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Natura Montenegrina journal for science and popular science
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Snežana DRAGIĆEVIĆ
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Prof. Dr Drago Marić
Prof. Dr Milivoje Purić
Dr Vlado Pešić
Ondrej Vizi
Marko Karaman M.Sc.
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Natura Montenegrina
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4

Dear Colleagues,
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5

Poštovane kolege,
Prirodnjački muzej Crne Gore od 2001. godine izdaje naučni i
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Marko G. Karaman
glavni urednik
6

Natura Montenegrina, Podgorica, 6/2007
CONTENTS:
NATURA MONTENEGRINA, PODGORICA, 6, 2007
EDITORIAL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
INSTRUCTIONS TO AUTHORS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Foreword by editor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Riječ urednika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
I Scientific part
BOTANY . . . . . . . . . . . . . . . . .
. . .
ZOOLOGY . . . . . . . . . . . . . . . .
. . .
BIOCHEMISTRY AND
GENETICS . .
II Short communications . . . .
. . .
I Naučni dio
9 BOTANIKA . . . . . . . . . . . . . . . . . . . . 9
63 ZOOLOGIJA . . . . . . . . . . . . . . . . . . . 63
123 BIOHEMIJA I GENETIKA . . . . . . . . . . 123
151 II Kratka saopštenja . . . . . . . . . . . . 151
BOTANY:
BOTANIKA:
Rodica BERCU
VARIATION IN THE ANATOMY OF THE VASCULAR SYSTEM OF Asplenium trichomanes-ramosum L.
Nada BUBANJA, Vladimir STEVANOVIĆ, Dmitar LAKUŠIĆ
Edraianthus dalmaticus A.DC. (CAMPANULACEAE) IN MONTENEGRO
Nada BUBANJA, Snežana VUKSANOVIĆ
Onosma pseudoarenaria Schur. ssp. tridentina (Wettst.) Br.- Bl., NEW TAXON IN FLORA OF
MONTENEGRO
9-17
19-25
27-30
Rajko T R I P I Ć
CONTRIBUTION TO THE KNOWLEDGE OF THE FLOWERING PHENOLOGY AND SEX EXPRESSION IN Acer 31-52
monspessulanum L. FROM MONTENEGRO
Snežana VUKSANOVIĆ & Danka PETROVIĆ
THE FLORA AND VEGETATION OF SALT WORKS IN ULCINJ
ZOOLOGY:
ZOOLOGIJA:
Vera VUKANIĆ
CONTRIBUTION TO KNOWLEDGE OF DISTRIBUTION AND SEASONAL DINAMIC OF THE PLANCTONIC
POLYCHETA IN SOUTH ADRIATIC WATERS
Bogić GLIGOROVIĆ, Vladimir PEŠIĆ & Aleksandra ZEKOVIĆ
A CONTRIBUTION TO THE KNOWLEDGE OF THE DRAGONFLIES (ODONATA) OF THE RIVER ZETA
(MONTENEGRO)
Jelena NIKČEVIĆ
THE ORTHOPTEROID FAUNA OF THE ĆEMOVSKO SEMI-DESERT FIELD NEAR PODGORICA,
MONTENEGRO
Natalija ČAĐENOVIĆ, Tanja VUKOV
MORPHOLOGICAL TRAITS OF COMMON TOAD Bufo bufo (Bufonidae) FROM BIOGRADSKO LAKE
Natalija ČAĐENOVIĆ
A CONTRIBUTION TO THE KNOWLEDGE OF DISTRIBUTION OF SPECIES Bufo bufo IN MONTENEGRO
53-61
63-71
73-89
91-100
101-109
111-114
Lidija POLOVIĆ, Katarina LJUBISAVLJEVIĆ
MORPHOLOGICAL CHARACTERISTICS OF A POPULATION OF THE MOSOR ROCK LIZARD (Dinarolacerta
mosorensis KOLOMBATOVIĆ, 1886) (SQUAMATA: LACERTIDAE) FROM LOVĆEN MOUNTAIN
(MONTENEGRO)
115-122
BIOCHEMISTRY AND GENETICS:
BIOHEMIJA I GENETIKA:
Nebojša JOCKOVIĆ, Milica PAVLOVIĆ, Marko SABOVLJEVIĆ and Nada KOVAČEVIĆ
CONTRIBUTION TO THE CHEMICAL CONSTITUENTS OF BALKAN BRYOPHYTES: PHENOLIC ACIDS,
FLAVONOIDS, TRITERPENES AND ALKALOIDS
123-129

Natura Montenegrina, Podgorica, 6/2007
Danko OBRADOVIĆ
PCR (Polymerase Chain Reaction) IN DETECTION OF Chlamydia trachomatis AND THE OTHER
METHODS – A COMPARATIVE SURVEY
Danko OBRADOVIĆ
INFLUENCE OF TRANSGENIC PLANTS ON ENVIRONMENT
131-136
137-149
II SHORT COMMUNICATIONS:
II KRATKA SAOPŠTENJA:
Vera BIBERDŽIĆ
Caldesia parnassifolia (L.) Parl NEW SPECIES IN MONTENEGRIN FLORA
Danijela STEŠEVIĆ & Nejc JOGAN
ADDITIONS TO THE FLORA OF MONTENEGRO: Setaria verticilliformis Dumort., Setaria viridis (L.) PB.
subsp. pycnocoma (Steud) Tzvel., Impatiens balsamina L. AND Catalpa bignoniodes Walt.
Igor TOMOVIĆ & Danijela STEŠEVIĆ
Duchesnea indica (Andr.) Focke, NEW ALIEN SPECIES IN THE FLORA OF MONTENEGRO
151-152
153-160
161-163

NATURA MONTENEGRINA, PODGORICA, 6: 9-17
VARIATION IN THE ANATOMY OF THE VASCULAR SYSTEM
OF Asplenium trichomanes-ramosum L.
Rodica B E R C U 1
1 Department of Botany, Faculty of Natural and Agriculture Sciences, “Ovidius” University,
Mamaia Str. 124, 900590, Constantza, Romania, E-mail: rodicabercu@yahoo.com
Key words:
anatomy,
variation,
vascular system,
estimation,
Asplenium trichomanesramosum
SYNOPSIS
The paper provides new data regarding the variation of the
vascular system in the vegetative organs of Asplenium trichomanesramosum
L. The anatomical structure of the adventitious root
(starting up directly from the underground stem), rhizome (stem)
and the frond’s petiole and rachis exhibit a diarch type root, a
dictyostelic rhizome and an X- and T-shaped petiole and rachis
vascular system up to the subterminal rachis. Based on the variation
of the vascular system in A. trichomanes-ramosum anatomy we
used a mathematical calculation to estimate the number of the
vascular bundles in the fern vegetative organs.
INTRODUCTION
Asplenium trichomanes-ramosum L. (syn. A. viride Huds.), known as bright
green spleenwort or simple green spleenwort, is a perennial member of family A
spleniaceae. The branching rhizomes are short, creeping to erect, bearing dark brown
to black scales (C i o c a r l a n , 2000). The evergreen compound fronds, simply divided
are tufted at the crown, 4–15 (-20) cm long; persistent to dying in the autumn and
circinnate. The frond’s rachis is green and not winged.
The petioles are often shorter than the blade. The petioles are brownish or
blackish only near the base and green above. It grows in spring and fructifies in
summer. The leaves disappear slowly during the next spring (W a t s o n . &
D a l l w i t z , 2004).
In Romania it grows in shaded crevices of limestone boulders, cool and moist
places, up to an altitude above 1800m (S a v u l e s c u , 1952). Knowledge on the
variations of the vascular system organization of the fern vegetative organs is quite
limited (O g u r a , 1938, 1972, S c h n e i d e r , 1996), and that of A. trichomanes-

10
Natura Montenegrina, 6/2007
ramosum L. is almost lacking. The goal of the research was to examine the anatomy
of the adventitious root, rhizome, and frond’s petiole and rachis in order to find a
mathematical distribution function to estimate the number of the vascular bundles in
the fern vegetative organs in accordance with plant morphology, habitat and the
cutting level.
MATERIAL AND METHODS
The plant was collected from the Bicaz Gorges in the Carpathians Mountains.
Cross sections of the adventitious root, rhizome and the frond’s petiole and rachis of
A. trichomnaes-ramosum were performed using a rotary microtome. The petiole and
rachis structure were analyzed by serial cross sections (2 to 2 cm), distributed from
the base up to the rachis tip. The samples were stained with alum-carmine and iodine
green and were embedded in Canada balsam. Observations were made with a
BIOROM-T bright field microscope, equipped with a TOPICA-6001A video camera.
The microphotographs were obtained from the video camera through a computer.
RESULTS AND DISCUSSION
Cross sections of the adventitious root revealed that the cortex is composed of
2-3 layers of large parenchyma cells (Fig. 1). The inner cells of the cortex are
modified and have thick walls. Kroemer (1903) has suggested that this wall thickening
is the result of cutinized blade superpositions. Some authors have noticed the
presence of such “curious cells” around the stele and have suggested that this tissue
belonged to the stele naming it “sclerenchymatous mass” (R u s s o w 1872,
B i e r h o r s t 1971) or a “stereomic sheath” (de B a r y 1877, O g u r a 1938, B e r c u
1998, 2006). This configuration has led S c h n e i d e r (1996) to ascribe this type of
root to that of Asplenium. The stele consists of xylem and phloem and is surrounded
by the pericycle (Fig. 1). The xylem vessels are joined together towards the center by
their metaxylem vessels (two for each bundle). The protoxylem vessels (three for each
bundle) are in an exarch position and face the pericycle. The phloem sieve cells,
lacking companion cells, are located on either side of the xylem string. Thus the
adventitious root of A. trichomanes-ramosum has a diarch structure. Remarkable are
the passing cells (Fig. 1).
Cross sections of the rhizome disclosed that the epidermis consists of a single
layer of cells not covered by cuticle (Fig. 2A). Below the epidermis is the cortex,
differentiated into a sclerenchyma cortex and the inner cortex, the latter consisting of
large parenchyma cells. Remarkable is the abundance of starch grains in the cortical
cells and endodermis as well (amiliferous sheath) (Fig. 2B). As O g u r a (1938) and
B i r (1957) reported for Aspleniaceae species, the stele is a dictyostele composed of
a variable number of meristeles (in accordance with the number of foliar traces) each
surrounded by its own endodermis (starch sheath) and pericycle. The latter is

11
B e r c u : VARIATION IN THE ANATOMY OF THE VASCULAR SYSTEM …
composed of parenchyma cells regularly arranged in one row, (locally in two or even
three rows). Each meristeles is hadrocentric with a binary structure (metaxylem
vessels towards the center and protoxylem elements facing the pericycle). The pith
rays among the meristeles occur. The pith occupies the central area of the rhizome
(Fig. 2B).
Fig. 1. Cross section of the adventitious
root, x 206: C- cortex; Mx- metaxylem; Pcpericycle;
Ph- phloem; Px- protoxilem; SSsclerenchyma
sheath.
Fig 2. Cross sections of the rhizome. (A). Portion of the epidermis, cortex and stele, x 160: C-
cortex, E – epidermis, GT – ground tissue, M- meristeles, Pi- pith, PiR- pith ray. (B) A stele
vascular bundle (meristele), x 212: Ed- endodermis, Mx- metaxylem, Pc- pericycle, Ph- phloem, Pxprotoxylem.
Cross section of the petiole base exhibits an epidermis, the cortex differentiated
into an external (sclerenchzmatous) and an inner cortex, followed by a ground tissue.
The vascular system of the leaf petiole base is composed of two meristeles, equal in
size (distele). Each meristele is surrounded by an endodermis and a “special
pericycle” (A n d r e i , 1978). The meristeles are composed of centrally located xylem
elements surrounded by phloem. Protoxylem vessels are in an exarch position
whereas the metaxylem vessels are in the centre. That is attributed to the stele a
hadrocentric and distelic structure (Fig. 3).

12
Natura Montenegrina, 6/2007
Fig. 3 Cross section of the leaf petiole
base, x 83: E- epidermis, GT- ground
tissue, IC- inner cortex, M- meristele,
SC- sclerenchyma cortex.
Fig. 4 Cross section of the leaf petiole (2 cm from the leaf base). (A) General view, x 64: C- cortex,
E- epidermis, St- stele. (B) Portion of epidermis and cortex, x 272: E- epidermis, IC- inner cortex,
SC-sclerenchymatic cortex. (C) The stele vascular bundle, x 266: Ed- endodermis, Pc- pericycle, Phphloem,
Mx- metaxylem, Px- protoxylem.

13
B e r c u : VARIATION IN THE ANATOMY OF THE VASCULAR SYSTEM …
Transverse sections of the petiole, cut from 2 cm above the leaf base, reveals a
single-layered epidermis, covered by cuticle, a cortex and a centrally located stele
(Fig. 4A). Below the epidermis of the base of the petiole is the sclerenchyma cortex
(hypodermis), which consists of a few layers of compactly arranged sclerenchyma
cells. The hypodermis is internally followed by a region of ground tissue (Fig. 4B). The
stele is monofascicular. The vascular bundle is composed of centrally located xylem
vessels surrounded by phloem. The stele X-shaped xylem elements possesses
themetaxylem vessels in a central position and the protoxylem elements in an exarch
arrangement, surrounded by phloem. That is attributed to the stele a hadrocentric
structure (Fig. 4C).
Cross sections of the rachis cut from 3-8 cm above the leaf petiole disclosed the
same one-layered epidermis composed of cutinized-walled cells, covered by a thick
cuticle, a cortex and a stele. The stele is monofascicular, composed of more or less T-
shaped xylem vessels (meta- and protoxylem vessels) characteristic to most of the
Aspleniaceae species (O g u r a 1972, B i r 1957; B e r c u 2004, 2005). Phloem
surrounds the xylem elements as well (Fig. 5). Toward the subterminal leaf rachis
remarkable is the reduced number of the vascular elements, surrounded by phloem
and embedded in a homogenous mesophyll (Fig. 5A, C). The vascular elements are in
a reduced number because of the veins marginal segments formation. Under the lower
and upper epiderms of the rachis, sclerenchyma cells are present. They provide
mechanical support to the weak and delicate rachis tip. A typical hypodermis is
absent.
Fig. 5 Cross section of the leaf rachis (3-8 cm
above the leaf rachis), x 300.
A. trichomanes-ramosum vegetative organs anatomy exhibits variations in the
vascular system organization. The even and odd number of the vascular bundles
(meristeles) may depend by the plant size, and organs diameter, the plant habitat
ecotope and the cutting level. The mathematical distribution function takes into
account of the ecological parameters such as soil humidity (U), temperature (T) and
soil reaction – pH (R) (Table 1) settled by P o p e s c u and S a n d a (1998). The
distribution function was based on S ă h l e a n u (1957), C e a p o i u (1968), S i r e ţ chi
(1985) and N a s t a s e s c u et al. (1988) works.

14
Natura Montenegrina, 6/2007
The root (r) – the diameter = 0,1-0,5 mm:
r
N Aspl. t-r . = (2n+1) K (a+b+c) πD 2 = 5 ·10 · 3,14 · 0,44 2 30,3954
= = 1,028 ≈ 1 stele
4e h 29,6 29,6
The rhizome (rz) – the diameter = 1-1,5cm:
rz
N Aspl. t-r = (2n+1) K (a+b+c) πD 2 3 ·10 · 3,14 · 1,26 2 149,55
=
= = 5,06 ≈ 5 meristeles
4e h 29,6 29,6
The rachis (rh) – the diameter = 0,5-1 mm:
pt. base
a. N Aspl. t-r. = 2n K (a+b+c) πD 2 = 2 ·2·10· 3,14 · 0,69 2 = 59,798
4e h 29,6 29,6
= 2,02 ≈ 2 meristeles
rh
b. N Aspl. t-r. = (2n +1) K (a+b+c) πD 2 = 3 ·10· 3,14 · 0,92 2 = 79,73 = 1,01 ≈ 1 meristele
4e h 4 · 2,71 3 78,732
Where:
r, rz, pt. base, rh
N Aspl. t-r.– the function distribution of the vascular bundles in root (r), rhizome
(rz), petiole base (pt. base) and rachis (rh);
h – the distance (in cm) between two succesive sections;
D – the vegetative organs diameter of the plant (in mm);
e – the natural logarithm base (e = 2, 718);
K (a+b+c) – the ecological constant of the plant depending of: humidity (a),
temperature (b) and soil reaction (c);
n – the multiplication order;
2n – the even number of the vascular bundles;
2n+1 – the odd number of the vascular bundles.
NOTE: it is sufficient the estimation of
r, rz, pt. base, rh
N As t-r.
only for n=1 sau n=2 value.

15
B e r c u : VARIATION IN THE ANATOMY OF THE VASCULAR SYSTEM …
Table 1: The ecological preferences of A. trichomanes-ramosum
The species
Ecological parameters
humidity
(H)
temperature
(T)
soil reaction
(R)
*The value of the
ecological constant
(K (a+b+c) )
Asplenium
trichomanesramosum
L.
mesohydrophyte
4
microtermous
2
Slightly neutroacidophilous
4
10
CONCLUSIONS
The successive cross sections, cut from the rhizome and the frond’s petiole and
rachis of A. trichomanes-ramosum L. disclose variations in the vascular system
structure. The root is of diarch type. The rhizome is a dictyostele with a variable
number of meristeles due to the number of the foliar traces. The petiole base vascular
system is bifascicular whereas from 2 cm above the petiole base up to the rachis tip
the vascular system is monostelic, first X-shaped and then T-shaped (to the tip).
These expected results based on any earlier research permited us to find a
mathematical distribution function to estimate the number of the vascular bundles in
the fern vegetative organs in accordance with the plant size, the organs diameter, the
main habitat plant factors and the cutting level. The distribution formula can be
extended to other members of this large group of plants.
LITERATURE
A N D R E I , M . (1978): Anatomia plantelor. - Didactic and Pedagogic Publishing House,
Bucharest, p. 227-229 (in Romanian).
B E R C U , R . (1998): Anatomical Modifications of the Corm Stele in Fern Asplenium
septentrionale (L.) Hoffm. - Forestry, Sumarstvo, Belgrade 1: 35-40.
B E R C U , R . 2004: Anatomical Aspects of Asplenium adiantum-nigrum L. - J. of Biol. Res.,
vol. 2: 57-61.
B E R C U , R . 2005: Anatomy of Asplenium ruta muraria L. - Studia bot. hung., vol. 36: 16-20.
B E R C U , R . 2006: Histoanatomy of the Vegetative Organs of some Polypodiales species. -
Ex Ponto, Constantza, 232p (in Romanian).
B I E R H O R S T , D . W . 1971: Morphology of Vascular Plants. Collier - Macmillan Ldt.,
Macmillan Co. London, New York, 296-298, 300-304.
B I R , S . S . 1957: Stelar anatomy of indian Aspleniaceae, Abstract. - Proc. Ind. Sci. Congr.
Association, Calcuta, 44: 232.

16
Natura Montenegrina, 6/2007
C E A P O I U , N . 1968: Statistics methods applied in agricultural and biological experiences.
- Agro-Silvica Publishing House, Bucharest, p.201-203 (in Romanian).
C I O C Â R L A N , V . 2000: Ilustred Flora of Romania - Pteridophyta et Spermatophyta. II
edition, Ceres, Bucharest, p.90-92 (in Romanian).
d e B A R Y , A . 1877: Verigleichende Anatomie der Vegetationsorgane der Phanerogamen
und Farne. - W. Engelmann, Leipzig, p.238-349 (in German)
K R O E M E R , K. 1903: Wurzelhaut, Hypodermis und Endodermis der Angiospermenwurzel. -
Biblioth. Bot., 12: 151 (in German).
NASTASESCU, C., TENA, M., ANDREI Gh. and OTRASANU, I.
1988: Problems of Algebraical Structures. - Reip. Pop. Romanian Acad., Bucharest,
p.122-125 (in Romanian).
O G U R A , Y . 1938: Anatomie der Vegetationsorgane der Pteridophyten. In: Handbuch der
Pflanzeanatomie, Gebrüder Borntraeger, Berlin, p.128-140 (in German).
O G U R A , Y . 1972: Comparative Anatomy of Vegetative Organs of Pteridophytes, II edition,
Gebrüder Borntraeger, Berlin, Stuttgart, p.313-512.
P O P E S C U , A a n d S A N D A , V . 1998: The Summary of the Spontaneous
Cormophytes from the Romanian Flora. - Acta Bot Horti., Bucharest, p.4-7 (in Romanian).
R U S S O W , E . 1872: Vergleichende Untersuchungen betreffend die Histologie der
vegetativen und sporenbildenden Organe und die Entwicklung der Sporen der Leitbündel-
Kryptogamen. Mém. Acad. Imp. Sc., St. Petersbourg, Sér. VII, 19: 1-207 (in German).
S A V U L E S C U , T . (editor in chief). 1952: Flora Reip. Pop. Romanicae, vol. 1, Editio Acad.
Reip. Pop. Romanicae, Bucharest, p.130-133 (in Romanian).
S C H N E I D E R , H . 1996: The Root Anatomy of Ferns: a Comparative Study. - Pteridology
in Perspective, Royal Botanical Garden, Kew, Whitstable, Litho Ltd., Whitstable, Kent:
271-283.
S I R E T C H I , G h . 1985: Differential and Integral Calculation (fundamental notions). -
Enciclopedical and Stiintifical Publisher, Bucharest, p.325-326 (in Romanian)
W A T S O N , L . a n d M . J . D A L L W I T Z 2004: The Ferns (Filicopsida) of the British
Isles. - Version: 28th November 2005. http://delta-intkey.com.
Summary
VARIATION IN THE ANATOMY OF THE VASCULAR SYSTEM
OF ASPLENIUM TRICHOMANES-RAMOSUM L.
In Romania Asplenium trichomanes-ramosum L. grows in shaded crevices of
limestone boulders, cool and moist places, up to an altitude above 1800m. The goal of
the research was to examine the anatomy of the adventitious root, rhizome, and
frond’s petiole and rachis in order to find a mathematical distribution function to

17
B e r c u : VARIATION IN THE ANATOMY OF THE VASCULAR SYSTEM …
estimate the number of the vascular bundles in the fern vegetative organs in
accordance with the plant morphology, habitat and the cutting level.
The root, in cross section possesses a primary structure and the stele is a diarh
one. Remarkable is the sclerenchimatous sheet around the stele (Fig. 1).
Cross section of the rhizome discloses a single layered epidermis (Fig. 2A),
cortex, differentiated into a sclerenchyma cortex and the inner cortex, the latter
consisting of large parenchyma cells consisting of numerous starch grains followed by
an amiliferous sheath (Fig. 2B). The stele is a dictyostele composed of a variable
number of meristeles (in accordance with the number of foliar traces) each surrounded
by its own endodermis and pericycle. Each meristeles is hadrocentric with a binary
structure (metaxylem vessels towards the center and protoxylem elements facing the
pericycle). The pith rays among the meristeles occur. The pith occupies the central
area of the rhizome(Fig. 2B, C). The frond’s petiole and rachis were analyzed by serial
cross sections (2 to 2 cm), distributed from the base up to the rachis tip. The petiole
base vascular system is bifascicular whereas from 2 cm above the petiole base up to
the rachis tip the vascular system is monostelic, first X-shaped and then T-shaped (to
the tip). These expected results based on any earlier research permited us to find a
mathematical distribution function to estimate the number of the vascular bundles in
the fern vegetative organs in accordance with the plant size, the organs diameter, the
main habitat plant factors and the cutting level.
Received: 26. 11. 2007.

18
Natura Montenegrina, 6/2007

NATURA MONTENEGRINA, PODGORICA, 6: 19-25
EDRAIANTHUS DALMATICUS A.DC. (CAMPANULACEAE)
IN MONTENEGRO
Nada BUBANJA 1 , Vladimir S T E V A N O V I Ć 2 , Dmitar L A K U Š I Ć 2
1 Natural History Museum of Montenegro, Podgorica, P.O. Box 374, 81000 Podgorica,
Montenegro
2 Institute of Botany and Botanical Garden, Faculty of Biology, University of Belgrade, Takovska
43, 11000 Belgrade, Serbia
Key words:
Edraianthus dalmaticus,
Budoške bare,
Montenegro,
flora,
distribution,
ecology
SYNOPSIS
Taxon Edrianthus dalmaticus A.DC. was found during
the floristic investigations of wetlands of Budoške bare
(reservoir Vrtac) locality in the vicinity of Nikšić, and it was
the first confirmed record of the this plant species in
Montenegro. Until now seven (sensu lato) or seventeen
(sensu stricto) taxa of genus Edrianthus at the species and
subspecies level has been found in Montenegro. Some
suspicious literature data on presence of Edrianthus
dalmaticus in Montenegro were cited in literature (Janchen
1910, Hayek 1930, Rohlena 1942). In this paper we also
presented the data about the characteristics of new locality,
habitat and ecological preferences of Edrianthus dalmaticus
A.DC. in Montengro.
Ključne riječi:
Edraianthus dalmaticus
A.DC., Budoške bare,
Crna Gora,
flora,
diverzitet,
ekologija
SINOPSIS
EDRAIANTHUS DALMATICUS A.DC.
(CAMPANULACEAE) U CRNOJ GORI
Tokom florističkih istraživanja na lokalitetu Budoške
bare (akumulacija Vrtac) u okolini Nikšića pronađen je takson
Edrianthus dalmaticus A.DC. čime je po prvi put zabilježeno i
potvrđeno prisustvo ove biljne vrste u Crnoj Gori. Do sada je
na teritoriji Crne Gore zabilježeno prisustvo 7 (sensu lato)
odnosno 17 taksona (sensu stricto) roda Edrianthus na nivou
vrsta i podvrsta. U starijoj florističkoj literaturi za Crnu Goru
se navode i neki sumnjivi podaci o prisustvu taksona
Edrianthus dalmaticus A.DC. (Janchen 1910, Lakušić 1974,
Hayek 1930, Rohlena 1942). U ovom radu dati su podaci o
karakteristikama lokaliteta, staništa i ekologiji Edrianthus
dalmaticusa A.DC. u Crnoj Gori.

20
Natura Montenegrina, 6/2007
INTRODUCTION
Genus Edraianthus DC. has its center of distribution in the Balkans and it
represents one of the most prominent groups of endemic plants in this region.
Additional disjunct parts of this group’s range are found in the Apennines, Sicily, and
the Southern Carpathians.
The widespread distribution of Edraianthus in the Balkans, and especially its
high variability and differentiation along latitudinal and altitudinal range, defines it as
one of the rare genera of the Balkan’s flora that was a subject of four monographs
(W e t t s t e i n 1887, B e c k 1893, J a n c h e n 1910, L a k u š i ć 1974). In the last
monograph and a couple of subsequent papers detailed systematic, phytogeographic
and especially ecological investigations of this genus were provided and a new system
of classification of edraianthoid campanulas was proposed (L a k u š i ć 1974, 1987,
1988). Additionally, the family Campanulaceae was segregated by L a k u š i ć (2001)
into a new family, Edraianthaceae, while the genus Edraianthus itself into several new
genera: Protoedraianthus, Visiania, Horvatia, Edraianthus, and Blecicia (Lakušić
1987, 1988, 2001).
According to different authors the genus comprises between 13 (J a n c h e n
1910) and 45 taxa (L a k u š i ć 1974, 1987, 1988, 1989) at the species and subspecies
rank. The most significant recent floristic accounts, Flora Europaea (K u z m a n o v
1976) and Med-Checklist (G r e u t e r et al. 1984), basically accept Janchen΄s concept,
with restriction of number of species and subspecies within the genus between 10
(K u z m a n o v 1976) and 14 (G r e u t e r et al. 1984).
Genus Edraianthus has its center of distribution in the region of Montenegro with
certainly recorded presence of 7 (sensu lato) that is 17 (sensu stricto) taxa at the
species and subspecies rank (L a k u š i ć 1974, 1987, 1988, 1989). From widely
comprehensible and generally accepted taxa on the territory of Montenegro these
ones are present: E. tenuifolius (Waldst. & Kit.) A. DC., E. graminifolius (L.) A. DC.,
E. tarae R. Lakušić, E. serpyllifolius (Vis.) A. DC., E. glisicii Černjavski & Soška,
E. wettsteinii Haláscy & Baldaccii subsp. wettsteinii and E. wettsteinii subsp.
lovcenicus E. Mayer & Blečić. Additionally, these subspecies are present on the
territory of Montenegro E. glisicii subsp. majae R. Lakušić and E. serpyllifolius
subsp. pilosulus (Beck) R. Lakušić, as well as taxa E. jugoslavicus R. Lakušić
subsp. jugoslavicus, E. jugoslavicus subsp. subalpinus R. Lakušić, E. caricinus
Schott, Nyman, & Kotschy, E. montenegrinus Horak, E. vesovicii R. Lakušić and E.
zogovicii R. Lakušić which according to some authors belong to the widely accepted
taxon E. graminifolius (L.) A. DC. sensu lato (J a n c h e n 1910, K u z m a n o v 1976,
G r e u t e r et al. 1984). In order to avoid nomenclature confusion the concept of
unique genus Edraianthus has been used in this supplement.
Except of the aforesaid taxa in older floristic literature, some suspected data
about the presence of the species E. dalmaticus A. DC in Montenegro are mentioned.
The first information on the presence of this species in Montenegro was published by

21
Bubanja, Stevanović, Lakušić: EDRAIANTHUS DALMATICUS A.DC. (CAMPANULACEAE)
J a n c h e n (1910: 8) which quotes the information from Reiser’s herbarium in
Sarajevo Museum: “Zweifelhafte Standortangaben: Duga-Pässe in Montenegro
(Reiser, H. Sarajev.)”, giving a commentary on these suspicious data. Lakušić in his
monograph (1974: 17), cited probably the same herbarium specimen, for this species
states “U Dugom polju (Reiser - Herb. SARA)”; However, he has correctly situated this
locality not in Montenegro but in Dugo polje in Herzegovina. The data about the
controversial presence of the species E. dalmaticus in Montenegro were mentioned
by H a y e k (1930), R o h l e n a (1942) and L a k u š i ć (1974). Based on Janchen’s
monograph, Hayek in his "Prodromus" states for this species “In rupibus subalpinis et
alpinis. Da. BH. Mt?” (H a y e k 1930: 561). In addition, Rohlena refering to Baldacci’s
data states ”In humidis silvat. Ad veterem typographiam pr. Rijeka (Ba). Sec. cl.
Hayek dubius” (R o h l e n a 1942: 353). Lastly, a very interesting statement on general
distribution of the species E. dalmaticus can be found in Lakušić’s monograph: ”...
Additionally, there was interfering of materials from quite distant places in some
herbarium sheets, which were probably compared and left in the same sheet, but they
belong to different species, as it is the case with Reiser’s material, which refers to this
species in the surrounding of Rijeka Crnojevića in Montenegro” (L a k u š i ć 1974: 18).
During the recent field investigations and collecting material for floristic study of
wet meadows in surrounding of Nikšić, E. dalmaticus was recorded and confirmed for
the first time in Montenegro.
RESULTS
DISTRIBUTION AND ECOLOGY
The species E. dalmaticus was discovered in the locality Budoške bare
(reservoir Vrtac) which is situated in the S.E. Montenegro that is in the south part of
Nikšićko Polje (Fig. 1, 2). Budoške bare occupies the area of 13,42 km², on the
altitude of 615 m.a.s.l.. To the west part of Budoške bare there is the reservoir Slano,
to its north part it is Kapino polje, to the south there is mountain Budoš and to the east
suburb Straševina. Waters which run through the channel from Slano reservoir and
Krupac reservoir as well as from the Zeta River have the main role in the hidrography
of this region. During the period of haevy precipitations, from October to the end of
May, huge amounts of waters from the channel and the Zeta river form a temporary
reservoir Vrtac which overflows the surrounding meadows. Budoške bare is influenced
by sumediterranean climate. Limnoglacial sediments occupy a wide space in Budoške
bare and they are represented with gravel, sand and clay. Deep, brown, clay-loamy,
lessivage meaedow soils and terrains are dominant in this region.

