37 (2): (2013) 113-120
Original Scientiic Paper
Sesleria serbica (Poaceae), a neglected species of the
Balkan Peninsula
Nevena Kuzmanović1✳, Snežana Vukojičić1, Zoltán Barina2
and Dmitar Lakušić1
1 Institute of Botany and Botanical Garden Jevremovac, Faculty of Biology, University of Belgrade, Takovska 43,
11000 Belgrade, Serbia
2 Department of Botany, Hungarian Natural History Museum, Könyves Kálmán krt. 40, 1476 Budapest, Hungary
ABSTRACT: Sesleria serbica is a neglected taxon traditionally wrongly synonymised with S. rigida. Our
multifaceted study conirmed its speciic status. Herewith we comprehensively describe its
morpho-anatomical and ecological features. It is an obligate serpentinophyte, and a protected
species in Serbia. Regarding its distribution, S. serbica is a Balkan endemic species (local
endemic of the eastern part of the Illyrian province). Nomenclatural and taxonomical notes
are also provided, as well as the national conservation status of this species.
Key words: Sesleria serbica, Poaceae, neglected species, morpho-anatomy, ecology, chorology
Received 25 May 2013
Revision accepted 30 July 2013
UDK 582.542.11(497)
INTRODUCTION
he species, treated now as Sesleria serbica (Adam.) Ujhelyi,
has been irst recognised by Adamović (1896) as a variety
of S. rigida Heuf. ex Rchb., and later raised to speciic level
by Ujhelyi (1959a). Traditionally, the taxon S. serbica
was included in the complex S. rigida (section Calcariae
Deyl, sub-section Rigida Deyl; Deyl 1946). It was mostly
treated as a lower infraspeciic taxon or just as a serpentine
ecotype of the species S. rigida (Deyl 1946, 1980; Diklić
& Nikolić 1986; Tatić 1976), and only rarely as a “good
species”, S. serbica as proposed by Ujhelyi (Ujhelyi 1959a;
Stevanović et al. 1995, 2003; Euro+Med 2006).
Following Ujhelyi, detailed molecular, morphoanatomical and chorological-ecological studies have
been carried out with the aim to describe and conirm
clear diferences that occur between S. serbica and other
members of S. rigida s.l. (Kuzmanović et al. 2009,
2012). he inal conirmation and circumscription of
S. serbica was published in Kuzmanović et al. (2013),
where it was shown that whithin the complex S. rigida
four species can be recognized – S. achtarovii, S. ilifolia,
correspondence: nkuzmanovic@bio.bg.ac.rs
✳
S. rigida and S. serbica. hese results have shown that
the genetic diferentiation pattern was relected by
morphological diferentiation, where leaf morphology and
especially anatomy play a primary role in morphological
discrimination of the constituents of S. rigida s.l. In
addition, further discrimination can be obtained using
the characters of overall habit such as the shape of tuts
(stoloniferous vs. compact) and spikes (lax and interrupted
vs. dense and compact).
he aim of the present work is to describe in
detail the anatomical, morphological, ecological and
phytogeographical specialty of the species S. serbica, as
well as to provide information regarding taxonomical and
nomenclatural issues.
MATERIAL AND METHODS
Plant material was collected during ive vegetation seasons
(2008–2012). Anatomical and morphometric analyses
were performed on dissected plant organs preserved in
50% ethanol (leaves) or in 1:1 glycerol/ethanol solution
(stems with spikes).
© 2013 Institute of Botany and Botanical Garden Jevremovac, Belgrade
vol. 37 (2)
114
Table 1. Accession used for morphometric analyses.
Locality
Coordinate
Altitude
Habitat
Voucher
Individuals
Serbia, Tara mountain,
Paljevine
43.8769 N, 19.4168 E
950
black pine forests
(Erico-Pinetum nigrae)
Lakušić, D. 27106
26
Serbia, Kopaonik,
Nebeske stolice
43.2605 N, 20.8298 E
1850
subalpine pasture
(Festuco-Seslerietea)
Lakušić, D. 27617
15
Serbia, Kopaonik,Treska
43.2604 N, 20.7854 E
1600
subalpine pasture
(Festuco-Seslerietea)
Jakovljević, K.,
Kuzmanović, N.
