides 2, Afghanistan, Prov.
Baghlan, Andarab-Tal, Fuβ des Koh-e Shindadara bei Shashan (NE von Deh Salah), 1900–2000 m, 35°47′ N 69°21′ E, 28.5.1972, O. Anders 9367 (MSB), –,
*MF401172; Acanthophyllum borsczowii Litv., Iran, Khorassan, Zarre & al. 41034 (TUH), KF924675, KF924727; Acanthophyllum bungei (Boiss.) Trautv.
1, Iran, E Azarbaijan, Rechinger 43834 (M), KF924634, KF924688; Acanthophyllum bungei 2, Turkey, A9 Kars, Kaĝizman-Tuzluça, 13 km W Tuzluça,
1060 m, 30.7.1984, Nydegger 19519 (MSB), *MF401121, *MF401169; Acanthophyllum cerastioides (D.Don) Madhani & Zarre 1, Pakistan, Hazar, Ewald
& Zetterlund 6227 (GB), KF924628, –; Acanthophyllum cerastioides 2, NW Pakistan, Swat, in valle Jabba E Kolalai, substr. Granit, 1600–2200 m, 4.6.1965,
Rechinger 30724 (M), *MF401122, *MF401168; Acanthophyllum grandiflorum Stocks, Afghanistan, Bamian, Podlech 1340 (MSB), KF924666, KF924718;
Acanthophyllum herniarioides (Boiss.) Madhani & Zarre, Tajikistan, N von Dushanbe Anzob-Pass im Gissar-Massiv, ca. 3400 m, 14.7.1975, MuellerDoblies 75083 (B), *MF401123, –; Acanthophyllum honigbergeri (Fenzl) Barkoudah, E Afghanistan, Gardes, in altoplanitie lapidosa vallis Logar 50 km
N Gardez, 33°37′ N, 69°09′ E, 2000 m, Rechinger 35371 (B), *MF401125, *MF401176; Acanthophyllum kabulicum Schiman-Czeika, Afghanistan, Ghazni,
Frey s.n. (GB), KF924663, KF924715; Acanthophyllum kandaharicum Gilli, Iran, Khorassan, Joharchi & Zangouei 36245 (FUMH), KF924662, KF924714;
Acanthophyllum laxiflorum Boiss., Afghanistan, Lugar, Ekberg W9184 (GB), *KF924659, *KF924711; Acanthophyllum mucronatum C.A.Mey. 1, Iran,
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Appendix 1. Continued.
W Azarbaijan, Assadi & Olfat 68668 (TARI), KF924652, KF924705; Acanthophyllum mucronatum 2, Armenia, Vayotsdzor Prov., Yeghegnadzor, vicinity
of Agarakadzor village, Azar gorge, 1320 m, 26.6.2002, Optima Iter XI/2050 (M), –, *MF401170; Acanthophyllum myrianthum (Rech.f.) Madhani &
A.Pirani, C Afghanistan, Deh Kundi: in saxosis 10 km w Shahrestan, 33°40′ N, 66°35′ E, versus Deh Kundi, 34°10′ N 66°07′ E, 2200 m, 1967, Rechinger
36812 (B), *MF401124, –; Acanthophyllum oppositiflorum Aytaç, Turkey, Sivas, Aytaç 7476 (GAZI), KF924651, KF924704; Acanthophyllum scapiflorum
(Akhtar) Schiman-Czeika, Afghanistan, Kabul, Podlech 31232 (MSB), KF924646, KF924699; Acanthophyllum sordidum Bunge ex Boiss., Iran, Isfahan,
Pirani & Moazzeni 2147 (TMRC), KF924644, KF924697; Acanthophyllum spinosum C.A.Mey., Iran, Isfahan, Pirani & Moazzeni 2150 (TMRC), KF924642,
KF924696; Acanthophyllum stocksianum Boiss., Afghanistan, Kandahar, Toncev s.n. (MSB), KF924639, KF924693; Acanthophyllum versicolor Fisch.