22
Natura Montenegrina, 6/2007
Fig 1. Herbarium specimen of E. dalmaticus A.DC. – coll. Montenegro, Nikšić, Budoške bare
(42º45.214 N, 18º53.163 E), wet meadows, 615 m (Bubanja, N. 26560, 14.6.2006, BEOU).

23
Bubanja, Stevanović, Lakušić: EDRAIANTHUS DALMATICUS A.DC. (CAMPANULACEAE)
Voucher specimens:
- Montenegro, Nikšić, Budoške bare (42º45.214 N, 18º53.163 E), wet meadows,
615 m (Bubanja, N., 14.6.2006, Seed plants collection Natural History Museum of
Montenegro)
- Montenegro, Nikšić, Budoške bare (42º45.214 N, 18º53.163 E), wet meadows,
615 m (Bubanja, N. 26560, 14.6.2006, BEOU)
Edraianthus dalmaticus is endemic species (Fig. 2) distributed in Croatia
(Dalmatia - Drniš, Dolac, Klis, Mosor, Povilo, Promina, Solin, Knin, Vrlika, Muć),
Bosnia and Herzegovina (Duvanjsko polje, Glamočko polje, Livanjsko polje, Drvar-
Drobnjak, Dugo polje, Čaprazlije, Donje Bare, Lištica – Posušje, Podklečani)
(B j e l č i ć , M a y e r 1983, J a n c h e n 1910, L a k u š i ć 1974, Flora Croatica
Database 2004) and Montenegro (Budoške bare near Nikšić). It could be designated
as dinaric floristic element of adriatic submediterranean province.
Fig. 2. Distribution of
Edraianthus dalmaticus (UTM
Grid zone 34; spot correspond
with basic square10 x 10 km).
Arrow indicate new locality in
Montenegro. ? indicate
suspicious literature data on
presence of E. dalmaticus in
the surrounding of Rijeka
Crnojevića in Montenegro
(Abbreviation: Cro – Croatia,
BH – Bosnia and Herzegovina,
Mtg – Montenegro)

24
Natura Montenegrina, 6/2007
THREAT
According to its endemic distribution, as well as the fact that species inhabits
very fragile periodically flooded karst meadows, it is as a Rare (R) species included in
Global IUCN Red List of Threatened Plants (W a l t e r , G i l l e t t 1998). Also as
European endemics (species for which the total (global) distribution is restricted to
Europe), as well as species included in Global IUCN Red lists, E. dlamaticus is
defined as “target species” or “species of European importance” (O z i n g a ,
S c h a m i n é e 2005).
In addition, E. dalmaticus is defined as highly protected species in Croatia
(Anonymus 2006), as well as in the category DD - data deficient included into Croatian
Red Book (N i k o l i ć , Topić , 2005), while according to Š i l i ć (1996) it has the
status of vulnerable (V) threatened species in Bosnia and Herzegovina.
LITERATURE
ANONYMUS 2006: Pravilnik o proglašavanju divljih svojti zaštićenim i strogo
zaštićenim (16. 1. 2006.), Narodne novine (7), Zagreb.
BECK G. 1893: Die Gattung Hedraeanthus. - Wiener Illustrierte Garten-Zeitung 18:
287-299.
BJELČIĆ, Ž., MAYER, E. 1983: Edraianthus A. DC. - In: Bjelčić, Ž. (ed.), Flora Bosnae
et Hercegovinae. IV Simpetalae. Pars 4: 54-56, Sarajevo, Štamparija Trebinje
188 pp.
Flora Croatica Database 2004: - Botanički zavod, PMF, Sveučilište u Zagrebu,
http://hirc.botanic.hr
GREUTER W. and T. RAUS. 1983: Med-Checklist Notulae, 8. - Willdenovia 13: 277-
288.
HAYEK A. 1930: Prodromus Florae peninsulae Balcanicae 2. Repertorium Speciorum
novarum Regni vegetabilis, Beihefte 30(2): 337-576.
JANCHEN E. 1910: Edraianthus-Arten der Balkanländer. - Mitteilungen des
Naturwissenschaftlichen Vereines an der Universität Wien 8: 1-40.
KUZMANOV B. 1976: Edraianthus A. DC. Pp. 99-100 in Flora Europea. Vol. 4,
Plantaginaceae to Compositae (and Rubiaceae), eds. Tutin T. G., V. H.
Heywood, N. A. Burges, D. M. Moore, D. H. Valentine, S. M. Walters and D. A.
Webb. Cambridge: Cambridge University Press.
LAKUŠIĆ R. 1974: Prirodni sistem populacija vrsta roda Edraianthus DC. Godišnjak
Biološkog Instituta u Sarajevu 26: 1-129.
LAKUŠIĆ R. 2001: Phytodiversity of the Order Campanulales Juss. in Montenegro.
Pp. 70 in Prirodni potencijali kopna, kontinentalnih voda i mora Crne Gore i
njihova zaštita. Plenarni referati i izvodi iz saopštenja sa naučnog skupa,
Žabljak 20 -23. 9. 2001. Book of Abstracts, ed. Regner S. Podgorica: Institut za

25
Bubanja, Stevanović, Lakušić: EDRAIANTHUS DALMATICUS A.DC. (CAMPANULACEAE)
biologiju mora Kotor & Republički zavod za zaštitu prirode Crne Gore -
Podgorica.
LOVRIĆ A. Š., RAC, M. 1987: Doprinos fitogeografiji Svilaje i njezina povezanot sa
Biokovom. Acta Biokovica 4: 189-204.
NIKOLIĆ, T., TOPIĆ, J. eds. 2005: Crvena knjiga vaskularne flore Hrvatske. -
Ministarstvo kulture, Državni zavod za zaštitu prirode, Zagreb. 693pp.
OZINGA, W.A. & SCHAMINÉE, J.H.J. (eds.). 2005: Target Species – Species of
European Concern. A database driven selection of plant and animal species for
the implementation of the Pan European Ecological Network. Wageningen,
Alterra, Alterra-report 1119. 193 pages; 30 figs.; 18 tables; 134 refs.
ROHLENA, J. 1942: Conspectus Florae Montenegrinae. - Preslia 20-21: 1-506.
ŠILIĆ, Č. 1996: Spisak biljnih vrsta (Pteridophyta i Spermatophyta) za Crvenu knjigu
Bosna i Hercegovine - Glasnik Zemaljskog muzeja Bosne i Hercegovine
(Prirodne nauke), Nov. ser. 31: 323-367.
WALTER, K.S. & H.J. GILLETT (eds.) (1998) 1997 IUCN Red List of Threatened
Plants. Compiled by the World Conservation Monitoring Centre. IUCN - The
World Conservation Union, Gland, Switzerland and Cambridge, UK.
WETTSTEIN R. 1887: Monographie der Gattung Hedraeanthus. Denkschr. kais. Akad.
Wissensch. math.-natur. Classe 53: 185-212.
Received: 27. 11. 2007.

26
Natura Montenegrina, 6/2007

NATURA MONTENEGRINA, PODGORICA, 6: 27-30
Onosma pseudoarenaria Schur. ssp. tridentina (Wettst.) Br.- Bl., NEW TAXON IN
FLORA OF MONTENEGRO
Nada BUBANJA¹, Snežana VUKSANOVIĆ ¹
¹Natural History Museum of Montenegro, Podgorica, Montenegro
Кеy words:
Nikšićka Župa,
Montenegro,
taxon
SYNOPSIS
During the floristic researches of Niksicka Zupa, a
taxon Onosma pseudoarenaria Schur.ssp.tridentina (Wettst.)
Br.- Bl. was found and it was the first time to find this taxon
in Montenegrin flora. The flora of Europe connects the
species Onosma tridentina Wettst. to Italy, but it suspects
that it might have spread out on the territory of ex-
Yugoslavia. Med-Check list, as well as the Flora of Europe,
suspects that this taxon can be found on the territory of ex-
Yugoslavia. In the electronic database of Croatian flora one
can find this taxon under the name Onosma pseudoarenaria
ssp.tridentina (highly protected species), and in that way its
presence has been confirmed on the territory of Croatia. By
this finding the presence of Onosma pseudoarenaria Schur.
ssp.tridentina (Wettst.) Br.- Bl. on the territory of Montenegro
has been confirmed and it represents the second finding of
this taxon on the territory of ex-Yugoslavia.
Ključne riječi:
Nikšićka Župa,
Crna Gora,
takson
SINOPSIS
Onosma pseudoarenaria Schur. ssp. tridentina (Wettst.) Br.-
Bl., NOVI TAKSON U FLORI CRNE GORE
Tokom florističkih istraživanja Nikšićke Župe pronađen
je takson Onosma pseudoarenaria ssp.tridentina (Wettst.)
Br.- Bl. što je prvi nalaz ovog taksona u flori Crne Gore. Flora
Evrope vrstu Onosma tridentina Wettst. navodi za Italiju, pri
čemu izražava sumnju u njeno rasprostranjenje na teritoriji
bivše Jugoslavije. Med-Check lista kao i Flora Evrope sumnja
u nalaz ovog taksona na području bivše Jugoslavije. U
elektronskoj bazi podataka flore Hrvatske naveden je ovaj
takson kao Onosma pseudoarenaria ssp.tridentina (strogo
zaštićena biljka) čime je potvrđeno njeno prisustvo na
teritoriji Hrvatske. Ovim nalazom potvrđeno je prisustvo
Onosma pseudoarenaria ssp.tridentina na teritoriji Crne Gore
što je ujedno I drugi nalaz ovog taksona za prostor bivše
Jugoslavije.

28
Natura Montenegrina, 6/2007
INTRODUCTION
Nikšićka Župa is situated in the south-west part of Montenegro. It occupies the
area of 40 km 2 , and with its broader mountain hinterland and brim 210 km 2 . A diversity
of flora and vegetation of this region is caused by the influence of Mediterranean and
continental climate as well as high mountains (Maganik, Stitovo and Prekornica) which
surround Nikšićka Župa. Nikšićka Župa is situated east of Nikšić.
MATERIAL AND METHODS
Floristic researches of Nikšićka Župa were carried out in period August, 2000 to
September 2003. The subspecies represented in this paper was collected in June
2002, and the vouchers are entrusted to the collection of Natural History Museum of
Montenegro. The basic literature used for determination of this taxon was: Flora
Europea (L.1972) and Med-Check list (G r e u t e r , B u r d e t & L o n g 1984).
Photographs of the species (fig.1) and habitation (fig.2) where the plant was collected
from are given in this paper .
Fig.1: Onosma pseudoarenaria Schur.
ssp. tridentina (Wettst.) Br.- Bl.

29
Bubanja, Vuksanović: Onosma pseudoarenaria Schur. ssp. tridentina (Wettst.) . . .
RESULTS AND DISCUSSION
During the floristic researches of Nikšićka Župa, a subspecies Onosma
pseudoarenaria ssp.tridentina (Wettst.) Br.- Bl was found. That is the first finding of
this taxon in Montenegro. A small population was recorded on only one locality
(Preserve of King Nikola-Morakovske bare). The plant grows in open, dry and rocky
habitation in beech forest. Genus Onosma L. is represented with 33 species in Flora
of Europe. According to the existing data, these species have been registered on the
territory of Montenegro: O. stellulatum W.K., O. Aucherianum DC. ssp. Javorkae
(SIMK.) HAY., O. arenarium W.K. and O. Visianii CLEM (R o h l e n a 1942).
H a y e k (1924-1933) did not record the presence of this species on the Balkan
Peninsula. Flora Europea as well as Med-Check List mentions this subspecies only for
Italy, doubting upon its spreading on the territory of ex-Yugoslavia. This subspecies
was not recorded in the regional flora of Croatia (D o m a c 1994), whereas in
electronic database of Croatian flora it is recorded as a highly protected species.
Fig.2: Habitation where the plant was collected from.
CONCLUSION
A finding of Onosma pseudoarenaria ssp.tridentina (Wettst.) Br.- BL. in
Nikšićka Župa clearly confirm its presence in the Balkan Peninsula. In relation to Flora
Europae and Med Check List this record significantly extends the subspecies range
towards the south-east.

30
Natura Montenegrina, 6/2007
LITERATURE
BUBANJA, N. 2004: Flora vlažnih i vodenih staništa Nikšićke Župe. Specijalistički
rad.Biološki fakultet,Beograd (manuscr.), 86pp.
GREUTER W., BURDET H. M., LONG G., 1984: Med-Check List. Vol. 1., Geneve &
Berlin, 109-110
PIGNATTI S. 1982: Flora d' Italia 2. - Edagricole, 400-402.
PULEVIĆ, V. 2005: Građa za vaskularnu floru Crne Gore. Dopuna "Conspectus florae
montenegrinae" J. Rohlene., knjiga 2, Republički zavod za zaštitu prirode Crne
Gore, Podgorica, 218pp.
ROHLENA, J. 1942: Conspectus florae Montenegrine. -Presila (Praha) 20-21: 254pp.
TUTIN, T.G., BURGES, N. A., CHATER, A. O., EDMONDS, J. R., HEYWOOD, V. H.,
MOORE, D. M., VALENTINE, D. H., WALTERS, S. M. & WEBB, D. A., 1993:
Flora Europaea. Vol. 3. - Cambridge, 89-94.
Received: 27. 11. 2007.

NATURA MONTENEGRINA, PODGORICA, 6: 31-52
CONTRIBUTION TO THE KNOWLEDGE OF THE FLOWERING PHENOLOGY AND
SEX EXPRESSION IN Acer monspessulanum L. FROM MONTENEGRO
Rajko T R I P I Ć 1)
1)
Republic Institute for the Protection of Nature of Montenegro, Podgorica, Trg vojvode Bećir
bega Osmanagića 16, Podgorica, Crna Gora
Кеy words:
Flowering phenology,
sex expression,
Acer monspessulanum
SYNOPSIS
The paper contains results of research of flowering
phenology and sex expression of flowers, inflorescences and
individual trees of Acer monspessulanum consecutively for
seven years on 40 marked trees from 2 local populations
near Nikšić. The flowers were morphologically false
bisexually and functionally unisexual (male or female). The
male flowers had very small, rudimentary pistil (unfunctional)
with normally developed stamens (yellow with functional
anthers on long filaments). The female flowers had normally
developed pistil and unripe stamens (the anthers are green,
hard and unfunctional, and filaments are very short).
Ključne riječi:
Fenologija cvjetanja,
polna ekspresija,
Acer monspessulanum
SINOPSIS
PRILOG POZNAVANJU FENOLOGIJE CVJETANJA I
POLNE EKSPRESIJE U ACER MONSPESSULANUM L. IZ
CRNE GORE
U radu su dati rezultati istraživanja fenologije cvjetanja
i polne ekspresije cvjetova, cvati i individualnih stabala Acer
monspessulanum uzastopnotokom sedam godina na 40
markiranih stabala iz dvije lokalne populacije blizu Nikšića.
Cvjetovi su bili morfološki lažno dvopolni a funkcionalno
jednopolni (muški ili ženski). Muški cvjetovi su imali vrlo mali,
rudimentarni tučak (nefunkcionalan) sa normalno razvijenim
prašnicima (žuti sa funkcionalnim anterama na dugačkim
filamentima), a ženski cvjetovi su imali dobro razvijen tučak i
nezrele prašnike (sa zelenim, tvrdim i nefunkcionalnim
anterama na vrlo kratkim filamentima).

32
Natura Montenegrina, 6/2007
INTRODUCTION
Morphological and functional variations in the sexuality of Acer flowers were
studied by many explorers. P a x (1885), monographer of Acer genus, said that flowers
of Acer monspessulanum are andro-monoecious. P o j a r k o v a (1933), also
monographer of Acer genus said that flowers of A. monspessulanum are male and false
bisexual. Z a m j a t n i n (1958) said that flowers of A. monspessulanum are male with
stamens and female with pistil and reduced stamens. G u d e s k i & D r e n k o v s k i
(1978) said that flowers of A. monspessulanum are functionally unisexual independently
from development of sexual organs of the opposite sex. Male flowers do not have pistil,
and if there is pistil it is reduced and sterile. Female flowers have well-developed
stamens but their anthers do not grow-up, half-anthers do not open and they produce
sterile pollen. P a l a m a r e v (1979) said that flowers of A. monspessulanum are
bisexual and unisexual.
MATERIAL AND METHODS
Flowering phenology and sexual expression of flowers, inflorescences and trees
of Acer monspessulanum were observed consecutively in period of 7 years on marked
trees (40 trees) in 2 local populations (SUBICA and GREBICE) near NIKŠIĆ. Sexuality
of the flowers was determined on base of their sexual organs functionality. During full
anthesis of the flowers all flowers had normally developed stamens and non-welldeveloped
pistils (rudimentary or abortive pistil) are considered to be male. Criteria for
female flowers is incompetence for releasing pollen, and in case of releasing pollen
such flowers are considered to be bisexual. The criteria for a female flower will be the
inability to release pollen. In case of pollen release, fertile or not, the flowers will be
considered as bisexual. Such the opinion represents contemporary monographer of
Acer genus J o n g (1976). The sexual expression of the inflorescences is determined
on base of classification of inflorescences according to type of their flowering (W i t t r o
c k 1886; modified by C o r r e n s 1928; taken from J o n g 1976) and numeral
relationships between male and female flowers in the inflorescences is observed in
specimen of 30 randomly taken inflorescences for every individual tree and every
specific year of observation. Phenophases: swelling of buds, opening of buds and
flowering were determined according to recommendations (B e j d e m a n 1979).

33
Tripić: CONTRIBUTION TO THE KNOWLEDGE OF THE FLOWERING PHENOLOGY . . .
RESULTS AND DISCUSSION
All analysed trees were monoecious plants. The male and female flowers are both
placed on the same inflorescence. The male flowers had very small, rudimentary pistil
(unfunctional) with normally developed stamens (yellow with functional anthers on long
filaments). The female flowers had normally developed pistil and unripe stamens (the
anthers are green, hard and unfunctional, and filaments are very short). We find similar
results with G u d e s k i & D r e n k o v s k i (1978). T r i p i ć (2006) presented very
similar results for Acer obtusatum Waldst. & Kit.. It is observed, but very rarely (on 5
trees), that during the flowering 1-2 almost normally developed female flowers were
transformed into functionally male flowers with illusory normal pistil but with abortive
ovary. Such flowers are morphological illusory bisexual but in fact they are functionally
unisexual male flowers.
Phenophase of flowering in observed populations begins in time interval from 9 th
March to 14 th April (Stubica) and 25 th March to 13 th April (Grebice) and lasts from 8 to 49
days in population (Stubica) and 27 to 47 days (Grebice), depending on weather
(especially temperature), number of flowering trees and volume of flowering of each
tree (Tab. 2., 3. and Graph. 2., 4.). The end of flowering varied from 8 th April to 5 th May
(Stubica) and 4 th May to 18 th May (Grebice). Massive and rich flowering of trees in this
populations was every the third year (Tab. 2. and Graph. 2., 4.) followed with 2 years of
weak or very little flowering (small number of flowering trees and often very small
number of flowering inflorescences and flowers). The annual flowering rhythmic is very
similar in observed populations while the rhythmic between observed trees was
significantly differentiated (Table 2. and Graph. 1., 2., 3., 4.). Individual tree flowered
depending on ecological conditions and richness of flowering from 3 to 28 days (
Stubica) and 2 to 29 days (Grebice) and all analysed trees of observed populations
depending also of ecological conditions, number of flowered trees and volume of
flowering of every tree lasted from 8 to 49 days (Tab. 2., Graph. 1., 2.,3.,4.). Also, for
individual trees it is determined that there is greater or lower variation of duration of
flowering in observed years, depending on ecological conditions and volume of
flowering of that tree (when volume of flowering is low when number of flowering
inflorescences on tree is small than phenophases of such trees starts later and lasts
significantly shorter, and their inflorescences have smaller number of flowering types,
so the tree has at that time narrower sexual expression than in years of rich flowering
(Graph. 1., 2., 3., 4.).

34
Natura Montenegrina, 6/2007
Table 1. The phenophases: Swelling of the buds and opening of the buds in observed
populations of Acer monspessulanum L.
Population
Year
Swelling of the buds
Opening of the buds
Start Massive Start Massive
STUBICA
1998.
1999.
2000.
2001.
2002.
2003.
2004.
18. 2. – 6. 4.
9.3. – 17. 4.
30. 3. – 19. 4.
14. 2. – 18. 3.
16. 3. – 15. 4.
27. 2. – 18. 4.
21. 2. – 1. 4.
1. 3. – 19. 4.
16. 3. – 22. 4.
15. 4. – 25. 4.
11. 3. – 29. 3.
6. 4. – 25. 4.
16. 3. – 30. 4.
13. 3. - 8. 4.
3. 3. – 13. 4.
24. 3. – 24. 3.
11. 4. – 24. 4.
15. 3. – 7. 4.
26. 3. – 26. 4.
26. 3. – 30. 4.
19. 3. – 6. 4.
12. 3. – 27. 4.
29. 3. – 26. 4.
19. 4. – 29. 4.
19. 3. – 7. 4.
13. 4. – 28. 4.
1. 4. – 3. 5.
23. 3. – 11. 4.
GREBICE
1998.
1999.
2000.
2001.
2002.
2003.
2004.
6. 3. – 24. 4.
25. 3. – 26. 4.
2. 4. – 21. 4.
18. 2. – 31. 3.
10. 3. – 13. 4.
6.3.(21.1.)– 22.4.
28. 2. – 29. 3.
20. 3. – 28. 4.
1. 4. – 29. 4.
19. 4. – 30. 4.
11. 3. – 7. 4.
28. 3. – 28. 4.
29. 3. – 2. 5.
20. 3. – 10. 4.
3. 4. – 1. 5.
3. 4. – 1. 5.
10. 4. – 28. 4.
18. 3. – 8. 4.
27. 3. – 27. 4.
8.4.(28.1.)- 28.4.
28. 3. – 11. 4.
7. 4. – 4. 5.
8. 4. – 4. 5.
21. 4. – 3. 5.
25. 3. – 18. 4.
6. 4. – 2. 5.
14. 4. – 5. 5.
30. 3. – 17. 4.
All male flowers of individual trees flowered at lest 2 days and at most 22 days,
and female flowering at least 1 day and at most 11 days (Tab. 3., Graph. 2., 4.). It is
determined that phenophases of flowering of male flowers at all protandrous trees and
at most of protogynous trees it is significantly longer from phenophases of flowering of
female flowers, while such phases at only several protogynous trees has similar period
of lasting or it is phenophases of flowering of female flowers were longer from
phenophases of flowering of male flowers (Graph. 2., 4.). T r i p i ć (2006) presented
very similar results for A. obtusatum while results for A campestre and A. platanoides
(in press) he presented that they had good and very good flowering intensity
consequently for several years followed with one or two years of weak and very weak
flowering. That shows that species which taxonomically belong to the same section
have very similar phenology of flowering while the phenology of flowering to the species
from different sections is different.

35
Tripić: CONTRIBUTION TO THE KNOWLEDGE OF THE FLOWERING PHENOLOGY . . .
Table 2. The phenophase of flowering in the observed populations of Acer
monspessulanum L.
Population
Year
Flowering of the individual trees
Flowering of the observed population
Start Massive The end Start Massive The end
STUBICA 1998.
1999.
2000.
2001.
2002.
2003
2004.
9. 3.– 14. 4.
30. 3.–18. 4.
14. 4.–20. 4.
19. 3.–10. 4.
1. 4. – 1. 4.
1. 4. –18. 4.
24. 3. – 9. 4.
14. 3.–16. 4.
1. 4. –10. 4.
- - - - - - - -
21. 3. – 1. 4.
- - - - - - - -
4. 4. –17. 4.
27. 3. – 8. 4.
5. 4. – 26. 4.
8. 4. – 24. 4.
16. 4.–22. 4.
4. 4. – 20. 4.
7. 4. – 8. 4.
20. 4.– 5. 5.
9. 4. – 23. 4.
9. 3.
30. 3.
14. 4.
19. 3.
1. 4.
1. 4.
24. 3.
30. 3.-13. 4.*
- - - - - - - - -
- - - - - - - - -
24. 3. – 13. 4.
- - - - - - - - -
- - - - - - - - -
1. 4. – 17. 4.
26. 4.
24. 4.
22. 4.
20. 4.
8. 4.
5. 5.
23. 4.
GREBICE 1998.
1999.
2000.
2001.
2002.
2003.
2004.
4. 4. –25. 4.
6. 4. –27. 4.
13. 4.–28. 4.
25. 3.–19. 4.
2. 4. – 1. 5.
13. 4. – 1. 5.
3. 4. –16. 4.
8. 4. –20. 4.
9. 4. –30. 4.
16. 4.–27. 4.
28. 3.–12. 4.
4. 5. – 4. 5.
16. 4.–28. 4.
6. 4. –18. 4.
14. 4. - 5. 5.
15. 4. – 9. 5.
27. 4.–14. 5.
13. 4. – 5. 5.
19. 4.–18. 5.
22. 4. – 9. 5.
16. 4. – 4. 5.
4. 4.
6. 4.
13. 4.
25. 3.
2. 4.
13. 4.
3. 4.
14. 4.–27. 4.*
14. 4.– 2. 5.*
21. 4.– 3. 5.*
1. 4. – 21. 4.
- - - - - - - - -
- - - - - - - - -
5. 4. – 25. 4.
5. 5.
9. 5.
14. 5.
5. 5.
18. 5.
9. 5.
4. 5.
Legend: * = semimassive; - - - - = non-flowering
Table 3. The duration of flowering of the observed individual trees and populations of
Acer monspessulanum L. (in days).
Population
Year
The individual tree
Population
All ♂ flowers All ♀ flowers All flowers The observed tree (Total)
STUBICA
1998.
1999.
2000.
2001.
2002.
2003.
2004.
3 – 22
4 – 21
2 – 5
2 – 19
3 – 3
4 – 19
6 – 15
2 – 10
1 – 11
1 – 3
2 – 8
3 – 3
1 – 9
2 – 7
6 – 28
5 – 23
3 – 7
3 – 26
7 – 8
8 – 26
12 - 22
49
26
11
33
8
35
31
GREBICE
1998.
1999.
2000.
2001.
2002.
2003.
2004.
2 – 18
2 – 20
4 – 17
9 – 24
2 – 14
3 – 11
6 – 18
1 – 9
1 – 8
1 – 7
2 – 14
2 – 5
1 – 8
1 – 10
6 – 28
3 – 29
6 – 23
14 – 28
2 – 20
5 – 22
11 – 25
32
34
32
42
47
27
32

36
Natura Montenegrina, 6/2007
Table 4. Level of flowering (number of flowers on tree) of individual trees of Acer
monspessulanum L. during a year
Population
Year
Level of flowering
N Non-flowering Very low Low Medium High Very high
1998.
20
1
3
4
2
4
6
1999.
20
10
2
3
1
3
1
STUBICA
2000.
20
11
6
3
-
-
-
2001.
20
-
1
1
-
5
13
2002.
20
18
2
-
-
-
-
2003.
20
2
6
1
1
2
8
2004.
20
-
1
1
1
8
9
1998.
20
4
2
3
-
5
6
1999.
20
3
3
2
-
7
5
2000.
20
7
1
-
1
4
7
GREBICE
2001.
20
-
-
-
-
11
9
2002.
20
13
4
1
-
2
-
2003.
20
1
8
2
9
-
-
2004.
20
1
4
7
8
Total number of flowers on single inflorescences varied from 3 – 28 (0 – 12 of the
female and 0 – 24 of the male flowers) (Table 5., 6.) . The inflorescences with only
female flowers were very rare (it were found only 2 of such inflorescences), while
inflorescences with only male flowers were very frequent on all trees. Quantitative
relationship of the male and female flowers on the single tree was from 50 %: 50 % (1
tree) to 99.9 % of the male flowers (6 trees) (Table 5., 6.).

