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Abstract and Figures

The phylogenetic positions of the genera Beesia and Eranthis were investigated with respect to seven species in two genera, representing the tribe Actaeeae and five species in five other Ranunculaceae genera as outgroup. Maximum parsimony analyses were performed separately on nuclear ribosomal DNA ITS, plastid trnL-F, and combined DNA sequence data. In these analyses the positions of both Beesia and Eranthis were well supported within the tribe Actaeeae by each analysis and Beesia calthifolia was sister to Anemonopsis macrophylla on a strongly supported clade. Tribe Actaeeae is redefined to include Actaea, Anemonopsis, Beesia, and Eranthis.
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693
http://journals.tubitak.gov.tr/botany/
Turkish Journal of Botany
Turk J Bot
(2015) 39: 693-707
© TÜBİTAK
doi:10.3906/bot-1405-34
Towards a new classication of Salvia s.l.: (re)establishing the genus Pleudia Raf.
Maria WILL*, Natalie SCHMALZ, Regine CLASSEN-BOCKHOFF
Institute of Systematic Botany and Botanical Garden, Johannes Gutenberg-University Mainz, Mainz, Germany
* Correspondence: willm@uni-mainz.de
1. Introduction
e approximately 945 species placed in Salvia L. are highly
diverse in distribution, ecology, life form, morphology,
and karyology (Will, 2013). is is reected by molecular
studies revealing that Salvia is nonmonophyletic (Walker
et al., 2004; Walker and Sytsma, 2007) and showing that
its phenotypic diversity largely reects parallel evolution
(Will and Claßen-Bockho, 2014). An example of parallel
evolution is the presence of a unique modication
of the androecium leading to the formation of lever-
like stamens. e lever was traditionally regarded as
a synapomorphic character supporting the genus and
was assumed to reect various degrees of diversication
(primitive vs. derived types) in certain subgeneric or
geographical groups (Himmelbaur and Stibal, 1932, 1933,
1934; Claßen-Bockho et al., 2004; Walker and Sytsma,
2007). Interestingly, staminal levers are also known from
two Australian genera, Hemigenia R.Br. and Microcorys
R.Br. (Guerin, 2005), reecting more apparent examples of
parallel stamen evolution within the Lamiaceae.
Salvia s.l. contains 4 distinct lineages (Clades I–
IV) that include all Salvia species as well as the genera
Dorystaechas Boiss. & Heldr., Meriandra Benth., Perovskia
Kar., Rosmarinus L., and Zhumeria Rech.f. & Wendelbo
(Figure1) (Walker et al., 2004; Walker and Sytsma, 2007;
Will and Claßen-Bockho, 2014). One might argue
that the small genera nested within Salvia s.l. represent
derived lineages, and probably emerged from obscured
evolutionary processes such as budding or a progenitor-
derivate relationship as reviewed by Hörandl (2006) or
addressed by Hörandl and Stuessy (2010). Consequently,
one might expect that the corresponding genera evolved
and/or diverged more recently than the Salvia lineages, a
pattern that might be reected in a calibrated phylogeny.
However, recently published estimates for divergence
times in Salvia s.l. (see Figure2 in Drew and Sytsma, 2012)
rather seem to support the relict hypothesis. According
to current data, the most recent common ancestor
(MRCA) of Rosmarinus and Perovskia is not found to
be younger than the one of the two species representing
Clade I. is approach is a rst approximation towards the
putative origin of the major lineages within Salvia s.l. and
might change when Salvia s.l. is represented with more
species. Today, in view of the preliminary calibration and
morphological data (e.g., primitive stamen morphology)
the relict hypothesis appears to be suciently supported.
We thus consider Dorystaechas, Meriandra, Perovskia,
Rosmarinus, and Zhumeria to be most probably ‘satellite
Abstract: Salvia L. in its traditional circumscription is the largest genus within the mint family. To date, the magnitude of the task has
rendered it dicult to provide a genus-wide revision based on morphological data. Current molecular investigations based on a dense
taxon sampling representing the whole phenotypic diversity and distribution range of Salvia conrmed that the genus is polyphyletic.
Salvia species fall in 4 distinct clades, although all of them, except Clade IV, also include non-Salvia genera. A taxonomic revision is
thus urgently needed with two dierent approaches that have to be considered: (1) to include the 5 morphologically distinct non-Salvia
genera in Salvia or (2) to split Salvia s.l. into Salvia s.s. and several additional genera. Since Salvia is already highly heterogeneous in
species distribution, morphology, and chromosome number, we prefer to split the genus into molecularly well-supported clades. is
new concept may facilitate monographic studies and more focused analyses of character evolution within or between the clades. Species
representing Salvia sect. Eremosphace Bunge (subclade III-A) were chosen exemplarily to provide arguments for elevating this particular
group to the level of genus (Pleudia Raf.).
Key words: Pleudia Raf., Salvia aegyptiaca-group, Salvia sects. Notiosphace Benth. and Eremosphace Bunge, phylogeny, North Africa,
Southwest Asia
Received: 14.05.2014 Accepted/Published Online: 27.02.2015 Printed: 30.07.2015
Research Article
WILL et al. / Turk J Bot
694
genera in the sense of Frodin (2004), who argued that a
single larger taxon would be accompanied by several,
oen quite small ‘satellite’ taxa. e supposed relict status
is also supported by the comparably narrow distribution
of, for example, Dorystaechas hastata in the Mediterranean
region of southern Turkey.
ese ndings demand taxonomic consequences
that should not be taken lightly. Maintaining Salvia in its
traditional circumscription would require the inclusion
of the 5 distinct genera and consequently lead to a
morphologically and karyologically more heterogeneous
group. is of course would involve fewer taxonomic
changes than the alternative approach, but morphological
characters have not been found that support all of the
major clades including, for example, Clade III (Salvia-
species plus Zhumeria). In contrast, splitting Salvia s.l.
into a narrowly dened Salvia s.s. plus several molecularly
highly supported clades might appear unpopular in terms
of the long taxonomic tradition and the use of Salvia in
horticulture or medicine. One might also argue that a new
classication should not be done before an exhaustive
taxonomic revision of Salvia s.l., which would probably
take decades.
So far, several genera of comparable size or with similar
horticultural importance as Salvia (e.g., orchids; Whitten
et al., 2007) have already been the subject of molecular
studies, which oen result in generic re-circumscriptions,
partial generic revisions, or at least proposals to split
polyphyletic genera (e.g., Frodin, 2004; Mansion, 2004;
Kress et al., 2005; Whitten et al., 2007; Kučera et al., 2013;
Dillenberger and Kadereit, 2014).
We are convinced that splitting Salvia will contribute to
comparably small, and thus manageable, genera and a natural
classication (see Frodin, 2004). Novel circumscriptions are
needed for the new genera, while Dorystaechas, Meriandra,
Perovskia, Rosmarinus, and Zhumeria are accepted with their
previous circumscriptions. Since the type species of Salvia,
S.ocinalis L. (Jarvis, 2007), is placed in a derived position
within Clade I, this lineage represents Salvia s.s.
As part of the ongoing revision of the polyphyletic
Salvia s.l., we suggest that well-supported clades that can
additionally be supported by distribution, morphology,
karyology, and/or ecology be recognized at generic level.
One of them is subclade III-A (Salvia aegyptiaca-group;
Will and Claßen-Bockho, 2014), as already proposed by
Ranesque (1837; gen. Pleudia Raf.).
100/1.00
94/1.00
98/1.00
96/1.00
89/1.00
100/1.00 S. aristata
S. areysiana
S. bariensis
S. aristata NCBI
S. herbanica
Zhumeria majdae
S. aegyptiaca
S. deserti (cp)
Z. majdae NCBI
100/1.00
100/1.00
100/1.00
100/1.00
85/1.00
77/1.00
100/1.00 Melissa
Clade II
Lepechinia
Clade IV
Clade I
Perovskia
Rosmarinus
92/1.00
99/1.00
100/1.00
Hyptis
Collinsonia
Salvia
-/0.96
S. deserti (nr)
S. geminata
99/1.00
100/1.00
97/1.00
100/1.00 100/1.00
88/1.00
Meriandra
-/0.98
Horminum
100/1.00
100/1.00
Dorystaechas
Salvia
rpl32-trnL ETS
III-A
III-B
III-A
III
Figure1. Simplied phylogenetic tree of Salvia s.l. (Will and Claßen-Bockho, 2014; modied). Molecular phylogeny based on
chloroplast data (le) compared to nuclear data (ETS) (right) with focus on SW Asian taxa nesting in subclade III-A (highlighted:
gray box). A highly supported incongruence within this clade is identied for the placement of S.deserti (dotted line).