28735
28
Serbia,
Mokra Gora 2
43.7319 N, 19.6431 E
800
black pine forests
(Seslerio-Pinetum nigrae)
Kuzmanović, N.
28739
26
Serbia,
Mokra Gora 1
43.8286 N, 19.5268 E
900
black pine forests
(Seslerio-Pinetum nigrae)
Kuzmanović, N.
28740
26
Serbia, Zlatibor,
Zlatiborska jezera
43.8142 N, 19.5132 E
1000
black pine forests
(Seslerio-Pinetum nigrae)
Kuzmanović, N.
28741
30
Serbia, Gornji Milanovac,
Brdjani’s gorge
43.9934 N, 20.421 E
300
rocky grasslands
(Festuco-Brometea)
Lakušić, D. et al.
28817
24
Serbia, Gornji
Milanovac,Vujan
43.9858 N, 20.4478 E
550
rocky grasslands
(Festuco-Brometea)
Lakušić, D. et al.
28830
28
Serbia, Maljen, Ljuti krš
44.1256 N, 19.9981 E
950
black pine forests
(Seslerio-Pinetum nigrae)
Kuzmanović, N.
29507
26
Bosnia and Herzegovina,
Gornja Maoča
44.7698 N, 18.6555 E
280
black pine forests
(Seslerio-Pinetum nigrae)
Kuzmanović, N. et al.
29515
22
Bosnia and Herzegovina,
Gornja Maoča
44.7606 N, 18.6526 E
300
rocky grasslands
(Festuco-Brometea)
Kuzmanović, N. et al.
29516
28
Analysis of 27 anatomical leaf characters was
performed on cross-sections of tiller leaves as described
by Kuzmanović et al. (2009). 23 macromorphological
characters were analysed following combined and
adjusted methods used for Festuca L. (Auquier 1974;
Lakušić 1999; Foggi et al. 1999, 2006) and Sesleria
Scop. (Alegro 2007; Di Pietro 2007). he anatomical
measurements were performed on the cross-sections
of 277 tiller leaves (each obtained from diferent
individuals, 11 populations). Anatomical analyses of
the leaves were done on permanent hand–made slides,
prepared by a standard method for light microscopy.
Cross sections of the tiller leaves were cleared in Parazone
and thoroughly washed before staining in safranin (1 %
w/v in 50 % ethanol) and alcian blue (1 % w/v, aqueous).
he measurements were performed using a Leica
Qwin (Leica Microsystem, Germany) and Digimizer
Image Analysis sotware (MedCalc Sotware, Belgium).
Statistical analyses were performed using Statistica 5.1
(StatSoft 1996). Voucher specimens were deposited at
BEOU. he information regarding populations used for
the morphometric analyses is provided in Table 1.
Chorological and ecological data were based on recent
ield studies, analysis of herbarium material deposited at
BEOU, BEO, BP, BUCA, GZU, PR, PRC, SOM, W and
WU (herbarium acronyms follow Thiers 2013), as well as
critically-evaluated literature data. All species-occurrence
data are stored in a Microsot Excel 2003 database.
Following the principles and methods of data cleaning
and data quality (Chapman 2005a, 2005b), each original
record was carefully reinterpreted, so that in addition
to primary species data which included taxonomic and
nomenclatural information and original spatial attributes
(verbatim data) every row in the database contained an
optimum of accurate and precise data on habitat, substrate,
altitude, aspect and Universal Transverse Mercator
(UTM)-coordinate in the Military Grid Reference System
(MGRS). Distribution data were mapped on the grid
N. Kuzmanović et al: Sesleria serbica (Poaceae), a neglected species of the Balkan Peninsula 115
map with squares of 10 km × 10 km, based on the UTM
projection according to Lampinen (2001).
RESULTS AND DISCUSSION
History of Sesleria serbica. he irst nomenclatural
element that can be associated with the name S. serbica is
the herbarium specimen collected in 1893 and deposited
in the Herbarium Generale collection in the Hungarian
Natural History Museum (BP 593596!) - “Sesleria rigida
Heuf. var: longifolia m. In saxosis ad Gornji Milanovac.
