Edraianthus canescens (Campanulaceae), a new species from the Central Balkan peninsula
Author
Lakušić, Dmitar
Department for Plant Ecology and Phytogeography, Faculty of Biology, University of Belgrade, Takovska 43, 11000 Belgrade, Serbia. E-mail: dlakusic @ bio. bg. ac. rs, tamaraz @ bio. bg. ac. rs, vstev @ bio. bg. ac. rs
Author
Niketić, Marjan
Natural History Museum, Njegoševa 51, 11000 Belgrade, Serbia. E-mail: mniketic @ nhmbeo. rs
Author
Rakić, Tamara
Department for Plant Ecology and Phytogeography, Faculty of Biology, University of Belgrade, Takovska 43, 11000 Belgrade, Serbia. E-mail: dlakusic @ bio. bg. ac. rs, tamaraz @ bio. bg. ac. rs, vstev @ bio. bg. ac. rs
Author
Stevanović, Vladimir
Department for Plant Ecology and Phytogeography, Faculty of Biology, University of Belgrade, Takovska 43, 11000 Belgrade, Serbia. E-mail: dlakusic @ bio. bg. ac. rs, tamaraz @ bio. bg. ac. rs, vstev @ bio. bg. ac. rs
text
Phytotaxa
2013
2013-07-19
118
1
22
28
http://dx.doi.org/10.11646/phytotaxa.118.1.3
journal article
10.11646/phytotaxa.118.1.3
1179-3163
5079260
Edraianthus canescens
D. Lakušić, Niketić & Stevanović
,
sp. nov
.
Figs. 1
&
2.
—
E. serbicus
sensu
Šmarda (1968: 45)
—
E. graminifolius
„
jugoslavicus
“ sensu Stefanovi ć
et al.
(2008: 470)
Type:
—
SERBIA
.
W
Serbia
:
Ov
č arsko-Kablarska gorge, foothill of
Mt. Kablar
,
Orlove
stene, rock crevices, limestone,
43°54.362' N
,
20°12.060' E
,
308 m
,
16 May 2008
,
D. Lakuši
ċ
26617
(
holotype
:
BEOU
!, isotypes:
BEOU
!,
NHMR
!)
.
FIGURE 1
.
Edraianthus canescens
D. Lakušić, Niketić & Stevanović
,
sp. nov.
(from the holotype). A. Habitus. B. Rosette leaf. C. Bracts of flowering capitula. D. Calyx. E. Anther. F. Style. G. Corolla. H. Capsule with axicorn (linedrawing V. Stevanović).
FIGURE 2
Edraianthus canescens
D. Lakušić, Niketić & Stevanović
,
sp. nov.
(from the locus classicus). A. Habitus. B. Inflorescence with involucral bracts. C. Flowers with calyx teeth. D. Rosette leaves. E. Cauline leaves. F. Inflorescence. G.. Capsules with basal lateral pores and axicorns. H. Capsule with axicorn. I. Seed. (photo D. Lakušić).
Whole plant densely greyish hirsute, closest to
E. graminifolius
, from which it differs by the indumentum on basal leaves (densely hirsute and greyish on both sides of leaves vs. glabrous with ciliate-fimbriate margins or fine deflexed-pilose on both sides of leaves), trichome orientation (trichomes raised directed to the leaf apex, or orientated in all directions except towards the leaf base vs. trichomes raised directed only to the leaf base) and particular capsule dehiscence (capsule with basal lateral porose dehiscence vs. capsule with irregular apical rupture dehiscence).
Caespitose perennial with herbaceous stems. Rhizome stout, somewhat woody, with a branched caudex. Stem simple (4−)9−17(−22) cm, erect to ascending, leafy, densely hairy, with a nested like structure of the dense dried leaves at the base; dried stems remain on the plants for several years. Leaves narrowly linear, with flat margin, pointed at the top, densely greyish hirsute on both leaf sides, entire, densely covered with raised slightly curved hairs directed to the leaf apex, or orientated in all directions except towards the leaf base, hairs (0.2−)0.3(−0.5) mm long; rosette leaves 50−97(−129) × (1.3−)1.7−3.2(−4) mm, rigid end erect, form a globular cushion from which several stems grow out; cauline leaves sessile, (17−)27−51(−71) × (2.3−)3.2− 6.2(−9) mm, usually with a wide amplexicaul base. Involucral bracts 6−12(−19), shorter or equal with the flower, densely hirsute and greyish above, entire, rarely slightly denticulate, whitish to pale green at the base, densely subtend the flower; inner bracts lanceolate to ovate-oblong, (8.3−)13.7−21.3(−27) × (4.3−)6−9.5(−13) mm; outer bracts subovate-lanceolate, slightly attenuate, (17.9−)18.7−32.5(41) × (4.5−)7.0−11.2(−14) mm. Inflorescence with (3−)5−9(−11) subsessile flowers in a terminal cluster. Calyx greyish to pale green, hirsute, (4.3−)5.0−6.8(−7) mm in diameter; calyx teeth narrowly lanceolate, pointed at the top, 2−3 times as long as the ovary, (6.2−)8.0−11.2(−14) × (0.7−)
0.8−1.2 mm
, commonly without setulose or ciliate appendages. Corolla narrowly campanulate, violet, glabrous, hirsute on veins and corolla lobes, (18.8−)21.1−28.5(−36) × (12.3−)13.6−18.3(−20) mm; corolla lobes (6.3−)8.3−12.2(−15) × (4.9−)6.1−8.3(9) mm. Style (18.9−)20.8− 26.2(−33) mm long, 2−3-lobed. Stamens 5, inserted on disc; anthers (5.9−)6.3−8.8(−11) mm long; filaments 0.9−2.0(−3.0) mm long, in lower part distinctly dilated to deltoid-shaped
1−4 mm
long structure. Capsule pale brownish, opening by basal lateral pores and with irregular apical rupture; axicorn very strong and prominent, with two strong terminal lobes. Seeds numerous, elliptic-ovate, light brown 1.4−1.8 ×
0.8−1.1 mm
.