WILL et al. / Turk J Bot
695
2. Materials and methods
Molecular data, distribution, and morphology were used
to circumscribe the highly supported Salvia aegyptiaca
-
group (Will and Claßen-Bockho, 2014) as a genus. For the
taxonomic treatment, morphological data and information
about species distribution were mainly obtained from the
literature (Hedge, 1966, 1974a; Bokhari and Hedge, 1977;
Hedge, 1982a, 1982b, 1982c; Santos and Fernández, 1986;
Hedge 1990; Scholz, 1993; ulin, 1993, 2009). Molecular
data are available for 8 of the 14 species (66.7%) (Table)
(Will and Claßen-Bockho, 2014). Chromosome numbers
for Clade III species are based on literature research and
the Index to Plant Chromosome Numbers (IPCN). e
incomplete taxon sampling for molecular studies is due to
Ye men
Oman
Saudi Arabia
Somalia
Ethiopia
Sudan
Egypt
Libya
Algeria
Tunesia
Morocco
W Sahara
Canary Islands
Mauritania Mali Niger
Tschad
Eritrea
UAE
Iran
Afg
Pakistan
Irak
Syria
Isr
Jord
Leb
macilenta
trichocalycina
eremophila
bariensis
areysiana
aegyptiaca
A
Ye men
Oman
Saudi Arabia
Somalia
Ethiopia
Sudan
Egypt
Libya
Algeria
Tunesia
Morocco
W Sahara
Canary Islands
Mauritania Mali Niger
Tschad
Eritrea
UAE
Iran
Afg
Pakistan
Irak
Syria
Isr
Jord
Leb
santolinifolia
bazmanica
tebesana
herbanica
chudaei
chudaei
B
hillcoatiae
geminata
deserti
deserti
?
Figure2. Distribution of Pleudia (S.aegyptiaca-group). Based on Hedge (1968, 1974a, 1982b, 1982c), Bokhari and Hedge (1977),
Santos and Fernández, (1986), and ulin (1993, 2009). Afg=Afghanistan, Isr=Israel, Jord=Jordan, Leb= Lebanon, UAE=United
Arab Emirates. A: Overlapping distribution of S.aegyptiaca, S.deserti, S.macilenta, S.eremophila, and S.trichocalycina vs. locally
restricted species on the Arabian Peninsula and in East Africa. B: Disjunct or scattered distribution of S.chudaei (Libya/Chad,
Algeria) and S.tebesana in contrast to locally restricted species, i.e. S. geminata, S.hillcoatiae, and S.herbanica (Fuerteventura,
Canary Islands).
WILL et al. / Turk J Bot
696
diculties in collecting the species in their natural habitats
in Northern Africa and Southwest Asia (Figure2) and the
restricted number of specimens available in collections
(herbaria and botanical gardens) for destructive sampling.
3. Results and discussion
3.1. Molecular data concerning the taxonomy of Salvia L.
Preliminary molecular investigations revealed that Salvia
is nonmonophyletic (Walker et al., 2004; Walker and
Sytsma, 2007). Bräuchler et al. (2010) assumed that the
genus was paraphyletic ‘with regard to at least Dorystoechas
[sic] and Perovskia’ (Bräuchler et al., 2010; p. 502). Drew
and Sytsma (2012) also considered Salvia as paraphyletic
but did not specify which of the ve non-Salvia genera
were nested within it. Our data (Will, 2013; Will and
Claßen-Bockho, 2014) similarly suggest that Salvia is
polyphyletic and therefore in need of revision. e strong
geographical signal detected in the ‘genus-wide’ study in
combination with morphological characters provides a
pragmatic approach to circumscribe monophyletic groups
that can be revised separately.
Major clades and subclades of Salvia s.l. supported in
phylogenetic studies are only partly in accordance with
the existing concepts of subgeneric classication (Will,
2013). Although the clades are not resolved at the species
level, it is reasonable to draw taxonomic conclusions at
this point, as new molecular data are only likely to resolve
relationships within the major clades, and are therefore
unlikely to lead to further taxonomic changes in the
future. As a rst example, we elevate S.aegyptiaca and its
Tab le . Species placed in Pleudia with some distinctive characters. Species highlighted (bold) were originally placed in Salvia sect.
Eremosphace Bunge (Bunge, 1873). Molecular data: based on Walker and Sytsma (2007), Will (2013), Will and Claßen-Bockho (2014);
leaf anatomy: Bokhari and Hedge (1977); distribution and morphology: Hedge (1966, 1974a, 1982b, 1982c), Santos and Fernández
(1986); ulin (1993, 2009); Afg= Afghanistan; Afr=Africa; C=Central; CI=Canary Islands; Co= Corolla, upper lip; E=East;
N=North/Northern; Pak=Pakistan; S=South/Southern; SE=Southeast; SW=Southwest; (×)= imperfect; !=dierent, lower thecae
fertile (Bokhari and Hedge, 1977) or sterile (Hedge, 1982b); −=no data available; p leaf margin lobed to pinnatid; *the maximal length
of the pedicel is generally based on a slight enlargement of the pedicel aer owering. We point to dierent descriptions of growth forms
in various oras, e.g., for S.aegyptiaca. While Hedge (1974a) refers to this species as ‘much branched suruticose herb, 10–20(–40 cm)’,
it is described as dwarf shrub in a later work by the same author (Sales et al., 2010).
Species [distribution] Morphology Phylogeny
Salvia species recognized in Pleudia
Height [in cm]
Simple leaves
Xeromorphic leaves
Co size [in mm]
Pedicel [in mm] *
Lower thecae fertile
ITS
ETS
rpl32- trnL
trnL-F
aegyptiaca L. [CI, E & N Afr to Pak, India] 6-20(40) × × 5-8 2-3.5
(-5) ×××××
areysiana Deers [S Yemen] 20-40(50) × 16 × × ×
bariensis ulin [Somalia, E Afr] 20-40 × 14-17 4 × × ×
bazmanica Rech.f. & Esfand. [S Iran] 20-30 × 10 1.5-4 ×
chudaei Batt. & Trab. [S Sahara] 30-60 × 7 1.5 ×
deserti Decne. [Sinai, Israel, Jordan, Arabia] ≤ 30 × × 6 0.6 × × ×
eremophila Boiss. [C & S Iran] 10-30 × × 4-5 1.5-5 !
geminata ulin [Yemen] ≤ 10 × 10 3 × ×
herbanica A.Santos & M.Fernández [Fuerteventura, CI] 12-15 × 14-16 (×) × × ×
hillcoatiae Hedge [Oman] 15 × 5-8 2 ×
macilenta Boiss. [Oman, S Iran, SW Afg] 15-50 × × 5-6 2-3.5 ×
santolinifolia Boiss. [S Iran, Pak, SE Afg, India] 10-30 × p× 5-6 1.5-2.5 × − − − ×
tebesana Bunge [SW Iran] 18-24 × × 5-6 1.5-3 ×
trichocalycina Benth. [E Afg, Pak] 10-25 × p× 10-12 1.5-5 × × − − ×
WILL et al. / Turk J Bot
697
allies, the former Salvia sect. Eremosphace Bunge, to the
level of genus, re-establishing the name Pleudia Raf. (see
taxonomic treatment; Table).
3.2. Taxonomic history of the species group centered on
Salvia aegyptiaca
Linnaeus (1753) already included S. aegyptiaca L. in
his Species Plantarum. Since then, various taxonomical
treatments have placed the species and its allies in
dierent sections (Bentham, 1832–1836, 1848; Bunge,
1873; Bentham, 1876; Briquet, 1897) and species groups
(Hedge, 1974, 1982b) or recognized it as its own genus
(Ranesque, 1837).
3.2.1. Pleudia Raf.
Ranesque (1837) provided one of the earliest concepts
for a taxonomic treatment of Salvia s.l. He suggested
that Salvia L. should be split into several small genera
(Ranesque, 1837; Appendix), among them Pleudia, which
he described as follows: ‘12. Pleudia, Galea brevissima
emarg. labio concavo ut Nepeta, stam. plerumque 4 [sic!]
fertilis ! S.egyptiaca [sic!] & c.’ (Ranesque, 1837; p. 94).
It is thus the oldest name proposed at the generic rank for
Salvia aegyptiaca and its relatives.
3.2.2. Salvia sect. Notiosphace Benth.
Bentham (1832–1836) established 14 sections in Salvia,
among them Salvia sect. Notiosphace Benth. According
to his circumscription, S.aegyptiaca and the other species
placed in this section are perennial herbs with either
fertile or sterile thecae at the lower lever arm. e section
contains exclusively small-owered species (Figure 3).