Solo serpentino. V. 1893 L. Adamović”. Since Lujo
Adamović did not publish this name with the description
or diagnosis, it can be considered as “nomen nudum”
(Art. 38 Ex. 1, McNeill et al. 2012). In the same year,
in April and May, he collected a number of specimens
that were deposited in more European herbaria mainly
under the name S. rigida Heuf. (WU 0042093!, GZU
259577!). Furthermore, in the special collection of
Josef Velenovský in PRC herbarium there is a specimen
collected in April 1893 by Adamović “In saxosis ad Gornji
Milanovac” that was originally determined by Adamović
only as Sesleria, and probably sent to Velenovský for
revision. Velenovský added the determination “Sesleria
rigida Heuf. var.”, and wrote probably to Adamović that
the plant represented a variety of S. rigida unknown to
him (J. Štepanek 2009, pers.comm.). Probably ater the
correspondence with Velenovský, and on the basis of
his own investigations, Adamović decided to describe
S. rigida var. serbica (Adamović 1896) - “Sesleria rigida
Heuf. v. serbica mihi, Rhizomate valde elongate tenui vix
dense caespitoso, culmis elatiborius, foliis pungentibus
infernos culmos aequantibus vel superantibus, spica
elongate laxa interrupta. In rupestribus calcareis montis
Vujan prope Gornji Milanovac, ca 500 m. Aprili lorens”.
From the locality it is apparent that this referred to the
same taxon that he had previously named S. rigida var.
longifolia, but with a diference on the geological substrate
on which the plant was collected. Another specimen was
found in W herbarium collected near Gornji Milanovac in
the year when the protologue reference was published (W
1897-0006747!) - “Sesleria serbica Adamov. In rupestribus
circa Gornji Milanovac. Loc. class. Solo calc. Apr. 1896 L.
Adamović”. he protologue was published in Allgemeine
Botanische Zeitschrit Syst. 2 (No. 7/8) – July and August,
so the specimen collected in the same year in April can
be unambiguously associated with the protologue and
treated as part of the original material. However, the
specimen from PRC appeared as the most appropriate for
the nomenclatural type of the name S. rigida var. serbica
(hence for the name S. serbica also), so it was designated
as the lectotype in Kuzmanović et al. 2013 (PRC451931!,
Figure 1).
Fig. 1. Lectotype of Sesleria serbica (Adam.) Ujhelyi (PRC 451931)
Ater Adamović, Ujhelyi (1959a) in his work Species
Sesleriae generis novae elevated this taxon to the species
level under the name S. serbica (Adam.) Ujhelyi: “Sesleria
serbica (Adam.) Ujhelyi pro specie.”, and indicated that
synonyms are Adamović’s S. rigida var. serbica and S.
rigida var. longifolia (in herb.). In this paper he gave an
extensive description, however, instead of typifying the
new combination by the basyonym type, he gave a new
holotype for the taxon, which was not in accordance with
Art 9.19 (McNeill et al. 2012). herefore, the “holotype”
he let in the Herbarium of the Natural History Museum
in Budapest (BP 734294!) cannot be considered as the
nomenclatural type for S. serbica. Regardless of this mistake
that he made, during his work on the genus Sesleria, József
Ujhelyi gave a signiicant contribution to the knowledge of
this complicated grass genus (Ujhelyi 1938, 1940, 1959a,
1959b, 1960; Ujhelyi & Felfoldy 1948).