37
Tripić: CONTRIBUTION TO THE KNOWLEDGE OF THE FLOWERING PHENOLOGY . . .
Table 5. Frequent of number of flowers in the individual inflorescences in Acer
monspessulanum L. (Stubica)
Tree
Year
Male flowers Female flowers All flowers in an
inflorescence
Min. Max. Frequently Min. Max. Frequently Min. Max. Frequently
% of ♂
flowers
on tree
1998. 3 16 7 , 8 0 6 1 , 2 5 16 11 , 9 81.0
1 2001. 4 14 8 0 6 2 , 1 6 15 10 84.3
2004. 5 14 7 , 6 0 9 1 , 0 5 18 10 80.0
1998. 6 14 9 0 2 0 6 14 14 , 13 96.3
2 1999. 6 16 9 , 8 0 5 0 , 1 6 16 9 , 10 93.3
2001. 6 13 8 , 7 0 4 0 6 14 8 94.7
2004. 5 14 9 , 8 1 6 0 , 2 6 18 10 , 12 88.9
1998. 4 11 6 , 7 0 6 4 , 5 6 15 8 , 9 70.8
3 2001. 4 13 9 , 12 0 5 4 , 5 5 17 14 , 15 79.6
2004. 6 14 9 , 10 1 7 5 7 20 14 66.7
4 2001. 4 11 7 , 8 0 9 3 , 2 6 18 10 , 11 71.7
2004. 5 13 9 , 7 1 7 3 , 6 8 17 12 66.8
5 2001. 4 13 8 , 6 1 5 1 , 3 6 16 7 , 9 85.1
2004. 3 12 7 , 8 1 7 3 6 17 9 , 7 63.7
6 2004. 4 10 7 1 7 4 6 17 8 , 12 60.2
7 2001. 3 10 6 , 5 1 5 1 , 3 6 15 8 , 9 62.9
2004. 2 7 5 1 6 1 , 5 5 13 7 , 6 63.8
8 2001. 6 16 9 , 13 0 1 0 6 16 9 , 13 99.9
2004. 7 18 12 , 10 0 1 0 7 18 12 , 10 99.99
9 2001. 4 13 6 , 7 0 2 0 4 13 6 , 7 97.3
2004. 4 13 6 0 5 0 5 15 6 92.5
1998. 2 10 5 , 7 0 5 3 , 1 4 14 6 , 9 69.1
10 2001. 3 10 6 , 4 0 6 3 5 13 9 , 7 65.8
2004. 3 10 7 , 6 1 6 4 , 5 6 15 9 , 11 64.1
1998. 5 17 11 , 10 0 2 0 5 17 11 , 10 99.9
11 1999. 6 14 10 , 11 0 4 0 6 14 11 , 10 99.9
2001. 5 14 7 0 3 0 5 14 7 99.99
2004. 5 17 13 , 9 0 3 0 5 17 13 , 9 99.99
1999. 4 15 8 , 11 0 3 0 4 15 8 , 11 99.9
12 2001. 5 14 7 , 6 0 5 0 5 15 7 99.2
2004. 5 21 8 , 11 0 5 0 5 21 8 , 11 99.7
1998. 3 16 9 0 7 5 , 4 8 22 14 , 12 67.6
13 2001. 2 17 9 1 7 5 , 4 7 23 15 , 7 63.8
2004. 6 16 8 , 14 1 7 4 , 6 11 21 14 , 16 72.9
14 2001. 5 14 7 0 2 0 6 14 7 , 8 94.0
1998. 2 8 5 , 3 1 5 2 , 4 4 13 7 , 8 60.4
15 1999. 2 7 3 1 7 1 , 2 4 10 5 , 6 54.6
2001. 2 8 4 , 5 1 5 3 5 12 7 , 8 58.7
2004. 4 10 6 , 7 1 7 5 , 7 5 16 14 58.4
16 2001. 5 15 8 , 9 0 3 0 5 16 8 , 9 90.0
2004. 6 18 10 , 11 0 5 0 7 18 10 , 14 99.3
1998. 3 11 7 0 2 0 4 12 8 99.9
17 2001. 3 12 7 0 3 0 5 13 7 , 8 99.5
2004. 4 15 7 , 10 0 2 0 4 15 7 , 10 99.4
18 2001. 4 12 6 1 5 3 5 14 7 , 10 63.3
2004. 5 14 8 , 9 0 7 5 , 1 8 19 11 , 15 69.2
19 2001. 3 20 5 , 6 0 3 0 4 22 6 , 5 99.99
2004. 6 14 8 , 10 0 4 0 6 14 8 , 10 99.99
1998. 0 14 7 0 5 3 , 2 4 17 9 , 8 69.7
20 2001. 1 10 3 , 4 1 5 5 , 4 5 14 8 , 7 49.4
2004. 4 12 8 1 7 5 , 3 6 18 13 , 12 58.7

38
Natura Montenegrina, 6/2007
Tab. 6. Frequent of number of flowers in the individual inflorescences in Acer
monspessulanum L. (Grebice)
Tree Year
Male flowers Female flowers Total of number flowers
in inflorescences
% of ♂
flowers
Min. Max. Frequently Min. Max. Frequently Min. Max. Frequently on tree
1999. 4 13 5 and 7 1 7 5 and 3 6 18 11 and 12 66.0
1 2000. 2 15 8 and 9 1 7 3 and 5 5 19 11 and 13 72.4
2001. 3 22 8 and 7 0 7 5 and 3 5 28 12 and 13 69.5
2002. 5 17 8 and 10 0 5 1 and 2 6 19 11 and 10 81.7
2004. 4 15 6 and 11 0 7 5 and 1 5 20 15 70.3
1999. 4 11 7 and 6 1 5 1 and 3 6 14 10 and 7 74.5
2 2001. 4 12 7 0 5 3 5 16 10 75.0
2004. 4 11 6 0 5 2 and 3 7 15 9 69.0
2000. 4 15 9 and 10 1 7 5 and 4 7 18 14 and 12 68.1
3 2001. 3 10 6 and 7 1 6 1 and 4 5 15 9 69.9
2004. 6 15 8 and 12 1 6 2 and 1 7 21 15 and 10 77.7
4 2001. 5 13 6 and 5 0 2 0 5 13 6 and 5 99.99
2004. 6 16 9 and 7 0 1 0 6 16 9 and 7 99.99
2004. 3 8 6 1 5 2 6 14 9 68.5
5 2001. 2 10 4 and 5 1 5 4 and 5 5 14 6 and 8 58.6
2004. 4 10 6 1 5 5 and 3 7 14 11 64.9
1999. 4 12 7 1 6 5 6 17 11 66.0
6 2001. 5 12 7 1 6 3 and 4 7 16 11 and 14 69.0
2004. 4 12 7 and 8 0 5 2 and 0 5 15 8 and 12 77.6
1999. 6 13 8 and 9 0 9 4 and 2 7 19 14 and 12 59.2
7 2001. 6 10 8 and 9 1 8 2 and 5 8 17 13 and 11 68.9
2004. 5 10 7 and 9 0 12 3 and 4 5 20 10 and 13 60.9
2000. 8 22 11 1 5 2 and 4 10 25 13 and 17 81.6
8 2001. 8 20 15 0 7 1 and 4 9 23 18 85.2
2004. 7 24 16 0 6 1 and 3 8 25 18 85.9
1999. 4 16 8 and 10 1 6 1 and 3 5 17 9 78.7
9 2000. 5 13 8 and 9 1 5 2 and 3 7 17 12 and 14 75.6
2001. 3 12 7 and 9 1 5 3 6 16 10 and 11 73.1
10 2001. 6 19 8 0 3 0 8 19 8 and 11 91.1
2004. 7 21 12 0 3 0 7 22 13 and 14 92.7
11 2001. 4 11 7 0 3 0 4 11 7 99.5
2004. 5 11 7 and 8 0 1 0 5 11 7 and 8 99.99
12 2001. 4 14 9 and 6 1 6 4 and 3 7 18 12 and 10 72.0
2004. 5 16 11 1 7 4 and 5 7 21 14 69.9
13 2000. 3 13 7 and 6 1 6 3 and 5 6 18 10 and 12 66.8
2004. 4 12 6 and 8 1 7 1 and 5 5 17 7 and 12 66.3
14 2001. 3 11 6 and 5 0 3 0 3 11 5 and 7 94.1
2004. 5 10 7 0 4 0 and 1 5 13 5 and 8 87.3
2001. 5 17 10 0 6 1 and 2 6 19 13 and 11 78.6
15 2004. 6 20 11 and 7 0 7 1 7 21 13 85.9
1999. 5 13 6 and 8 1 4 1 and 2 6 14 7 and 9 83.7
16 2000. 3 17 12 and 9 1 7 3 and 6 6 20 12 and 16 68.7
2004. 5 21 6 and 14 1 7 1 and 2 6 22 7 and 16 87.8
17 2001. 7 17 12 and 9 0 6 0 and 4 7 17 12 84.7
2004. 7 16 10 0 5 2 and 0 11 18 12 and 14 82.4
1999. 0 15 7 and 10 0 8 0 and 5 6 16 10 and 11 92.4
18 2001. 7 16 11 and 9 0 7 0 and 1 8 17 92.6
2004. 6 18 9 and 11 0 7 1 and 2 7 18 16 and 11 82.8
19
2001. 4 11 6 and 7 1 5 1 and 3 5 15 7 and 10 74.4
2004. 5 12 7 and 11 0 5 1 and 0 7 16 8 and 7 89.0
20 1999. 4 11 7 and 9 0 6 0 and 1 6 16 10 and 8 81.4
2000. 5 12 6 and 8 0 5 0 and 2 6 17 8 and 13 85.6

39
Graph. 1. The phenophases: swelling of buds, opening of buds and flowering of the individual
trees of Acer monspessulanum L. (Stubica)
Year
1998
1999
2000
Tree February March April May
10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10.
o C 2.3 8.0 7.8 5.5 2.1 3.0 11.0 7.2 11.2 13.3
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
o C 0.4 -1.3 2.2 5.6 5.7 6.5 9.7 8.2 10.9 14.9
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
o C 3.1 2.5 0.8 3.6 3.4 7.2 7.3 13.3 14.9 14.5
1
2
3
4
5
6

40
Graph. 1.
Year
2000
2001
2002
Tree February March April May
10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10.
o C 3.1 2.5 0.8 3.6 3.4 7.2 7.3 13.3 14.9 14.5
7
8
9
10
11
12
13
14
15
16
17
18
19
20
o C 3.7 4.7 2.8 6.2 9.3 10.2 8.4 6.4 11.9 14.7
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
o C 5.6 6.0 4.6 9.8 8.9 5.9 7.7 10.8 12.3 14.6
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16

41
Year
2002
2003
Graph . 1.
Tree February March April May
10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10.
o C 5.6 6.0 4.6 9.8 8.9 5.9 7.7 10.8 12.3 14.6
17
18
19
20
o C -2.0 -20 0.9 4.8 4.3 6.9 4.4 11.1 10.9 19.9
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
1
2
3
4
5
6
7
8
9
2004
10
11
12
13
14
15
16
17
18
19
20
Legend:
20
o C -3.4 0.5 0.8 0.4 8.5 7.5 10.5 9.7 10.6 10.8
- phenophase of the swelling of the buds;
- phenophase of the opening of the buds;
- phenophase of the flowering

42
Graph. 2. The phenophase of the flowering of the observed individual trees of Acer
monspessulanum L. (Stubica)
Year
1998.
1999.
2000.
Tree
March April May
5. 10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10.
o C 5.5 2.1 3.0 11.0 7.2 11.2 13.3
1
2
3
4
5
6
7
8
9
10
11
12 Non flowered
13
14
15
16
17
18
19
20
o C 5.6 5.7 6.5 9.7 8.2 10.9 14.9
1
2
3
4 Non flowered
5 Non flowered
6 Non flowered
7 Non flowered
8 Non flowered
9 Non flowered
10 Non flowered
11
12
13
14 Non flowered
15
16 Non flowered
17
18
19
20 Non flowered
o C 3.6 3.4 7.2 7.3 13.3 14.9 14.5
1
2
3
4 4 – 7 Non flowered

43
Graph. 2.
Year
2000.
2001.
2002.
Tree
March April May
5. 10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10.
o C 3.6 3.4 7.2 7.3 13.3 14.9 14.5
8
9
10 Non flowered
11 Non flowered
12
13
14 Non flowered
15
16 Non flowered
17
18
19 Non flowered
20 Non flowered
o C 6.2 9.3 10.2 8.4 6.4 11.9 14.7
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
o C 9.8 8.9 5.9 7.7 10.8 12.3 14.6
1 Non flowered
2 Non flowered
3 Non flowered
4 Non flowered
5 Non flowered
6 Non flowered
7 Non flowered
8 Non flowered
9 Non flowered
10 Non flowered
11 Non flowered
12 Non flowered
13 Non flowered
14

44
Year
2002.
2003.
2004.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
Legend:
Graph. 2.
Tree
March April May
5. 10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10.
o C 9.8 8.9 5.9 7.7 10.8 12.3 14.6
15
16 16 – 20 Non flowered
o C 4.8 4.3 6.9 4.4 11.1 10.9 19.9
1
2
3
4
5
6 Non flowered
7
8
9
10
11
12
13
14 Non flowered
15
16
17
18
19
20
o C 0.4 8.5 7.5 10.5 9.7 12.6 10.8
- phenophase of the flowering of the male flowers
- phenophase of the flowering of the female flowers

45
Graph. 3. The phenophases: swelling of buds, opening of buds and flowering of the
individual trees of Acer monspessulanum L. (Grebice)
Year
1998
1999
2000
Tree February March April May
10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10.
o C 2.3 8.0 7.8 5.5 2.1 3.0 11.0 7.2 11.2 13.3
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
o C 0.4 -1.3 2.2 5.6 5.7 6.5 9.7 8.2 10.9 14.9
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
o C 3.1 2.5 0.8 3.6 3.4 7.2 7.3 13.3 14.9 14.5
1
2
3
4
5
6
7
8
9

46
Graph. 3.
Year
2000
2001
2002
Tree February March April May
10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10.
o C 3.1 2.5 0.8 3.6 3.4 7.2 7.3 13.3 14.9 14.5
10
11
12
13
14
15
16
17
18
19
20
o C 3.7 4.7 2.8 6.2 9.3 10.2 8.4 6.4 11.9 14.7
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
o C 5.6 6.0 4.6 9.8 8.9 5.9 7.7 10.8 12.3 14.6
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20

47
Year
2003
Graph. 3.
Tree February March April May
10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10.
o C -2.0 -20 0.9 4.8 4.3 6.9 4.4 11.1 10.9 19.9
1
2
3
4
5
6
7
5
9
10
11
12
13
14
15
16
17
18
19
1
2
3
4
5
6
7
2004 8
9
10
11
12
13
14
15
16
17
18
19
20
Legend:
20
o C -3.4 0.5 0.8 0.4 8.5 7.5 10.5 9.7 12.6 10.8
- phenophase of the swelling of the buds;
- phenophase of the opening of the buds;
- phenophase of the flowering

48
Graph. 4. The phenophase of the flowering of the observed individual trees of Acer
monspessulanum L. (Grebice)
Year
1998
1999
2000
Tree
March April May
5. 10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10. 15. 20.
o C 5.5 2.1 3.0 11.1 7.2 11.2 13.3 14.4
1
2
3
4
5
6
7
8
9
10
11 Non flowered
12
13
14 Non flowered
15 Non flowered
16
17 Non flowered
18
19
20
o C 5.6 5.7 6.5 9.7 8.2 10.9 14.9 15.8
1
2
3
4
5
6
7
8
9
10
11
12
13
14 Non flowered
15 Non flowered
16
17 Non flowered
18
19
20
o C 3.6 3.4 7.2 7.3 13.3 14.9 14.5 17.1
1
2
3
4 Non flowered
5 Non flowered
6
7
8
9

49
Graph. 4
Year
2000
2001
2002
Tree
March April May
5. 10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10. 15. 20.
o C 3.6 3.4 7.2 7.3 13.3 14.9 14.5 17.1
10
11 Non flowered
12 Non flowered
13
14 Non flowered
15
16
17 Non flowered
18
19 Non flowered
20
o C 6.2 9.3 10.2 8.4 6.4 11.9 14.7 14.4
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
o C 9.8 8.9 5.9 7.7 10.8 12.3 14.6 15.8
1
2 Non flowered
3
4 Non flowered
5 Non flowered
6
7 Non flowered
8
9
10 Non flowered
11 Non flowered
12 Non flowered
13 Non flowered
14 Non flowered
15 Non flowered
16
17 Non flowered
18 Non flowered
19 Non flowered
20

50
Year
2003
Graph. 4.
Tree
March April May
5. 10. 15. 20. 25. 5. 10. 15. 20. 25. 5. 10. 15. 20.
o C 4.8 4.3 6.9 4.4 11.1 10.9 19.9 17.8
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
1
2
3
4
5
6
7
8
9
2004
10
11
12
13
14
15
16
17
18
19
20
Legend:
20 Non flowered
o C 0.4 8.5 7.5 10.5 9.7 10.6 12.6 12.3
- phenophase of the flowering of the male flowers
- phenophase of the flowering of the female flowers

51
Tripić: CONTRIBUTION TO THE KNOWLEDGE OF THE FLOWERING PHENOLOGY . . .
According to G u d e s k i & D r e n k o v s k i (1978) relationship between female
and male flowers in individual trees of A. monspessulanum during a year there are only
or almost female flowers or only or almost male flowers, however, in the next year the
situation was contrary or a proximally equal. Our investigation shown different results –
in each year (in seven years of observing) relationship between female and male
flowers of individual trees was similar (Tab. 5. and 6.).
The flowering of the single tree lasted from 2 – 29 days and of the entire
population from 23 – 59 days (the most frequent 30 – 34 days) depending of the
weather conditions, especially temperature, but also, from number of flowered trees,
richness of flowered trees and numeral relationship of male and female flowers on the
tree (Graph. 2., 4.). Phenophases of flowering of male and female flowers on the same
trees were totally separated in time with 1/2 to 7 days of pause between them. During
phenophases of flowering on individual trees, series of male flowers are shown followed
with series of female flowers (protandrous trees) or when there are series of female
flowers they are followed with series of male flowers (protogynous trees). Very rarely
(in case of very good flowering) on same trees another serie of male flowers (♂ - ♀ -
♂) is shown and also very rerely there is (only with one tree during one year) other
serie of female flowers (♀ - ♂ - ♀ - ♂) during period of flowering.
T r i p i ć (2006) presents very similar results for A. obtusatum but for A.
campestre and A. platanoides he presents results that claim that all trees protandrous
trees always had two series of male flowers (♂ - ♀ - ♂) and all protogynous trees
always had only one serie male flowers and one serie female flowers (♀ - ♂).
Protogyny is found on 8 (40 %) trees in population STUBICA and 11 (55 %) trees
in population GREBICE, and protandry is found on 12 (60 %) trees in population
STUBICA and 9 (45 %) trees in population GREBICE (Graph. 2., 4.). The same trees
always had protandry or protogyny during all years of observation. These are flowering
types of inflorescences that found: A type (♀) – very rare, B type (♀ - ♂) – frequent, D
type (♂ - ♀) – frequent, E type (♂) – frequent.
CONCLUSION
The flowers were morphologically false bisexually and functionally unisexual. The
male flowers had very small (rudimentary) pistil and well-developed stamens (long
filaments and yellow functional anthers), and the female flowers had well-developed
pistil and unripe stamens (green, hard and non-functional anthers on very short
filaments). Protogyny is found on 8 (40 %) trees in population STUBICA and 11 (55 %)
trees in population GREBICE, and protandry is found on 12 (60 %) trees in population
STUBICA and 9 (45 %) trees in population GREBICE (Graph. 2., 4.). With 19 of
observed trees, during every year of observing, protogyny is established, while 21 trees

52
Natura Montenegrina, 6/2007
always had protandry. Total number of the flowers in individual inflorescence varies
from 3 - 28 (male flowers 0 - 24 and female flowers 0 - 12). Quantitative relationship of
the male and female flowers on single trees is 49.46 % - 99.99 % of the male flowers.
The flowering types of inflorescence that were found: A type (♀) – very rare, B
type (♀ - ♂) – frequent, D type (♂ - ♀) – frequent, E (♂)– frequent.
LITERATURE
B E J D E M A N, I. N. 1974: Metodika izučenija fenologiji rastenij i rastitel'nyh
soobšćestv. ANSSSR, Sibirskoe otdelenije, Limnologičeskij Institut, 1-155 pp.
Č A J L A H J A N, M. H. 1988: Reguljacija cvetenija vysših rastenij. AN SSSR,
''Nauka'', Moskva, 3-559 pp.
C H A I L A K H Y A N, M. KH., K H R I A N I N, V. N. 1982: Sex of plants and its
hormonal regulation. AN SSSR, ''Nauka'', Moskva.
G U D E S K I, A. & D R E N K O V S K I, R. 1978: Morfologija i pol na cvetovite kaj
nekoi vidovi od rodot Acer L. Godišen zbornik na Šumarskiot fakultet na
Univerzitet vo Skopje, knj. 28: 6-24 (1977/78), Skopje.
J O N G, P. C. De 1976: Flowering and sex expression in Acer L. Mededelingen,
Landbouwhogeschool, Wageningen, Nederland, 76-2.
P A L A M A R E V, E. 1979: Acer L. In: Flora na Narodna Republika B’lgarija, VII: 221
– 240, Izdatelstvo na B’lgarskata Akademija na naukite, Sofija,
P A X, F. 1885: Monographie der Gattung Acer L. Engler's botanischen Jahrbüchern,
Band VII, Heft 2: 177- 263, Leipzig.
P A X, F. 1902: Aceraceae. In: A. Engler, Das Pflanzenreich IV, 163, 8: 1-89, Leipzig.
P O J A R K O V A, A. I. 1933: Botaniko-geografičeskij obzor klenov SSSR v svjazi s
istoriej vsego roda Acer L. Trudy Botaničeskogo instituta AN SSSR, Ser. I: 225-
374, Izdatel'stvo AN SSSR, Leningrad.
T R I P I Ć, R. 2006: Flowering phenology and sex expression in Acer obtusatum
Waldst. & Kit. from Montenegro. Natura Montenegrina N o 5: 15-24, Natural History
Museum of Montenegro, Podgorica.
T R I P I Ć, R. 2006: Flowering phenology and sex expression in Acer campestre L.
from Montenegro. Natura Montenegrina N o 5: 25-36, Natural History Museum of
Montenegro, Podgorica.
T R I P I Ć, R. 2006: Contribution to the knowledge of flowering phenology and sex
expression in Acer platanoides L. from Montenegro. (In press).
Z A M J A T N I N, B. N. 1958: Acer L. In: Derev’ja i kustarniki SSSR, IV: 405 – 499,
Izdatel’stvo Akademii nauk SSSR, Moskva – Leningrad.
Received: 25. 12. 2007.

NATURA MONTENEGRINA, PODGORICA, 6:53-61
THE FLORA AND VEGETATION OF SALT WORKS IN ULCINJ
Snežana V U K S A N O V I Ć 1 & Danka PETROVIĆ 2
1 Natural History Museum of Montenegro, Trg vojvode Bećir bega Osmanagića 7, Podgorica; Montenegro,
E-mail: prmuzej@cg.yu; vukss@cg.yu
2 The University of Montenegro Faculty of science, Department of Biology, Cetinjski put b.b., Podgorica;
Montenegro, E-mail: danka.petrovic@cg.yu
Кеy words:
Ulcinj Salts,
taxon,
flora,
vegetation types,
species,
habitats
Ključne riječi:
Ulcinjska Solana,
takson,
vegetacijski tipovi,
vrste,
stanista (habitati)
SYNOPSIS
The results two years’ research of flora and vegetation
of Ulcinj Salts have been presented in this paper. A rare
taxon Beta vulgaris ssp. maritima noted until now in flora of
Montenegro only at one locality, was found. The paper
consists of a review of vegetation types present in the Salt
Works with a list of species which form them. A table with
names of species and habitats they inhabit has also been
presented.
SINOPSIS
FLORA I VEGETACIJA ULCINJSKE SOLANE
U radu su predstavljeni rezultati istraživanja flore i
vegetacije ulcinjske Solane tokom dvije godine. Pronađen je
rijetki takson Beta vulgaris ssp. maritima koji se u flori Crne
Gore bilježi još na samo jednom lokalitetu. U radu je dat
pregled tipova vegetacije prisutnih na prostoru Solane uz
spisak vrsta koje ih izgrađuju. Na kraju je predstavljena
tabela sa imenima vrsta i habitata koje naseljavaju.
INTRODUCTION
The Ulcinj Salt Works are located in the far south of Montenegro. Built in the
region with the biggest number of sunny days, the greatest insolation and the biggest
number of tropical days in the former SFRY (M i c e v et all, 1995), they occupy around
15 km 2 of salt pools. In the past, at the area of today’s salt pans there was Zoganjsko
blato (mud), swamp with brackish water, which started assuming anthropogenic
infrastructural form by end thirties of the century before last (H a d ž i b r a h i m o v i ć ,

54
Natura Montenegrina, 6/2007
2002). Today, these are artificial, men-directed ecosystems, where the time for filling
the pools with the sea water, the level of water therein and water salinity are set in
advance. The immediate vicinity of the sea, the Bojana River, Šasko Lake, Velipolje in
Albania, and of the Lake of Skadar, makes this “Cultural Laguna”, as V a s i ć called it
(1989), very interesting for ornithology and the science in general.
MATERIAL AND METHODS
The investigations of flora and vegetation of Salt Works, presented in this
paper, were carried out in 2003 and 2004. During the first year, only one field visit
was made (June), while in 2004 field visits were taking place during the entire
vegetation season (March-October). The material collected on the filed was
subsequently determined, put into herbaria and deposited with the Natural History
Museum of Montenegro. The borders of vegetation entities were recorded and the
alteration thereof depending on the time of year and the water level in the basins was
monitored. The paper includes an overview of the types of vegetation present in the
area of Salt Pans along with a list of species that make up these types of vegetation.
HALOPHYLOUS VEGETATION
Grounds with a large quantity of salt are unfavourable for plant life, and thus
they are inhabited only with the species that are specifically adapted to cruel living
conditions in these habitats. Such plants, known as halophytes, form floristically poor,
but very interesting plant cover. Typical or genuine halophytes or euhalophytes can be
found in the habitats with high salinity degree and significant moistness. Their
communities present a predominant type of vegetation in the Salt Works. They were
found on the muddy-clayish boundary places of the pools, Lake I, Lake II, canals, as
well as on the embankments. They partially have an emerged character, i.e. they
inhabit shallow waters. The zonation of halophytic vegetation in the Salt Works is inter
alia determined by the composition of canals and embankments.
The association of Salicornietum herbaceae Jank. & Stev., is developed on
very salty and constantly flooded sites, and some associations are located in shallow
waters. The species Salicornia herbacea L. is predominating species of this
community. In addition to this taxon, the association is dominated by Sueda maritima
Dum, while less numerous are the species Limonium angustifolium (Tausch) Degen
and Atriplex portulacoides L. As accompanying species there appear Salsola soda L.
and Atriplex prostrata Boucher ex DC. The community emerges in the month of May,
and right before the end of summer and in autumn, when Salicornia blossoms and
gets a characteristic red colour, the community reaches the peak of its development.
The community of Arthrocnemetum fruticosi Br. – Bl., which appears on the salt
grounds very often in combination with the community of Salicornietum herbaceae
Jank. & Stev., is present only in fragments on the Salt Works. It develops at

55
Vuksanović & Petrović: THE FLORA AND VEGETATION OF SALT WORKS IN ULCINJ
somewhat distant, always flooded and very salty places. It is of the same floristic
composition, but there is a difference in the level of presence of certain species.
These two communities are at some places mosaically combined. In the dry and less
salty areas the characteristic species in the community is Limonium angustifolium
(Tausch) Degen.
The best developed species of euhalophyta are located in the beginning zone
of the lake I (in the vicinity of blue houses). These are real meadows on which one
can notice two zones of halophytic vegetation. The first one, developed along the
water, is made up of the community of Salicornietum herbaceae Jank. & Stev .
Going toward the dams and canal, the second, more complexly built halophytic zone,
appears. It is formed by constituents of diverse floristic composition, which alternate
mosaically among each other. These are: pure constituents of the species Limonium
angustifolium (Tausch) Degen; constituents wherein along with Limonium
angustifolium (Tausch) Degen appear: Atriplex portulacoides L., Sueda maritime Dum.
and Carex sp.; the dominant species Sueda maritima Dum. and Carex sp; constituents
wherein Sueda maritime Dum. and Limonium angustifolium (Tausch) Degen appear
along Carex sp.,; pure constituens of Carex sp. As accompanying species there
appear: Spergularia salina Presl, Plantago coronopus L. and Coronopus squamatus
(Forsskal) Asch.
In the remaining parts of the Salt Works (Lake II, First Evaporation, pools of
Zoganj, Second Evaporation, dams) there are also two halophytic zones wherein the
euhalophytic vegetation is dispersed in the same way, although it is less developed. In
the crystallisation basins and along the margins of the canal we find individual
specimens, which do not form communities.
Going further from the water, the terrain rises (soil becomes deeper), salinity
and humidity of the ground decrease. In this area the dominant species become those
which appear in the second zone of the halophytic vegetation as accompanying
species (Spergularia salina Presl., Plantago coronopus L. i Coronopus squamatus
(Forsskal) Asch). This type of vegetation is best developed during April and in the
beginning of May, when dominant species blossom. By the passing of the vegetation
season, there start to emerge the elements of meadow and ruderal vegetation which
become the dominant ones.
On the dams, which divide the basins, the species Inula crithmoides L., an
important element of halophytic vegetation, appears.
MEADOW VEGETATION
The communities of meadow flora spread in deeper layers of the ground with
less salinity. Meadow type of vegetation exists on the raised land along the channel
leading to Lake I. On the meadow, different aspects change each other during the
vegetation season. During March geophyte are predominant, such as: Narcissus
tazetta L., Romulea bulbocodium (L.) Sebast. & Mauri, Hyacinthus orientalis L. and
Ornithogalum sp. Later, the following species dominate in the meadow: Avena barbata

56
Natura Montenegrina, 6/2007
Pott ex Link, Dactylis glomerata L., Phragmites comunis Trin., Carex sp., and during
the autumn Limonium angustifolium L. is also present. The facies with dominant
species Trifolium nigricens Viv. exists only in fragments. Accompanying elements of
such communities are Anemone hortensis L., Anagalis arvensis L., Bellis perennis L.,
Bituminaria bituminosa (L.) Stirton, Echium vulgare L., Silene gallica L. etc.
In the central part of meadow situated between First evaporation and pools of
Zoganj, typical meadow vegetation is developed during the spring, with dominating
species: Petrorhagia saxifraga (L.) Link, Petrorhagia prolifera (L.) P.W.Ball &
Haywood, Moenchia mantica (L.) Bartl., Kickxia commutata (Bernh. ex Reichenb.)
Fritsch etc. But, in autumn, at the same space, predominant species is Limonium
angustifolium (Tausch) Degen. The community with distinctive taxon Juncus acutus L.
cover the rest of meadow. Like the first-mentioned meadow the same geophyte are
developed in March, and here are noted the represents of family Orchidaceae: Orchis
laxiflora Lam., Serapias lingua L., and S. vomeracea (Burm) Briq.
REEDS
Under natural conditions, the reeds reach Phragmites communis Trin. Its most
intesive development is on swampy, boggy and peat-boggy soils with a sufficient
water layer above the soil surface 60-80 cm. On dry or drying-up habitats the reed can
only develop if the ground water level is high (not deeper than 3-4 cm below the
surface)(N i k o l a j e v s k i j , V. G. 1971.).
Uniform constituents of reed by Phragmition communis W. Koch are located in
pools and canals, in places, formed the islands that raised above the water. Specially,
constituens of reed are well developed in area of Second evaporation. We believe that
reed will continue to expand and occupy new water surface areas. This species is
present as an accompanying element in the vegetation of wet meadows as well.
In muddy coastal area, the community Juncetum maritime-acuti Hić. exists
only in fragments. The component species of such community are Juncus acutus L.
and J. maritimus Lam. Individual trees of taxon Tamarix africana Poiret can also
frequently be found.
RUDERAL VEGETATION
Ruderal plants belong to a widespread type of vegetation inhabiting areas
exposed to the intensive human influence. They can be found in the areas of
permanent or temporary settlements of people and domestic animals, around roads,
railroads, embankments … On the land around Salt-Work, ruderal vegetation is
present along the whole road, on embankments along the railroad, and the elements
of this flora are mixed with halophytes vegetation and on the embankments separating
individual basins.