Except for the widespread species S. plebeia R.Br.
(Southwest to East Asia and Northern Australia), all other
representatives of the section are restricted to Southwest
(SW) Asia and Northern Africa. Bentham maintained
Salvia sect. Notiosphace in his later treatments (Bentham
1848, 1876) and placed it in subg. Leonia Benth. e name
Notiosphace is derived from the Greek ‘notio’ (south) and
‘sphace’ (sage) (Sales et al., 2010).
3.2.3. Salvia sect. Eremosphace Bunge
Paying special attention to species distribution, Bunge
(1873) split Salvia sect. Notiosphace Benth. into Salvia
sects. Eremosphace Bunge and Notiosphace Benth. p.p.
e former contained only Salvia species restricted to
SW Asia, i.e. S.aegyptiaca (also occurring on the Arabian
Peninsula, in Northern Africa, and on the Canary Islands),
S. santolinifolia, S. eremophila, and the newly described
species S. tebesana. ese species were characterized by
having fertile lower thecae (except S.eremophila; Hedge,
1982b; see Table), and were separated from species with
completely sterile lower thecae, which were maintained
in Salvia sect. Notiosphace. Briquet (1897) followed this
classication and added S.deserti, S. trichocalycina, and
S.macilenta to Salvia sect. Eremosphace.
e sectional name Eremosphace is derived from the
Greek ‘eremos’ (ερεμος) referring to lonely, desolate, and
uninhabited places, and the Latin ‘sphaceo’, which comes
from the Greek ‘elelisphacos’ (ἐλελίσφκος referring to
S.triloba) and ‘sphacos’ (σφάκος describing S.calycina, the
so-called sage-apple) used by Pliny the Elder (Carvahlo,
1850).
3.2.4. Species groups sensu Hedge
Not aiming to provide a new subgeneric classication, but
instead contributing to a more natural system for the genus,
Hedge (e.g., 1974a) introduced informal ‘species groups’
in his treatments of Old World Salvia. ese groups were
largely dened by distribution and morphology (Hedge,
1974a, 1982a, 1982b). ereby, problems arose from the
lack of comparability between local oras (Hedge, 1974a,
1982a, 1982b). In his revision of African Salvia, Hedge
(1974a) placed S.aegyptiaca in ‘species group F’ together
with the African species S.chudaei and S.deserti. In the
Flora Iranica (Hedge 1982b), S. aegyptiaca was part of a
species group with 8 SW Asian species (‘Grex A’), among
them S.viridis L. and S.plebeia R.Br. ese two species are
morphologically clearly distinct from the S. aegyptiaca-
group. ey also have minute to small owers but are
the only annuals placed in the group (Hedge, 1982b).
Molecular data highly support the placements of S.plebeia
in Clade IV (Will, 2013) and S.viridis in Clade I (Walker
et al., 2004). All species previously placed in Salvia sect.
Eremosphace are highly supported as members of Clade III
(Figure1).
Bokhari and Hedge (1977) conducted a comparative
anatomical study of S.aegyptiaca and its relatives (Table).
ey recognized 11 species but argued that some of them
might be conspecic with one another, e.g., S. gabrieli
Rech.f. and S. aegyptiaca L. or S. tebesana Bunge and
S. santolinifolia Boiss. Since this study, 4 species tting
morphologically into the S. aegyptiaca-group sensu
Bokhari and Hedge (1977) have been described (Hedge,
1982c; Santos and Fernández, 1986; ulin, 1993, 2009).
Today, 14 species clearly belong to this species group
(Table) (Hedge, 1966, 1974a; Bokhari and Hedge, 1977;
Hedge, 1982a, 1982b, 1982c; Santos and Fernández, 1986;
Hedge, 1990; Scholz, 1993; ulin, 1993; Walker and
Sytsma, 2007; ulin, 2009; Will and Claßen-Bockho,
2014). Another taxon, S.halaensis Vicary (Vicary, 1847),
is also assumed to belong to this group, but further studies
are needed to conrm this placement (Hedge, 1982b,
1990).
3.3. Circumscription of Pleudia Raf. based on molecular
data
As we provide nomenclatural changes at the end of
this article and in order to facilitate comparisons with
previously published phylogenies, we retain the names
(Salvia spp.) as provided in the corresponding studies.
WILL et al. / Turk J Bot
698
Figure3. Habit and morphology of Pleudia Raf. (Salvia aegyptiaca-group). (A) S.geminata; dwarf shrub approx. 10cm in
height; specimen: M. ulin, A. Beier & Mohammed A. Hussein no. 9629; K00248959; isotype, (B-C) S.areysiana; specimen:
ulin, Erikson, Gifri & Långstöm no. 8472; V64083 (Kew); (B) simple, revolute leaves from below with dense indumentum;
(C) ower: approx. 17mm excluding pedicel; (D–F) S.aegyptiaca, minute to small owers; (D) frontal view; (E) lateral view,
(F) drawing of S.aegyptiaca L. (as S.pumila Benth.) (Jacquemont, 1844); (G) S.chudaei (Battandier and Trabut 1907), with
straight upper lip and exposed stamens; (H) habit of S.geminata, a much-branched dwarf shrub up to 10cm (photo: M.
ulin); (I–K) S.herbanica from a natural population on Fuerteventura; (I) ower, lateral view; (J) dissected ower (MJG
009888); stamen with upper and lower lever arm; (K) close-up of the lower lever arm; fertile thecae with pollen grains; 16
owers (2 populations) from Fuerteventura were dissected; size and shape of the thecae of the lower lever arm largely vary
(fertile and sterile thecae).
WILL et al. / Turk J Bot
699
Eight (57%) of the 14 species recognized in Pleudia
are represented in molecular studies (Walker and Sytsma,
2007; Will, 2013; Will and Claßen-Bockho, 2014).
According to plastid and nuclear data (trnL-F, psbA-trnH,
rpl32-trnL-F, ITS, ETS) (Figures 1 and 4), they are well
supported as members of Clade III (subclade III-A, C;
Figures1 and4). According to ETS data, 6 of these species
are placed in subclade III-A (1.00PP/100%BS) (Figure1)
(Will and Claßen-Bockho, 2014). Walker and Sytsma
(2007) included only three Pleudia species, S.santolinifolia,
S.trichocalycina, and S.aegyptiaca, which were supported
in a polytomy (Figures4A–4C). e former were placed
in (Bunge, 1873; Briquet, 1897) or near (Hedge, 1966,
1974a, 1982a, 1982c) Salvia sect. Eremosphace. Since
morphological and molecular data are not conicting, we
also include species that were not sampled in molecular
studies in the new genus.
Chloroplast data show Pleudia (subclade III-A) to be
part of a trichotomy with Zhumeria and subclade III-B
(Figure4D). Within the latter, S.aristata Aucher (Northeast
Iran; Southeast Turkey), S. pterocalyx Hedge (Northeast
Afghanistan), S. vvedenskii Nikitina, and S. margaritae
Botsch. (both Central Asia) are highly supported. Nuclear
data support a slightly dierent position for S.margaritae
as sister species to Zhumeria (Figure4B). However, this
relationship is only moderately supported (78%BS).
Based on molecular data, one might recognize the
whole Clade III including Zhumeria majdae (Rechinger
and Wendelbo, 1967) as one genus. Bokhari and Hedge
(1976) investigated the anatomy, taxonomy, and anities
of Zhumeria. e authors described the genus as ‘most
isolated genus […] with some links with the genera of
the tribe Meriandreae. is particular tribe was already
recognized as an articial assemblage of isolated relict
genera, i.e. Dorystaechas, Meriandra, and Perovskia
(Bokhari and Hedge, 1971, 1976; Henderson et al., 1986).
Indeed, molecular, morphological (e.g., pollen; Jamzad et
al., 2006), and anatomical (Bokhari and Hedge, 1971) data
support the hypothesis that these genera are neither closely
related to Zhumeria nor to each other, except probably
Dorystaechas and Meriandra.
To recognize Clade III as one genus requires the
acceptance of a quite heterogeneous taxon. Although
some characters are shared by all Salvia representatives
nesting in Clade III, i.e. clearly bilabiate corollas bearing
2 fertile stamens with signicantly enlarged connectives
and 2 staminodes, characters to support the inclusion of
Zhumeria have as yet not been identied (Bokhari and
Hedge, 1976; Harley et al., 2004). is monospecic genus
rather diers from all other species nesting in Clade III
in: (1) the lack of an enlarged connective tissue separating
both thecae (the latter are only somewhat separated and
do not form a lever; see Figure1 in Bokhari and Hedge,
1976), (2) staminodes exserted from the corolla (Harley et
al., 2004), (3) an indistinctly bilabiate corolla with subequal
lips, (4) few owers borne singly in the uppermost leaf
axils, and (5) an extraordinary diversity of trichomes not
recorded for any other genus of the family (for details see
Bokhari and Hedge, 1976). Considering morphology and
the previously discussed relict status of Zhumeria, it seems
most parsimonious that this genus is a basally branching
lineage within Clade III. Species with lever-like stamens
are most probably derived, although the underlying
evolutionary processes leading to their development and
radiation remain unclear based on current data.