& C.A.Mey., Turkey, Kars, Nydegger 43597b (MSB), KF924633, KF924687; Arenaria serpylloides Gay, KP148897, KP148997; Balkana spergulifolia
(Griseb.) Madhani & Zarre, Serbia, Altserbien, Zlatibor, südlich von Kremna, Umgebung von Gaj, Cigota Höhen, 1020–1475 m, 15.9.2004, Kalheber 041558 (M), *MF401126, *MF401185; Bolanthus cherlerioides (Bornm.) Bark., Turkey, B3 Isparta, Akşehir-Şarkikaraağaç, 5.1 km NE Örkenez an der
Paßauffahrt, Schutt, 1480 m, 12.8.1975, K.P. & E. Buttler 19986 (M), *MF401128, *MF401183; Bolanthus confertifolius (Hub.-Mor.) Madhani & Heubl,
Turkey, Antalya, between Fethiye and Kalkan, Pinus brutia forest, ca. 200 m, 5.2002, Özkan Eren 4362 (B), *KX834007, –; Bolanthus huber-morathii
Simon, Turkey, A2 Bursa, Soĝukpinar-Keles, 4 km nach Soĝukpinar, 860 m, 5.7.1980, Nydegger 15138 (MSB), *KX834006, *MF401184; Bolanthus minuartioides (Jaub. & Spach) Hub.-Mor., Turkey, Denizli, Muğla, 6.6.1955, Walter 201 (B), *KX834005, –; Bolanthus ortegioides (Fisch. & C.A.Mey.) Madhani
& Rabeler, Turkey, B5 Kayseri, above Talas, Ağida mt., 1700 m, 8.8.1997, Zarre 42 (MSB), *KX834008, *MF401182; Cerastium fontanum Baumg.,
AY936241, FJ404899; Cyathophylla chlorifolia (Poir.) Bocquet & A.Strid, Turkey, C3 Antalya, SW Anatolien, offener Steinschutt über Salikent, 2050 m,
Exp. N, 2.9.1995, Ulrich s.n. (M), –, *MF401186; Cyathophylla viscosa (C.A.Mey.) Madhani & Rabeler, Armenia, Vayotsdzor Prov., Vajk Distr, road
Vajk–Kochbek, ca. 8 km ENE Vajk, gorge of Darab river, 1380 m, sandy area, 1380 m, 26.6.2002, Optima Iter XI/1846 (M), *MF401117, *MF401165;
Dianthus andrzejowskianus Kulcz., JN589032, –; Dianthus armeria L., JN589087, FJ404903; Dianthus candicus (Ball & Heywood) Madhani & Heubl,
Greece, Ep. Sfakia, in declivibus australibus montis Akones ad orientem pagi Imvros, 1100–1150 m, 8.10.1966, Greuter 7679 (M), –, *MF401178; Dianthus
carthusianorum L., EF407941, EF674194; Dianthus chinensis L., JN589157, –; Dianthus cyri Fisch. & C.A.Mey., GU440808, –; Dianthus deltoides L.,
JN589027, –; Dianthus recticaulis Ledeb., Armenia, Vayotsdzor Prov., Yeghegnadzor Distr., ca. 12 km N Yeghegnadzor, around village Eghegis, 1540 m,
45°22′ E, 39°52′ N, humid meadows along river, forest with Quercus macranthera, dry slopes and rocks, 27.6.2002, Fayvush & al., OPTIMA Iter XI/2199
(M), –, *MF401177; Dianthus tunicoides (Ser.) Madhani & Heubl,, Greece, Chios, ca. 2 km nordöstlich Volissos am Rand der Straβe nach Potamia,
Grauwacke, 200 m, 16.6.1966, Lüdtke 581 (M), *MF401129, *MF401179; Diaphanoptera afghanica Podlech, Afghanistan, Baghlan, Podlech 21075 (MSB),
KF924632, –; Diaphanoptera ekbergii Hedge & Wendelbo 1, Afghanistan, Takhar, Podlech 11848 (MSB) & 11760 (MSB), KF924631, KF924686;
Diaphanoptera ekbergii 2, Afghanistan, Prov. Takhar, Khost-o-Fereng, oberes Khaush-Tal, Granitfelsen, 3800 m, 11.7.1965, Podlech 11760 (MSB), –,
*MF401173; Diaphanoptera lindbergii Hedge & Wendelbo, Afghanistan, Fariab, Hedge & al. W8336 (GB), KF924630, KF924685; Diaphanoptera stenocalycina Rech.f. & Schiman-Czeika 1, Iran, Golestan, Attar & Mehdigholi 24422 (TUH), KF924629, KF924684; Diaphanoptera stenocalycina 2, Iran,