57
Vuksanović & Petrović: THE FLORA AND VEGETATION OF SALT WORKS IN ULCINJ
CONCLUSIONS
The communities of halophyte plants are floristically poor, but ecologically very
interesting, these plants are specifically adapted to cruel living conditions at their
habitats. Development of tourism and strong urbanization in coastal part of
Montenegro, cause for disappearing of several habitats of this type of vegetation
(Topolica-Bar, Igalo). Preserved and not fragmentary communities of typical
halophytes exist only on land around the Salt Works in Ulcinj and Tivatsko polje.
Because that, Salt Works in Ulcinj are very important for preservation of these
communities.
Among the plants growing on Salt Works, Ophrys bertolonii Moretti, species
from family Orchidaceae, is particularly interesting and important, and it is protected in
Montenegro. Many representatives of this family are included in Red Lists in most of
the countries of Europe, and there are some suggestions to protect them also in
Montenegro.
It is very important to mention the species of this family that are presented in the
meadow flora on Salt Works, such as: Orchis laxiflora Lam., Serapias lingua L.i S.
vomeracea (Burm.) Briqu..
The finding concerning the subspecies Beta vulgaris ssp. maritime, up to now
has been noted only at one locality (A d a m o v i c 1913, sand betwen Bar and Ulcinj).
We consider that the most important sites of Salt Works regarding the flora and
vegetation are localities with the best developed belt of halophyte and where we found
the most interesting plants (species). The best development stands of euhalophyte are
located by the Lake I, as well as on the embankment that separates Lake I from Lake
II. The taxon Beta vulgaris ssp. maritime was found on this embankment, too, so we
believe that area should be preserved from any intervention intended for the
rehabilitation of Salt Works. On meadow situated between First evaporation and pools
of Zoganj we found mentioned representatives of family Orchidaceae, and taxon
Ophrys bertolonii Moretti was noted at the beginning of the path that separates
Knete from Lake II.
Therefore these localities should not be subjected to any interventions regarding
the rehabilitation of Salt-Works.
Domination of ruderal vegetation on Salt Works (except ground with large
quantity of salt) shows that there is a strong influence of man and cattle. But
halophyte vegetation is not exposed to grazing, because the cattle avoid plants with
high contents of salt.
SPECISMENS
Ajuga chamaepitys (L.) Schreb
Alkana tinctoria (L.) Tausch
Anemone hortensis L.
Anagalis arvensis L.
Anchusa officinalis L.
HABITAT
near road, meadow
near road
near road, meadow
near road, meadow
meadow

58
Natura Montenegrina, 6/2007
Anthemis arvensis L.
near road
Aristolochia rotunda L.
embankment with Tamarix (behind Zoganjski pools)
Arum maculatum L.
meadow
Asphodelus microcarpus Viv.
near pools of Zoganj
Aster tripolium L.
embankment with Tamarix (behind Zoganjski pools)
Atriplex portulacoides L.
around lake I, embankments
Atriplex prostrata Boucher ex DC.
around lake I, embankments
Avena barbata Pott ex Link
meadow near lake I
Bellis perennis L
near road, meadow
Beta vulgaris ssp. maritima
embankment between lake I and lake II
Bidens tripartita L.
near road, embankment behind Zoganjski pools
Bituminaria bituminosa (L.) Stirton
near road, meadow near lake I
Blackstonia perfoliata (L.) Huds
embankment behind Zoganjski pools
Calepina irregularis (Asso) Thell
near road, meadow near lake I
Capsella bursa-pastoris (L.) Medicus
near road, meadow near lake I
Cardamine hirsuta L.
near road, meadow near lake I
Carex sp.
meadow
Centaurea alba L.
embankment behind Zoganjski pools
C. calcitrapa L. near road
C. solstitialis L. near road
Centaurium erythraea Rafn.
meadow
Cichorium intybus L
near road
Cirsium arvensis (L.) Scop
near road, embankments
Clematis viticella L.
embankment behind pools of Zoganj
Clinopodium vulgare L.
near road, meadow
Convolvulus arvensis L.
near road, meadow
Conyza canadensis (L.) Cronquist
near road, meadow
Coronopus squamatus (Forsskal) Asch. raised land near lake I
Cynoglossum creticum Miller
near road, meadow
Dactylis glomerata L.
meadow near lake I
Daucus carota L.
embankment behind pools of Zoganj
Delphinium peregrinum L.
near road above lake I
Ditrichia viscosa (L.) Greuter
near road, embankment behind pools of Zoganj
Dorycnium hirsutum (L.) Ser.
embankment behind pools of Zoganj
Echium vulgare L.
near road, meadow
Epilobium hirsutum L.
near canal behind pools of Zoganj
Erigeron annuus (L.) Pers.
near road, meadow
Erodium cicutarium (L.) L’Her
near road
E. malacoides (L.) L’Her near road
Eryngium amethistinum L.
embankment behind pools of Zoganj
Eupatorium cannabinum L.
embankment behind pools of Zoganj
Euphorbia helioscopia L.
near road, meadow
E. peplis L. near canal along lake I

59
Vuksanović & Petrović: THE FLORA AND VEGETATION OF SALT WORKS IN ULCINJ
E. peplus L. near road
E. terracina L. near road toward Kneta
E. seguieriana Necker near road toward Kneta
Filago vulgaris Lam.
near road
Geranium columbinum L.
near road
G. dissectum L. near road
G. robertianum L. near road, meadow
G. brutim Gasparr. meadow
Hyacinthus orientalis L.
near road, meadow
Helichrysum italicum (Roth.) G. Don
near road
Heliotropium europaeum L.
near road, embankments
Hordeum vulgare L.
near road, meadow
Hypericum perforatum L.
near road, meadow
Inula brittanica L.
embankment behind pools of Zoganj
I. crithmoides L. embankments
Juncus acutus L.
muddy places near pools of Zoganj
J. maritimus Lam. muddy places near pools of Zoganj
Kickxia commutata (Bernh. ex Reichenb.) meadow, embankment behind pools of Zoganj
Fritisch
Lamium purpureum L.
near road
Lathyrus cicera L.
near road, meadow
Limonium angustifolium (Tausch) Degen near pools, canals, on medow
Linaria vulgaris Miller
near road, meadow
Linum nodiflorum L.
near road, meadow
L. usitatissimum L. meadow
Lotus corniculatus L.
near road, meadow
Medicago minima (L.) L.
near road, meadow
Melilotus officinalis (L.) Pallas
embankment, near road
Moenchia mantica (L.) Bartl.
meadow, embankment behind pools of Zoganj
Muscari comosum (L.) Miller
meadow, embankment between lake I and lake II
Narcissus tazetta L.
meadow
Nigella damascena L.
embankment behind pools of Zoganj
Ophrys bertolonii Moretti
road between first evaporation and Kneta
Orchis laxiflora Lam.
meadow
Ornithogalum sp.
near road, meadow
Oxalis corniculata L.
embankment behind pools of Zoganj
Parentucellia latifolia (L.)Caruel
meadow
Petrorhagia prolifera (L.) P.W.Ball &
near road, meadow
Heywood
Petrorhagia saxifraga (L.) Link.
meadow
Phragmites communis Trin
pools, canals, humid medow
Picris hieracioides L
near road
Plantago coronopus L.
raised salt land

60
Natura Montenegrina, 6/2007
P. lanceolata L. near road
P. major L. near road
Polygonum aviculare L.
along the road, embankment
Portulaca oleracea L.
embankment between lake I and lake II
Prunella laciniata (L.) L.
near road, meadow
P. vulgaris L. near road, meadow
Pulicaria dysenterica (L.) Bernh
embankment behind pools of Zoganj
Reseda phyteuma L.
embankment behind pools of Zoganj
Romulea bulbocodium (L.) Sebast.& Mauri near road, meadow
Rosa canina L.
embankment behind pools of Zoganj, near road
toward Kneta
Rubus idaeus L.
embankment behind pools of Zoganj
Salicornia herbacea L.
salt, humid places
Salsola soda L.
salt, humid places
Salvia verbenaca L.
near road, meadow
S. verticillata L. near road, meadow
Scandix pectin-veneris L.
near road
Scolymus hispanicus L.
near road, embankments
Senecio rupestris Waldst. & Kit.
near road
Serapias lingua L.
meadow between first evaporation and pools of
Zoganj
S. vomeracea (Burm.) Briqu. meadow between first evaporation and pools of
Zoganj
Sherardia arvensis L.
meadow, near road
Silene conica L.
meadow
Silene gallica L.
meadow, near road
Silene nocturna L.
meadow
Solanum nigrum L.
embankments, near road
Sonchus arvensis L.
near road
Spergularia salina
raised salt land
Stellaria media (L.) Vill.
meadow, near road
Suaeda maritime Dum.
salt, humid places
Tamarix africana Poir.
humid medow
Taraxacum officinale Weber
meadow
Teucrium chamaedrys L.
meadow, near road
Trifolium campestre Schreber
meadow, near road
T. incarnatum L. meadow
T. nigricens Viv. meadow, near road
T. resupinatum L. meadow
T. subterraneum L. meadow
Trigonella esculenta Willd.
near road
Verbascum sinuatum L.
near road
Verbena officinalis L.
embankment behind pools of Zoganj, near road

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Vuksanović & Petrović: THE FLORA AND VEGETATION OF SALT WORKS IN ULCINJ
Veronica arvensis L.
meadow
V. chamaedrys L. meadow, near road
Vicia grandiflora Scop.
meadow, near road
Vicia sativa L. subsp. nigra (L.) Ehrh.
meadow, near road
Vincetoxucum hirundinaria Medicus
embankment behind pools of Zoganj
LITERATURE
D O M A C , R. 1967: Ekskurzijska flora Hrvtske i susjednih područja. - Institut za
botaniku sveučilišta u Zagrebu. Zagreb, 543pp.
JANKOVIĆ , M., S T E V A N O V I Ć , V. 1983: Prilog poznavanju slatinske vegetacije
Boke Kotorske. - Zbornik Roberta Visianija Šibenčanina, Muzej grada Šibenika
10:377-396
HORVATIĆ , S. 1934: Flora i vegetacija otoka Paga. - Prir. istraž. Jugosl. Akad. znan. i
umjetnosti, Zagreb. 19: 116-372.
HORVATIĆ , S. 1963: Vegetacijska karta otoka Paga s općim pregledom
vegetacijskih jedinica Hrvatskog primorja. Prir. istraz., Acta Biol. IV/33: 5 - 187.
JAZU, Zagreb
J A V O R K A , S., C S A P O D Y , V. 1934: Iconographia florae Hungaricae, Budapest.
576 pp.
K A R A M A N , V. 1997: Flora istočnog dijela Bokokotorskog zaliva. Magistarski rad. - Biološki
fakultet, Beograd ( manuscr.). 186 pp.
NIKOLAJEVSKIJ, V. G. 1971: Research into the Biology of the Common Reed
(Phragmites communis Trin.) in the U.S.S.R. - Folia-geobotanica Phytotaxonomica
6/2: 221 – 230. Czechoslovak Academy of Sciences, Praha.
P A R O L L Y , G. 1992: Die Orchideenflora Montenegros. - AHO Mitteilungsblatt,
Baden Wuertenberg 24 (2): 141 -391
P I G N A T T I , S. 1982: Flora d' Italia. 1-3. - Edagricole, Bologna
PULEVIĆ , V. 2005: Građa za floru Crne Gore. - Republički zavod za zaštitu prirode,
Podgorica, 218 pp.
R O H L E N A , J. 1942: Conspectus florae Montenegrinae 20-21: 1-506.- Preslia,
Praha.
T U T I N , T. G., HEYWOOD, V. H., BURGES, N. A. MOORE., VALENTINE, D. H.,
WALTERS, S. M., WEBB, D. A. (eds) 1964-1980: Flora Europaea 1-5
Cambridge.
T U T I N , T. G., BURGES, N. A., CHATER, A. O., EDMONDS, J. R., HEYWOOD, V.
H., MOORE, D. M., VALENTINE, D. H., WALTERS, S. M. & WEBB, D. A. 1993:
Flora Europaea. Vol. 1. - Sec. ed. Cambridge. 581pp.
Received: 24. 11. 2007.

62
Natura Montenegrina, 6/2007

NATURA MONTENEGRINA, PODGORICA, 6:63-71
CONTRIBUTION TO THE KNOWLEDGE OF DISTRIBUTION AND SEASONAL
DYNAMICS OF THE PLANCTONIC POLYCHETA IN SOUTH ADRIATIC WATERS
Vera VUKANIĆ 1
1
Faculty of Technical Science, Departmant of Technics, Chemistry and Biology, The University of Novi
Pazar, Vuka Karadzica bb, Novi Pazar, Serbia; v_vukanic@yahoo.com
Key words:
polycheta,
Tomopteris elegans,
Adriatic,
Bay of Boka Kotorska,
distribution
Synopsis
This paper presents the first data about seasonal
overveiw of horisontal distribution and abundance for the
genus Polycheta in Bay of Boka Kotorska. The only
determined species was Tomopteris elegans Chun, which is
present throughout the Adriatic Sea, but as it has a quite
narrow ecological valence its abundance is much greater in
deep open waters of the Adriatic than in the coastal waters.
Also presented are seasonal oscillations and quantitative
presence of Polycheta population during 2002.
Ključne riječi:
Polycheta,
Tomoperis elegans,
Jadran,
Bokokotorski zaliv,
distribucija
Sinopsis
PRILOG POZNAVANJU DISTRIBUCIJE I SEZONSKE
DINAMIKE PLANKTONSKIH POLIHETA U JUŽNOM
JADRANU
U radu iznosimo prve pregledne podatke o sezonskoj
horizontalnoj distribuciji i abundanciji roda Polycheta u
Bokokotorskom zalivu. Determinisana je samo vrsta
Tomopteris elegans Chun, koja je prisutna u cijelom Jadranu.
Ova vrsta ima prilično usku ekološku valencu, tako da ima
mnogo veću abundanciju u pučinskim dubokim vodama
Jadrana nego u priobalnim vodama. Takođe smo prikazali
sezonske oscilacije i kvantitativno prisustvo cijele grupe
Polycheta tokom 2002. godine.

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Natura Montenegrina, 6/2007
INTRODUCTION
Geographic position and specific ecological conditions make Bay of Boka Kotorska
a distinct biotope. The interactive relationships of its wildlife are influenced by influx of
freshwater from the mainland as well as the currents from the open sea. Therefore, we
tried to use the data collected in one-year cycle of study in order to determine the
main regularities in oscillations of ecological parameters, important for fluctuations in
diversity of Polycheta biocenosis. Research on pelagic polychaetes in Adriatic had
started alredy in the 19 centry. The first data were collected in Northern Adriatic
(B u s c h , 1851; V e j d o v s k i , 1876; A p s t e i n , 1893; B a l d a s s e r o n i , 1914) and
Otrant strata (R o s a , 1912). Data for eastern coast and South Adriatic are mentioned
only in recent times (K n e ž e v i ć , 1942; G a m u l i n , 1948, 1979; H u r e , 1961;
Vukanić ,D., 1975.; B a t i s t i ć et al. 2004; V u k a n i ć ,V., 2004). H u r e (1961) cited
the following species for the open waters of South Adriatic: Loperorhynchus
uncinatus, Alciopa cantrainii, Vanadis formosa, Tomopteris helgolandica, Tomopteris
elegans. Tomopteris elegans is the most common Adriatic species cited by H u r e
(1955). V u k a n i ć ,D. (1975) presented data on participation of polychaetes in total
zooplankton of coastal waters of open South Adriatic with 0.80% and at 6 NM off the
shore with 0.17%. G a m u l i n (1979) cited the species recorded by Zei in 1956:
Tomopteris elegans, T. helgolandica, Alciope contrainii, Vanadis formosa and V.
crystallina. Batistić et al. (2004) when discussing material collected on a cruise
from April 1993 to February 1994, cited the following species for the South Adriatic:
Tomopteris elegans, Callizonella lepidota, Callizona nasuta, Pelagobia longicirrata,
Sagitella kowalevskii, Travisiopsis lanceolata i Typhloscolex muelleri.
MATERIALS AND METHOD
Study area - Boka Kotorska Bay is situated in the southeastern part of Adriatic.
Its geographical position is determined by key points: 42º31'N, 42º23'32''S,
18º46'32''E and 18º30'29W. It penetrates into the mainland for 28 km and is composed
of four interconnected bays: Herceg Novi Bay (which directly communicates with the
open sea), Tivat Bay, as well as the inner bays, Risan and Kotor. The surrounding
mainland is a karstic area, arid during summer, while in autumn, winter and early
spring it has the most plentiful precipitation in Europe (6000 mm). Boka Kotorska Bay
is under a strong influence of waters from deep southern Adriatic and the entering
Eastern Mediterranean current from one side, while on the other side there is a huge
influx of freshwater from the surrounding mainland in autumn and winter-spring
season, making it a very dynamic system with specific physical-chemical and
biological characteristics. At the southwest, in front of the entrance to Boka Kotorska
Bay, there is a point where the greatest depth of Adriatic Sea was measured (1330m).

65
Vukanić: CONTRIBUTION TO THE KNOWLEDGE OF DISTRIBUTION AND . . .
Fig.1. Map of sampling area in southern Adriatic - Bay of Boka Kotorska: 1. P-IBM
(Sampling point near the Marine Biology Institute); 2. P-K (Sampling point in central part of
Kotor’s Bay); 3. P-O (Sampling point near Orahovac); 4. P-R (Sampling point in central part of
Risan’s Bay); 5. P-M (Sampling point near Morinj); 6. P-T (Sampling point in central part of
Tivat’ Bay); 7. P-HN (Sampling point in central part of Herceg Novi’s Bay).
Sampling - Zooplankton was sampled in 2002, on 25 th each month, at seven
stations. The plankton nets used were of Nansen type, either surface area of the
opening 1/4m 2 - length 2.5m, or surface area of the opening 1m 2 - length 3.5m. The
density of the mesh was 150µm and 200µm. Vertical samples were taken from 10m to
0m at the shallow stations of each bay. The collected zooplankton material was
fixated on the ship, in 2.5% formaldehyde-sea water. The qualitative quantitative
analyses were done in the laboratory from the representative sample of 1/25 of total
catch, under the stereomicroscope and binocular lens. After that, the whole catch was
carefully analyzed in order to record any rare species. The quantity was presented as
number of individuals under m -2 below sea surface (ind.m -2 ). Temperature, salinity,

66
Natura Montenegrina, 6/2007
pH, oxygen saturation and percent oxygen saturation were measured in situ with a
probe Multiline P-4, at 0.5m, 2m, 5m and 10m in the shallow part of the Bay near the
coast, and at 0.5 m, 10m, 15m, 20m and 30m of depth at central deep stations. The
transparency of the sea was measured with Secchi disk, 30 cm in diameter, of white
color. Color of the sea was measured according to Forell from I to XXI.
RESULTS AND DICUSSION
Environmental conditions - The maximal temperature (27.7ºC) was recorded in
July, while the medium maximal temperature of all layers was 25.3ºC. The minimal
temperature (10ºC) was recorded in Risan Bay (P 2 -R) in February. The salinity of the
sea varied within the values 7.9‰ - 38.71‰, and the lowest value ever recorded in the
Bay was registered at the shallow station P-IBM in the surface layer 0-5m, 2.30‰ in
September. Oxygen saturation varied from 4.68mL/L at the surface of Herceg Novi
Bay (P 4 -HN) in June and July to 33.7mL/L at the 10 m depth at shallow station P-O in
June. The percent of oxygen saturation was over 100%, and the greatest value was
recorded at the shallow station P-O in June (220%), while at the surface at the middle
of Kotor Bay (P 1 -K) it was 117%. The high values of oxygen saturation, which
exceeded 100% throughout the year, show that the Bay is a biotope with a high
degree of trophic activity. The whole Bay had relatively low values of transparency,
from 4 m in Kotor Bay (P 1 -K) in September, to 15 m in Herceg Novi Bay (P 4 -HN) in
January. The color of the sea according to Forell ranged from II in June to VI in
September.
Policheta general distribution – Polychaetes have occasionally been recorded
in small numbers in Bay of Boka Kotorska. Their percentage participation within the
whole zooplankton was: in Bay of Kotor 0.11%, in Bay of Risan 0.22%, in Bay of Tivat
0.21% in Bay of Hercegnovi 0.73%. They are the most abundant during the autumn
season. We have determined the species Tomopteris elegans Chun, which was
occasionally recorded at all stations.
18000
16000
14000
12000
10000
8000
6000
4000
2000
0
Polycheta [TOTAL]
I II III IV V VI VII VIII IX X XI XII
Fig.1. Oscillations of abundance of Polycheta group in Bay of Boka Kotorska during the study

67
Vukanić: CONTRIBUTION TO THE KNOWLEDGE OF DISTRIBUTION AND . . .
Postaje
Sezone
P-IBM P-O P-M P 1 -K P 2 -R P 3 -T P 4 -HN
Zima 200 200 200 600 1200 400 100
Proljeće 300 1100 1100 900 1000 100 600
Ljeto 100 800 600 700 200 200 600
Jesen 200 3100 600 4100 2300 2700 12600
% 0.1 0.5 0.34 0.11 0.22 0.21 0.73
Tab.1. Percentage participation of Polycheta group (ind/m 2 ) in total zooplankton during the study
Polycheta
Total zooplankton
18000
16000
14000
12000
10000
8000
6000
4000
2000
0
I II III IV V VI VII VIII IX X XI XII
5000000
4000000
3000000
2000000
1000000
0
Fig.2. Logarithm presentation of oscillations in abundance
of total zooplankton and the polychaete group
Tomopteris elegans Chun was the most abundant pelagic polychaete in the
Adriatic. It was occasionally recorded at all stations in Boka Kotorska Bay, in
insignificant number or as single individuals; it was more common and abundant in the
Bay of Herceg Novi in autumn season. The percentage participation of this species
within the total number of polychaete in the Bay was 3.27%. This species does not
tolerate large variations in hydrographic factors, has a quite narrow ecological valence
and therefore much larger abundance in the open sea waters of the Adriatic.
400
300
200
100
0
Tomopteris elegans
I II III IV V VI VII VIII IX X XI XII
Fig.3. Oscillations of abundance of species Tomopteris elegans in
Bay of Boka Kotorska during the study

68
Natura Montenegrina, 6/2007
The larval stages of polychaetes were very abundant. The smaller maximum of
abundance was recorded in March and April and the larger maximum in October and
November. The percentage participation of larval stages in total number of recorded
polychaete specimens was 71.27%. The other individuals of this group participated
with 20.20% in total number of polychaetes. These are the first data on this group of
organisms for Bay of Boka Kotorska.
Polycheta larvae
30000
25000
20000
15000
10000
5000
0
I II III IV V VI VII VIII IX X XI XII
Fig.4. Oscillations of abundance of Polycheta larvae in Bay of Boka Kotorska during the study
Bay of Kotor
Bay of Risan
Policheta larvae
Policheta sp.
Tomopteris elegans
Policheta larvae
Policheta sp.
Tomopteris elegans
Bay of Tivat
Bay of Hercegnovi
Policheta larvae
Policheta sp.
Tomopteris elegans
Policheta larvae
Policheta sp.
Tomopteris elegans
Fig.6. Percentage participation of certain taxa from Polycheta group, for each bay separately

69
Vukanić: CONTRIBUTION TO THE KNOWLEDGE OF DISTRIBUTION AND . . .
Fig.5. Oscillations of abundance of Polycheta group throughout the year, for each bay separately
Bay of Kotor
4000
3500
3000
2500
2000
1500
1000
500
0
I II III IV V VI VII VIII IX X XI XII
Polychaeta larvae
Tomopteris elegans
Polycheta sp.
1800
1600
1400
1200
1000
800
600
400
200
0
Bay of Risan
I II III IV V VI VII VIII IX X XI XII
Polychaeta larvae
Tomopteris elegans
Polycheta sp.
2000
1800
1600
1400
1200
1000
800
600
400
200
0
Bay of Tivat
I II III IV V VI VII VIII IX X XI XII
Polychaeta larvae
Tomopteris elegans
Polycheta sp.

70
Natura Montenegrina, 6/2007
Bay of Hercegnovi
12000
10000
8000
6000
4000
Polychaeta larvae
Tomopteris elegans
Polycheta sp.
2000
0
I II III IV V VI VII VIII IX X XI XII
CONCLUSIONS
Bay of Boka Kotorska is a typical neritic area of the eastern coast of Southern
Adriatic. In the zooplankton biocenosis of the Bay we have recorded a great
abundance of larval and juvenile forms. A large number of species was recorded in
Polychaete group longitudinally from the inner bay (Bay of Kotor and Bay of Risan)
toward the other bays (Bay of Tivat and Bay of Herceg Novi), together with a high
abundance of larval forms toward the inner bays.
REFERENCES
APSTEIN,C., 1893: Die Alciopiden der Berliner Zool. Sammlung. – Arch. Naturg.
59(1).
B A L D A S S E R O N I ,V., 1914: Nota sui Tifloscolecidi raccolti della R.N. ‘’Ciclope’’
nelle crociere III-IV. – R.Com. Tal. It. Mem. 38, 1-19.
BATISTIĆ ,M., K R Š I N I Ć , F., JASPRICA, N., CARIĆ , M., V I L I Č I Ć ,
LUČ O Ć , D.& D. 2004: Gelatinous Invertebrate Zooplankton of the South
Adriatic: Species Composition and Vertical Distribution. Journal of Plankton
Research, Vol.26. (3). p.1-16.
B U S C H ,W., 1851: Beobachtungen über Anatomie und Entwicklung einiger
werbelloser Seetiere. 1 – 143. Berlin.
G A M U L I N ,T., 1948: Prilog poznavanju zooplanktona srednjedalmatinskog otočnog
područja. Acta Adriatica, 3(7), 159-194.
G A M U L I N , T., 1979: Zooplankton istočne obale Jadranskog mora. Acta Biologica,
8/j – 10, prir. istr., 43,p.177-217.

71
Vukanić: CONTRIBUTION TO THE KNOWLEDGE OF DISTRIBUTION AND . . .
H U R E , J., 1955: Distribution annuelle verticale du zooplancton sur une station de
l'Adriatique meridionale.–Acta Adriatica, 7(7),1–72.
H U R E , J., 1961: Dnevna migracija i sezonska vertikalna raspodjela zooplanktona
dubljeg mora.–Ibid., 9(6),1–60).
KNEŽEVIĆ , M.,1942: Prilog poznavanju geografske rasprostranjenosti Tomopterida
u Jadranskom moru. – Vter. Arh., 12(11), 495-496.
R O S A , D., 1912: Nota sui Tomopteridi dell Adriatico raccoleti Navi ‘’Montebello’’ e
‘’Ciclope’’. – R.Com. Tal. Ital., Mem. 26,1-10.
V E J D O V S K I , F., 1878: Beiträge der Tomopteriden. – Zeitschr. Wiss. Zool. 31.
VUKANIĆ ,D., 1975: Prilog poznavanju zooplanktona obalnih voda južnog Jadrana.
Ekologija/Acta Biologica Jugoslavica/.10,1 , 79–106.
VUKANIĆ ,V., 2004: Ekološka studija zooplanktona u funkciji analize stanja i zaštite
biodiverziteta Bokokotorskog zaliva; (Magistarska teza) Biološki fakultet,
Univerziteta u Beogradu, SiCG p.1-128.
Z E I ,M., 1956: Pelagic Polychaetes of the Adriatic. – Thalassia Jugosl., 1,2, 33-68.
Received: 02. 10. 2007.