Molecular data clearly distinguish Pleudia (subclade
III-A) from the remainder of Clade III, which have their
westernmost distribution in SW Asia (Figure 2). Species
nesting in subclade III-B also occur in dry habitats, but
were traditionally separated from Pleudia based on
morphology and distribution (Bunge, 1873; Briquet, 1897;
Pobedimova, 1954). More broadly sampled molecular
96 S. aristata
Zhumeria majdae
S. aegyptiaca
100
S. tetrodonta
S. santolinifolia
S. trichocalycina
trnL-F + psbA-trnH + nrITS
Zhumeria majdae
97 S. aristata
S. tetrodonta
S. trichocalycina
S. aegyptiaca
95
trnL-F + nrITS nrITS
-/78
*/* Zhumeria majdaa
S. margaritae
*/*
S. vvedenskii
S. aristata NCBI
*/*
III-A
III-B
III
S. trichocalycina NCBI
*/* S. aegyptiaca
S. aegyptiaca NCBI
S. herbanica
*/98
*/* Zhumeria majdae
S. margaritae
S. vvedenskii*/94
S. pterocalyx
S. aristata
S. aristata NCBI
S. aristata
*/*
*/94
III-B
*/92
S. aegyptiaca
S. herbanica
III-C
S. bariensis
S. areysiana
S. deserti
*/98
III */*
*/93
III-A
rpl32-trnL
ABCD
Figure4. Placement of species representing Pleudia (S. aegyptiaca-group) based on various molecular markers in previous
molecular studies. Asterisks above branches indicate maximal support values (100%BS, 1.00PP); species representing Pleudia are
highlighted by light gray branches; A and C: strict consensus trees, based on maximum parsimony analysis of combined datasets
(plastid and nuclear markers) (Walker and Sytsma, 2007; modied); B and D: topology of Clade III; based on nuclear (nrITS; B)
and plastid data (rpl32-trnL; D); maximum likelihood analysis and bayesian inference (Will, 2013; modied). e placement of
S.santolinifolia and S.trichocalycina in a basal polytomy within Clade III (Figures4A–4C) is not in conict with the recognition/
circumscription of Pleudia.
WILL et al. / Turk J Bot
700
studies of Old World Salvia-species support S. aristata,
S.pterocalyx, S.vvedenskii, and S.margaritae as one highly
supported clade (subclade III-B; Figure4D; Will, 2013).
Based on similar morphology and distribution, 16 species
from Central (former USSR) and SW Asia are expected
to belong to this group representing sect. Physosphace
Bunge and subg. Macrosphace Pobed. (Pobedimova, 1954).
Although this group needs to be studied in more detail,
some supporting characters might be corolla (≥25mm) and
calyx (rarely <15 mm) size; relatively long pedicels (≥5mm
up to 20mm); large, occasionally lobed or even compound
leaves; and large seeds (Pobedimova, 1954; Hedge, 1960;
Nikitina, 1962; Hedge, 1974b, 1982b; Behçet and Avlamaz,
2009). Concerning the proposed split of Clade III, the
name Polakia Stapf (1885) has to be considered for the
subclade III-B species. Briquet (1897) adopted this genus
distinct from Salvia based on Polakia paradoxa Stapf (syn.
Salvia aristata Aucher ex Benth.; WCSP, 2014). If subclade
III-B is shown to form a monophyletic group with Pleudia,
the latter name would have priority at the generic rank.
us, accepting Pleudia for the members of subclade III-A
would not entail further nomenclatural changes if they
are found to be congeneric with the members of subclade
III-B. Further hypotheses on the relationships of these
large-owered relatives of Pleudia are discussed in more
detail by Hedge (1960, 1974b).
3.4. Genetic diversity vs. morphological uniformity
ough Pleudia is well-supported and characterized,
morphological characters separating the species from
each other are largely lacking. According to Bokhari and
Hedge (1977), leaf shape, degree of revolution of the
leaf margin, calyx shape, and indumentum are the most
suitable characters for species delimitation. However,
character states vary even within taxa. Hedge (1974a)
observed a slightly changed ratio of leaf length to width as
a function of distribution, with the leaves of S.aegyptiaca
being generally narrower in North African specimens. In
addition, plants collected in the easternmost distribution
range have more glandular hairs than those from N Africa
and on the Canary Islands (Bokhari and Hedge, 1977). As a
consequence, the indumentum might reect adaptation to
microclimatic dierences (locally more mesic conditions;
Hedge 1974a), instead of being a suitable character for
species delimitation (see also Bokhari and Hedge, 1977).
3.5. Local endemics vs. widespread species
Within the new genus, the distribution area of
S. aegyptiaca (Figure 2A) almost completely overlaps
with the distribution areas of taxa restricted to Africa
and Saudi Arabia (Figure2). e latter are geographically
well isolated from each other. A general trend in the new
genus seems to be a locally restricted distribution except
for S. aegyptiaca. It would be worthwhile to investigate
whether the corresponding species are adapted to certain
edaphic conditions. Salvia herbanica for example is
restricted to only a few populations growing on basaltic
clis on Fuerteventura (Canary Islands). A decline of
their populations in size was observed and is most likely
explained by the restricted number of suitable, geological
oen instable habitats, land use (grazing goats and sheep),
and an infection/consumption of the fruits by parasitic
insects (Scholz, 1993). Today, S. herbanica is critically
endangered (Scholz, 1993; Gobierno de Canarias, 2004;
Scholz and Santos Guerra, 2004; Gangoso et al., 2006;
Moreno, 2008; Scholz and Santos Guerra, 2011; Rodríguez
González et al., 2013). Such detailed information is not
available for further Pleudia species (IUCN, 2014) and
thus the conservation status of these taxa is not easy to
evaluate.
3.6. Nuclear vs. chloroplast sequence data—hybridization
discovered
Plastid and nuclear data suggest the hybrid origin of
one species nesting in subclade III-A, i.e. S. deserti. e
nding is based on the highly supported conict observed
between nuclear and chloroplast data (Figure 1, dotted
branches). In the plastid dataset, S.deser ti is found in a
basally branching position, sister to the remainder of
subclade III-A (Figure 1, le). is molecular marker
is inherited maternally. erefore, the seed parent was
expected to be the sister species to S.deserti. Since no sister
species was found, we conclude that the maternal parent
was not sampled or is an extinct species. In contrast, ETS
data support S.deserti as a sister species to S.aegyptiaca
(Figure1, right). According to this bi-parentally inherited
marker, we assume S.aegyptiaca is most likely the pollen
parent of S.deserti.
Hybridization is an important factor triggering
speciation (Arnold, 1992; Abbott et al., 2013 and literature
cited therein). Several aspects such as intermediate
morphological characters, aberrant chromosome
numbers, and the lack of isolation mechanisms to avoid
hybridization have been repeatedly discussed for Salvia s.l.
(Hrubý, 1933, 1935; Epling, 1938; Steward, 1939; Hrubý,
1941; Epling, 1947; Grant and Grant, 1964; Emboden,
1971; Wu and Huang, 1975; Hedge, 1982a; Meyn and
Emboden, 1987; Fernández Alonso, 1991; Reiseld, 1993;
Hihara et al., 2001; Van Jaarsveld, 2002; Reales et al., 2004;
Zhiyun et al., 2004; Wester and Claßen-Bockho, 2006;
Wood, 2007; Wester and Pauw, 2009; Bercu et al., 2012).
However, the hypothesis of a hybrid origin has only been
tested for a limited number of taxa and has not yet been
conrmed by molecular studies (Sudarmono, 2007; Jenks,
2008; Will and Claßen-Bockho, 2014; P. Wester, pers.
comm.).