Prov. Golestan, Golestan National Park, Almeh, 1600 m, 19.5.1975, Firuznia 1174 (M), –, *MF401174; Eremogone aculeata (S.Watson) Ikonn., JN589018,
FJ404882; Eremogone picta (Sm.) Dillenb. & Kadereit, KP148933, KP149035; Graecobolanthus fruticulosus (Bory & Chaub.) Madhani & Zarre, Greece,
Insula Euboea septentrionalis, in saxosis serpentinicis et manesiticis ad litus a pago Mantudi orientem versus, 30.6.1958, Rechinger 19439 (M), –, *MF401180;
Graecobolanthus graecus (Schreb.) Madhani & Rabeler, Greece, Epirus, Tal des Venetikos nördlich Eleftherokhori, an der Straβe Kalambaka-Grevena,
500 m, 11.10.1975, Merxmüller & Podlech 31173 (MSB), *KX834004, *MF401181; Gypsophila acantholimoides Bornm., Iran, Kuh-i Karkas (Kuh-i Kargiz),
in declivibus supra Tar, 2300–2500 m, 27.5.1974, Rechinger 46581 (MSB), *MF401083, *MF401141; Gypsophila acutifolia Fisch., Russia, Podkumok-Tal
bei Kislovodsk, 21.7.1967, Quasdorf 67 (B), *MF401100, *MF401156; Gypsophila antari Post, Iraq, Distr. Basra, Desertum meridionale (Southern Desert)
Jabal Sanam, ca. 30°10′ N, 47°30′ E, 18.3.1967, Rechinger 8568 (M), *MF401089, *MF401134; Gypsophila arabica Barkoudah, Israel, Negev Highlands:
Makhtesh Hazera (Makhtesh Katan), sandy alluvium, pebbly wadis and limestone outcrops, Altim. 10 m, 10.3.1989, Danin & al. 35.036 (B), *MF401082,
–; Gypsophila aretioides Boiss., Iran, Gachsar, Hezar Band mountain, alt. 3200 m, 30.7.2015, 36°03′ N 51°17′ E, Madhani 47116 (TUH), *MF401090, –;
Gypsophila arrostii Guss, JN589043, –; Gypsophila arrostii var. nebulosa (Boiss. & Heldr.) Greuter & Burdet, Turkey, C3 Afyon, Isparta-Denizli, 21 km
SE Dinar, 990 m, 5.8.1978, Nydegger 13410 (MSB), –, *MF401155; Gypsophila aucheri Boiss. 1, JN589077, –; Gypsophila aucheri 2, Turkey, B8 Erzincan,
Erzerum und Tercan, 18 km östlich Tercan), 1860 m, Steilbord, 2.8.1983, Nydegger 18633 (MSB), *MF401098, *MF401147; Gypsophila bermejoi G.Lopez,
Spain, Prov. Segovia, Vallelado, UTM 30t UL 78, alt. 750 m, 26.8.1983, Ladero & Casaseca 12107 (B), *MF401106, –; Gypsophila bicolor Grossh., JN589151,
–; Gypsophila bicolor, Iran, Prov. Ghazvin, Abgarm to Avaj, 12 km to Avaj, 1 km to tunnel, beside the road, 1420–1500 m, 10.5.2004, Zarre, Mashayekhi,
Taeb, Pirani & Moazzeni 35136 (MSB), –, *MF401149; Gypsophila bucharica B.Fedtsch. 1, JN589057, –;Gypsophila bucharica 2, Tadzhikistania, jugum
Chodzha-Kazian, declivibus australis montis Koipioztau, 1000 m, 8.5.1976, Kinzikaeva & Koczkareva 6663 (M), *MF401102, *MF401162; Gypsophila
capillaris (Forssk.) C.Chr. 1, KJ021878, –; Gypsophila capillaris 2, Egypt, Sinai Peninsula, Jebel Maghara, 8 km N of Bir el Hamma, 270 m, siliceous
rocks and flats, 33°30′ E, 30°40′ N, 3.5.1991, Podlech 50067 (MSB), *MF401092, *MF401135; Gypsophila capitata M.Bieb., Russia: Caucasus, Dagestan,
Distr. Chunzach, inter pag, Golotl et Kachib, vallis fl. Avarskyi Koissu, 17.7.1961, Tzvelev, Czerepanov, Bobrov & Dogadova 7559 (B), *MF401103,
*MF401161; Gypsophila capituliflora Rupr. 1, JN589143, –; Gypsophila capituliflora 2, Tadzhikistan: Pamir orientalis, Czeczekty, prope Stationem
Biologicam, fundus siccus valleculae, 3850 m, 21.8.1959, Ikonnikov 4365 (M), *MF401111, *MF401157; Gypsophila cephalotes (Schrenk) F.N.Williams 1,