72
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NATURA MONTENEGRINA, PODGORICA, 6:73-89
A CONTRIBUTION TO THE KNOWLEDGE OF THE DRAGONFLIES (ODONATA) OF
THE RIVER ZETA (MONTENEGRO)
Bogić GLIGOROVIĆ 1) , Vladimir PEŠIĆ 2) & Aleksandra Z E K O V I Ć 1)
1
Department of Biology, Faculty of Sciences, University of Montenegro, Cetinjski put b.b., 81000 Podgorica,
Montenegro (bogic1@cg.yu)
2
Department of Biology, Faculty of Sciences, University of Montenegro, Cetinjski put b.b., 81000 Podgorica,
Montenegro (pesicv@cg.yu)
Key words:
dragonflies,
new records,
faunistics,
the Zeta River,
Montenegro.
Synopsis
An updated list of the dragonflies (Odonata) of the
River Zeta (Montenegro) is given, including 27 species.
Three of them: Aeshna afinis Vander Linden, Lestes barbarus
(Fabricius) and Gomphus vulgatisimus (Linneus) are new for
fauna of Montenegro.
Ključne riječi:
vilini konjici,
novi nalazi,
faunistika,
rijeka Zeta,
Crna Gora.
Sinopsis
PRILOG POZNAVANJU ODONATA (Odonata) RIJEKE ZETE
(CRNA GORA)
U radu je data lista vilinih konica (Odonata) rijeke Zete
(Crna Gora), koja uključuje 27 vrsta. Tri vrste: Aeshna afinis
Vander Linden, Lestes barbarus (Fabricius) and Gomphus
vulgatisimus (Linneus) nove su za faunu Crne Gore.
INTRODUCTION
Although the order Odonata is an important group among the insects, studies of
the Odonata fauna in Montenegro are too few (e.g., A d a m o v i ć , 1949, 1996;
Adamović et al., 1996). The aim of our studies is both to give a new insight in the
fauna of the country, and to obtain valuable documentation of the occurence of
dragonflies in the previously unstudied area of the River Zeta. In the course of this
survey, we detected 27 species, among them 3 are new for the fauna of Montenegro.

74
Natura Montenegrina, 6/2007
FIGURE 1. Map of study area
with the sampling sites.
MATERIALS AND METHODS
In 2006-2007 the dragonflies fauna from 8 sampling sites on the River Zeta were
studied (Fig. 1). The River Zeta is the main tributary of the River Morača and belongs
to the drainage basin of the Skadar Lake, which is the largest lake of the Balkan
Peninsula, with a surface area varying between 370 and 600 km 2 . Lakes Skadar and
Ohrid and the River Drim, compose the largest river system in the western Balkan
zoogeographic region (B ă n ă rescu, 1992), and drains in to the Adriatic Sea basin.

75
Gligorović & a l . A CONTRIBUTION TO THE KNOWLEDGE OF THE DRAGONFLIES . . .
It starts near Nikšić, under the Planinica hill with a mean discharge in its mouth of 74
m 3 s -1 , (M a r t i n o v i ć -Vitanović & Kalifatić , 1995), flows eastwards for 86 km
until it confluxes into the Morača River near Podgorica.
Odonata specimens were collected with aerial nets. Specimens were put into
envelopes and transported to the lab. All the specimens were deposited in the
collection of the first author. B e s c h o v s k i (1994) were used for identification of the
specimens. Unless stated otherwise, all material has been collected by the first
author. The composition of the material is given as: (males/females/larvae).
RESULTS
Subordo Zygoptera
Family Calopterygidae
Calopteryx virgo (Linneaus, 1758)
Distribution: Palaearctic.
Material: Studenci, 5 May 2007 (0/0/1), 30 June 2007 (2/0/0); Slap Zete, 9
March 2007 (0/0/2), 16 April 2007 (0/0/1), 5 May 2007 (2/0/0), 13 July 2007 (2/1/1);
Danilovgrad, 21 July 2007 (2/1/0), 26 August 2007 (2/0/0); Martinići, 18 May 2007
(0/0/3), 21 June 2007 (3/2/1), 13 July 2007 (1/1/0); Dobro Polje, 21 July 2007 (2/2/0);
Tunjevo, 21 June 2007 (2/1/2), 30 June 2007 (2/0/0), 13 July 2007 (2/1/0); Glava
Zete, 9 March 2007 (0/0/2), Vidrovan.13 July 2007 (2/1/0).
Calopteryx splendens ( Harris, 1782)
Distribution: Palaearctic.
Material: Studenci, 9 March 2007 (0/0/1), 30 June 2007 (1/2/1); Danilovgrad, 21
June 2007 (3/1/0); Martinići, 16 April 2007 (0/0/2), 5 May 2007 (2/0/0), 30 June 2007
(2/1/0); Tunjevo, 13 July 2007 (3/2/0), 26 August 2007 (1/1/3); Glava Zete, 21 June
2007 (3/1/1); Vidrovan, 14 July 2007 (2/1/1).
Family Platycnemidae
Platycnemis pennipes (Pallas, 1771)
Distribution: Mediterranean.
Material: Martinići 13 July 2007 (2/2/1).
Family Lestidae
Lestes sponsa (Hensemann, 1823)
Distribution: Palaearctic.
Material: Vidrovan, source 14 July 2007 (2/1/0).
Remarks: New for Monenegro.

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Natura Montenegrina, 6/2007
Lestes dryas Kirby, 1890
Distribution: Holarctic.
Material: Vidrovan, source, 14.07.2007 (2/0/0).
Lestes barbarus (Fabricius, 1798)
Distribution: Holomediterranean.
Material: Studenci, 21.06.2007 (3/1/2); Danilovgrad, 26.08.2007 (1/1/0);
Martinići, 21.07.2007 (1/1/0); Dobro Polje, 18.05.2007 leg. Zeković (1/1/0);Tunjevo,
30.06.2007 (1/1/0),
Remarks: New for Montenegro.
Family Coenagrionidae
Ischnura elegans (Vander Linden, 1820)
Distribution: Palaearctic.
Material: Martinići, 21 June 2007 (2/2/0); Dobro Polje, 30 June 2007 (2/2/0);
Vidrovan, 14 July 2007 (2/0/0).
Ischnura pumilo (Charpentier, 1825)
Distribution: Mediterranean.
Material: Tunjevo 30 June 2007 (3/2/0); Glava Zete, 30 June 2007 (2/2/0).
Coenagrion ornatum (Sélys, 1850)
Distribution: Euro-mediterranean.
Material: Slap Zete, 21 June 2007 (2/1/0); Danilovgrad, 18 May 2007 (2/1/0), 30
June 2007 (2/2/0); Martinići, 26 August 2007 (2/1/0); Dobro Polje, 21 July 2007
(2/1/0); Tunjevo, 13 July 2007 (2/2/0); Glava Zete 13 July 2007 (2/2/0).
Coenagrion hastulatum (Charpentier, 1840)
Distribution: Euro-siberian.
Material: Slap Zete, 5 May 2007 (2/2/0), 21 June 2007 (3/2/0); Vidrovan, 14 July
2007 (3/2/0).
Coenagrion puella (Linneus, 1758)
Distribution: Euro-mediterranean.
Material: Martinići, 21 June 2007 (1/1/0); Dobro Polje, 26 August 2007 (1/1/0);
Tunjevo, 13 July 2007 (2/1/0), Glava Zete, 30 June 2007 leg. Zeković (2/2/0);
Vidrovan, 14 July 2007 (1/1/0).
Erythromma viridulum (Charpentier, 1840)
Distribution: Holomediterranean.
Material: Tunjevo, 30 June 2007 (2/2/0), 13 July 2007 leg.Zeković (1/1/0).

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Gligorović & a l . A CONTRIBUTION TO THE KNOWLEDGE OF THE DRAGONFLIES . . .
Subordo Anisoptera
Family Aeshnidae
Aeshna afinis Vander Linden, 1820
Distribution: Euro-mediterranean.
Material: Slap Zete, 26 August 2007 (2/1/0); Dobro Polje, 18 May 2007 leg.
Zeković (0/0/1), 13 July 2007 (1/1/0).
Remarks: New for Monenegro
Aeshna cyanea (Müller, 1764)
Distribution: Europe.
Material: Vidrovan, izvor, 14 July 2007 (1/1/2).
Family Libellulidae
Orthetrum anceps (Schneider, 1845)
Distribution: East Mediterranean.
Material: Danilovgrad, 30 June 2007 (2/1/0); Martinići, 21 July 2007 (3/0/0);
Dobro Polje, 30 June 2007 (1/2/2); Tunjevo, 26 August 2007 (0/1/0); Vidrovan, 14 July
2007 (2/1/0).
Orthetrum cancellatum ( Linnaeus, 1758)
Distribution: Euro-siberian.
Material: Slap Zete, 13 July 2007 (2/1/0); Martinići, 30 June 2007 leg. Zeković
(1/1/2); Tunjevo, 30 June 2007 (1/0/0).
Crocothemis erythraea (Brullé, 1823)
Distribution: Mediterranean.
Material: Danilovgrad, 30 June 2007 (2/1/2); Martinići, 26 August 2007 (2/0/0).
Libelulla depressa (Linnaeus, 1758)
Distribution: Europe
Material: Danilovgrad, 30 June 2007 (4/2/4); Martinići, 21 July 2007 (3/0/0);
Dobro Polje, 21 June 2007 (3/1/3); Tunjevo, 13 July 2007 (3/2/0); Glava Zete, 9 March
2007 leg. Zeković (0/0/4), 26 August 2007 (4/0/0); Vidrovan, 14 July 2007 (2/1/2).
Sympetrum flaveolum (Linnaeus, 1758)
Distribution: Palaearctic.
Material: Tunjevo, 13 July 2007 (2/0/0), 21 July 2007 (2/1/1).
Sympetrum meridionale (Sélys, 1841)
Distribution: Mediterranean.

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Natura Montenegrina, 6/2007
Material: Studenci, 18 May 2007 (2/0/0), 13 July 2007 (2/1/1); Slap Zete, 26
August 2007 (2/0/0),
Sympetrum striolatum (Charpentier, 1840)
Distribution: Palaearctic.
Material: Studenci, 18 May 2007 (2/1/0); Slap Zete, 26 August 2007 (2/2/2);
Danilovgrad 21 July 2007 (2/1/1), 26 August 2007 (2/0/0); Glava Zete, 5 May 2007
leg.Zeković (2/0/0), 30 June 2007 (2/2/1); Vidrovan 14 July 2007 (1/0/0).
Sympetrum sanquineum ( Müller, 1764)
Distribution: Euro-mediterranean.
Material: Martinići, 21 July 2007 (2/1/2).
Family Cordulegastridae
Cordulegaster bidentatus (Sélys, 1843)
Distribution: Mediterranean.
Material: Studenci, 16 April 2007 (0/0/2), 5 May 2007 (0/0/1), 13 July 2007
(2/1/0), 21 July 2007 (2/0/0), 26 August 2007 (0/1/0); Tunjevo, 21 June 2007 (2/2/2),
30 June 2007 (2/0/0), 13 July 2007 (2/1/0); Vidrovan, 14 July 2007 (3/1/2).
Family Cordulidae
Somatochlora metallica (Vander Linden, 1825)
Distribution: Euro-siberian subregion.
Material: Slap Zete, 30 June 2007 (2/1/0), 13 July 2007 (2/1/0), 21 July 2007
(2/0/0); Danilovgrad, 13 July 2007 leg. Zeković (2/3/0); Dobro Polje, 21 July 2007
(2/0/0); Tunjevo, 30 June 2007 (2/1/1); Glava Zete, 21 June 2007 (2/2/0); Vidrovan,
14July 2007 (2/1/2).
Family Gomphidae
Gomphus flavipes (Charpentier, 1825)
Distribution: Euro-siberian subregion.
Material: Studenci, 30 June 2007 (3/3/4), Slap Zete, 21 June 2007 (2/1/3), 30
June 2007 leg. Zeković (2/2/2); Danilovgrad, 30 June 2007 (2/0/0), Martinići, 21 June
2007 (2/2/4), 26 August 2007 (0/1/0), Dobro Polje, 21 July 2007 (2/0/0), Tunjevo, 30
June 2007 (1/0/0), 13 July 2007 (3/2/3), Glava Zete, 30 June 2007. (2/1/3), Vidrovan,
14 July 2007 (3/1/2).
Gomphus vulgatissimus (Linnaeus, 1758)
Distribution: European subregion.

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Gligorović & a l . A CONTRIBUTION TO THE KNOWLEDGE OF THE DRAGONFLIES . . .
Material: Studenci, 21 June 2007 (2/1/2); Slap Zete, 21 July 2007 (3/3/4); Dobro
Polje, 5 May 2007 (1/0/0), 18 May 2007 (1/1/2), 21 June 2007 leg. Zeković (2/0/0);
Tunjevo, 21 June 2007 (1/2/2); Glava Zete, 21 July 2007 (3/1/1); Vidrovan, 14 July
2007 (1/1/2).
Remarks: New for Montenegro
Onychogomphus forcipatus (Linnaeus, 1758)
Distribution: Holomediterranean subregion.
Material: Studenci, 16 April 2007 (2/0/0), 21 June 2007 (3/2/3); Tunjevo, 30 June
2007 (1/1/3), 13 July 2007 (1/2/3); Glava Zete, 18 May 2007 (3/1/4), 21 June 2007
(3/2/2); Vidrovan, 14 July 2007 (2/1/2).
DISCUSSION
In the freshwater habitats of the River Zeta 27 species of dragonflies (Odonata)
belonging to 15 genera and 9 families were recorded at 8 sites. Of these, 3 species:
Aeshna afinis Vander Linden, Lestes barbarus (Fabricius) and Gomphus vulgatisimus
(Linneus) are recorded from Montenegro for the first time.
The total number of specimens of dragonflies collected on the River Zeta
amounts to 486. Eight species are dominant (> 5% total abundance): Calopterix virgo
(Linneus), C. splendens (Harris), Cordulegaster bidentatus (Sélys), Somatochlora
metalica (Vander Linden), Libelulla depressa (Linnaeus), Gomphus vulgatisimus
(Linneus), G. flavipes (Charpentier) and Onychogomphus forcipatus (Linneus). Six
species are subdominant (abundance 5-2%): Lestes barbarus (Fabricius), Coenagrion
ornatum (Sélys), C. hastulatum (Charpentier), C. puella (Linneus), Orthetrum anceps
(Schneider), Sympetrum meridionale (Sélys) and S. striolatum (Charpentier). The
remaning 12 species have abundance less than

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Natura Montenegrina, 6/2007
Table 1. Abundance of the collected species. D = dominant (> 5% total abundance), SD =
subdominant (2-5% total abundance), R = rare (< 2% total abundance).
Species
Number of
collected
specimens
Relative
abundance
%
Dominancy
1. Calopterix virgo (Linneus, 1758) 46 9.46% D
2. Calopteryx splendens (Harris, 1782) 34 6.99% D
3. Lestes sponsa (Hensemann, 1823) 3 0.61% R
4. Lestes dryas Kirby, 1890 2 0.41% R
5. Lestes barbarus (Fabricius, 1798) 14 2.88% SD
6. Platycnemis pennipes (Pallas, 1771) 4 0.82% R
7. Ischnura elegans (Vander Linden, 1820) 8 1.64% R
8. Ischnura pumilo (Charpentier, 1825) 9 1.85% R
9. Coenagrion ornatum (Sélys, 1850) 24 4.93% SD
10. Coenagrion hastulatum (Charpentier, 1840) 11 2.26% SD
11. Coenagrion puella (Linneus, 1758) 13 2.67% SD
12. Erythromma viridulum (Charpentier, 1840) 6 1.23% R
13. Aeshna cyanea Müller, 1764 4 0.82% R
14. Aeshna afinis Vander Linden, 1820 6 1.23% R
15. Cordulegaster bidentatus (Sélys, 1843) 26 5.34% D
16. Somatochlora metalica (Vander Linden,1825) 27 5.55% D
17. Libelulla depressa (Linnaeus, 1758) 38 7.81% D
18. Orthetrum cancellatum (Linnaeus, 1758) 8 1.64% R
19. Orthetrum anceps (Schneider, 1845) 15 3.08% S
20. Crocothemis erythraea (Brullé, 1823) 7 1.44% R
21. Sympetrum flaveolum (Linnaeus, 1758) 6 1.23% R
22. Sympetrum meridionale (Sélys, 1841) 17 3.49% SD
23. Sympetrum striolatum (Charpentier, 1840) 20 4.11% SD
24. Sympetrum sanquineum (Müller, 1764) 5 1.02% R
25. Gomphus vulgatisimus (Linneus, 1758) 36 7.4% D
26. Gomphus flavipes (Charpentier, 1825) 56 11.1% D
27. Onychogomphus forcipatus (Linneus, 1758) 41 8.43% D
ACKNOWLEDGEMENTS: We are grateful to Branimir Gligorović and Igor Temelkovski
for their logistic support.

81
Gligorović & a l . A CONTRIBUTION TO THE KNOWLEDGE OF THE DRAGONFLIES . . .
APPENDIX 1. Photographs of the collected species.
FIGURE 2.
Calopteryx virgo ♀
FIGURE 3.
Calopteryx virgo ♂
FIGURE 4.
Calopteryx splendens ♂
FIGURE 5.
Calopteryx splendens ♀
FIGURE 6.
Lestes sponsa (Hensemann) ♂
FIGURE 7.
Lestes dryas Kirby ♂

82
Natura Montenegrina, 6/2007
FIGURE 8.
Lestes barbarus (Fabricius) ♀
FIGURE 9.
Platycnemis pennipes (Pallas) ♂,♀
FIGURE 10.
Ischnura elegans (Vander Linden) ♂
FIGURE 11.
Ischnura pumilo (Charpentier) ♂,♀
FIGURE 12.
Coenagrion ornatum (Sélys) juvenile ♀
FIGURE 13.
Coenagrion ornatum (Sélys) larvae

83
Gligorović & a l . A CONTRIBUTION TO THE KNOWLEDGE OF THE DRAGONFLIES . . .
FIGURE 14.
Coenagrion puella (Linneus) juvenile ♀
FIGURE 15.
Coenagrion puella (Linneus) ♂
FIGURE 16.
Coenagrion hastulatum (Charpentier) ♂
FIGURE 17.
Erythromma viridulum (Charpentier) ♂
FIGURE 18.
Aeshna afinis Vander Linden ♂
FIGURE 19.
Aeshna afinis Vander Linden ♀

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Natura Montenegrina, 6/2007
FIGURE 20.
Aeshna afinis Vander Linden larvae
FIGURE 21.
Aeshna cyanea larvae
FIGURE 22.
Aeshna cyanea ♂
FIGURE 23.
Aeshna cyanea ♀
FIGURE 24.
Orthetrum anceps (Schneider) ♀
FIGURE 25.
Orthetrum cancellatum (Linnaeus) ♀

85
Gligorović & a l . A CONTRIBUTION TO THE KNOWLEDGE OF THE DRAGONFLIES . . .
FIGURE 26.
Crocothemis erythraea (Brullé) ♂
FIGURE 27.
Sympetrum meridionale (Sélys) ♀
FIGURE 28.
Libelulla depressa (Linnaeus) ♂
FIGURE 29.
Libelulla depressa (Linnaeus) ♀
FIGURE 30.
Sympetrum flaveolum (Linnaeus) ♀
FIGURE 31.
Sympetrum striolatum (Charpentier) ♂

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Natura Montenegrina, 6/2007
FIGURE 32.
Sympetrum sanquineum (Müller) ♂, ♀
FIGURE 33.
Cordulegaster bidentatus (Sélys) ♂
FIGURE 34. Somatohlora
metalica ♀ and ♂
FIGURE 35.
Gomphus vulgatissimus (Linnaeus) ♀
FIGURE 36.
Gomphus vulgatissimus (Linnaeus) ♂

87
Gligorović & a l . A CONTRIBUTION TO THE KNOWLEDGE OF THE DRAGONFLIES . . .
FIGURE 37.
Onychogomphus forcipatus ♀
FIGURE 38.
Gomphus flavipes (Charpentier) ♂
APPENDIX 2. Photographs of the sampling sites.
FIGURE 39. The River Zeta – Studenci
FIGURE 40. The River Zeta – Martinići
FIGURE 41. The River Zeta – Danilovgrad
FIGURE 42. The River Zeta – Slap Zete

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Natura Montenegrina, 6/2007
FIGURE 43. The River Zeta – Dobro Polje
FIGURE 44. The River Zeta – Tunjevo
FIGURE 45. The River Zeta – Glava Zete
FIGURE 46. The River Zeta – Vidrovan
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Adamović , Ž. 1949 : La liste des Odonates du Museum d'historie naturelle du Pays
Serbe. - Glasnik Prirodnjačkog muzeja srpske zemlje, Beograd, 1–2: 275– 293.
Adamović , Ž. 1996: Odonata taken and observed in Donji Ceklin, Montenegro. -
Acta Entomologica Serbica, 1(1/2): 39-48.
Adamović , Ž., Anđ us, Lj. & Mihajlović , L j . 1996: Odonata (Insecta) in:
K a r a m a n , G. (Ed.). The Fauna of Durmitor. - The Montenegrin Academy of
Sciences and Arts, 5, 43 – 80, Podgorica.
B ă n ă r e s c u P. M. 1992: Zoogeography of Freshwaters. Vol. 2. Distribution and
Dispersal of Freshwater Animals in North America and Eurasia. - AULA-Verlag,
Wiesbaden, 517–1091.
B e s c h o v s k i , V. 1994: Fauna Bulgarica. 23 Insecta, Odonata. - Bulg. Acad. Sci.,
Sofia. 372pp.

89
Gligorović & a l . A CONTRIBUTION TO THE KNOWLEDGE OF THE DRAGONFLIES . . .
Martinović -Vitanović V. & Kalafatić V. 1995: Basic Hydrogeological
Characteristics of Yugoslavian Freshwaters. In S t e v a n o v i ć V & V a s i ć V.
(eds), Biodiverzitet Jugoslavije. - Ecolibri, Biološki Fakultet, Beograd, 97–115 (in
Serbian).
Received: 22.11.2007

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NATURA MONTENEGRINA, PODGORICA, 6:91-100
PRELIMINARY INVESTIGATIONS OF ORTHOPTEROID FAUNA OF
THE ĆEMOVSKO POLJE, SEMI-DESERT FIELD NEAR PODGORICA, MONTENEGRO
Jelena N I K Č EVIĆ 1)
1)
Institute for the Protection of Nature of Montenegro, 81000 Podgorica, Trg Bećir bega Osmanagića 16,
Montenegro, E-mail: jnikcevic@cg.yu
Key words:
fauna,
orthoptera,
Ćemovsko polje field,
semi-desert,
ecological
characteristics
SYNOPSIS
Orthopteroid fauna of Ćemovsko polje, a semi-desert
field in Montenegro, is unknown and insufficiently studied.
Owing to the investigations in period 2006-2007 in the area
of Ćemovsko polje 37 species were determined and grouped
in 4 superorders, 2 orders, 8 families and 27 genera. Data
about ecological chara-cteristics, distribution, zoogeography
and type of endemism for each species are presented.
Ključne reči:
fauna,
pravokrilci,
Ćemovsko polje,
polupustinja,
ekološke karakteristike
SINOPSIS
PRELIMINARNA ISTRAŽIVANJA FAUNE
ORTOPTEROIDNIH INSEKATA POLUPUSTINJSKOG PODRUČJA
ĆEMOVSKOG POLJA, CRNA GORA
Fauna ortoptera Ćemovskog polja u Crnoj Gori je
nepoznata i neistražena. Zahvaljujući istraživanjima u
periodu 2006-2007. godine na prostoru Ćemovskog polja
determinisano je 37 vrsta svrstanih u 4 nadreda, 2 reda, 8
familija i 27 rodova. U radu su takodje dati podaci o
distribuciji prisutnih vrsta, zoogeografskoj pripadnosti, tipu
endemizma i ekološkim karakteristikama.
INTRODUCTION
Throughout southern Europe, 14000 years ago the predominant picture was of
an arid, semi-desert like steppe perhaps similar to the cool and arid mountain steppes
found today in the Pamirs of southern Russia. A rapid warming and moistening of
climate all across Europe occurred shortly before this time, with insect communities in

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NW Europe suggesting conditions as warm as or warmer than the present-day for
about the first 500 14C years (Atkinson et al. 1987). That sudden event was the basis
for today’s picture of orthopteroid fauna and their evolution.
The suddenness of this warming left ecosystems out of equilibrium. Trees had
not yet had time to spread back over Europe, and a steppe cover predominated. The
ice sheets in the north began to melt, but that had lasted for thousand years less than
in the normal warm climate period. Some open woodland cover appeared in western
Russia quite rapidly – within a few hundred years - after the initial warming events and
woodland cover began increasing in the mountainous areas of southern Europe where
trees had survived the glaciations.
Data for orthoipteroid insects from region of Ćemovsko polje, a semi-desert
field near Podgorica, Montenegro, are unknown in the literature. In course of
investigation in period 2006-2007 of the orthopteroid fauna of this field, the detailed
data about ecological characteristics and distribution in the semi-desert condition
were established.
MATERIAL AND METHODS
Material was collected during 2006/2007. Depending of differences in the
taxonomic group sampling was realised by application of different methods (collecting
with mowing and method with individual catching). Collected material was identified by
using keys: H a r z , K. (1969): The Orthoptera of Europe. Vol. I. The Hague; H a r z , K.
(1975): The Orthoptera of Europe. Vol. II. The Hague; H a r z , K., K a l t e nbach, A.
(1976): The Orthoptera of Europe. Vol. III. The Hague; U s , P. (1992): Favna
Ortopteroidnih insektov Slovenije, Slovenska Akademija Znanosti in Umetnosti,
Razred za prirodoslovne nauke, Biološki Institut Jovana Hadžija, br. 12., Ljubljana.
INVESTIGATED AREA
The Ćemovsko semi-desert field is the part of Zeta valey in Montenegro. It is
situated southeast of the capital town of Montenegro – Podgorica, between the two
rivers Morača and Cijevna. The soil subastrate of this field is made of fluvioglacial
sand. In the field there are hills: Ljubović (112m), Zelenika (166m) and Srpska Gora
(95m). The climate is modified Mediterranean one. The greater part of the filed is used
for human activities. Some parts of the field, nearer to Podgorica, are under the
great antropogenic influences.

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Nikčević: PRELIMINARY INVESTIGATIONS OF ORTHOPTEROID FAUNA OF THE ĆEMOVSKO POLJE . . .
Picture 1. Ćemovsko field (photo:J. Nikčević)
Biogegrafic charactetistics of vegetation:
This type of semi-desert habitats biogeograficaly belongs to Mediterranean
semi-desert province (Š i m i ć , 1987 in L a k u š i ć , R.) with discontinueted areal from
northwest Africa and Pirinei peninsula in the West, to Iran and Irak to the East. This
type of geological soil, climate features and ecological factors are basic for xeroterm
vegetation of semi-desert type. According to Černjavski (1949) this type of vegetation
communities is predominated by Satureia subspicata-Poa bulbosa which spread all
over the field.
RESULTS
The results of the investigations from 2006 and 200. at Ćemovsko polje field are
presented on the Table 1. with remarks on distribution, endemic and ecological
characteristics. In accordance to these investigations 37 species belonging to 27
genera classified into 4 superordo, 2 ordo and 7 families of Orthopteroidea have been
determined.