Morphological description. Plant perennial, broadly
caespitosae (up to 1m in diameter), rhizome very
elongated and without reticulate basal leaf sheaths (Figure
1A). Leaf sheaths yellowish-green, glabrous. Culms (15.7) 23.8-42.6 (-55.9) cm tall, glabrous. he uppermost culm
116
vol. 37 (2)
Fig. 2. Sesleria serbica (Adam.) Ujhelyi. A. Plant on the locus
classicus (Mt. Vujan; photo D. Lakušić); B. Leaf transverse section,
C. Spikelet, D. Seed
Fig. 3. Distribution of Sesleria serbica (Adam.) Ujhelyi (UTM Grid
zone T34, spots correspond to the basic square of 10 × 10 km)
leaf is 0.20-1.66 cm long. Ligules short, ciliate. Tiller
leaves 4.5-45.3 cm long, setaceous, prostrate or slightly
upright, never erect or rigid, dark green, shiny. he leaves
are rolled around the central nerve (convolute), and their
leaf blade form varies from the oval to elliptical (Figure
2B). Surface of the leaf blades ranges from 152.01 to
523.05 mm2. Width of the leaf blades ranges from 0.60 to
1.42 mm. hickness of the leaf blades in the zone of the
central vein varies from 0.18 to 0.29 mm, and the largest
thickness from 0.14 to 0.28 mm. On the adaxial side of the
leaf there is only a central rib whose width varies from 0.09
to 0.19 mm. he mesophyll is not diferentiated to spongy
and palisade tissues. It is built of the chlorenchyma cells
which fulill all the space not covered by sclerenchyma
or vascular bundles and their sheath layer. he vascular
bundles make just one row and they are located in the
mesophyll, close to the epidermis of the adaxial side of the
leaf. hey are diferent in size, so it is easy to distinguish
the “major” (big) and “minor” (small) bundles. All the
bundles have elliptic form and are surrounded by one
layer of cells, making the sheath of the vascular bundle.
he major vascular bundles have clearly developed big
tracheas. Number of major vascular bundles varies from
3 to 7. Minor vascular bundles are small, without or with
hardly noticeable big tracheas. Number of minor vascular
bundles varies from 3 to 12. Height of the central vascular
bundle varies from 0.05 to 0.11 mm. Width of the central
vascular bundle varies from 0.04 to 0.09 mm. Height of
the largest lateral vascular bundle varies from 0.05 to 0.11
mm and width of the largest lateral vascular bundle varies
from 0.04 to 0.08 mm. Sclerenchyma is interrupted, with
a tendency to form a continual subepidermal layer in the
older leaves. It is organized in the form of sclerenchyma
girders (from the adaxial to the abaxial side of the leaf they
descend to the vascular bundles) and strands (they do not
touch the vascular bundles). he sclerenchyma strands
on the adaxial side of the leaf are generally absent, while
on the abaxial side of the leaf they are always present. he
number of sclerenchyma strands on the abaxial side of
the leaf varies from 1 to 6. Sclerenchyma girders appear
both on adaxial and abaxial sides of the leaf in a similar
number - number on adaxial side of the leaf varies from 6
to 14, and on the abaxial side from 5 to 14. Occasionally,
the sclerenchyma strands and girders on the abaxial side
of the leaf are mutually connected, forming sclerenchyma
strands extending in a parallel way with the epidermis of
the leaf. Extremely rarely, the sclerenchyma strands are
registered on the adaxial side of the leaf as well. Height
of the sclerenchyma strand of the central vascular bundle
varies from 0.02 to 0.06 mm. Within the zone of the central
vascular bundle the sclerenchyma is organized exclusively
in the form of a sclerenchyma strand, located on the
abaxial side of the leaf. he surface of the sclerenchyma
N. Kuzmanović et al: Sesleria serbica (Poaceae), a neglected species of the Balkan Peninsula 117
varies from 13.29 mm2 to 98.69 mm2. Occasionally, in the
mesophyll some colorless cells can be observed, within the
zone of sclerenchyma girders on the adaxial side of the leaf.
On the adaxial side of the leaf the presence of moderately
to densely distributed simple hairs are observed, whose
length varies from 0.02 to 0.10 mm. hinned out hairs
are present on the abaxial side of the leaf, whose length
varies from 0.01 to 0.08 mm. Bulliform cells are present
in the highest number of the analyzed leaves, and their
dimension (expressed as a relative ratio of the lengths of
bulliform cells and neighboring cells of the epidermis of
the adaxial side of the leaf) varied in a range from 0.88 to
4.01. Spike cylindrical, elongated and slightly interrupted
(16-) 19-30 (-44) mm Í (4-) 5-7 (-8) mm, with (7-) 1116 (-20) spikelets. Spikelets on prominent pedicels, with
2-3 lowers (Figure 2C). Glumes unequal, membranous,
with a single vein, obtuse, glabrous. Lower glumes 3.47.2×1.1-2.8 mm, upper glumes 3.6-7.8×1.4-2.9. Lemma
oblong, sparsely pubescent to glabrous between the veins,
membranous, with 3-5 veins and 3-5 awns (middle awn
the largest), 3.8-6.3×1.7-3.4 mm with 0.4-1.4 mm long
awn. Palea as long as or shorter than the lemma, twoveined, shortly awned, pubescent on the veins, obtuse,
3.7-6.1×1.3-3.0 mm. Anthers 2.44-4.48 mm long. Seed
lanceolate, pubescent in the upper part, 2.05-3.24 mm
long, 0.96-1.26 mm wide, hilume 0.46-0.96 mm long
(Figure 2D).