JN589105, –; Gypsophila cephalotes 2, Afghanistan, Prov. Badakhshan, Wakhan, unteres Waghjir Tal bis Zemestan-e Tikili, 37°06′ E, 74°05′ N, 3950–400 m,
21.7.1971, Anders 7613 (MSB), *MF401105, *MF401158; Gypsophila curvifolia Fenzl 1, JN589159, –; Gypsophila curvifolia 2, Turkey, C4 Antalya, Orta
Toroslar, zwischen Anamur und Kazanci, Friedhof bei Akpinar, Hügel aus (Kreide-)Kalk, offene Ruderalflur, 1630 m, 36°20′ N, 32°50′ E, 22.7.1992, P. Hein
52-2 (B), *MF401099, *MF401159; Gypsophila elegans M.Bieb. 1, JN589130, –; Gypsophila elegans 2, Iran, Prov. Azarbaijan Sharqi, 19 km SE of Asheeqli
(Asheglou) in the Aras valley at road to Kaleibar, Arasbaran Protected Area, 1850 m, 46°48′55″ E, 38°53′54″ N, 1850 m, 20.6.2001, Podlech & Zarre 55293
(MSB), –, *MF401143; Gypsophila elegans 3, Germany, Bayern, Oberpfalz, Weiden, Mülldeponie/Bauschuttdeponie, offene Erde, 410–420 m, MTB 6338/2,
28.8.1991, Weigend 1895 (M), *MF401081, *MF401144; Gypsophila fastigiata L. 1, JN589144, –; Gypsophila fastigiata 2, Germany, Rheinland-Pfalz,
Rheinhessen, Mainz, Autobahnböschung am NSG Großer Sand, 17.7.1988, Kalheber 88-2892 (M), *MF401097, *MF401152; Gypsophila globulosa Stev.
Russia, Caucasica, Pjatigorsk, Felshügel über den Mineralquellen, ca. 4–500 m 11.7.64, Köhler (61) Bm 4306210 (B), *MF401108, –; Gypsophila glomerata
Pall. ex M.Bieb., Bulgaria, Bezirk Kolarovgrad, Kalkfelsen bei Madara, 17.8.1968, Merxmüller & Zollitsch 24599 (M), *MF401109, –; Gypsophila gypsophiloides (Fenzl) Blakelock, Iran, Prov. Luristan, Dow Rud, in declivibus aridis ad intoritum faucium fluvii Dez, substr. Calc., 1500–1600 m, 17.6.1974,
Rechinger 48149 (M), *MF401086, *MF401138; Gypsophila heteropoda Freyn 1, JN589110, –; Gypsophila heteropoda 2, Georgia, Caucasus, peripheria
urbis Tbilisi, haud procul a lacu Lisi, Alt. 600 m s. m., 17.5.1985, Vašák s.n. (B), *MF401085, –; Gypsophila laricina Schreb., Turkey, C5 Adana, Aladaglari,
Kayacik Deresi, entrance to Narpiz Bogazi gorge, northern slope, alt. 2450 m, 7.8.1999, Doring, Parolly & Tolimir 1231 (B), *MF401112, *MF401145;
Gypsophila leioclada Rech.f., Iran, Prov. Azarbaijan Sharqi, near At Darrehsi, ca. 70 km SE Bostanabad toward Mianeh, 1434 m, 47°23′23″ E, 37°30′54″ N,
18.6.2001, Podlech & Zarre 55219 (MSB), *MF401104, *MF401148; Gypsophila linearifolia (Fisch. & C.A.Mey.) Boiss., Iran, Prov. Ilam, 36 km from
Dehloran towards Mehran, gypsum hills, Pteropyrum community, 320 m, 4.5.1992, Akhani 8509 (MSB), *MF401091, *MF401136; Gypsophila nabelekii
Schischk., Iran, Azerbaijan occidentalis, in monte Chalil Kuh prope Razhan, 2600–3200 m, 1974, Rechinger 48847 (B), *MF401088, *MF401142; Gypsophila
oblanceolata Bark., Turkey, Provinz Nigde, bei Eregli nordlich von Ulukila, 1150 m ü. m., Steppe, Bodenoberfläche mit Trockenrissen und Salzabscheidungen,
27.9.1984, Hagemann, Binder & Schwarz 2144 (B), *MF401115, *MF401160; Gypsophila sp. (Fisch. & C.A.Mey.) Boiss., Turkey, Akdagh bei Amasya,
Manissadjian 1165 (B), *MF401087, –; Gypsophila pacifica Komarov, JX274528, –; Gypsophila paniculata L., JN589150, FJ404908; Gypsophila patrinii
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Appendix 1. Continued.