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Table1: The preliminary list of orthopteroid fauna at the semi-desert field Ćemovsko
polje, with remarks on distribution, endemic and ecological characteristics.
EU: European, M: Mediterranean, MS: Mediterranean-submediterranean, OM:
oromediterranean-mediterranean, TM: tropic-mediterranean, TPM: tropicpontomediterranean
PM: pontomediterranean, E: endemic characteristics, EC: ecological characteristics
EU M MS OM TM TPM PM E EC
MANTODEA
Mantidae
1.Mantis religiosa
Linneus, 1758
Empusidae
2.Empusa fasciata -
Brulle 1836
GRYLLOPTERA
TETTIGONIOIDEA
Tetigonidae
3.Leptophyes
punctatissima
(Bosc, 1792)
4.Phaneroptera
nana
Fieber, 1853
5.Tylopsis liliifolia
(Fabricius, 1973)
6.Saga pedo
(Pallas, 1771)
7.Eupholidoptera
chabrieri schmidti
(Charpentier, 1825)
8.Metrioptera
ambitiosa
Uvarov, 1923
9.Metrioptera
oblongicollis
(Bruner, 1882)
10.Pholidoptera
dalmatica
(Krauss, 1878)
11.Pholidoptera
littoralis
(Fiaber, 1853)
12.Pholidoptera
maritima
Zeuner, 1931
13.Rhacocleis
germanica (Herrich +
+ phitophil, thermophil
+ thermophil
+ thermophil
phytophil
silvicol
+ thermophil
arbusticol
arboricol
+ thermophil
graminicol
arbusticol
+ praticol
graminicolarbusticol
pratinicol
thermophil
+ mediterran/
submediterran
silvicol, terricol
arbusticol
thermophil
+ Balcan praticol higrophil
phitophil
+ Balcan praticol xeromesophil
phitophil
+ mediterran thermophil,
xerophil,
graminicol
+ mediterran silvicol arbusticol
thermophil
+ thermophil
xerophil
graminicol
thermophil

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Nikčević: PRELIMINARY INVESTIGATIONS OF ORTHOPTEROID FAUNA OF THE ĆEMOVSKO POLJE . . .
Schaeffer, 1840)
14.Ephgippiger
ephippiger
(Fiebig, 1784)
GRYLLODEA
Gryllidae
15.Oecanthus
pellucens
(Scopoli) 1763
Gryllotalpidae
16.Gryllotalpa
gryllotalpa
(Linneus, 1758)
ORTHOPTERA
TETRIGODEA
Tetrigidae
17.Depressotetrix
depressus
Brisout, 1848
ACRIDOIDEA
Catantopidae
18.Pezottetix
giornae
(Rossi, 1794)
19.Anacridium
aegyptum
(Linneus 1764)
20.Schistocerca
gregaria
(Forskal, 1775)
21.Calliptamus
italicus
(Linneus 1758)
Acrididae
22.Acrida turrita
(Linnaeus, 1758)
23.Acrida ungarica
ssp. mediterranea
(Herbst 1786)
24.Acrotylus
insubricus
(Scopoli, 1786)?
25.Acrotylus
longipes
(Charpentier,1845)
+ pratinicol silvicol
arbusticol
+ silvicol herbicol
arboricol
+ geobiont
higrophil mesophil
+ xerophil
+ thermophil
pratinicol
terricol
herbicol
+ arbusticolarboricol
+ arbusticol
arboricol
+ thermophil,
pratinicol
tericol
herbicol xerophil
+ thermophil
pratinicol
graminicol
+ thermophil
hygrophil
pratinicol
graminicol
+ pratinicol
termophil
tericol,
psamophil
xerophil
+ Pratinicol
Termophil
tericol, psamophil,
xerophil

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26.Acrotylus
patruelis (Herrisch-
Schaeffer, 1838)
27.Aiolopus
strepens
(Latreille) 1804
28.Aiolopus
thalassinum
(Fabricius 1781)
29.Oedipoda
coerulescens
(Linnaeus, 1758)
30.Oedipoda
germanica
(Latreille, 1804)
31.Oedipoda
miniata
(Pallas, 1771)
32.Psophus
stridulus
(Linne, 1758)
33.Sphingonotus
coerulans
(Linneus, 1767)
34.Aeropus
sibiricus
(Linneus) 1767
35.Dociostaurus
genei
(Ocskay) 1832
36.Dociostaurus
marrocanus
(Thunberg 1815)
37.Omocestus
rufipes
(Zetterstedt, 1821)
+ pratinicol,
thermophil, terricol,
psamophil, xserophil
+ thermophil,
pratinicol,
geophil,
phitophilmesophil
+ thermophil, pratinicol,
geophilphitophil,
higrophil
+ pratinicolterricol,
xerophil
+ pratinicolterricol,
litophil, xerophil
+ xerophil, thermophil
+ thermophil, pratinicol,
terricol
xerophil
+ terricol,
psamophil,xerophil
+ hygrophilstenotherm,
thermophob
+ thermophil, pratinicol
+ thermophil, pratinicol,
geophilphitophil
+ xerophil
DISCUSSION
Discussion in this chapter is about diversity and connections among the
orthopteroid insects within different taxonomic groups. The zoogeographic
characteristics of species are also presented, with remarks on endemic species.
MANTODEA
Fam- Mantidae - 1 species
Mantis religiosa
Fam. Empusidae – 1 species
Empusa fasciata

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Nikčević: PRELIMINARY INVESTIGATIONS OF ORTHOPTEROID FAUNA OF THE ĆEMOVSKO POLJE . . .
TETTIGONIOIDEA
Fam. Tettigonidae – 12 species
Leptophyes punctatissima, Phaneroptera nana, Tylopsis liliifolia, Saga pedo,
Eupholidoptera chabrieri schmidti, Metrioptera ambitiosa, Metrioptera oblongicollis,
Pholidoptera dalmatica, Pholidoptera littoralis, Pholidoptera maritima, Rhacocleis
germanica , Ephgippiger ephippiger
Fam. Gryllidae - 2 species
Oecanthus pellucens, Gryllotalpa gryllotalpa
TETRIGODEA
Fam. Tetrigidae - 1 species
Depressotetrix depressus
ACRIDOIDEA
Fam. Catantopidae - 4 species
Pezottetix giornae, Anacridium aegyptum, Schistocerca gregaria, Calliptamus italicus
Fam Acrididae – 16 species
Acrida turrita, Acrida ungarica ssp. mediterranea, Acrotylus insubricus, Acrotylus
longipes, Acrotylus patruelis, Aiolopus strepens, Aiolopus thalassinum, Oedipoda
coerulescens, Oedipoda germanica, Oedipoda miniata, Psophus stridulus,
Sphingonotus coerulans, Aeropus sibiricus, Dociostaurus genei, Dociostaurus
marrocanu, Omocestus rufipes
The largest diversity of species belongs to family Acrididae (16), after which there
follow the diversity of family Tettigonidae (12). The other types of species belong to
families with smaller diversity (Catantopidae 4, Gryllidae 2, and families Mantidae,
Empusidae and Tetrigidae with 1 species).
Zoogeographic analysis of faunistic elements:
European species: 13
Mantis religiosa, Leptophyes punctatissima, Saga pedo, Rhacocleis germanica,
Gryllotalpa gryllotalpa, Pezottetix giornae, Aiolopus strepens, Aiolopus thalassinu,
Oedipoda coerulescens, Oedipoda germanica, Psophus stridulus, Aeropus sibiricus,
Omocestus rufipes
Mediterranean species: 2
Pholidoptera dalmatica, Pholidoptera littoralis
Mediterranean – submediterranean species: 11
Empusa fasciata, Tylopsis liliifolia, Eupholidoptera chabrieri schmidti, Metrioptera
ambitiosa, Metrioptera oblongicollis, Pholidoptera maritima, Oecanthus pellucens,
Depressotetrix depressus, Calliptamus italicus, Dociostaurus genei, Dociostaurus
marrocanus

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Picture 4. Ćemovsko field (photo:J. Nikčević)
Oromediterranean-mediterranean species: - no data
Tropic-mediterranean species: 2
Acrotylus patruelis, Sphingonotus coerulans
Tropicpontomediterranean species: 5
Anacridium aegyptum, Schistocerca gregaria, Acrida turrita, Acrida ungarica ssp.
mediterranea, Acrotylus insubricus
Pontomediterranean species: 4
Phaneroptera nana, Ephgippiger ephippiger, Acrotylus longipes, Oedipoda miniata
Endemic species: 5
Balcan endemic species: 2
Metrioptera ambitiosa , Metrioptera oblongicollis
Mediterranean endemic species: 2
Pholidoptera dalmatica , Pholidoptera littoralis
Mediteranean-submediterranean species: 1
Eupholidoptera chabrieri schmidti

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Nikčević: PRELIMINARY INVESTIGATIONS OF ORTHOPTEROID FAUNA OF THE ĆEMOVSKO POLJE . . .
The greatest number of species (13) belongs to European type of distribution,
followed by 11 species of Mediterranean-submediterranean distribution. Five species
belongs to tropic-pontomediterranean zoogeographic distribution and four species to
pontomediterranean species. The lowest number (2) of species are those with
mediterranean and tropic-mediterranean type of distribution. There are no available
data for oromediterranean-mediterranean species distribution.
There are 5 endemic species with Balkan endemic type and 2 species with
Mediterranean endemic type and 1 species of Mediterranean-submediterranean type
of endemism.
In this discussion we can see that the greatest number of species have
thermophil, xerophil, graminicol ecological characteristics.
CONCLUSIONS
During the investigation of ortopteroid insects 37 species belonging to 27 genera
classified into 4 superordo, 2 ordo and 7 families were determined from collected
material. The largest diversity of species belongs to family Acrididae (16), followed by
the diversity of family Tettigonidae (12). The other types of species belong to families
with smaller diversity (Catantopidae 4, Gryllidae 2, and families Mantidae, Empusidae
and Tetrigidae with 1 species).
The greatest number of species (13) belongs to European type of distribution,
then follow 11 species of mediterranean-submediterranean distribution. Five species
belong to tropic-pontomediterranean zoogeographic distribution, 4 species to
pontomediterranean ones. The lowest number (2) of species belongs to species with
mediterranean and tropic-mediterranean type of distribution. For oromediterraneanmediterranean
species there are no data so far.
We can see that the greatest number of species have thermophil, xerophil,
graminicol ecological characteristics in every type of taxonomical belonging.
In Ćemovsko polje filed, according to this investigation (in literature there are no
data about orthopteroid fauna on that field), the diversity of orthopteroid fauna is
relatively small owing to a low diversity of vegetation cover, but abounding in relict
and endemic species and species with low number specimens. Last characteristic is
present in some rare taxa. This „small number“of orthopteroid insects has an origin in
still insufficiently studied conditions in the history of orthopteroid evolution for that
area. Some species and some habitats were on the same place at the same time with
different ecological characteristics. Vegetation communities were displaced from one
type to the other, opposite to the natural way of evolution of origin and succession.
REFERENCES
A D A M S , J.: Environmental Sciences Division, Oak Ridge National Laboratory, Oak
Ridge, TN 37831, USA
Č E J C H A N, A. 1984: Catalogue of the Orthopteroid Insects of Montenegro - Acta
faunistica entomologica musei nationalis Pragae sv. XVII: 9-22, Pragae.

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Č ERNJAVSKI, P., GREBENŠČ IKOV, O. & PAVLOVIĆ , Z. 1949: O flori i
vegetaciji Skadarskog područja. - Glasn. Prir. Muz. Srpske Zemlje, Ser. B, Biol.
Nauke 1-2: 4-91. Beograd.
H A R Z, K. 1969: The Orthoptera of Europe. Vol. I. The Hague.
H A R Z, K. 1975: The Orthoptera of Europe. Vol. II. The Hague
H A R Z, K., KALTENBACH, A. 1976: The Orthoptera of Europe. Vol. III. The Hague.
I N G R I S C H, S., KÖEHLER, G. 1998: Die Heuschrecken Mitteleuropas, Die Neue
Brehm-Bucherei Bd.629: 27-41, Westarp Wissenchaften - Magdeburg.
ŠIMIĆ , S. 1987: Syrphidae (Insecta, Diptera), Biogeografska i ekološka analiza
faune osolikih muva Durmitora sa osvrtom na faunu osolikih muva Crne Gore,
Fauna Durmitora Sv. 2., CANU, Posebna izdanja, knj. 21, Odeljenje prirodnih
nauka, knj. 13: 11-154, Titograd.
U S , P. (1992): Favna Ortopteroidnih insektov Slovenije, Slovenska Akademija
Znanosti in Umetnosti, Razred za prirodoslovne nauke, Dela 32, Biološki institut
Jovana Hadžija, 12:61-285.
Received: 25.10.2007.

NATURA MONTENEGRINA, PODGORICA, 6:101-109
MORPHOLOGICAL TRAITS OF COMMON TOAD Bufo bufo (Bufonidae) FROM
BIOGRADSKO LAKE
Natalija Č A Đ ENOVIĆ ¹, Tanja V U K O V ²
¹ Natural History Museum of Montenegro, Podgorica, Crna Gora, E-mail: lazo@cg.yu
² Institute for Biological Research “Siniša Stanković”, Beograd, Srbija
Key words:
Bufo bufo,
morphometric,
qualitative characters,
differences between
the sexes.
SYNOPSIS
In this paper we have presented the results of the
analysis of morphological traits of Common Toad Bufo bufo
population, from Biogradsko Lake. On the samples taken
from this locality univariant and multivariant statistical
analyses have been performed. A large number of
morphometric and qualitative characters have been done for
the first time. For morphometric characters basic parameters
of descriptive statistics have been calculated, and for
qualitative characters three qualitative traits with three
conditions at four body regions of Common Toad separately
by sexes, have been treated. As for the morphometric
characters it has been established that there is a significant
statistical difference between the sexes.
Ključne riječi:
Bufo bufo,
morfometrijski,
kvalitativni karakteri,
razlike među
polovima.
SINOPSIS
MORFOLOŠKE KARAKTERISTIKE OBIČNE KRASTAVE ŽABE
BUFO BUFO (BUFONIDAE) SA BIOGRADSKOG JEZERA
U ovom radu su prikazani rezultati analize morfoloških
odlika populacije krastave žabe Bufo bufo, sa Biogradskog
jezera. Na uzorcima uzetim sa ovog lokaliteta urađene su
univarijantne i multivarijantne statističke analize. Po prvi put
je obrađen veliki broj morfometrijskih i kvalitativnih karaktera.
Za morfometrijske karaktere izračunati su osnovni parametri
deskriptivne statistike, a za kvalitativne karaktere obrađene
su tri kvalitativne karakteristike sa tri stanja na četiri tjelesna
regiona obične krastave žabe odvojeno po polovima. Što se
tiče morfometrijskih karaktera utvrđena je statistički značajna
razlika između polova.

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INTRODUCTION
Common Toad has a large distribution and popultes mainly the entire palearctic
area. It populates almost the entire Europe, with the exception of far north, Ireland,
Corsika, Sardinia, Balear's Islands, Malta, Crete, as well as some smaller
islands (B o r k i n and V e i t h , 1997). It also populates the northwestern Africa
(P a s t e u r and B o n s , 1959). Although common at lower altitudes, it may also be
found also on altitudes exceedeing 2500 m and on Iberian Peninsula (A n g e l , 1946;
B o r k i n and V e i t h , 1997). In Montenegro it is mainly widely distributed. Owing to
wide distribution, climatic conditions in various parts of areal of distribution are very
different. Thus, in northern part of the areal the winters are long, cold and humid, so
that the period favorable for growth is rather short (J o h a n n e s s e n , 1970). On
higher altitudes the conditions are even more extreme. On the other hand, southern
populations of this species are exposed to much more favorable conditions of outer
environment; therefore the period favorable for growth is much longer.
Species Bufo bufo is a polytypic one. From the scope of this species a taxon
from the Far East has been separated on the level of species Bufo gargarizans
(A n a n j e v a et al. 1998, K u z m i n 1999a, 1999b), and the character of polytype is
determined to it in addition to nominotype subspecies three other ones B. b.
grediscola, B. b. verrucosissimus, B. b. spinosus. If we recognize the fact that
Caucasus Toad (Bufo verrucosissimus) is a separate species (e.g. O r l o v a &
T u n i y e v 1989, A n a n j e v a et al. 1998), than the taxonomic differentiation in scope
of Common Toad is even more limited. Nominotype subspecies is the most widely
distributed (Northern and Southern Europe). The two first subspecies have a limited
geographical area (mountains Gredos and Caucasus), nominotype subspecies is the
most widely distributed (Northern and Central Europe), whereas the areal of
subspecies B. b. spinosus is limited to the region of Mediterranean (northwestern
parts of Africa, southeast Europe, southern Switzerland, Italy as well as the southern
parts of France and Spain).
MATERIAL AND METHODS
A total of 41 specimens has been analyzed, 26 males and 15 females. The
largest part of the material has been collected in mating period. The material has
mainly been manually collected and meredov (a net with a handle) has also been
used.
The samples belong to herpetological collection of the Natural History Museum
of Montenegro; they are kept in 70 % alcohole.
Morphometric analysis has been done on 21 traits which determine size and
form of body and head of tailless amphibians. Measured traits are: L – longthe of body

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Čađenović & Vukov: MORPHOLOGICAL TRAITS OF COMMON TOAD BUFO BUFO . . .
from the top of head to the opening of the cloaca; Lpa – length of front exstremity; F –
length of femur measured fromxthe opening of cloaca to knee joint; T – lengthj of
tibiofibula from knee joint to tibiotarsal joint; P – distance from tibiotarsal joint to the
tip of the longest finger (IV); n – length of metatarsus from the internal metatarsal
ridgelet to the tip of the longest finger (IV); DpPa – length of the first finger off front
exstremity; DpPp – length of the first finger of back extremity; Cint – the biggest
length of the internal metatarsal ridgelet; Lc – length of head from the top of head to
jaw joint; Ltc – width of head measured between the jaw joints; Spp – betweeneyes
space between the internal edges of eye lids; Lc – lengh of head from the top of head
to jaw joint; Ltc – width of head measured between jaw joints; Spp – betweeneyes
space between the internal edges of eye lids; Spi – distance betwnn the outer nasal
openings; Spcr – distance between front angles of eye openings; Lo – the largest
length of eyeball; Ltp – the largest width of upper eye lid; Dro – distance from the top
of head to the eye edge; Dno- distance of nasal opening from eye; Lh - length of
cutaneous folds on heels; Lg - length of partoid salivary glands; Wg - width of
salivary glands and weight.
Three qualitative traits (Table 1) with three conditions on four body regions of
toad: head part - dorsal side, dorsal part-dorsal side, head part -ventral side,
abdominal part-ventral side.
The conditions of qualitative traits have been coded in the following manner:
Dorsal side – head part:
1a - round warts, 1b - oval warts, 1c – kidneylike warts; 2a – very protuberant
warts, 2b – protuberant warts, 2c – drawn in warts; 3a – very marked thornlike ends of
warts; 3b – medium marked thornlike ends of warts; 3c – weakly marked thornlike
ends of warts;
Dorsal side - dorsal part:
4a - round warts, 4b - oval warts, 4c - kidneylike warts; 5a – very protuberant
warts, 5b – protuberant warts, 5c – drawn in warts; 6a – very marked thornlike ends of
warts; 6b – medium marked thornlike ends of warts; 6c – weakly marked thornlike
ends of warts;
Ventral side – head part:
7a - round warts, 7b - oval warts, 7c - kidneylike warts; 8a – very protuberant
warts, 8b – protuberant warts, 8c – drawn in warts; 9a – very marked thornlike ends of
warts; 9b – medium marked thornlike ends of warts; 9c – weakly marked thornlike
ends of warts;

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Ventral side – abdominal part:
10a - round warts, 10b - oval warts, 10c - kidneylike warts; 11a – very
protuberant warts, 11b – protuberant warts, 11c – drawn in warts; 12a – very marked
thornlike ends of warts; 12b – medium marked thornlike ends of warts; 12c – weakly
marked thornlike ends of warts;
Trait
Warts shape
Condition
a) round
b) oval
c) reniform
a)very convex
Warts convexity
b) convex
c) inverted
Prominence of thornlike
warts endings
a)very prominent
b)averagely prominent
c)weakly prominent
Table 1. Analysed qualitatie traits in Bufo bufo
Statistical Analysis
Programme package STATISTICA (version 5.0) has been used for statistical
analysis of data. For morphometric characters basic parameters of descriptive
statistics have been separately calculated by genders: medium value ( x ), error of
medium value (SE), standard deviation (SD), minimal (min) and maximal (max) values
traits as well as coeficient of variation (CV). Analysis of sexual dimorphism has been
performed on basic data (untransformed measures for every property), as well as on
standardized residuals on eight traits (L, Lpa, F, T, P, n, Lc, Ltc). Regression of every
property on body length (L) and use of standardized residuals from such a regression
provides for the elimination of the impact of body size. Comparison of differences
between the genders on basic data and on standardised residuals has been performed
by the use of Tukey Test for the samples of uneven size.
For the establishment of the significance of differences in respect of
morphometric characters between the genders analysis of variance (ANOVA) has
been used.
Absolute and relative frequences (%) of the conditions of qualitative traits have
been established. Qualitative traits have been analysed also through the
corresponding analyses.

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RESULTS AND DISCUSSION
Morphometric Characters
Former data on values of morphometric characters of the species Bufo bufo are
rather insufficient, or, some of them have been given only descriptively. That is why
we have presented here the average values for a large number of characters that
have not been previously analyzed and data about that do not exist in the literature.
Basic parameters of morphometric characters descriptive statistics have been
presented separately by sexes in the table 2. The smallest male of the sample of the
population from Biogradsko Lake is 64.28, and the largest one 82.54 mm. The
smallest female is 90.94 mm, and the largest 109.94 mm. Obtained values of body
size of common toad from Biogradsko Lake are within the limits of the so far recorded
values from other parts of areal of the species. (R a d o v a n o v i ć , 1951; A r n o l d and
B u r t o n , 1978; Đ urović et al, 1979 etc).
Sexual dimorphism has been analyzed in scope of population (localities) by
application of Tukey test for samples of unequal size (“Unequal N HSD Tukey” test).
This test has shown an expressed sexual dimorphism, where the females have higher
average values of analyzed traits (Table 2). On basis of calculated parameters of
descriptive statistics, separately by sexes and on basis of compared differences
between the sexes, it has been established that the females have statistically higher
values of all the characters, especially when the word is about L – body length, Lpa –
length of front extremity, Lc – head length, Ltc – head width, Lg – length of salivary
glands, weigth.
With the population of Biogradsko Lake a great number of traits that have been
demonstrating sexual dimorphism by the analysis of basic data where the size has
been included, indicate the absence of sexual dimorphism for the largest number of
characters when body size is excluded. The charactereistics which demonstrate the
presence of sexual dimorphism also after the eliminating body size are T- length of
tibiofibula and F – length of femur.
Females are bigger than males in a large number of amphibian species
(S h i n e , 1979; M i a u d et al., 1999; K h o n s u e et al.., 2001a, 2002b; M o n n e t and
C h e r r y , 2002). That has been especially expressed with tailless amphibians in
which females are bigger than males with 90% of species (S h i n e , 1979). This has
also been established in this paper for studied population from Biogradsko Lake, as
well as for the populations of the remaining part of the areal of distribution of this
species (G i t t i n s et al., 1980; H e m e l a a r , 1988). The most common explanation of
sexual dimorphism in body size is the advantge the females have in respect of eggs
production (H a l l i d a y and V e r r e l l , 1986; C v e t k o v i ć et al., 2003). Namely,
there is a positive correlation in many groups between body length and fecundity
(C u m m i n s , 1986; B e r v e n , 1988; S i n s c h , 1998; G i b b o n s and M c C a r t h y ,
1986). Assumption is that the selection, owing to a pointed correlation of body length

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Males Females sexual dimorphism
Unequal N HSD (Tukey)
Trait N Means SD SE Min Max N Means SD SE Min Max basic data/standard results
L 26 73.71 4.1447 0.8128 64.28 82.54 15 100.64 5.0865 1.3133 90.94 109.94 ***
Lpa 26 54.62 3.6473 0.7153 46.68 63.17 15 71.64 3.7562 0.9699 62.23 77.60 ***/ns
F 26 32.53 3.1809 0.6238 25.32 38.86 15 40.67 4.9746 1.2844 31.98 46.95 ***/***
T 26 24.10 2.0556 0.4031 20.52 27.25 15 28.82 2.4023 0.6203 25.55 34.46 ***/**
P 26 36.65 4.2990 0.8431 27.03 42.91 15 42.98 2.8349 0.7320 37.06 47.03 **/ns
n 26 51.90 4.6742 0.9167 43.30 60.54 15 62.52 4.8611 1.2551 56.26 72.83 ***/ns
n-P 26 15.25 4.0443 0.7931 7.26 22.45 15 19.54 4.4970 1.1611 12.24 27.70 **
DpPa 26 6.58 0.7584 0.1487 5.10 7.99 15 10.44 1.5234 0.3933 7.48 13.45 ***
DpPp 26 6.74 0.8867 0.1739 5.29 8.63 15 8.46 1.2908 0.3333 5.85 10.79 ***
Cint 26 4.53 0.5892 0.1155 3.37 5.56 15 6.34 0.9030 0.2331 4.51 7.74 ***
Lc 26 17.50 1.1857 0.2325 14.28 19.46 15 24.69 1.6894 0.4362 22.04 27.83 ***/ns
Ltc 26 22.41 1.5435 0.3027 19.67 24.73 15 31.19 2.0888 0.5393 28.09 36.37 ***/ns
Spp 26 8.75 0.9524 0.1868 6.72 10.92 15 12.55 1.6316 0.4213 9.75 15.07 ***
Spi 26 4.03 0.5097 0.1000 2.85 4.92 15 5.32 0.6099 0.1575 4.39 6.47 ***
Spcr 26 7.49 0.5751 0.1128 6.28 8.62 15 11.82 0.9789 0.2528 10.28 14.21 ***
Lo 26 6.25 0.6112 0.1199 4.95 7.43 15 9.28 0.7010 0.1810 7.61 10.27 ***
Ltp 26 5.88 0.6321 0.1240 4.32 6.71 15 7.80 0.8845 0.2284 5.72 9.61 ***
Dro 26 7.54 0.5957 0.1168 6.27 8.75 15 10.72 0.8648 0.2233 9.42 12.33 ***
Dno 26 3.22 0.3864 0.0758 2.59 4.08 15 4.49 0.5739 0.1482 3.54 5.41 ***
Lh 26 3.56 0.5723 0.1122 2.65 4.77 15 5.09 0.3795 0.0980 4.49 5.81 ***
Lg 26 14.86 1.7528 0.3438 11.78 18.10 15 20.77 2.2236 0.5741 17.04 24.34 ***
Wg 26 5.80 0.7387 0.1449 4.53 7.50 15 8.20 1.0614 0.2741 6.48 10.06 ***
Weight 26 44.50 6.4576 1.2664 33.00 56.00 15 141.07 31.1367 8.0395 97.00 230.00 ***
Table 2. Basic parameters of descriptive statistics of males and females of Biogradsko Lake and Tukey (HSD) test with gender factor

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and fecundiy, favorises several values for body length of females. Greater eggs
production on the other hand is explained by larger internal space for their storage,
which larger females have (A n d e r s o n , 1994).
The direction and magnitude of dimorphism in body length depend of various
selective pressures the specimens of male and female sex are exposed to. D a r w i n
(1871) assumed that sexual selection (though the competition among the specimens
of the same sex or through an active selection of partner), may lead to sexual
dimorphism. For instance, in species with pointed struggles among the males, males
are frequently bigger than females.
However, it is important to underline that common toad is one of rare secies in
which males are a smaller sex, although there are struggles for females among them
(H a l l i d a y and V e r r e l l , 1986; A r a k , 1988).
Qualitative Analysis
By a comparison of rerlative frequency of condition of monitored qualitative traits
it has been noted that the same conditions, as a rule, have the greatest frequency
both in the sample of males and in the sample of females (Tabele 3). The exception is
the convexity of of warts in head region (in males the largest relative frequency have
the warts which are indented, and with females warts are very protuberant) and the
appearance of thornlike endings of ventral side of head region (most frequently weakly
expressed in males and averagely expressed in females). The table of relative
frequency of the conditions indicates that with females the variability of condition is
higher than with males.
By a correspondant analysis we have monitored the impact of qualitative traits
on ordination of specimens of both sexes. The first corresponding axis has separated
24.69% of variability, and the other one 17.03%. There is no clear sex separation
(Figure 1). The sample of males indicates an uniformity in respect of monitored traits.
Only one male has been separated during the second coresponding axis from the
„main cloud” of male specimens on basis of two traits (11b and 12b) (Figure 2). One
may observe a higher variability in scope of the sample of females. Traits 1c, 4b and
5a separate two females along the first correspoding axis from other females, where it
should be mentioned that these two traits as well as the traits 2a, 3b and 6b influence
separating of females from males (Figures 1 and 2).
The significance of this paper is in presenting for the first time the analysis of a
larger number of morphometric and qualitative characters of population of the species
Bufo bufo from Biogradsko Lake. These data make the grounds for further
investigations of the populations of the species Bufo bufo both from Biogradsko Lake
and the populations of this species from the territory of entire ontenegro.

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LITERATURE
ANANJEVA, N., BORKIN, L., DAREVSKIJ, I. & ORLOV, N. 1998: Zemnovodnie i
presmikajuščiesja. - Enciklopedija prirodi Rosii. ABF, Moskva, pp. 574.
ARNOLD, E.N., BURTON, J.A. 1978: A Field Guide to the Reptiles and Amphibians of
Britain and Europe. – Collins, London
ARAK, A. 1988: Sexual Dimorphism in Body Size: A Model and a Test. - Evolution 42:
820-825.
ANDERSON, S. 1994: Sexual Selection. - Princeton University Press, Princeton, New
Jersey.
BERVEN, K.A. 1988: Factors Affecting Variation in Reproductive Traits within a
Population of Wood Frogs Rana sylvatica. - Copeia, 1988: 605-615.
BORKIN, L.J., VEITH, M.1997: Bufo bufo (Linnaeus, 1758). In: GASC, J. P., GABELA,
A., CRNOBRNJA-ISAILOVIĆ, J., DOLMEN, D.,GROSSENBACHER, K.,
HAFFNER, P., LESCURE, J., MARTENS, H., MARTINEZ RICA, J.P., MAURIN,
J., OLIVIERA, M.E., SOFIANIDOU, T.S., VEITH, M., ZUIDEREWIJK, A. (eds),
1997. Atlas of Amphibians and Reptiles in Europe. Pp. 118-119. Societas
Europaea Herpetologica & Muséum National d'Historie Naturelle (IEGB/SPN),
Paris.
CVETKOVIĆ, D., ALEKSIĆ, I., CRNOBRNJA – ISAILOVIĆ, J. 2003: Reproductive
Traits in Common Toad Bufo bufo from the Vicinity of Belgrade. - Archives of
Biological Sciences, Belgrade 55: 25P-26P.
CUMMINS, C.P. 1986: Temporal and Spatial Variation in Egg size and Fecundity in
Rana temporaria. - Journal of Animal Ecology 55: 303 – 316.
DARWIN, C. 1871: The Descent of Man and Selection in Relation to Sex. – Murray,
London.
ĐUROVIĆ, E., VUKOVIĆ, T., POCRNJIĆ, Z. 1979: Vodozemci Bosne i Hercegovine. -
Zemaljski muzej BiH, Sarajevo.
GITTINS, S.P., PARKER, A.G., SLATER, F.M. 1980: Population Traits of the Common
Toad (Bufo bufo) Visiting a Breeding Site in mid- Wales. - Journal of Animal
Ecology, 49: 161-173.
HALLIDAY, T.R., VERRELL, P.A. 1986: Sexual Selection and Body Size in
Amphibians. - Herpetological Journal, 1: 86 – 92.
HEMELAAR, A.S.M. 1988: Age, Growth and other Population Traits of Bufo bufo from
Different Latitudes and Altitudes. - Journal of Herpetology, 22: 369-388.
JOHANNESSEN, T.W. 1970: The Climate of Scandinavia. In: C. C. Wallen (ed), World
KUZMIN, S.L. 1999a. Zemnovodnie bivšego SSSR. Tovariščestvo naučnih izdanij
KMK, Moskva.
KUZMIN, S.L. 1999b: The Amphibians of the former Soviet Union. Vol. I. - Pensoft,
Sofia/Moscow.