Etymology:
—The specific epithet is derived from the main and prominent characteristic of the plant⎯densely hirsute and greyish indumentum on the stem as well as on the both sides of leaves and involucral bracts.
Chromosome number:
—2n
=
32 (counted on plants from locus classicus).
Distribution and ecology:
—
Edraianthus canescens
is distributed in a very narrow area restricted to the Ovčarsko-Kablarska gorge only (western
Serbia
). Closest populations of the
E. graminifolius
complex (
E. jugoslavicus
) are situated c.
30 km
west-, south- and northwards from the
type
locality (
Fig. 3
).
E. canescens
belongs to the eastern Dinaric (Illyrian) local endemic species.
The new species primarily inhabits the calcareous south-facing exposed rocky crevices at elevations between 300 and
750 m
.
Edraianthus canescens
is one of the most abundant rock dwellers growing together with
Asplenium ruta-muraria
L.,
A. trichomanes
L.,
Leontodon crispus
DC. ex Nyman
,
Helianthemum canum
(L.) Baumg.,
Micromeria thymifolia
(Scop.) Fritsch
,
Scabiosa graminifolia
L.,
Seseli rigidum
Waldst. & Kit.
,
Genista triangularis
Willd.
etc. Except in the rocky crevices, rare individuals of
E. canescens
were recorded in vegetation of the calcareous screes, growing together with
Carex humilis
Willd. ex Kunth
,
Euphorbia subhastata
Vis. & Pančić
,
Seseli rigidum
Waldst. & Kit.
,
Fraxinus ornus
L.,
Jurinea mollis
Rchb.
,
Leontodon crispus
DC. ex Nyman
,
Clematis vitalba
L.,
Silene vulgaris
(Moench) Garcke
,
Sesleria juncifolia
Suffren
s.l.
,
Laserpitium siler
L.,
Lembotropis nigricans
(L.) Griseb., etc.
Conservation status:
—
Edraianthus canescens
is known only from its
type
locality. Its population size is estimated to be less than 2000 mature individuals while the area of occupancy is smaller than
1 km
2
. Therefore according to the
IUCN (2001)
Criteria it should be regarded as Critically Endangered, CR B1 i, ii, iv; B2a.
Additional specimens examined (
paratypes
):
—
SERBIA
.
W
Serbia
:
Ovčarsko-Kablarska
gorge, in rupium fissuris prope
Ovčar Banja
ad radices montis
Kablar
, solo calcareo,
300 m
.
s.m,
20 May 1974
,
E
.
Mayer
,
D
.
Trpin
,
T
.
Wraber
39602
(
LJU
!);
Ovčarsko-Kablarska
gorge, limestone rocks,
24 August 1978
,
V
.
Nikoli
ċ
,
N
.
Dikli
ċ
,
S
.
Mladenovi
ċ (
BEO
!);
Ovčarsko-Kablarska
gorge, limestone rocks,
21 Jun 2004
,
M
.
Niketi
ċ,
20040606/017
(
BEO
!);
Mt Kablar
, screes in stands of
Ostrya carpinifolia
woodland, limestone,
12 Jun 2005
,
M
.
Niketi
ċ
20050602/008
(BEO!),
M
.
Niketi
ċ
19958
(
BEOU
!);
Ovčarsko-Kablarska
gorge,
Orlove
stene (“
Eagle’s
cliffs”),
43°54.362' N
,
20°12.06' E
,
Asplenietea rupestris
, limestone,
308.2 m
,
17 July 2006
,
V
.
Stevanovi
ċ,
D
.
Lakuši
ċ
20960
(
BEOU
!);
Ovčarsko-Kablarska
gorge,
Orlove
stene (“
Eagle’s
cliffs”),,
43°54.362' N
,
20°12.06' E
,
Asplenietea rupestris
, limestone,
308.2 m
,
17 May 2007
,
D
.