Hybrids might become isolated from the parental
species by geographic isolation or by establishing eective
barriers to avoid back-crossing. For sympatric species,
WILL et al. / Turk J Bot
701
the latter is possible via polyploidization. Interestingly,
the chromosome number of S.deserti is relatively high
(2n = 48; Al-Turki et al., 2000). Several chromosome
numbers for S. aegyptiaca have been reported in the
literature, most frequently 2n= 26 or 28 (Hedge, 1974a;
Borgen, 1980; Haque and Ghoshal, 1980; Haque, 1981;
Siddiqi, 1985; Dalgaard, 1986; Al-Turki et al., 2000), but
dierent studies have found 2n=12 (Díaz Lifante et al.,
1992), 2n=38 (Löve, 1971), and 2n=42 (Humphries et al.,
1978). Except for S.chudaei (2n=28; Haifa and Joumena,
1991), other chromosome numbers have not been reported
for Pleudia or for subclade III-B and Zhumeria.
We cannot exclude the possibility that the observed
incongruence reects incomplete lineage sorting (Joly
et al., 2009), but based on an overlapping distribution
(Figure2A) and the relatively high chromosome number
of S.deserti (2n=48; Al-Turki et al., 2000), we prefer the
hybrid hypothesis and assume that S. deserti arose via
allopolyploidization. Further chromosome counts in the
new genus could support our hypothesis.
4. Taxonomic and nomenclatural consequences
Pleudia Raf.
Basionym: Pleudia Raf., Fl Tell 3:94 (1837).
Synonym: Salvia sect. Notiosphace Benth. p.p. (Labiat.
Gen. Spec. p. 309 (1832–36); A. De Candolle Prodr. 12 p.
354 (1848); Gen. Pl. 2, p. 1167 (1876); non Bentham 1833
Hook. Bot. Misc. 3 p. 374); Salvia sect. Eremosphace Bunge
(Bunge 1873, Lab. Pers. in Mém. Acad. St. Petersbg., ser.
7, XXI, Nr. 1, 51 (1873); species group F (Hedge, 1974a;
Notes Roy. Bot. Gard. Edinburgh 33(1): 1–121).
Type: S.aegyptiaca L.
Low growing shrubs or suruticose herbs, oen
appearing as dwarf shrubs, usually <40cm, rarely up to
50 or 60cm (Table); stems much branched. Leaves small,
simple, narrow linear-elliptic, rarely obovate-oblong
(P. bariensis, occasionally in P. aegyptiaca) and pinnatid,
leaf margins usually revolute, thick textured. Flowers small,
up to 16mm, usually less than 10mm. Verticillasters up
to 10, few-owered [(1)–2(–8)], with short-lived owers.
Calyces somewhat enlarging in fruit, slightly reexed
upper lip. Corolla upper lip ± straight, occasionally shorter
than or equal to lower lip; white, pale violet to pink; tube
with annulus, subannulate (P. macilenta) or exannulate
(P. tebesana). Stamens at least partly exposed; connective
very short; lower thecae fertile, sometimes reduced
(very small or even sterile), e.g., in P.eremophila; within
P. herbanica both fertile and sterile lower thecae appear.
Nutlets small (up to 2 × 1.4mm), black, mucilaginous on
wetting.
Distribution and ecology: typical elements of the
Saharo-Sindian phytogeographical region, usually locally
restricted to arid habitats on the Canary Islands, in North
Africa, on the Arabian Peninsula (Oman, Saudi Arabia,
Yemen), and in SW Asia (Syria, Lebanon, Israel, Jordan,
Palestine, Iran, Afghanistan, Pakistan); generally on sandy
soil, gravel wadi beds and basalt rocks; on open limestone
(P. bariensis) and basaltic slopes (P. herbanica); well-
adapted to intense sunlight and water deciency during
the summer, e.g., by reduced number and size of leaves,
small, ± linear leaves with revolute margin; xeromorphic
features (well-developed cuticula, lower surface of the
leaves with stomatal grooves, mesophyll of the leaves
composed of palisade parenchym only, well-developed
chlorenchyma in the wings and anks of petioles and
cortex of the stems); indumentum variable with trichomes
of dierent types: simple, eglandular, retrorse hairs and oil
globules; short spiky eglandular hairs; very short to long
spreading eglandular hairs; stalked glands; 1- to 5-celled
trichomes.
Referring to the characteristic growth form of Pleudia
(e.g., Figure 3A), we propose the vernacular name
dwarf-sage’ for the new genus. e trivial name ‘desert-
sage’, used for dierent representatives of Salvia s.l. that
are also restricted to dry or desert-like habitats, is not
advised. It refers to a group of species molecularly and
morphologically clearly distinct from Pleudia, i.e. S.dorrii
(Kell.) Abrams, S.eremostachya Jeps., S.pachyphylla Epl. ex
Munz (all restricted to America; Clade II), and S.deserta
Schang (Eurasia; Clade I) (Quattrocchi, 1999).
A taxonomic revision of Pleudia based on eldwork
is still needed to clarify species delimitations. Due to the
lack of comprehensive studies on morphology, anatomy,
karyology, and ecology, we do not provide a key to the
species here.
Pleudia aegyptiaca (L.) M.Will, N.Schmalz & Class.-
Bockh., comb. nov.
Basionym: Salvia aegyptiaca L., Sp. Pl. 1: 23 (1753).
non S.aegyptiaca L. Mant. Pl. 26 (1767).
Type: ‘Habitat in Aegypto’; Herb. Cliord 13, Salvia 17
(BM-000557609) (photo!); lectotype dened by Hedge in
Notes Roy. Bot. Gard. Edinburgh 33(1): 89 (1974a).
Icon.: Hedge in: Rechinger, Fl Iranica (Tabulae), Vol.
150; Tab. 468 (1982).
Synon.: Pleudia aegyptiaca (L.) Raf., Fl. Tellur. 3: 94
(1837).
= Melissa perennis Forssk., Fl Aegyptiaca: LXVIII no.
296; Descriptiones plantarum: 108; Cent. IV no. 30 (1775).
Type: syn. C; C10002589 (photo!), C10002590
(photo!); C10002591 (photo!); Fl. Aeg. Arab. no. 296, p.
108 Cent. 30; Herb. Forskål no. 341, ‘in desertis Kahirinis’
C10002592 (photo!)
= Salvia arida Salisb., Prodr. Stirp. Chap. Allerton: 73
(1796).
WILL et al. / Turk J Bot
702
= ymus hirtus Viv., Fl. Libyc. Spec.: 30 (1824), nom.
illeg.
Type: Africae-borealis, in collibus arenosis Magnae
Syrteos, 1817; P. Della Cella s.n.
Icon.: Viviani, Fl. Libyc. Spec., Tab. XIV, g. 1 (1824).
ymus syrticus Spreng., Syst. Veg. 2: 697 (1825). nom.
nov. for . hirtus Viv.
= Salvia pumila Benth., Labiat. Gen. Spec.: 726 (1835).
Type: caulibus erecti, oribus diluti violaceis, violaceo
punctatis, odore teucrii botrydis, herb. V. Jacquemont 74
K0090121 (photo!); ‘Hab. in collib. gypsosis et salinis juxta
PindadenKhan 6. Apr.’ P00714654 (photo!); MPU (!).
Icon.: Jacquemont, Descriptions des collections
botanique, t. 133, (1844). Salvia aegyptiaca var. pumila
(Benth.) J.D. Hook., Fl. Br. Ind. Vol. IV: 656 (1885) p.p.
Salvia aegyptiaca var. pumila (Benth.) Aschers. &
Schweinf. ex I.Löw Sitzungsber. Kaiserl. Akad. Wiss., Phil.
Hist. Cl. Vol. 161(3), p. 28 (1909).
= Salvia aegyptiaca var. glandulosissima Bornm. &
Kneuck., Allg. Bot. Z. Syst. 22(1-4): 4 (1916).
Type: Sinaihalbinsel, im NW der Halbinsel, im Wâdi
Fîran, 25. März 1904, Hans Guyot s.n.; Sinaihalbinsel, ‘zw.
den Tälern Sahâra, es-Sahîr, Cscheib ect. im SW. am 24.
April 1904 und im SO. zw. den Tälern ab-Orta und Chreise
am 2. Mai 1904 gesammelt.’; A. Kneucker s.n.
Salvia aegyptiaca f. colorata Maire, Bull. Soc. Hist. Nat.
Afrique N. 23: 205 (1932).
Type: ‘rocailles calcaires à Agadir-n-Ighir’
= Salvia aegyptiaca var. intermedia E.Peter, Repert.
Spec. Nov. Regni Veg. 39: 182 (1936).
Type: Punjab; Drummond 14429
= Salvia gabrieli Rech.f., Bot. Jahrb. Syst. 71: 538 (1941).
Type: Iranisch-Baločistan: Bashakird, Paß Pohki
zwischen Anguhran und Ispand, 1635 m; a 1928, A.