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MIAUD, C., GUYETANT, R., ELMBERG, J. 1999: Variations in Life-history Traits in the
Common Frog Rana temporaria (Amphibia: Anura): a Literature Review and New
Data from the French Alps. - Journal of Zoology, 249: 61-73.
MONNET, J. M., CHERRY, M. 2002: Sexual Size Dimorphism in Anurans. -
Proceedings of the Royal Society of London, Series B-Biological Sciences 269:
2301 – 2307.
ORLOVA, V.F. & TUNIYEV, B.S. 1989: On the Taxonomy of the Caucasian Common
Toads Belonging to the Group Bufo bufo verrucosissimus (Pallas) (Amphibia,
Anura, Bufonidae). - Biol. MOIP, Otd. biol., 94 (3): 13-24.
PASTEUR, G. & BONS, J. (1959): Les Batraciens du Maroc. - Travaux de l'Institut
Scientifique Cherifien, Serie Zoologique No 17, pp. 240. Rabat.
RADOVANOVIĆ, M. 1951. Vodozemci i gmizavci naše zemlje. - Naučna knjiga,
Beograd.
SINSCH, U. 1998. Biologie und Okologie der Kreuzkrote (Bufo calamita). - Laurenti,
Bochum.
SHINE, R. 1979. Sexual Selection and Sexual Dimorphism in the Amphibia. – Copeia,
2: 297 – 306.
Received: 20.10.2007.

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NATURA MONTENEGRINA, PODGORICA, 6:111-114
A CONTRIBUTION TO THE KNOWLEDGE OF DISTRIBUTION OF SPECIES Bufo
bufo IN MONTENEGRO
Natalija Č A Đ ENOVIĆ ¹
¹Natural History Museum of Montenegro, Podgorica, Crna Gora, E-mail: lazo@cg.yu
Key words:
Bufo bufo,
distribution,
Montenegro
SYNOPSIS
During the herpetological investigations in period 2001-2004
a voluminous material was collected. On various localities in
Montenegro we have collected the species Bufo bufo (Common
Toad), for which there are no relevant literature data for the territory
of Montenegro. This taxon is only reported for Bukumirsko jezero
(lake) by prof. Radovanović (1951) za Durmitor, Skadarsko jezero,
Lovćen, Biogradska gora (Džukić, 1991; 1995).
A list of localities on which Common Toad has been
collected as well asd UTM map with marked localities have been
presented in the paper.
Ključne riječi:
Bufo bufo,
rasprostranjenje,
Crna Gora
SINOPSIS
PRILOG POZNAVANJU RASPROSTRANJENJA
VRSTE Bufo bufo (Linnaeus, 1758) u CRNOJ GORI
Tokom herpetoloških istraživanja u periodu od 2001-2004.
godine sakupljen je obiman materijal. Na različitim lokalitetima u
Crnoj Gori sakupili smo vrstu Bufo bufo (krastavu žabu), za koju ne
postoje relevantni podaci u literaturi za teritoriju Crne Gore. Ovaj
takson navodi se jedino za Bukumirsko jezero od strane prof.
Radovanovića (1951.)
U radu je naveden spisak lokaliteta na kojima je sakupljena
krastava žaba kao i UTM karta sa unešenim lokalitetima.
INTRODUCTION
Bufo bufo is the biggest frog both in our region and in entire Europe
(Radovanović , 1951). Adults of this species are to 15 cm long, with marked
geographical variability in size. Females are significantly bigger than males. In course
of multipliation migrations Bufo bufo covers large distances and it is considered tht it
is one of the most migratory speces of tailless amphibians which may cover a
distance up to 3000 m, what indicates an markedly dispersion potential of the

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specimens of this species (S e m l i t c h and B o d i e , 2003). For the time being,
common Common Toads are the only amphibians which, at the level of adult
specimens may also undertake vertical migrations ofseveral hundreds meters
(S z t a t e c s n y and S c h a b e t s b e r g e r , 2005).
LIST OF LOCALITIES ON WHICH COMMON TOAD HAS BEEN RECORDED
1. Skadar Lake CM 5.7
2. The Crnojevića River CM 3.9
3. Lješanska nahija CM 4.9
4. Velje Brdo CN 5.0
5. Kuči CN 7.0
6. Piperi CN 5.1
7. Bjelopavlići CN 4.1
8. Prekornica CN 5.3
9. Mateševo CN 7.3
10. Biogradsko Lake CN 8.4
MATERIAL AND METHODS
Field investigations have been discontinued, as they mainly depended on
weather conditions.
The greatest part of the materiala has mainly been collected in mating period
since at that time a large number of specimens gather at one spot imediately after the
rain, or when the humidity is high. Material has mainly been collected by hand and a
meredov (a net with a handle).
This material has been subjected to morphometric analyses which will be
described in some other paper.
The material has been deposited in herpetological collection of the Natural
History Museum in Podgorici and it is kept in 70% alcohol.
CONCLUSION
So far there have been no relevan data in the literature about the distribution
of this species on the territory of Montenegro. This taxon is reported only for
Bukumirsko Lake by R a d o v a n o v i ć (1951). On basis of the list of localities we may
state that this species is widely distibuted on the territory of Montenegro, but it is also
endangered.

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C M 0 9
BP
CP
DP
BN
CN
DN
BM
CM
DM

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LITERATURE
DžUKIĆ , G. 1991: Vodozemci i gmizavci. – Građa za faunu vodozemaca i
gmizavaca Durmitora. In: Nonveiller, G. (ed.) Fauna Durmitora. CANU, Posebna
izdanja 24, Odjeljenje prir. Nauka 15, Sveska 4: 9-78.
DžUKIĆ , G., V U Č KOVIĆ , M., N I K Č EVIĆ , J., M A R I Ć , D., V I Z I , O. 1995:
Revizija bazne studije za prostorni plan. Bazna studija "Flora i Fauna". -
Republički zavod za zaštitu prirode, Podgorica.
RADOVANOVIĆ , M. 1951: Vodozemci i gmizavci naše zemlje. - Naučna knjiga,
Beograd: 46-48.
S E M L I T S C H , R.D., B O D I E , J.R. 2003: Biological criteria for buffer zones around
wetlands and riparian habitats for amphibians and reptiles. - Conservation
Biology, 17: 1219-1228.
S Z T A T E C S N Y , M., S C H A B E T S B E R G E R , R. 2005: Into thin air: vertical
migration, body condition, and quality of terrestrial habitats of alpine common
toads, Bufo bufo. - Cannadian Journal of Zoology, 83: 788-796.
Received: 20.10.2007.

NATURA MONTENEGRINA, PODGORICA, 6:115-122
MORPHOLOGICAL CHARACTERISTICS OF A POPULATION OF THE MOSOR
ROCK LIZARD (Dinarolacerta mosorensis KOLOMBATOVIĆ, 1886) (SQUAMATA:
LACERTIDAE) FROM LOVĆEN MOUNTAIN (MONTENEGRO)
Lidija P O L O V I Ć 1 , Katarina LJUBISAVLJEVIĆ 2
1
The Natural History Museum of Montenegro, 81000 Podgorica, Montenegro, e-mail: lidijapolo@cg.yu
2
Department of Evolutionary Biology, Institute for Biological Research „Siniša Stanković“, 11060
Belgrade, Serbia
Key words:
Dinarolacerta
mosorensis,
morphological
characteristics
Synopsis
External morphological traits of the Mosor rock lizard,
Dinarolacerta mosorensis (Kolombatović, 1886) from Lovćen
mountain (Montenegro) were examined. The results of
Descriptive statistics for 14 morphometric, 21 meristic and
percentages of states for eight qualitative traits were
presented. Results are discussed in comparison with the
literature data for other populations.
Ključne riječi:
Dinarolacerta
mosorensis,
morfološke
karakteristike
Sinopsis
MORFOLOŠKE KARAKTERISTIKE POPULACIJE
MOSORSKOG GUŠTERA (DINAROLACERTA MOSORENSIS
KOLOMBATOVIĆ, 1886) (SQUAMATA: LACERTIDAE) SA
PLANINE LOVĆEN, CRNA GORA
U ovom radu analizirane su spoljašnje morfološke
odlike populacije mosorskog guštera Dinarolacerta
mosorensis (Kolombatović, 1886) sa planine Lovćen.
Predstavljeni su rezultati deskriptivne statističke analize 14
morfometrijskih, 21 merističkog i procentualna zastupljenost
stanja osam kvalitativnih karaktera. Rezultati su upoređeni sa
postojećim literaturnim podacima za ostale populacije ove
vrste.

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INTRODUCTION
The Mosor rock lizard Dinarolacerta mosorensis (Kolombatović , 1886),
previously Lacerta mosorensis (see A r n o l d et al., 2007) represents both a relict
and a steno-endemic species of the Balkan Peninsula (e.g. C r n o b r nj a –
Isailović and Dž u k i ć, 1997). Its distribution is restricted to the south-western
Dinaric mountain karst in Croatia, Bosnia & Herzegovina and Montenegro, exposed to
the influence of the Mediterranean climate. There, it is patchily distributed, restricted
to altitudes ranging between 450 and 1900 m (Dž u k i ć, 1989; C r n o b r nj a –
Isailović and Dž u k i ć, 1997).
The Lovćen population of the Mosor rock lizard is located at the south-eastern
boundary of the distribution range. Here it occupies altitudes from 1200 to 1350 m
inhabiting the cliffs and blocks of rocks surrounded by subalpine beech forest (plant
community Fagetum montenegrinum subalpinum) with whitebark pine (Pinus
heldreichii) as the differential species (T o m i ć – S t a n k o v i ć, 1970;
Ljubisavljević et al., in press).
Up until very recently D. mosorensis was one of the least studied
representatives of the European herpetofauna (Dž u k i ć, 1989; O d i e r n a and
A r r i b a s , 2005). However, the latest studies have revealed substantial morphological
and genetic differences among populations of the Mosor rock lizard (C a r r a n z a et
al., 2004; Lj u b i s a v lj e v i ć et al., 2007) and certain peculiarities in life-history
traits (Lj u b i s a v lj e v i ć et al., in press).
Although specimens from the Lovćen population were included in some of these
studies, complete descriptive data on morphological characters have not been
published until now.
Dinarolacerta mosorens-adult specimen

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Polović, Ljubisavljević: MORPHOLOGICAL CHARACTERISTICS OF A POPULATION OF THE . . .
MATERIAL AND METHODS
Analyses were carried out on the samples collected from localities Ivanova
korita (42° 22’ N, 18°50’ E) and Međuvršje (42°24’ N, 18°50’ E) in the Lovćen mt. A
total of 22 adult males, 23 females, 2 immature males and 5 immature females were
examined. Specimens preserved in 70% ethanol were from Dr Georg Džukić's
Herpetological Collection of the Institute for Biological Research, Belgrade (Međuvršje
locality), and the Herpetological Collection of the Natural History Museum of
Montenegro, Podgorica (Ivanova korita locality). Specimens were examined for the
following characters:
Morphometric characters: Tot – total length, Lcor – snout-vent length, Lcap –
head length, Ltcap - head width, Altcap - head height, Lfo – mouth length, Ltfo –
mouth width, Lpa – forelimb length, Lpp – hindlimb length, Ldg – length of fourth toe
on hindlimb, Lpil – pileus length, Ltpil – pileus width, Doa – orbit to ear distance, Pap
– distance between fore and hind limbs.
Meristic characters included the numbers of: SOC – supraocular scales, CIL -
supraciliary scales, GRA - supraciliary granules, POC – postocular scales, TMP –
temporal scales, STM - supratemporal scales, PNS - postnasal scales, 1LO - first
loreal scales, 2LO second loreal scales, PROC - praeocular scales, SLB- supralabial
scales anterior to subocular, SUB - sublabial scales, SMX - submaxilar scales, GUL -
gular scales along the throat midline, COL – large collar scales, VENT – inner ventral
scales counted longitudinally, DOR – dorsal scales around mid-body, PAN – praeanal
scales surrounding anteriorly the anal plate, FPO – femoral pores, FEM – femural
scales, SDG – lamellar scales under the fourth toe.
Qualitative characters: (I) masseteric plate: a—single, b—divided, c—indistinct.
(II) row of supraciliary granules: a—complete; b—incomplete (included cases when
only one granula was separated from the rest). (III) additional scale between the first
postocular 1POC and the last supraocular 4SOC scale: a—present, b— absent. (IV)
multiplication of supralabials anterior to subocular scale: a—by insertion of additional
small scales, b—by vertical splitting of scales, c—state of four supralabials. (V) type
of dorsal pattern: a—diffuse, scattered spots, b—diffuse, interconnected spots in more
or less reticulate pattern, c—banded pattern, spots arranged in a single vertebral
and/or two narrow juxtaposed paravertebral bands, d—banded pattern, broad
vertebral band consisting of scattered spots, e—absent. (VI) spots of the dorsal
pattern: a—large, b—medium, c—small, d—absent. (VII) areas of background color
free of dark pattern: a—broad, b—narrow, c—absent. (VIII) lateral bands: a—distinct,
b—indistinct. The band names of dorsal pattern were according to A r n o l d and
B u r t o n (1978).
Symetrical characters were taken from both sides of the body. Data processing
concerned the mean of the right and left values for quantitative traits, while for
qualitative traits a combination of both sides was used.
The body and head dimensions were taken with digital callipers to the nearest
0.01 mm. Scale counts were taken under a stereoscopic microscope. For
morphometric and qualitative characters, statistical analyses included only mature

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Natura Montenegrina, 6/2007
individuals, while for analysis of the scalation, adults and immature individuals were
pooled.
Descriptive statistics (mean, standard error, range) for quantitative traits, and
percentages of states for each qualitative trait were calculated. Statistical analyses
were carried out using the computer package Statistica (STATISTICA for Windows.
StatSoft, Inc., Tulsa, OK, USA).
RESULTS AND DISCUSSION
Quantitative traits
Descriptive statistics of morphometric and meristic characters of adult males and
females are presented in Tables 1 and 2. Maximum total length recorded was 205 mm
and 197 mm for males and females, respectively. These values are smaller than those
previously reported for the species as a whole (up to 220 mm, B i s c h o f f, 1984),
and Herzegovinian population (222 mm, V e i t h, 1991). However, it should be noted
that due to the great number of regenerating individuals, we could analyse total length
on a small sample. Our measures showed that snout-vent length (Lcor) values for
females were between 56.3 – 70.7 mm ( x = 63.8 mm ± 0.8), and for males were
between 59.5 – 71.1 mm ( x = 65.5 mm ± 0.7). While female data were mainly in
accordance with literature data (57 - 64.4 - 68 mm, Bischoff, 1984), males showed
greater mean Lcor than previously reported (51 – 62.3 - 68 mm Bischoff, 1984). No
concrete literature data are available regarding the other morphometric characters
here analysed.
males
females
Character N Mean Min Max SE N Mean Min Max SE
Tot 5 192.00 179.00 205.00 4.54 6 176.17 160.00 197.00 5.63
Lcor 22 65.53 59.53 71.08 0.67 23 63.76 56.34 70.72 0.82
Lcap 22 16.85 15.67 17.62 0.14 23 14.41 13.46 15.71 0.12
Ltcap 22 10.37 9.03 11.68 0.15 23 8.79 7.80 9.74 0.10
Altcap 22 6.75 5.54 8.33 0.16 23 5.49 5.02 6.06 0.07
Lfo 22 12.77 11.62 13.59 0.12 23 11.07 10.35 12.05 0.10
Ltfo 22 9.82 8.67 10.90 0.15 23 8.30 7.30 9.33 0.09
Lpa 22 22.79 20.94 24.58 0.19 23 20.23 18.72 22.10 0.18
Lpp 22 36.06 32.70 40.57 0.39 23 31.39 29.73 34.37 0.25
Ldg 22 11.39 10.63 12.80 0.11 23 10.15 9.15 11.08 0.10
Lpil 22 15.66 14.37 16.43 0.14 23 13.41 12.60 14.58 0.11
Ltpil 22 6.85 6.30 7.30 0.06 23 6.01 5.50 6.60 0.06
Doa 22 5.12 4.60 5.65 0.07 23 3.96 3.50 4.60 0.05
Pap 22 29.01 25.85 31.96 0.36 23 32.16 25.82 38.24 0.63
Table 1. Descriptive statistics of 14 morphometric characters of adult male and female
Dinarolacerta mosorensis of the Lovćen population studied. Sample size (N), mean value (in mm),
range, standard error (SE). Abbreviations of characters are given in “Material and Methods”.

119
Polović, Ljubisavljević: MORPHOLOGICAL CHARACTERISTICS OF A POPULATION OF THE . . .
Concerning meristic characters, the mean number of four supraocular scales
and variation range of supraciliary scales were in agreement with literature data (CIL:
5 – 8, Bischoff, 1984), while the number of supraciliary granules varied (especially in
females) in a broader range than it was previously described for this species (GRA: 8
– 12, Bischoff, 1984). The most frequently detected number of two postnasals (PNS)
in our sample was in agreement with previous studies (M é h e l y, 1903;
Radovanović , 1951; B i s c h o f f , 1984; A r n o l d and O v e n d e n, 2002; A r n o
l d et al., 2007). The number of loreal (1LO, 2LO) and praeocular scales (PROC)
varied from 1 – 2, although individuals with single-scale state were the most frequent
in our sample. Previous literature data (B i s c h o f f, 1984) also showed single state
of these scales as characteristic for this species. The usual number of four or five
supralabial scales (SLB) in front of the subocular (with possibility of an asymmetric
condition), as well as usual number of six sublabial (SUB) and submaxilar scales
(SMX) reported for D. mosorensis by other authors (e.g. R a d o v a n o v i ć , 1951; B i s
c h o f f, 1984; A r n o l d et al., 2007) were in agreement with our results. However,
the number of gular (GUL), dorsal (DOR) and subdigital scales (SDG) varied in
broader range than previously recorded (GUL: 23 - 30; DOR: 36 - 40 - 45; SDG: 22 –
23,4 - 25 in B i s c h o f f, 1984). The mean values and/or variation range for other
analysed meristic characters were mainly in agreement with those previously reported
for this species (e.g. R a d o v a n o v i ć, 1951; B i s c h o f f, 1984).
males
females
Character N Mean Min Max SE N Mean Min Max SE
SOC 24 4.02 4.00 4.50 0.02 28 4.00 4.00 4.00 0.00
CIL 24 5.94 4.50 7.00 0.15 28 6.09 5.50 8.00 0.12
GRA 24 9.17 7.00 11.50 0.25 28 8.65 5.00 12.00 0.29
POC 24 3.81 2.50 4.00 0.08 28 3.52 3.00 4.50 0.10
TMP 24 42.23 24.50 73.00 2.07 28 39.09 25.00 54.00 1.69
STM 24 2.52 1.50 3.50 0.11 28 2.35 2.00 3.00 0.07
PNS 24 1.88 1.00 2.00 0.06 28 1.96 1.50 2.00 0.03
1LO 24 1.00 1.00 1.00 0.00 28 1.02 1.00 1.50 0.02
2LO 24 1.00 1.00 1.00 0.00 28 1.07 1.00 2.00 0.05
PROC 24 1.13 1.00 2.00 0.06 28 1.07 1.00 2.00 0.04
SLB 24 4.69 4.00 5.50 0.12 28 4.57 3.50 5.00 0.08
SUB 24 6.02 4.50 7.00 0.11 28 6.04 5.00 7.00 0.10
SMX 24 5.96 5.50 6.00 0.03 28 5.78 5.00 6.00 0.08
GUL 24 25.88 20.00 32.00 0.52 28 24.48 19.00 28.00 0.40
COL 24 8.83 6.00 10.00 0.21 28 8.67 7.00 11.00 0.22
VENT 24 25.25 23.50 27.50 0.23 28 26.72 25.00 29.00 0.21
DOR 24 39.06 33.00 44.00 0.66 28 38.46 30.50 43.00 0.59
PAN 24 7.71 6.00 9.00 0.13 28 7.30 6.90 9.00 0.18
FPO 24 18.35 16.50 21.00 0.23 28 17.19 12.00 20.50 0.32
FEM 24 4.00 3.00 5.00 0.11 28 3.67 3.00 4.50 0.10
SDG 24 22.29 20.00 24.50 0.24 28 21.96 20.00 25.50 0.26
Table 2. Descriptive statistics of 21 meristic characters of adult male and female Dinarolacerta
mosorensis of the Lovćen population studied. Sample size (N), mean value (in mm), range,
standard error (SE). Abbreviations of characters are given in “Material and Methods”.

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males
(N = 22)
females
(N = 23)
males
(N = 22)
females
(N = 23)
character % % character % %
Iaa 68 65 VIa 73 83
Iab 5 4 VIb 9 13
Iac 9 22 VIc 9 4
Ibb 0 0 VId 9 0
Ibc 0 0 VIIa 5 4
Icc 18 9 VIIb 0 9
IIaa 68 57 VIIc 95 87
IIab 14 13 VIIIa 5 35
IIbb 18 30 VIIIb 95 65
IIIaa 0 0
IIIab 5 4
IIIbb 95 96
IVaa 0 0
IVab 5 4
IVac 5 9
IVbb 41 35
IVbc 23 26
IVcc 27 26
Va 23 46
Vb 64 46
Vc 5 13
Vd 0 0
Ve 9 0
Table 3. Percentages of states of qualitative traits (in %) in Dinarolacerta mosorensis of
the Lovć en population. For symmetrical traits combination of states for both body sides
were given. Abbreviations of characters are given in “Material and Methods”.
Qualitative traits
Percentages of states for qualitative traits of adult males and females are
presented in Table 3. In general, analysed individuals are characterised by the
presence of single masseteric plate (Ia), complete row of supraciliary granules (IIa),
absence of additional scale between the first postocular and the last supraocular
scale (IIIb), and symmetrical multiplication of supralabials by vertical partition of
scales (IVbb). Concerning the type of dorsal pattern, they are distinguished by
predominance of diffuse type consisting of large size spots coupled with indistinct
lateral bands (Va,b; VIa; VIIIb) and the absence of areas of background color free of
dark pattern (VIIc). The diffuse pattern with interconnected spots in more or less
reticulate pattern was predominant in males (Vb), while in females reticulate pattern
(Vb) and scattered spots (Va) were present in the same percentage. A certain
percentage of individuals of both sexes with symmetrical presence of an indistinct
masseteric plate (Icc), and complete absence of dorsal pattern (males) (Ve, VId) were
also recorded from Lovćen population.

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Polović, Ljubisavljević: MORPHOLOGICAL CHARACTERISTICS OF A POPULATION OF THE . . .
Concerning the qualitative traits, the prevalence of specimens with distinct
masseteric plate, complete row of supraciliary granules and diffuse reticulate pattern
found in this study were also considered characteristic for the Mosor rock lizard (R a d
o v a n o v i ć, 1951; B i s c h o f f, 1984; A r n o l d et al., 2007). On the other hand,
we rarely found unmarked (uniform) specimens, the feature frequently referred to D.
mosorensis by other authors (T o m a s s i n i, 1889; B i s c h o f f, 1984; V e i t h,
1991; A r n o l d and O v e n d e n, 2002).
CONCLUSION
Most of the values or percentages of occurrences obtained from the present
analysis of external morphology of the Mosor rock lizard are in accordance (or within
the variation range) with previous studies of the species. However, the number of
supraciliary granules, supralabial, gular, dorsal and subdigital scales varied in
somewhat broader range than it was previously reported for a species as a whole.
ACKNOWLEDGEMENTS
We are grateful to Georg Džukić for useful comments and making part of literature and
sample collection available to us. We also thank Miloš Kalezić for helpful suggestions. Katarina
Ljubisavljević was supported by the Serbian Ministry of Science (grant no. 143052, “Patterns of
amphibian and reptile diversity on the Balkan Peninsula”).
LITERATURE
A R N O L D, E. N., and J. A. B U R T O N. (1978). A Field Guide to the Reptiles and
Amphibians of Britain and Europe. - Harper Collins Publishers, London, UK, 1-272.
A R N O L D, E. N., and D. O V E N D E N (2002). A Field Guide to the Reptiles and
Amphibians of Britain and Europe. Second Edition. - HarperCollins Publishers, London,
UK, 1-288.
A R N O L D, E. N., A R R I B A S, O., and S. C A R R A N Z A (2007). Systematics of the
Palaearctic and Oriental Lizard Tribe Lacertini (Squamata: Lacertidae: Lacertinae), with
Descriptions of Eight New Genera. - Zootaxa 1430, 1-86.
B I S C H O F F, W. (1984). Lacerta mosorensis Kolombatović, 1886, Mosor-Eidechse, In:
Handbuch der Reptilien und Amphibien Europas, 2-1, Echsen II, Lacerta (Ed. W. Böhme),
290-300. Aula -Verlag, Wiesbaden, Germany.
C A R R A N Z A, S., A R N O L D, E. N., and F. A M A T (2004). DNA Phylogeny of Lacerta
(Iberolacerta) and other Lacertine Lizards (Reptilia:Lacertidae): Did Competition Cause
Long-term Restriction?. - Systematics and Biodiversity 2, 57-77.
C R N O B R N J A – I S A I L O V I Ć, J., and G. DŽ U K I Ć (1997). Lacerta mosorensis. In:
Atlas of the Amphibians and Reptiles in Europe (Eds. J. P. Gasc, A. Cabela, J. Crnobrnja-

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Isailović, D. Dolmen, K. Grossenbacher, P. Haffner, J. Lescure, H. Martens, J. P.
Martýnez Rica, H. Maurin, M. E. Oliveira T. S. Sofianidou, M. Veith, and A. Zuiderwijk),
250-251. Societas Europaea Herpetologica & Museum Nationall d’ Histoire Naturelle,
Paris.
DŽ U K I Ć, G. (1989). Remarks on Distribution and Protection Problems of the Mosor Rock
Lizard, Lacerta mosorensis Kolombatović, 1886 (Reptilia, Lacertidae). - Biologia Gallohelenica
15,185-190.
LJ U B I S A V LJ E V I Ć, K., A R R I B A S, O., DŽ U K I Ć, G. and S. C A R R A N Z A
(2007). Genetic and Morphological Differentiation of Mosor Rock Lizards, Dinarolacerta
mosorensis (Kolombatović, 1886), with the Description of a New Species from the
Prokletije Mountain Massif (Montenegro) (Squamata: Lacertidae). - Zootaxa 1613,1-22.
LJ U B I S A V LJ E V I Ć, K., P O L O V I Ć, L., T O M A Š E V I Ć – K O L A R O V, N.,
DŽUKIĆ , G., and M. L. K A L E Z I Ć. Female Life-history Characteristics of the Mosor
Rock Lizard, Dinarolacerta mosorensis (Kolombatović, 1886) from Montenegro
(Squamata: Lacertidae). - J. Nat. Hist., in press.
M É H E L Y, L. (1903). Lacerta mosorensis Kolomb., a Magyar királyság új gyikja,
származástani kapcsolatában. - Állattani Közlemények, Budapest, 212 pp.
O D I E R N A, G. and O. A R R I B A S (2005). The Karyology of ‘Lacerta’ mosorensis
Kolombatovic, 1886, and Its Bearing on Phylogenetic Relationships to other European
Mountain Lizards. - Ital. J. Zool. 72, 93-96.
R A D O V A N O V I Ć, M. (1951). Vodozemci i gmizavci naše zemlje. - Naučna knjiga,
Beograd, 1-249.
T O M A S S I N I, O. (1889). Crtice o životu gmazova koji žive u Bosni i Hercegovini. - Glasnik
Zem. muz. u B. i H. 1, 46-60.
T O M I Ć – S T A N K O V I Ć, K. (1970). Flora Lovćena I. - Zbornik Filozofskog Fakulteta u
Prištini, 7,1-39.
V E I T H, G. (1991). Die Reptilien Bosniens und der Herzegowina, teil I. Herpetozoa 3, 97-196.
Received: 12. 11. 2007.

NATURA MONTENEGRINA, PODGORICA, 6:123-129
CONTRIBUTION TO THE CHEMICAL CONSTITUENTS OF BALKAN BRYOPHYTES:
PHENOLIC ACIDS, FLAVONOIDS, TRITERPENES AND ALKALOIDS
Nebojša J O C K O V I Ć 1 , Milica P A V L O V I Ć 1 , Marko SABOVLJEVIĆ 2, * and
Nada K O V A Č EVIĆ 1
1
Department of Pharmacognosy, Faculty of Pharmacy, University of Belgrade, Vojvode Stepe 450, 11000
Belgrade, Serbia
2
Institute of Botany and Garden, Faculty of Biology, University of Belgrade, Takovska 43, 11000
Belgrade, Serbia
* corresponding author: marko@worldpost.eu
Key Words:
Bryophyte chemistry,
mosses,
liverworts,
phenolic acids,
flavonoids,
triterpenes,
alkaloids
Synopsis
In this paper preliminary study of chemical constituents
of three bryophyte species will be presented. Gametophytes
of Polytrichum formosum, Eurhynchium hians (mosses) and
Pellia endiviifolia (liverworth) have been collected in the
native habitats in Petnica near Valjevo (W. Serbia). TLC
technique was applied for the preliminary study of petroleum
ether and methanol extracts of three investigated species.
According to this assay the presence of flavonoids (aglycones
and glycosides) in all examined extracts was confirmed. In extracts
of two species phenolic acids were detected. Besides, in the
petroleum ether extract of all three species triterpenes were
detected. Alkaloids were absent in the extracts of investigated
bryophyte species.
Ključne reči:
hemija briofita,
mahovine,
jetrenjače,
fenolne kiseline,
flavonoidi,
triterpeni,
alkaloidi
Sinopsis
PRILOG POZNAVANJU HEMIJSKOG SASTAVA
BALKANSKIH BRIOFITA: FENOLNE KISELINE,
FLAVONOIDI, TRITERPENI I ALKALOIDI
U ovom radu su dati preliminarni rezultati hemijskih
konstituenata tri vrste briofita. Gametofiti Polytrichum
formosum, Eurhynchium hians (mahovine) i Pellia endiviifolia
(jetrenjača) su sakupljeni u prirodnim staništima okoline
Valjeva (Zap. Srbija). Primenjena je TLC tehnika (tankoslojna
hromatografija) u karakterisanju petroleimskog i metanolnog
ekstrakta tri odabrane vrste.
Prisustvo flavonoida (aglikona i glikozida) je potvrdjeno
u svim analiziranim ekstraktima. U ekstraktima dve vrste
detektovane su fenolne kiseline. Osim toga u petroleumskom
ekstraktu sve tri vrste detektovani su triterpeni. Alkaloidi nisi
pronadjeni kod ni jedne od tri istraživane vrste.