Ser. 1, JN589076, –; Gypsophila patrinii 2, Russia, Altay Republic, Ongudayskiy Rayon, N side of Chuya valley near Belyy Bom 31 km SSE of Inya, Dry
shrubland with Artemisia, S-exposed, limestone, 980 m, 50°21′ N, 87°03′ E, 15.7.2002, Raab-Straube 020105 (B), *MF401110, *MF401150; Gypsophila
perfoliata L., Turkey, B4 Ankara, Yassihuyuk (ancient site of Gordion), ancient of Gordion proper, 39°36′ N 32°02′ E, alt. ca. 1100 m, 30.9.1999, Doring,
Parolly & Tolimir 7438 (B), *MF401114, *MF401139; Gypsophila petraea (Baumg.) Rchb., Romania, Reg. Ploiesti Bucegi, vale Jepilor zwischen Busteni
cab. Caraman, Felsen, 1600–2000 m, 30.7.1965, Buttler & Dietrich 8953 (B), *MF401095 (ITS1), *MF401151; Gypsophila pilosa Huds. 1, Spain, Murcia,
Provincia de Albacete, Abzweigung von der Straße Hellin–Cieza in Richtung Agramón kurz vor Minateda, Felsen aus miozänen Kalken und Gipsflächen,
ca. 520 m, 19.5.1983, Bayer & Grau 34 (M), *MF401093, *MF401140; Gypsophila pilosa 2, Iran, Prov. Isfahan, at the entrance of Ghamsar to Ghohroud,
ruderal vegetation of main Boullevard 5 km after Ghamsar, alt. 1667 m, 33°45′33.09″ N, 51°28′ 31.50″ E, 27.05.2015, Zarre & Madhani 34287 (TUH)
*MF401094, –; Gypsophila pilulifera Boiss. & Heldr., JN589132, –; Gypsophila pinifolia Boiss. & Hausskn. ex Boiss. 1, JN589050, –; Gypsophila pinifolia
2, Turkey, B6 Malataya, (Straße O, Malataya–36,5–Gürün, 140) ca. 4 km E Sarĭhacĭ köyü an der Straße nach Malataya Felshang, 1480 m, 29.8.1971, Buttler
5774 (M), *MF401116, *MF401163; Gypsophila repens L. 1, KF737521, –; Gypsophila repens 2, Austria, Tirol, Stubaier Alpen, E-Hänge der Serlesscharte
SW Maria Waldrast bei Matrei a. Brenner, 2100 m, 1.8.1983, Podlech 38401 (MSB), *MF401101, *MF401153; Gypsophila scorzonerifolia Ser., JN589100,
–; Gypsophila silenoides Rupr., JN589049, –; Gypsophila stevenii Fisch. ex Schrank, JN589022, –; Gypsophila tomentosa L., Spain, Alicante, El Salobrar,
12.6.1986, Molero 30SWJ90 (33) (M), *MF401113, *MF401146; Gypsophila uralensis Less., KF317641, –; Gypsophila venusta Fenzl, Turkey, Anatolia,
B6, Sivas, Zwischen Zara und Sivas, 15 km E Sivas, alt. 1250 m, Steppe auf Marmor, 22.7.1981, Nydegger 16995 (B), *MF401096, *MF401154; Gypsophila
virgata Boiss., Iran, Prov. Azerbaijan orient., in saxosis faucium 38 km NNW Marand versus Jolfa, 1100 m, Rechinger 43614 (B), *MF401107, –; Gypsophila
viscosa Murray, Turkey, B5 Nevsehir, Goreme Tarihi Milli Parkim Goreme Valley, slopes SE of the Open Air Museum, ca. 1050–1100 m, tuff, secondary
steppe and ruderal vegetation, 25.5.2006, Bircan & Parolly 110 (B), *MF401084, *MF401137; Heterochroa desertorum (Bunge) Fenzl 1, JN589021, –;