Gabriel no. 57; (holo. W)
= Salvia aegyptiaca f. albiora Sauvage, Mém. O. Nat.
Anti-Acridien 2: 34 (1947).
Type: Mauritanie septentrionale; Zemmour, Bir
Moghrein, Kedia Kheneijat; 5. Nov. 1942, ‘. entièrement
blanches’, coll. Ch. Runge & Sauvage, R.S. no. 30;
MPU005370 (photo!)
Pleudia areysiana (Deers) M.Will, N.Schmalz &
Class.-Bockh., comb. nov.
Basionym: Salvia areysiana Deers, Bull. Soc. Bot.
France 43: 229 (1896).
Type: P.D.R.Y. (Peoples Democratic Republic of
Yemen; South Yemen), ‘Bilad Fodhil, ad fauces australes
montis el-Areys, prope Serreya, 27. April 1893’, Deers
1041; (holo. P)
holo.: P00714656 (photo!), P00714657 (photo!),
P00714658 (photo!)
iso.: MPU (!)
Pleudia bariensis (ulin) M.Will, N.Schmalz &
Class.-Bockh., comb. nov.
Basionym: Salvia bariensis ulin, Opera Bot. 121: 145
(1993).
Type: Somalia, Bari Region, Al Miskat Mts, Bahaya;
Hab.: stony limestone slope; shrublet, 0.2–0.4m; owers
blue; 11°18N, 49°49E; 26. Nov. 1986, ulin & Warfa
6059; (holo. UPS)
holo.: UPS:BOT:V-041526 (photo !)
iso.: K, MOG
Icon.: ulin, Opera Bot. 121: 146, g. 1 (1993).
Pleudia bazmanica (Rech.f. & Esfand.) M.Will,
N.Schmalz & Class.-Bockh., comb. nov.
Basionym: Salvia bazmanica Rech.f. & Esfand., Oesterr.
Bot. Z. 99: 61 (1952).
Type: Persiae prov. Balučistan (Makran): Bazman, 11.
March 1949, Sharif 1150 E, (holo. W).
Icon.: Rechinger, Oesterr. Bot. Z. 99: 61, g. 10a (1952);
Hedge in: Rechinger, Fl Iranica (Tabulae), Vol. 150; Tab.
469 (1982).
Placement of this species in Pleudia is based on
distribution and habitat preferences, growth form, and the
combination of dierent morphological characters, such
as ower morphology; it was not sampled in a molecular
study.
Pleudia chudaei (Batt. & Trab.) M.Will, N.Schmalz &
Class.-Bockh., comb. nov.
Basionym: Salvia chudaei Batt. & Trab., Bull. Soc. Bot.
France 53 (Sess. extrao.): XXX (1906 publ. 1907).
Type: Sahara Central; Algeria: Ahaggar mts., Tit, 6.
Aug. 1909, Chudeau; (holo. MPU) holo.: MPU010317!
Icon.: Bull. Soc. Bot. France 53 (Sess. extrao.): t. 10
(1906 publ. 1907).
Synon.: Salvia chudaei var. typica Bull. Soc. Hist. Nat.
Afrique N. 23: 205 (1932).
= Salvia chudaei Batt & Trab. var. lanuginosa Maire.
Bull. Soc. Hist. Nat. Afrique N. 34(6): 138 (1943)
Type: Sahara méridional, Tibesti, Sommet du mont
Toussidé, 3000m, coll. 1939-1940, . Monod s.n.
= Salvia tibestiensis A.Chev., Bull. Soc. Bot. France 78:
322 (1931).
Type: Plantes du Borkou. Tibesti récoltées par M. Jean
Tarrieux s.n., Nov. 1930; Tibesti, station dans les Oueds,
endroits hum., renseignts divers-Hauteur P00541281
(photo!), P00541282 (photo!); Salvia chudaei var.
tibestiensis (A.Chev.) Maire, Bull. Soc. Bot. France 78: 322
(1931).
= Salvia chudaei var. tefedestica Maire, Bull. Soc. Hist.
Nat. Afrique N. 23: 205 (1932).
Type: Algeria; Hab.: in arenosis ad radices montium
Tefedest Saharae centralis: Tehi-n-Beidigen inter montes
Tefedest & Ahaggar-n-Deren; ad alt. c. 1200m; 12. Apr.
1928; Maire 953; (holo. MPU)
WILL et al. / Turk J Bot
703
holo.: MPU002988 (photo!)
Placement of this species in Pleudia is based on
distribution and habitat preferences, growth form, and the
combination of dierent morphological characters, such
as ower morphology; it was not sampled in a molecular
study.
Pleudia deserti (Decne.) M.Will, N.Schmalz & Class.-
Bockh., comb. nov.
Basionym: Salvia deserti Decne., Ann. Sc. Nat. Paris
sér. 2, 2: 248 (1834).
Type: Rabeja Arab. désert du Sinaï, June 1832, N. Bové
s.n.; (holo. P)
holo.: P00714698 (photo!)
Pleudia eremophila (Boiss.) M.Will, N.Schmalz &
Class.-Bockh., comb. nov.
Basionym: Salvia eremophila Boiss., Diagn. Pl. Orient.
Ser. 1,5: 12 (1844).
Type: In desertis Persiae australis, P.M.R. Aucher-Eloy
5194; (holo. G, iso. K, G);
holo.: G00156032 (photo!)
iso.: G00098506 (photo!); G00098507 (photo!);
K000479419 (photo!); P00714729 (photo!); P00714730
(photo!)
Icon.: Hedge in: Rechinger Fl Iranica (Tabulae), Vol.
150; Tab. 467 (1982).
Placement of this species in Pleudia is based on
distribution and habitat preferences, growth form, the
combination of dierent morphological characters, such as
ower morphology, and leaf anatomy; it was not sampled
in a molecular study.
Pleudia geminata (ulin) M.Will, N.Schmalz &
Class.-Bockh., comb. nov.
Basionym: Salvia geminata ulin, Nordic J. Bot. 27:
336 (2009).
Type: Yemen, Al Mahrah Region, Ras Fartak, above Al
Wadi, 15°40N, 52°11E; Hab. rocky slope; 7. Nov. 1998, M.
ulin, B.-A. Beier & Mohammed A. Hussein 9629; (holo.
UPS, iso. K, para. K, UPS)
holo.: UPS:BOT:V-095774 (photo !)
iso.: K00248959 !
para.: Yemen, Al Mahrah Region: 35 km northeast
of Itab along road to Qishn; Hab. Rocky hillside; plant
forming small cushions; owers pale blue; 15°24N,
51°35E, 5. Nov. 1998, M. ulin, B.-A. Beier & Mohammed
A. Hussein 9569, UPS:BOT:V-095713
Icon.: ulin, Nordic J. Bot. 27: 337, g. 1 (2009).
Pleudia herbanica (A.Santos & M.Fernández) M.Will,
N.Schmalz & Class.-Bockh., comb. nov.
Basionym: Salvia herbanica A.Santos & M.Fernández,
Lazaroa 9: 52 (1986 publ. 1988).
Type: Fuerteventura; circa Vigán, 350m s.m. 22. Febr.
1985, F. de la Roche no. 29239; Rarissima; (holo. ORT).
Icon.: Santos and Fernández, Lazaroa 9: 53, g. a–f
(1986).
Pleudia hillcoatiae (Hedge) M.Will, N.Schmalz &
Class.-Bockh., comb. nov.
Basionym: Salvia hillcoatiae Hedge, Notes Roy. Bot.
Gard. Edinburgh 40(1): 69 (1982). Type: Sultanate of
Oman; Dhofar, Wadi Shibun, ‘Ramaida, 11. Febr. 1947,
esinger s.n.; (holo. BM)
Icon.: Hedge, Notes Roy. Bot. Gard. Edinburgh 40(1):
70, g. 3 (1982).
Placement of this species in Pleudia is based on
distribution and habitat preferences, growth form, and
morphology; it was not sampled in a molecular study.
Pleudia macilenta (Boiss.) M.Will, N.Schmalz &
Class.-Bockh., comb. nov.
Basionym: Salvia macilenta Boiss., Diagn. Pl. Orient.
Ser. 1,5: 13 (1844).
Type: Oman, ad radices montium regni Mascatensis,
P.M.R. Aucher-Eloy 5210, s.d. (holo. G, iso. W)
holo: G00156030 (photo!)
iso.: G00098509 (photo!); P00714705 (photo!);
P00714706 (photo!); MO-149605 (photo!)
syn.: BM001125654 (photo!)
Placement of this species in Pleudia is based on
distribution and habitat preferences, growth form, leaf
anatomy, and the combination of dierent morphological
characters, such as ower morphology; it was not sampled
in a molecular study.