Distribution and ecology. Sesleria serbica represents a
local endemic taxon (Balkan endemic), distributed on the
Inner Dinarides of central and eastern Bosnia (Mitrovići,
Tajan, Krivaja, Varda, Konjuh, Gostović, Gornja Maoča)
and western and central Serbia (Vujan, Brđanska gorge,
Tučkovo, Maljen, Tara, Mokra Gora, Zlatibor, Studena
planina mountain , Stolovi, Goč, Čemerno, Ibar valley,
Studenica valley, Raška, Kopaonik, Jadovnik, Stari vlah,
Ozren, Rogozna). A doubtful record from the serpentine
areas in Banija (Croatia, Sekulić et al. 1988) could not
be conirmed in the course of the present study (N.
Kuzmanović, ield observations). Chorologically, it can be
classiied as an East Illyrian endemic, i.e. local endemic
of the eastern part of the Illyrian province. Distribution
is presented on the map with squares of 10 km × 10 km
(Figure 3).
Sesleria serbica occurs only on serpentine bedrock,
and by this feature it belongs to the group of obligate
serpentinophytes (Stevanović et al. 2003). However,
in the protologue (and on some herbarium specimens),
Adamović (1896) recorded the species on limestones
of Mt. Vujan near Gornji Milanovac. Our recent ield
investigations could not conirm the presence of S. serbica
on calcaerous bedrock on mt. Vujan (where limestone
and serpentine areas have a wide contact zone); it was
exclusively on serpentine outcrops conirming its belonging
to serpentinophytes. Its populations occur at altitudes
ranging from c. 300 m a.s.l. (Brdjani gorge, Gostovićka
river gorge) to 1900 m a.s.l. (Kopaonik-Nebeske stolice),
mostly on north and northeast expositions, on the slopes
ranging from 15 up to 85 degrees. According to Deyl
(1946) it belongs to the eurythermal group of species,
which are blooming in early spring.
Regarding phytosociological features, S. serbica has an
important role in forming several types of primary and
secondary communities. On the summits of mountains
that it inhabits, it forms the primary community type
Seslerietum serbicae (Seslerietum ilifoliae Z. Pavlović 1955,
Luzulo-Seslerieteum rigidae D. Lakušić 1987) that can be
included in the class Elyno-Seslerietea Br.-Bl. 1948. On
the steep slopes, in northern expositions, on the lower
altitudes, it dominates the understory of the speciic forest
community Querco-Pinetum Z. Pavlović 1964, Seslerio
rigidae-Pinetum nigrae Gajić 1954 and Pinetum nigraesylvestris Z. Pavlović 1951 (Erico-Pinetea Horvat 1959), as
well as Seslerio-Ostryetum, Seslerio-Fagetum and SeslerioBetuletum community types (Querco-Fagetea Br.-Bl. et
Vlieg. 1937). Furthermore, on places where the forests are
degraded, it builds up the speciic secondary community
Erico-Seslerietum rigidae R. Jovanović et S. Jovanović 1985
and communities where S. serbica dominates on the open
rocky grasslands that can be included in the class FestucoBrometea Br.-Bl. et Tüxen ex Soó 1947. On the territory of
Serbia, S. serbica was recorded in many other communities,
e.g.
in
Euphorbieto
(cyparissias)-Brachypodietum
pinnati E. Vukićević 1965, Ostryeto-Quercetum petraeae
serpentinicum E. Vukićević, Helleboro-Querco-Ostrytetum
B. Jovanović 1967, Pinetum nigrae Gajić 1954, Silenetum
serbicae D. Lakušić 1987, etc. (Lakušić et al. 2005).