Lakuši
ċ,
T
.
Raki
ċ,
24020
(BEOU!)).
FIGURE 3.
Distribution of taxa of the
Edraianthus graminifolius
complex in C. &. S Balkan Peninsula. Arrow indicates stenoendemic range of
E. canescens
. Country abbreviations: GR: Greece, AL: Albania, MA: Macedonia, BU: Bulgaria, SR: Serbia, BH: Bosnia and Hercegovina.
Discussion
:—The newly described species,
Edraianthus canescens
belongs to the
E. graminifolius
complex (sensu Stefanović
et al.
2008) which includes plants with long linear basal leaves, sessile flowers, either solitary or in terminal cluster, closely subtended by large leaf like or ovate bracts, narrowly lanceolate calyx teeth, which are pointed at the top, 2-3 times as long as the ovary and commonly with setulose or ciliate appendages; leaves and bracts are ciliate-fimbriate at margins only or fine deflexed-pilose on both sides, with hairs directed to the leaf base.
Edraianthus canescens
is easily distinguished from its closest relatives in the
E. graminifolius
complex by the trichome orientation and the particular capsule dehiscence (Tab. 1,
Fig. 2
).
An exceptionally dense greyish-green indumentum of leaves, stems, bracts and calyx, at first glance, is the most striking feature of the new species. However, unlike all the other representatives of the complex
E. graminifolius
,
which have trichomes raised directed to the leaf base, the new species has trichomes raised directed to the leaf apex, or orientated in all directions, except towards the leaf base. The significance of trichome orientation in differentiating
Edraianthus
taxa has been recently discussed by Surina
et al.
(2009).
In addition, unlike other representatives of the
E. graminifolius
complex with pronounced indumentum (specially in the southern distribution range⎯
E. siculus
Strobl 1883: 551
,
E. australis
(
Wettstein 1887: 201
) Lakušić ex
Meyer 2011: 154
,
E. horvatii
Lakušić 1974: 44
,
E. pubescens
Meyer 2011: 156
whose indumentum density may vary substantially within the same population (from densely hairy to completely glabrous plants), all plants from the Ovčarsko-Kablarska gorge have a very dense gray indumentum.
Molecular data indicate that plants from the Ovčarsko-Kablarska gorge are clearly distinguished from other taxa with exceptionally dense indumentum from Southern Balkan and Sicily. Parsimony bootstrap supports (BS) and Bayesian posterior probabilities (PP) strongly indicates that
E. horvatii
(S.
Macedonia
⎯BS 89 %, PP 100 %),
E. australis
(S.
Greece
⎯BS 66 %, PP 100 %) and
E. siculus
(Sicily⎯BS 85 %, PP 100 %), are phylogenetically separated from the plants from the Ovčarsko-Kablarska gorge, representing “
molecularly distinct lineages
” (Stefanović
et al.
2008).
The capsule with the basal lateral porose dehiscence is undoubtedly the most important character that separates the new species from all other representatives of the
E. graminifolius
complex. The importance of fruit dehiscence in
Edraianthus
and its position within
Campanulaceae
is observed and recently discussed in detail by Stefanović
et al.
(2008). In that publication it was stated that in some individuals of
Edraianthus
some fruits open apically while other fruits open laterally, reminiscent of those found in
Campanula
. Taking into account that different states of capsule dehiscence can be found even within a single individual, it is clear that the taxonomic significance of the capsule dehiscence is limited for the higher-level family classification in
Campanulaceae
. Basal pores and prominent axicorn are extremely rare features, registered only in two strictly endemic
Edraianthus
species
(
E. tare
Lakušić 1987: 108
from sister group of the
E. graminifolius
complex and
E. glisicii
Č ernjavski & Soška 1937: 88 from
E. serpyllifolius
complex, according to Stefanović
et al.
2008). These characters have therefore high taxonomic significance at species level.
Furthermore, the overall habitus of the plants with a nested like structure of dense dried leaves at the base and dried stems that remain on the plants for several years, is a very specific morphological feature of
E. canescens
.
In addition to the above-described morphological characteristics, the plants from the Ovčarsko-Kablarska gorge are separated from other representatives of the
E. graminifolius
complex from the central Balkans by the shape of the bracts. Correspondence Analysis (CA) showed that, in respect to the qualitative characters analyzed,
E. canescens
is significantly distinct from all studied populations within the
E. graminifolius
complex (Rakić
et al.
2012). Except for the extremely dense indumentum, resulting in the greyish color of the plant surface, these plants are characterized by the broad and short apex of the involucral bracts.
Taking all this into account, we conclude that the plants from the Ovčarsko-Kablarska gorge are morphologically clearly defined and can be easily distinguished from all the other species from
E. graminifolius
complex, and hence, they merit rank of new species.