Pleudia santolinifolia (Boiss.) M.Will, N.Schmalz &
Class.-Bockh., comb. nov.
Basionym: Salvia santolinifolia Boiss., Diagn. Pl.
Orient. Ser. 1,5: 13 (1844).
Type: Persia, in desertis ad sinum Persicum, P.M.R.
Aucher-Eloy 5214 s.d. (holo. G, iso. W, G, P).
holo.: ‘desert ad sin. Persicum‘ G00156029 (photo!)
iso.: G00098510 (photo!); P00714786 (photo!); ‘Salvia
a. S. deserti, 931, desert ad sin. Persicum’ P00714787
(photo!); P00714788 (photo!); MO-149608 (photo!)
Icon.: Hedge in: Rechinger Fl Iranica (Tabulae), Vol.
150; Tab. 462, 463 (1982).; Hedge in: Ali & Nasir Fl.
Pakistan, Vol. 192 (1990), accessed online: Tropicos.org.
Missouri Botanical Garden. 5. March 2014; http://www.
tropicos.org/Image/100165672.
Pleudia tebesana (Bunge) M.Will, N.Schmalz & Class.-
Bockh., comb. nov.
Basionym: Salvia tebesana Bunge, Labiat. Persic.: 52
(1873) in: Mém. Acad. Scienc. Petersbg. Sér. 7, 21: 52
(1873).
Type (syn.): [Iran] in praeruptis calcareis supra Tebbes
et prope Meibut inter Jezd et Isfahan, Mai 1859, Bunge &
Bienert; (syn. G, P); nec non in rupe Tacht-i Rustam prope
Isfahan Bode (G)
WILL et al. / Turk J Bot
704
A.A. von Bunge s.n., 30. April 1859 G00098508
(photo!); inter Jezd et Isfahan, Mai 1859, A.A. von Bunge
s.n. G00156031 (photo!); syn.: inter Jezd et Isfahan, Maj
1859, A.A. von Bunge s.n. P00714813 (photo!)
Icon.: Hedge in: Rechinger Fl Iranica (Tabulae), Vol.
150; Tab. 466 (1982).
Synon.: = S.lacei Mukerjee, Notes Roy. Bot. Gard.
Edinburgh 19: 304 (1938).
Typus: Baluchistan, Chappar Ri, Lace No. 3976; in
Cal et co-typus in E;
Iso.: E00301998 (photo !), E00301999 (photo !)
Placement of this species in Pleudia is based on
distribution and habitat preferences, growth form, leaf
anatomy, and the combination of dierent morphological
characters, such as ower morphology; it was not sampled
in a molecular study.
Pleudia trichocalycina (Benth.) M.Will, N.Schmalz &
Class.-Bockh., comb. nov.
Basionym: Salvia trichocalycina Benth., in: de Candolle
A, Prodr. 12: 356 (1848).
Type: Herb. Late East India Comp. 3984, Grith 791
(holo. K); K000929791 (photo!); K000929792 (photo!)
Icon.: Hedge in: Rechinger Fl Iranica (Tabulae), Vol.
150; Tab. 464, 465 (1982).
Taxon of uncertain status probably belonging to
Pleudia:
Salvia halaensis Vicary, Journ. As. Soc. Bengal 16: 1165
(1847).
No specimen is cited in the original description (Hedge,
1982b, 1990) and no specimens bearing the name have
been traced so far. e description most likely refers to a
representative of the Salvia aegyptiaca-group. According
to Hedge (1990), P. aegyptiaca and P. santolinifolia antedate
P.halaensis. Further ambiguity arises from the assumption
that P. halaensis might be the earliest valid name for
P.  t e b es a n a Bunge (Hedge, 1982b). Without further
material it is not possible to decide whether it is a distinct
species or synonym to one of the corresponding Pleudia
species.
Acknowledgments
We thank the herbaria GOET, K, and especially MPU
for providing plant material and Prof M ulin (Uppsala
University, Sweden) for contributing plant material
and photographs. We thank S Scholz (Fuerteventura)
for information about P. herbanica; Prof D Albach
(Oldenburg University, Germany), Dr K Challis (Kew,
UK), Dr T Oliver (SANBI, South Africa), Prof J Reveal
(Cornell University, NY, USA), and especially Prof J
McNeill (Edinburgh, Scotland, UK) for their assistance
with taxonomic questions; Dr AJ Moore (Providence, RI,
USA) for proof reading; and the three reviewers for their
critical comments on the manuscript.
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The family Ranunculaceae, a member of early-diverging eudicots that is increasingly being used as a model for the study of plant developmental evolution, has been the focus of systematic studies for centuries. Recent studies showed that the family can be divided into 14 tribes, with Glaucideae, Hydrastideae, and Coptideae being the successive basal-most lineages. The relationships among the remaining 11 tribes, however, remain controversial, so that a clear picture of character evolution within the family is still lacking. In this study, by sequencing, assembling and analyzing the chloroplast (cp) genomes of 35 species representing 31 genera of the 14 tribes, we resolved the relationships among the tribes and genera of the Ranunculaceae and clarified several long-standing controversies. We found that many of the characters that were once widely used for taxonomic and systematic considerations were actually results of parallel, convergent or even reversal evolution, suggestive of unreliability. We also found that the family has likely experienced two waves of radiative evolution, through which most of the extant tribes and genera were generated. Notably, both waves of radiation were correlated with the increase in the temperature of the earth, suggesting that global warming may have been the driving force of the radiation events. Based on these observations, we hypothesize that global warming and the associated decrease in the type and number of animal pollinators may have been the main reason why taxa with highly elaborate petals as well as those without petal were generated during each of the two waves of radiation.
... An examination of interfamilial relationships within family Ranuncuaceae using 26S rDNA by Ro et al. (1997) showed that Actaea/Cimicifuga are sister groups to Eranthis based on MP analysis. Later Compton and Culham (2002) included Eranthis in the tribe Actaeeae along with Actaea (including Cimicifuga and Souliea), Anemonopsis and Beesia. The sister-group relationship of Eranthis and Actaea was also discussed by Kosuge et al. (1995); more interestingly, Hoot (1995), on the basis of atpB, rbcL and 18S nuclear rDNA sequence data, nested Eranthis within the Actaea/Cimicifuga clade. ...
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A comprehensive morphological and anatomical study was carried out on seeds of 28 species from three tribes and eight genera of subfamily Helleboroideae (Aconitum, Actaea, Caltha, Cimicifuga, Delphinium, Eranthis, Megaleranthis and Trollius) and two putatively related genera in Ranunculaceae (Adonis and Ranunculus) using scanning electron and light microscopy to evaluate seed characteristics for use in the examination of systematic relationships. Considerable differences were found in seed coat morphology and anatomy both among and within genera of the subfamily. There are four major types of seed coat surface: striate, lineate, colliculate and irregularly wrinkled. The shape of testal cells was either elongated rectangular, rectangular chiseled, irregular or polygonal to subpolygonal. The wall ornamentation was predominantly smooth and either without any ornamentation or having finely granulated or some ribbon like appendages. The mechanical layer of the seed coat was of the exotestal type except in all species of Eranthis, in which the seed coat mechanical layer was absent; such a seed coat was referred to as being an 'undifferentiated seed-coat'. Maximum parsimony analysis of morphological features establishes three groupings within the studied genera: Aconitum/. Delphinium, Actaea/. Cimicifuga, and Caltha/. Eranthis/. Trollius/. Megaleranthis. This study is congruent with the earlier groupings of the Helleboroideae based on morphology and also agrees in part with recent molecular studies. Our data convincingly support a close relationship between Caltha- Trollius- Megaleranthis and between Actaea and Cimicifuga. Another group supported strongly by the results of this study is Aconitum-Delphinium.
... з триби Helleboreae до триби Actaeae Spach (≡ Cimicifugeae Torr. & A. Gray) (Compton et al. 1998a(Compton et al. , 1998bCompton 2002). Серед інших новин сучасної систематики жовтицевих слід відмітити зведення родів Actaea L. та Souliea Franch. ...
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This paper is a continue of previous work (Новіков 2013) and represents new key for identification of genera from Ranunculaceae family of Western Ukraine (Lviv, Transcarpathian (Zakarpattia), Rivne, Volhynia, Ivano- Frankivsk, Chernivtsi, Ternopil and Khmelnitsk regions (oblasts)). This key is consensual and do not corresponds to contemporary taxonomical tendencies in all its points because it is build for non-specialists and/or young scientists which need easy identification tool for their routine field and herbarium work. However, most of modern taxonomical changes and related publications are shortly introduced here and in my previous paper (Новіков 2013). Hence, in present work I suggest that family Ranunculaceae in Western Ukraine is represented by 2 subfamilies, 10 tribes, 22 genera and 102 species. Corresponding consensual taxonomical system is represented. Short characteristics of genera are completed by information about species number and its distribution.