In Bosnia and Herzegovina, in the eastern parts, the
species S. serbica builds up communities with Viola
beckiana Fiala ex Beck (belonging to the class FestucoBrometea Br.-Bl. et Tüxen ex Soó 1947) - Sesleria rigidaViola beckiana=Sesleria serbica-Viola beckiana (Krause
& Ludwig 1957, Ujhelyi 1959a), while in central and
northwest parts the community Seslerio serbicae-Pinetum
Rt. 1970 is recorded (belonging to class Erico-Pinetea
Horvat 1959, Lakušić et al. 1977).
Conservation status. According to national legislation,
Sesleria serbica is a protected species in Serbia (Sl_
RS_5/2010), and following Stevanović et al. (1995)
it was included in the Review of the vascular lora of
Yugoslavia of international importance, evaluated as rare.
Based on our recent studies, the high number of actually
know populations, and the fact that S. serbica is usually
a dominant species of various plant communities, we feel
that the nature conservation status of the species should
be reconsidered and that it is not threatened or a red list
species, even in Serbia and Bosnia and Herzegovina, while
its occurrence needs conirmation in Croatia.
CONCLUSION
Our comprehensive study employing morpho-anatomical,
chorological and ecological data conirmed the specialty of
Sesleria serbica that deserves to be treated as a species. It can
be easily distinguished from other members of the complex
S. rigida primarily for being an obligate serpentinophyte,
while all other species are strictly calcicole, but also by
having stoloniferous, broadly caespitose tuts, spikes that
are elongated and lax (4-5 times longer than wide), and
tiller leaf hairy on both sides. Additionally, S. serbica is
restricted to the Illyrian province (Dinaric Mts. of eastern
Bosnia and western Serbia), whereas S. rigida belongs to
the Carpathian (Carpathians in Romania), S. ilifolia to
the Moesian (Balkan Mts. in eastern Serbia and western
and northern Bulgaria and Munţii Banatului in Romania)
and S. achtarovii to the Macedonian-hracian province
(Rhodope Mts. in southern Bulgaria and northeastern
Greece) with one isolated population on the island of
hassos in the Aegean province of the MacaronesianMediterranean region.
Acknowledgements — We gratefully acknowledge the
inancial support provided by the Serbian Ministry of
Science and Technological Development (project no.
173030). Support from the European SYNTHESYS
programme (Project No. HU-TAF 2956) granted to N.K.
is also acknowledged. he work of the third author was
supported by grant OTKA104443. We also acknowledge
the herbarium curators L. Pignotti (W), W. Till (WU), L.
Somlyay (BP), O. Šida (PR), J. Štepanek and M. Štefanek
(PRC), Anton Drescher (GZU) for help with searching
for the original material of the investigating taxon. Our
friends – G. & D. Mandić, J. Georgijev-Kuzmanović, I.
Arsić (Belgrade) and colleagues – M. Niketić, G. Tomović
(Belgrade), S. Redžić (Sarajevo) and A. Alegro (Croatia)
helped with ield work and collecting of the plant samples.
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120
REZIME
Sesleria serbica (Adamović) Ujhelyi (Poaceae),
zapostavljena biljna vrsta sa Balkanskog poluostrva
Nevena Kuzmanović, Snežana Vukojičić, Zoltán Barina, Dmitar Lakušić
S
esleria serbica je zapostavljen takson koji je tradicionalno sinonimiziran sa vrstom S. rigida. Naša sveobuhvatna
istraživanja potvrdila su njen status vrste. U radu je dat detaljan opis njenih morfo-anatomskih, horoloških i
ekoloških osobina. Takodje, detaljno su razmatrana i pitanja vezana za nomenklaturu i taksonomiju, kao i status
ugroženosti ove vrste. Ustanovljeno je da je vrsta S. serbica obligatna serpentinoita, rasprostranjena u istočnom delu
Ilirske provincije, u zapadnoj Srbiji i istočnoj Bosni, te da u tom smislu predstavlja lokalno endemičnu (istočnoilirsku) vrstu Balkanskog poluostrva.
Ključne reči: Sesleria serbica, Poaceae, zapostavljena vrsta, morfo-anatomija, ekologija, horologija