Heterochroa desertorum 2, Russia, Tuviskaja ACCP, Tuva, Distr., Ovjur, prope pagum Ak-Czyra Cleistogeneto-Nanphyteta Stepposa, 3.8.1973, Timokhina
& Daniljuk 6371 (M), *MF401118, *MF401171; Heterochroa violacea Fenzl, JN589068, –; Moehringia lateriflora (L.) Fenzl, JX274536, FJ404924; Petroana
montana (Balf.f.) Madhani & Zarre, Aden peninsula, Upper Crater, plateau SE of the Tower of Silence, ca. 120–170 m, 12°47′ N, 45°02′ E, on fine-gravelly
Pozzolane, 22.3.1997, Kilian, Hein & Smalla NK 4487 (B), *MF401119, *MF401167; Petroana montserratii (Fern.Casas) Madhani & Zarre 1, JN589155,
–; Petroana montserratii 2, Spain, Albacete: pr. Ferez, 800 m, in rupi bus calcareis verticalibus, 19.8.1972, Casas s.n. (B), *MF401120, *MF401166;
Petrorhagia prolifera (L.) P.W.Ball & Heywood, GU440883, –; Petrorhagia saxifraga (L.) Link, JQ307895, FJ404930; Petrorhagia thessala (Boiss.) P.W.
Ball & Heywood, GU440885, –; Petrorhagia dubia (Raf.) G.López & Romo, AY857974, –; Psammophiliella muralis (L.) Ikonn. 1, JN589037, –;
Psammophiliella muralis 2, Germany, Bayern, MTB 6831/1: Acker nahe Mohrhof bei Poppenried/Höchstädt, 25.8.1987, E. Dörr s.n. (M), *MF401127,
*MF401186; Psammosilene tunicoides W.C.Wu & C.Y.Wu, JN589122, –; Saponaria glutinosa M.Bieb., HE602400, –; Saponaria griffithiana Boiss.,
Afghanistan, Prov. Badakhshan, Shewa valley, 37°16′ N 70°38′ E, 2640 m, 29.5.2008, Schloeder & Jacobs 1757 (M), *MF401080, *MF401133; Saponaria
ocymoides L. 1, AY936271, FJ404936; Saponaria ocymoides 2, Spain, Prov. Teruel, Monte Sierra de Javambre, 1650–1700 m, 9.7.2002, Šída & Vagnerová
3658 (M), *MF401077, *MF401130; Saponaria officinalis L. 1, AY594313, FJ404937; Saponaria officinalis 2, Japan, Kamite, Toyoshina-machi, Minamiazumi-gun, Nagano Pref., 550 m, 9.9.2002, dry bed river, Sugawara 2080906 (M), *MF401078, *MF401131; Saponaria prostrata Willd., Turkey, B6
Kahramanmaraş, 30 km to Göksun from Sariy, Bin Boğa Daği, above Yalak village, 2100–2400 m, 11.8.1997, Zarre 122 (MSB), *MF401079, *MF401132;
Saponaria pumila Hayek, AY594311, –; Saponaria sicula Raf., –, Z83153; Silene alexandri Hillebr., EF060222, EF061382; Silene gallica L., U30959
(ITS1), U30985 (ITS), JX560214; Silene italica (L.) Pers., AY936258, KF305909; Silene repens Patrin, JX274527, DQ908842; Silene viscosa (L.) Pers.,
FN821148, FN821316; Silene vulgaris (Moench) Garcke, AY857967, EF674192; Stellaria media (L.) Vill., KF737498, FJ404953; Vaccaria hispanica (Mill.)
Rauschert 1, X86896.1; Vaccaria hispanica 2, JF421553.1 (ITS2); Vaccaria hispanica 3, X83847.1 (ITS2); Velezia rigida L. 1, AY936269, –; Velezia rigida
2, GU440888, –.
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