... 62:1). Clade D corresponds to tribe Cimicifugeae of Compton and Culham (2002). A synapomorphy for this clade is lack of benzylisoquinoline alkaloids (char. ...
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Previous phylogenetic analyses of Ranunculales, which have mostly been focused on an individual family and were based on molecular data alone, have recovered three main clades within the order. However, support for relationships among these three clades was weak. Earlier hypotheses were often hampered by limited taxon sampling; to date less than one-tenth of the genera in the order have been sampled. In this study, we used a greatly enlarged taxon sampling (105 species, representing 99 genera of all seven families in the order). Our study is, furthermore, the first to employ morphology (65 characters) in combination with sequence data from four genomic regions, including plastid rbcL, matK and trnL-F, and nuclear ribosomal 26S rDNA to reconstruct phylogenetic relationships within Ranunculales. Maximum parsimony and Bayesian inference were performed on the individual and combined data sets. Our analyses concur with those of previous studies, but in most cases provide stronger support and better resolution for relationships among the three main clades retrieved. The first, comprised solely of the monogeneric family Eupteleaceae, is the earliest-diverging lineage. The second clade is composed exclusively of taxa of Papaveraceae, which is sister to the third clade, the core Ranunculales, comprising the other five families of the order. Circaeasteraceae and Lardizabalaceae form a strongly supported clade. Pteridophyllum is supported as sister to Hypecoum, contradicting the viewpoint that the former is the earliest-diverging genus in Papaveraceae. Glaucidium is basalmost in Ranunculaceae. Within this phylogenetic framework, the evolution of selected characters is inferred and diagnostic morphological characters at different taxonomic levels are identified and discussed. Based on both morphological and molecular evidence, a classification outline for Ranunculales is presented, including the proposal of two new subfamilies, Menispermoideae and Tinosporoideae in Menispermaceae and a new tribe, Callianthemeae, for the genus Callianthemum (Ranunculaceae).
... Wang & al., , 2009). Several analyses focus on selected subfamilies (W.) or tribes (e.g.,Ro & al., 1999;Compton & Culham, 2002;Emadzade & al., 2010) within the family. Furthermore, circumscriptions of selected genera have been improved, such as Clematis (Miikeda & al., 2006) and Anemone (Meyer & al., 2010, and therein). ...
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Calathodes and Megaleranthis are two eastern Asian endemic genera of Ranunculaceae with controversial tribal position and/or taxonomic status. In this study, we used matK sequences to determine the tribal position of Calathodes and Megaleranthis within the family, and three molecular loci (matK, trnL-F, ITS) and morphological data to further clarify their phylogenetic relationships and taxonomic status. All analyses show that Calathodes and Megaleranthis belong in the tribe Adonideae, and that Calathodes is monophyletic and closely related to Trollius. The latter two genera share rhizomes not well developed and pollen grains with striate exine ornamentation, but markedly differ in petals, follicles, receptacles, pollen grains, and chromosomes. Trollius contains two major clades with different geographical distributions. Megaleranthis is embedded in clade I and sister to Trollius macropetalus, with which it shares fruits with long beaks and a 7 bp deletion in the trnL-F dataset. Adonideae are redelimited as consisting of three genera, Adonis, Calathodes and Trollius. Morphological synapomorphies for the tribe are lacking.
Article
Calathodes and Megaleranthis are two eastern Asian endemic genera of Ranunculaceae with controversial tribal position and/or taxonomic status. In this study, we used matK sequences to determine the tribal position of Calathodes and Megaleranthis within the family, and three molecular loci (matK, trnL‐F, ITS) and morphological data to further clarify their phylogenetic relationships and taxonomic status. All analyses show that Calathodes and Megaleranthis belong in the tribe Adonideae, and that Calathodes is monophyletic and closely related to Trollius. The latter two genera share rhizomes not well developed and pollen grains with striate exine ornamentation, but markedly differ in petals, follicles, receptacles, pollen grains, and chromosomes. Trollius contains two major clades with different geographical distributions. Megaleranthis is embedded in clade I and sister to Trollius macropetalus, with which it shares fruits with long beaks and a 7 bp deletion in the trnL‐F dataset. Adonideae are redelimited as consisting of three genera, Adonis, Calathodes and Trollius. Morphological synapomorphies for the tribe are lacking.
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The use of DNA sequence data in plant systematics has brought us closer than ever to formulating well-founded hypotheses about phylogenetic relationships, and phylogenetic research keeps on revealing that plant genera as traditionally circumscribed often are not monophyletic. Here, we assess the monophyly of genera documented in Rothmaler's "Exkursionsflora von Deutschland" (Gefäßpflanzen: Grundband, 19th Ed.; Jäger 2005). Using a survey of the phylogenetic literature, we discuss which classifications would be consistent with the phylogenetic relationships found and could be followed, provided monophyly is accepted as the primary criterion for circumscribing taxa. We indicate whether and which names are available when changes in generic assignment are made (but do not present a comprehensive review of the nomenclatural aspects of such names). Among the 840 genera examined, we identified c. 140 where data quality is sufficiently high to conclude that they are not monophyletic, and an additional c. 20 where monophyly is questionable but where data quality is not yet sufficient to reach convincing conclusions. While it is still fiercely debated how a phylogenetic tree should be translated into a classification, our results could serve as a guide to the likely consequences of systematic research for the taxonomy of the German flora and the floras of neighbouring countries.
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Caltha is a widely distributed genus in the buttercup family (Ranunculaceae) showing interesting distribution patterns in both hemispheres. Evolutionary history of Caltha was examined by means of phylogenetic, molecular dating, and historical biogeographic analyses with a more comprehensive sampling than previous studies. The internal transcribed spacer from the nuclear genome and trnL-F and atpB-rbcL regions from the plastid genome were used and analyzed using parsimony and Bayesian methods. Divergence time was estimated using Bayesian dating analyses with multiple fossil calibrations. Historical biogeography was inferred using the Bayes-DIVA method implemented in RASP. We obtained a well-resolved and well-supported phylogeny within the Caltha lineage. Caltha natans Pall. diverged first from the genus and the other species grouped into two clades. Our expanded sampling scheme revealed a complicated evolutionary pattern in the C. palustris complex. Caltha sinogracilis W. T. Wang was resolved to be a member of the C. palustris complex, rather than closely related to C. scaposa Hook. f. & Thomson. Caltha rubriflora B. L. Burtt & Lauener was also revealed to be not just a red-flower form of C. sinogracilis. The diversification of the genus began at 50.5 mya (95% high posterior density: 37.1–63.9 mya), and its ancestral range was very probably in the Northern Hemisphere. The South American species may derive from western North American ancestors that dispersed along the western American Cordillera during the Cenozoic era. The vicariance model of the Southern Hemisphere species proposed by a previous study was rejected in this study.
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Cimicifugeae is one of the rich sources for various active components and the health promoting and therapeutic values of the components have been corroborated by long-term use in folk medicine and traditional Chinese medicine. Increasing interest in Cimicifugeae pharmaceutical resources has led to the further discoveries of triterpenoid saponins, phenolic compounds, chromones, and many other compounds in various species of Cimicifugeae, and to the investigations on their chemotaxonomy, molecular phylogeny, and bioactivities. Based on our pharmacophylogenetic studies, the progress in phytochemistry, chemotaxonomy, molecular biology, and phylogeny of Cimicifugeae had been summarized since 2007, especially Cimicifuga L. ex Wernisch. and Actaea L., and their relevance to therapeutic efficacy. An exhaustive literature survey is used to characterize the global scientific effort in the phytochemical and biological studies of Cimicifugeae. More triterpenoid saponins have been found in various species, among which the cimigenol type (type A) is predominant. The versatile bioactivities of saponins and extracts, as well as those of phenolics and other ingredients, were summarized and discussed. The morphology-based five-genus classification of Cimicifugeae is not supported by molecular phylogeny. Molecular phylogeny based on nuclear and chloroplast DNA sequences tends to merge Cimicifuga Wernisch., Souliea Franch., and Actaea L. into a single genus. It is indispensable to integrate the emerging technologies into Cimicifugeae research for both the sustainable utilization of Cimicifugeae pharmaceutical resources and finding novel compounds with potential clinical utility and less adverse effects. Systems biology and omics technologies would play an increasingly important role in booming pharmaceutical research involving bioactive compounds of Cimicifugeae.