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INTRODUCTION
Chemical studies of the bryophytes were neglected for a long time. They have
now been shown to be a storehouse of naturally occurring materials, including some
with novel chemical structures. Many of these materials display considerable
biological activity. Investigations are hampered frequently by too small amounts of
plant material. The resulting low yields of components are then generally inadequate
to permit testing for biological activity. In vitro culture and appropriate chemical
synthesis on a preparative scale are being undertaken to overcome this difficulty.
However, bryophytes are the second biggest group of land plants after flowering
plants and a source of chemically new and unknown compounds (e.g. A s a k a w a ,
1994, 2001; S a b o v l j e v i ć & al., 2001; Z i n s m e i s t e r & al., 2003.). Studies of
chemical constituents of bryophytes are recently being performed, but still inadequate
and neglected (A s a k a w a , 1995; T o y o t a & al., 1998; E d e l m a n n & al., 1998;
S p e i c h e r & al., 2000, 2001; K l i n k & al., 2002; P o pper & Fry, 2003;
H e r t e w i c h & al., 2003). These data help in systematic of hardly morphologically
classified bryophytes (e.g. A s a k a w a , 2004). The data on biological activities of
bryophyte extracts and/or chemical constituents are even hard to find (e.g. B a s i l e &
al., 1998a, b, 1999; D u l g e r & al., 2005; S a b o v l j e v i ć & al., 2006,
Sabovljević & Sabovljević , 2007)
MATERIAL AND METHODS
Three selected bryophyte species were used for phytochemical screening using
TLC technique in order to show the presence of phenolic components, triterpenes and
alkaloids in the their extracts. One liverwort and two mosses, Pellia endiviifolia
(Dicks.) Dumort. (talous liverwort), Eurhynchium hians (Hedw.) Sande Lac.
(pleurocarpous moss) and Polytrichum formosum Hedw. (acrocarpous moss) were
analysed. Only the gametophytes were used for the chemical analyses. The
specimens were collected in the native habitats in Petnica near Valjevo (W. Serbia) in
August 2002, dried at the room temperature and stored in paper bags at 4°C till the
beginning of the experiments.
Preparation of extracts
3.0 g of ground material was extracted by maceration with 30 ml of petroleum
ether during 48h at room temperature, the extract was filtered and taken into dryness
under reduced pressure. The dry residue was dissolved in 0.2 ml of petroleum ether.
The same material used in previous step was left to dry, and extracted afterward with
30 ml of methanol at the sonic bath during 30 minutes. The extract was filtered, the
filtrate concentrated under reduced pressure and residue taken up in 0.2 ml of
methanol.

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Jocković; Pavlović, Sabovljević, Kovačević: CONTRIBUTION TO THE CHEMICAL CONSTITUENTS . .
Thin layer chromatography
Thin layer chromatography was performed on silica gel 60 F 254 plates (DC
Alufolien, Merck, Germany). The mobile phases used were: toluene-ethyl acetate
(70:30, v/v) for flavonoid aglycones and triterpenes, ethyl acetate-formic acid-glacial
acetic acid-water (100:11:11:26, v/v/v/v) for phenolic acids and flavonoid glycosides
and toluene-ethyl acetate-diethylamine (70:20:10, v/v/v) for alkaloids.
Chromatograms were evaluated under UV light at 254 and 365 nm before and
after spraying with NP-reagent for flavonoids, after spraying with vanillin-sulphuric
acid reagent (VS) for triterpenes and Dragendorff reagent for alkaloids (W a g n e r &
B l a d t , 1996).
RESULTS
The lipophilic compounds of Eurhynchium hians, Pellia endiviifolia and
Polytrichum formosum were isolated by petroleum ether and analysed by TLC in
toluene-ethyl acetate (70:30, v/v). The chromatogram of petroleum ether extract of
Eurhynchium hians, Pellia endiviifolia and Polytrichum formosum showed the
presence of several zones that correspond to flavonoid aglycones. They were
detected by quenching fluorescence under UV-245 nm and by yellow fluorescence
under UV light at 365 nm before and after spraying with NP-reagent.
Comparing the positions of the zones of yellow fluorescence of all three
bryophyte species on the chromatogram, it may be concluded that they contain some
substances in common, having at least one substance present in all three petroleum
ether extracts (Fig. 1)
After developing the chromatogram in the mobile phase toluene-ethyl acetate
(70:30, v/v), and spraying with vanillin-sulphuric acid reagent triterpenes were shown
as bluish-violet zones in the petroleum ether extracts of the examined bryophyte
species (Fig. 2).
The zones in extracts of Pellia endiviifolia and Polytrichum formosum that are
the closest to the start line might derive of the same compound. Similarly, the zones in
Eurhynchium hians and Polytrichum formosum that are the closest to the solvent front
are likely to represent the same substance. Regarding the zones at the half of the
chromatogram, it may be concluded that petroleum ether extracts of all three
bryophytes have one compound in common from triterpene group (Fig. 2).
Methanolic extracts of Eurhynchium hians, Pellia endiviifolia and Polytrichum
formosum were analysed by TLC in ethyl acetate-formic acid-glacial acetic acid-water
(100:11:11:26, v/v/ v/v) as developing solvent. Flavonoid glycosides were detected in
the chromatogram of methanol extracts of all three species as yellow fluorescing
zones (UV-365 nm) before and after spraying with NP reagent. Chromatogram of
methanol extracts of Pellia endiviifolia and Polytrichum formosum, near the solvent
front showed the presence of blue fluorescing zones in UV light at 365 nm which
correspond to the phenolic acids.

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Natura Montenegrina, 6/2007
Figure 1. Chromatogram of petroleum ether
extracts of Eurhynchium hians (1), Pellia
endiviifolia (2) and Polytrichum formosum (3),
obtained in toluene-ethylacetate (70:30, v/v),
after spraying with NP reagent, under UV light
at 365 nm.
Figure 2. Chromatogram of petroleum ether
extract of Eurhynchium hians (1), Pellia
endiviifolia (2) and Polytrichum formosum (3),
obtained in toluene-ethyl acetate (70:30, v/v),
after spraying with vanillin-sulphuric acid
reagent.

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Jocković; Pavlović, Sabovljević, Kovačević: CONTRIBUTION TO THE CHEMICAL CONSTITUENTS . .
In order to screen methanolic extracts of analysed bryophyte species for
presence of alkaloids, the chromatograms were developed in toluene-ethyl acetatediethylamine
(70:20:10, v/v/v), and sprayed with Dragendorff reagent. Under given
conditions of analysis, no orange-brown or brownish zones that correspond to alkaloid
compounds could be noticed.
CONCLUSION
The study gives first insight into chemistry of bryophytes from the Balkan
Peninsula. The compounds detected are already known among bryophytes. Generally,
liverworts are more often used for chemical studies due to their oil bodies and so
other species of Pellia was subject of chemical content studies. However, it remains
unknown to the authors whether there have been previously chemical analyses of
selected moss species for this study at all.
Acknowledgement
Many thanks to Dr. Rayna Natcheva (Sofia) for valuable comments to manuscript.
LITERATURE
ASAKAWA, Y. (1994): Highlights in Phytochemistry of Hepaticae - Biologically
Active Terpenoids and Aromatic Compounds. - Pure and Applied Chemistry,
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ASAKAWA Y. (1995): Chemical Constituents of the Bryophytes. In: Herz W., Kirby
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ASAKAWA, Y. (2001): Recent Advances in Phytochemistry of Bryophytesacetogenins,Terpenoids
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623-669.
BASILE, A., SORBO, S., GIORDANO, S., LAVITOLA, A. & CASTALDO-
C O B I A N C H I , R. (1998a): Antibacterial Activity in Pleurochaete squarrosa
extract. - International Journal of Antimicrobial Agents, 10: 169-172.

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BASILE, A., GIORDANO, S., SORBO, S., VUOTTO, M. L., IELPO, M. T. L.
& C A S T A L D O - C O B I A N C H I , R. (1998b): Antibiotic Effects of Lunularia
cruciata (Bryophyta) Extract. - Pharmaceutical Biology, 36 (1): 25-28.
BASILE, A., GIORDANO, S., LÓPEZ-SÁEZ, J. A., CASTALDO-
C O B I A N C H I , R. (1999): Antibacterial Activity of Pure Flavonoids Isolated
from Mosses. - Phytochemistry, 52: 1479-1482.
DULGER, B., TONGUÇ-YAYINTAS, Ö. & GONUZ, A. 2005. Antimicrobial
Activities of some Mosses from Turkey. - Fitoterapia, 76: 730-732.
E D E L M A N N , H., N E I N H U I S , C., J A R V I S , M. C., E V A N S , B., F I S C H E R ,
E. & B A R T H L O T T , W. (1998): Ultrastructure and Chemistry of the Cell Wall
of the Moss Rhacocarpus purpurascens (Rhacocarpaceae): a Puzzling
Architecture among Plants. - Planta, 206: 315-321.
HERTEWICH U.M., ZAPP J. & BECKER H. (2003): Secondary Metabolites
from the Liverwort Jamesoniella colorata. - Phytochemistry, 63 (2): 227-233.
KLINK, J.W.VAN, ZAPP, J. & BECKER, H. (2002): Pinguisane-type
Sesquiterpenes from the South American Liverwort Porella recurva (Taylor)
Kuhnemann. - Zeitschrift für Naturforschung, 57: 413-417.
POPPER, Z. A. & FRY, S.C. (2003): Primary Cell Wall Composition of Bryophytes
and Charophytes. - Annals of Botany, 91: 1-12.
SABOVLJEVIĆ , M., BIJELOVIĆ , A. & G R U B I Š I Ć , D. (2001): Bryophytes as
a Potential Source of Medicinal Compounds. - Lekovite Sirovine, 21: 17 – 29.
SABOVLJEVIĆ , A., S O K O V I Ć , M., SABOVLJEVIĆ , M. & G R U B I Š I Ć , D.
(2006): Antimicrobial Activity of Bryum argenteum. – Fitoterapia, 77: 144-145.
SABOVLJEVIĆ , A. & SABOVLJEVIĆ , M. (2007): Bryophytes, a Source of Bioactive
and New Compounds. – In: G o v i l , J. N. (ed.). Phytopharmacology and Therapeutic
Values IV, the Series "Recent Progress in Medicinal Plants". Pp. 9-25.
SPEICHER, A., HOLLEMEYER, K. & H E I N Z L E , E. (2000): Rapid Detection
of Multiple Chlorinated Bis(bibenzyls) in Bryophyte Crude Extracts Using Laser
Desorption/ionization time-of-flight Mass Spectrometry. - Rapid Communications
in Mass Spectrometry, 15 (2): 124 – 127.
SPEICHER, A., HOLLEMEYER, K. & HEINZLE, E. (2001): Rapid Detection
of Chlorinated Bisbenzyls in Bazzania trilobata Using MALDI-TOF Mass
Spectrometry. – Phytochemistry, 57: 303-306.
TOYOTA, M., MASUDA, K. & ASAKAWA, Y. (1998): Triterpenoid Constituents
of the Moss Floribundaria aurea subsp. nipponica. - Phytochemistry, 48 (2): 297-
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Chromatography Atlas. Springer-Verlag Berlin.

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Jocković; Pavlović, Sabovljević, Kovačević: CONTRIBUTION TO THE CHEMICAL CONSTITUENTS . .
Z I N S M E I S T E R , H. D., B E C K E R , H. & E I C H E R , T. (2003): Bryophytes, a
Source of Biologically Active, Naturally Occurring Material. - Angewandte
Chemie, 30(2): 130-147.
Received: 07.05.2007.

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NATURA MONTENEGRINA, PODGORICA, 6:131-136
PCR (Polymerase Chain Reaction) IN DETECTION OF Chlamydia trachomatis AND THE
OTHER METHODS – A COMPARATIVE SURVEY
Danko OBRADOVIĆ 1
1
University of Montenegro, Faculty of Sciences, Department of Biology, P. O. Box 211, 81000 Podgorica,
Montenegro
Key words:
PCR,
Chlamydia
trachomatis
Klučne riječi:
PCR,
Chlamydia
trachomatis
Synopsis
Polymerase chain reaction (PCR) based methods are the
most advanced methods for detection of Chlamydia trachomatis.
There are various tests for detection of C. trachomatis. Many
people, including specialists, do not know about reliability of tests
and a comparative survey is welcome for overcoming this problem.
Lack of sensitivity, specificity, or both, sensitivity and specificity, and
some other disadvantages are common problems of those tests.
However, polymerase chain reaction based methods do not have
any of those common problems. They are the best methods for
detection of C. trachomatis, which provide the most reliable results.
Samples can be processed in just 5-6 hours. Disadvantage of PCR
based methods is high price and high technology which makes them
unavailable especially for people in developing countries.
Sinopsis
PCR (REAKCIJA POLIMERIZACIJE LANCA) U DETEKCIJI
Chlamydia trachomatis I DRUGE METODE – UPOREDNA
ISTRAŽIVANJA
Metode bazirane na reakciji polimerizacije lanca (PCR) su
najsavremenije metode za detekciju Chlamydia trachomatis. Postoje
različiti testovi za detekciju C. trachomatis. Mnogi ljudi, uključujući
specijaliste, nisu upoznati sa pouzdanošću testova, pa je jedan
uporedni pregled važan za prevazilaženje ovog problema.
Nedostatak osetljivosti, specifičnosti, ili oboje, osetljivosti i
specifičnosti, i neke druge mane su česti problemi vezani za ove
testove. Ipak, ni jedan od tih problema nije prisutan kod metoda
baziranih na reakciji polimerizacije lanca. To su najbolje metode za
detekciju C. trachomatis, koje daju najpouzdanije rezultate. Uzorci
mogu biti obrađeni za samo 5-6 časova. Mana metoda baziranih na
PCR-u je visoka cijena i visoka tehnologija koje su uzrok
nedostupnosti metode, naročito za ljude u zemljama u razvoju.

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INTRODUCTION
Diseases caused by bacterium Chlamydia trachomatis are pretty common
worldwide. Actually, it is one of the most common sexually transmitted diseases. In
2005, according to statistics, 976,445 C. trachomatis diagnoses were reported (up
from 929,462 in 2004) in the USA. However, most of C. trachomatis cases remain
undiagnosed and it is estimated that there are 2.8 million new cases annually
(Weinstock et al., 2004). Direct and indirect costs of cure of the disease exceed 2.4
billion US dollars annually (Washington and Katz, 2000). Increase in cases is probably
partially caused by advance in diagnostic laboratory tests. C. trachomatis is an
intracellular bacterium and cannot be grown on agar plates. The infectious,
extracellular form is an elementary body, typically 0.2-0.6 μm in diameter. Elementary
bodies that have been endocytosed by eukaryotic cells typically remain in vacuolar
inclusions and transform into reticulate bodies. Reticulate bodies range up to 1.5 μm.
If untreated, chlamydial infections can progress to serious reproductive and other
health problems with both short-term and long-term consequences. One of the main
reasons that this disease is very common is the fact that it is mostly asymptomatic
(Stamm, 1999). An asymptomatic disease can be diagnosed only by laboratory tests
because of lack of symptoms. This is the reason that efficient cure and eradication of
the disease is possible only if laboratory tests are highly reliable.
A COMPARATIVE SURVEY OF METHODS
There are several basic methods in laboratory detection of C. trachomatis.
They have various specificities, and all of them (except PCR) have low sensitivity or
cannot be efficiently applied for large scale analyses. In the case of asymptomatic
diseases, which can be diagnosed only with a laboratory test, low sensitivity of the
test is unacceptable. A survey of sensitivity of basic methods for detection of C.
trachomatis is given in table 1 (Ossewaarde, 1995).
DIAGNOSTIC METHOD
SENSITIVITY (PERCENT)
PCR 90-100%
Cell Culture (McCoy) 60-80%
Fluorescent Microscopy with Fluorescent 55-75%
Antibodies
ELISA 50-70%
Hybridization with DNK Probes 50-70%
Fast Tests 40-60%
Table 1. Basic methods used in laboratory diagnostics of C. Trachomatis
and their diagnostic sensitivity.

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Cell culture. Analysis in cell culture of McCoy cell line is one of methods for
detection of C. trachomatis (Black, 1997). Its sensitivity is 60-80% and it takes 4-7
days. Disadvantage of this method is that is not suitable for large scale application.
Maintenance of cell culture is expensive and labor and time consuming. Sterile
conditions are essential for cell culture and contamination is a very common problem
especially in case of processing of many samples. Thus cell culture is not suitable for
routine laboratory diagnostics. Actually, the relatively low sensitivity, long turnaround
time, difficulties in standardization, labor intensity, technical complexity, stringent
transport requirements, and relatively high cost are the primary disadvantages of cell
culture isolation of C. trachomatis. This is the reason that this method has not been
applied in large scale diagnostics.
Fluorescent microscopy. Fluorescent microscopy can be with monoclonal and
polyclonal antibodies. Sensitivity and specificity are 55-75% and 85-95% respectively.
People who do analysis have to be well trained and the accuracy of results largely
depends of their experience. If little amount of bacteria and large amount of cells and
cell debris is present in the sample, result cannot be clear and depends of subjective
opinion of person who does the test. This is the reason that this method largely
depends of experience and degree of training, and sensitivity and specificity can be
even lower in case of bad specialist. Actually, well trained specialists and with good
experience are rare. Another disadvantage of this method is possibility of reaction of
applied antibodies with antigens of other species of genus Chlamydia, which would
lead to false positive results. Antibodies available on the market sometimes can react
even with species that do not belong to genus Chlamydia. Monoclonal antibodies are
expensive and polyclonal give worse results.
ELISA. ELISA is another method applied in diagnostics of C. trachomatis. Its
sensitivity is 50-70%. It is often applied in detection of specific antibodies in patients’
blood that are synthesized under influence of C. trachomatis. In primary immune
response first are synthesized antibodies of class IgM, and later IgG and IgA are the
last. In secondary immune response just IgG and IgA are present, and IgM is absent.
Because presence of various antibodies during the infection vary, it is necessary to
perform test with at least 2 classes of antibodies, which additionally increase
expenses. Immunity of a patient can be decreased for various reasons (the other
disease, stress, treatment with immunosuppressors in case of patients with
transplanted organ etc.), which can decrease concentration of antibodies below the
sensitivity threshold of the method and increase amount of false negative results.
Besides, antibodies begin to be synthesized always after an infection, and only after a
given time amount of antibodies is increased above sensitivity threshold of the
method. If concentration of antibodies is below the threshold, diagnostics is not
possible. Very serious problem of this method is the fact that antibodies are present in
the body of healthy patients after a treatment. This test, in this case, would be positive
in spite of the fact that patient is healthy. This is a problem of all indirect methods that

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detect presence of specific antibodies instead of causative agent of the disease.
There is ELISA that detects antigens. These antigens are extracted by bacteria into
the environment which can increase possibility to detect them because permanent
extraction increases their concentration in the environment. In this case the method
does not detect bacteria, but antigens that are extracted into environment. ELISA has
the same problems with false positive reactions with bacteria of other species and
genera as fluorescent antibodies methods, because it is based on antibodies, too.
This method should not be used for detection of rectal samples because of increased
reaction with fecal bacteria that can give more false positive results. Reaction of
ELISA antibodies with Escherichia coli and Klebsiella pneumoniae is often reason of
false positive results (Demaio et al., 1991; Anderson et al., 1993). Some ELISA tests
(which detect presence of lipopolysaccharides of Chlamydia) do not distinguish
species of genus Chlamydia, and positive result does not mean always that detected
bacterium is C. trachomatis. Good characteristics of ELISA tests are high degree of
automation, which made possible employment of low qualified personal, and low cost
of the test. Bad characteristics are low reliability because of low sensitivity and
possible false positive results.
Hybridization with DNA probes. Hybridization with DNA probes is a molecular
biology test. This kind of test is complicated, time and labor consuming, and
expensive. Its sensitivity is just (50-70%) which is substantially less than sensitivity of
the other kind of molecular test (PCR).
Fast tests. This kind of tests is possible to bye in drug stores and patients can
apply it by themselves following directions submitted with the test. Results can be
obtained in 10-15 minutes. They can be based on immunochromatographic or specific
enzyme reaction. They have a low sensitivity (40-60%) and specificity (Suchland et
al., 1997). This kind of tests has just 1 good characteristic which is expressed in their
name – they are fast and simple for application. Thus, they can be applied just for
very rough screening.
Polymerase chain reaction (PCR) based tests. PCR has been discovered by
American scientist Kary Mullis in 1984, and shortly, it became a widely applied method
in scientific research, and as well as in various applications (diagnostic of infectious
and genetic diseases, forensic etc.). Kary Mullis became Nobel Prize winner in 1993
for polymerase chain reaction. PCR based methods are the most advanced tools for
diagnostic of C. trachomatis. It has a number of advantages in comparison to the
other methods.
• High sensitivity is one of the most important characteristics of PCR. PCR is
based on replication (amplification) in vitro of a DNA sequence that is highly
specific for infectious agent. A sequence that is present in a few or even just one
copy can be copied (replicated) to produce a huge amount (thousands of billions

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copies or even more). Such large amount of DNA molecules can be detected
easily, and this is the main reason that PCR is more sensitive than the other
methods. Analytical sensitivity of PCR is in most cases 1-10 elementary bodies
of C. trachomatis. Sensitivity of fluorescent antibodies microscopy is 10-500
elementary bodies, and ELISA 5000-100 000 elementary bodies (Black, 1997).
Diagnostic sensitivity of PCR based tests for C. trachomatis is 90-100%.
• PCR based methods are direct methods. They directly detect presence of the
bacterium. Many other methods detect specific antibodies, or a product of the
bacterium which is extracted into the environment. PCR based methods detects
a specific DNA which is present in the bacterium.
• High specificity of PCR based methods is possible because they amplify and
detect a sequence highly specific for the bacterium. The sequence can be highly
conservative that makes possible detection of causative agents with high
antigenic changeability. Specificity of PCR based tests for C. trachomatis is
99.6% (Black, 1997).
• PCR methods can process samples in just 5-6 hours and provide highly reliable
results.
• Samples for PCR based methods can be processed from all sources including
noninvasive ones, as urine.
Disadvantages of PCR based methods are high price and complicated
processing of the samples. This makes them unsuitable for large scale screenings.
However, high reliability and other advantages are good compensation for mentioned
disadvantages.
CONCLUSION
Polymerase chain reaction based methods are the most reliable methods for
detection of C. trachomatis infections. Very important characteristics of a laboratory
diagnostics method are sensitivity and specificity. PCR based methods are the most
superior in this sense. They are direct methods based on detection of a DNA
sequence specific for C. trachomatis, which insure that positive result means
presence of the bacterium. These methods make possible processing of samples in
one working day and they can be obtained from all sources including noninvasive,
which make them very suitable for eradication of the disease. In contrast, low
sensitivity of the other methods makes them unsuitable for reliable diagnostics.
However, high expenses and application of high technology makes PCR based
methods less accessible to patients.

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REFERENCES
A N D E R S O N , J. R., M U M T A Z , G., M U L H A R E , P., P O D D A R , J. and
R I D G W A Y G. L. 1993: Mandatory Use of Confirmation Stage with
Chlamydiazyme During Urinary Sediment Analysis. - J Clin Pathol 46: 896-897.
B L A C K , C.M. 1997: Current Methods of Laboratory Diagnosis of Chlamydia
trachomatis Infections. - Clinical Microbiology Reviews 10: 160–184
D E M A I O , J., B O Y D , R.S., R E N S I , R. and C L A R K , A. 1991: False-positive
Chlamydiazyme Results During Urine Sediment Analysis due to Bacterial
Urinary Tract Infections. - J Clin Microbiol 29: 1436-8.
O S S E W A A R D E , J. M. 1995: Journal of the European Academy of Dermatology
and Venerology 5: 111-123.
S T A M M , W.E. 1999: Chlamydia trachomatis Infections of the Adult. In: Holmes KK,
Sparling PF, Mardh P-A, et al., eds. Sexually Transmitted Diseases. 3 rd ed. New
York, NY: McGraw-Hill,: 407-22.
SUCHLAND, K.L., COUNTS, J.M. and S T A M M , W.E. (1997): Laboratory
Methods for Detection of Chlamydia trachomatis: Survey of Laboratories in
Washington State. - J Clin Microbiol 35: 3210-4.
W A S H I N G T O N , A.E. and K A T Z , P. 1991: Cost of and Payment Source for Pelvic
Inflammatory Disease. Trends and Projections, 1983 through 2000. - JAMA 266:
2565-9.
W E I N S T O C K , H., B E R M A N , S. and C A T E S , W. Jr. 2004: Sexually
Transmitted Diseases among American Youth: Incidence and Prevalence
Estimates, 2000. - Perspectives on Sexual and Reproductive Health 36: 6-10.
Received: 27.04.2007.

NATURA MONTENEGRINA, PODGORICA, 6:137-149
INFLUENCE OF TRANSGENIC PLANTS ON ENVIRONMENT
Danko OBRADOVIĆ 1
1
University of Montenegro, Faculty of Sciences, Department of Biology, P. O. Box 211, 81000 Podgorica,
Montenegro
Key words:
transgenic plants,
commercial use
Klučne riječi:
transgene biljke,
komercijalna
primena
Synopsis
In 1983, transgenic plants were introduced by four groups
of authors. However, commercial use of transgenic plants dates
from 1996. Since its first application, cultivation area of
transgenic crops around the world has been permanently
increasing. Transferring of a transgene from microorganisms,
animals, and distant plants, transgenic plants can obtain novel
treats that would probably never be introduced by conventional
breeding. New treats that plants can get in nature by
spontaneous mutations and breeding occur slowly. During the
evolution organisms have been accommodated to changes in
their environment, in contrast to revolutionary changes
introduced by molecular biology methods applied on production
of transgenic crops. This can make disbalance among organisms
and cause some ecological problems. Such problems can be
solved by regulations which forbid use of transgenic crops
without vigorous assays for safe application.
Sinopsis
UTICAJ TRANSGENIH BILJAKA NA ŽIVOTNU SREDINU
Četiri grupe autora su 1983. godine po prvi put dobile transgene
biljke. Ipak, komercijalna primena transgenih biljaka datira od 1996.
godine. Od svoje prve primene, obradiva površina pod transgenim
kulturama širom sveta je u stalnom porastu. Transferom nekog gena iz
mikroorganizama, životinja i nesrodnih biljaka, transgene biljke mogu
dobiti nove osobine koje verovatno ne bi nikada dobile konvencionalnim
ukrštanjem. Nova svojstva koja biljke mogu dobiti u prirodi spontanim
mutacijama i ukrštanjem događaju se sporo. U toku evolucije organizmi
su se prilagođavali na promene u njihovoj okolini za razliku od
revolucionarnih promena koje su uvele metode molekularne biologije
primenjene u proizvodnji transgenih kultura. Ovo može dovesti do
disbalansa među organizmima i uzrokovati neke ekološke probleme.
Takvi problemi mogu biti rešeni propisima koji zabranjuju primenu
transgenih biljaka bez prethodne temeljne provere bezbednosti primene.

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INTRODUCTION
Plant breeding has been a basic method for obtaining plants of particular traits
for several thousand years. People have used this method to improve some
characteristics of plants (sweeter and larger fruits, aridity and disease resistance,
faster growth) in spite of the fact that they did not understand breeding process and
its background. In 1865, Gregor Mendel showed that the inheritance of traits follows
particular regularities, but the significance of Mendel's work was not recognized until
1900. Traditional plant breeding is very difficult and time and labor consuming
process. By artificial plant crossing, plant breeders try to obtain plants of desirable
traits. To achieve this goal, they have to travel all around the world to find plants of
particular traits that are suitable for breeding. Variations of treats among plants arise
as result of mutations. However, these mutations, and as well as variations, are of
limited number. In 1926, Hermann Joseph Muller found that X-rays irradiation can
induce mutations (in 1946 he was awarded Nobel Prize for this discovery) (M u l l e r ,
1926). After this discovery, it was found out that irradiation of seed can greatly
increase amount of mutations in next generation. Since the end of World War II,
induced mutation have been widely applied in plant breeding. Such method was
named mutation breeding. Mutations, and thus, variations of traits, can be induced
with ionizing radiation (X-rays, gamma rays, alpha particles, beta particles, neutrons,
protons) or chemical agents (sodium azide, ethyl methanesulphonate). This is a way
to increase variability and to get plants of desirable traits that are not available in
nature. This method is named mutation breading. It became popular after World War
II, and it is popular at the present time as well.
Classical plant breeding uses deliberate interbreeding of closely or distantly
related species to produce new crops with desirable properties. Breeding is possible
between plants within the same species, genus and, less commonly, between plants
of different genera. Plants that are evolutionally distant have larger amount of different
properties. Thus, amount of plant property variations that are available for breeding
and obtaining of new crops is limited, because breeding between plants that are
evolutionally distant is not possible. Even application of mutation breading produces
new crops with limited variations in treats. Additionally, induced mutations which are
obtained by mutation breeding produce new treats that cannot be predicted, because
mutations are random events. All these features of classical and mutation breeding
limit their application.
In 1983, four groups of authors, almost simultaneously, introduced a novel
method for production of new crops. They worked independently, and three of them
announced their work at a conference in Miami, USA in January 1983 (Framond et al.,
1983; Schell et al., 1983; Fraley et al., 1983). Their researches enable gene transfer
from bacteria into plants. These works were published in scientific journals as well
(B e v a n , et al. 1983; H e r r e r a - E s t r e l l a et al., 1983; F r a l e y et al., 1983). The
fourth had transferred a plant gene from one species into another species. They
announced their research at a conference in Los Angeles, USA, in April 1983, and

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later it was published in a journal (M u r a i et al., 1983). Plants obtained this way are
transgenic plants. This method enables gene insertion of one or more genes from a
plant which can be evolutionally very distant into another plant. The genes can be
transferred from any organism (animals, bacteria, viruses etc.) into a plant. This
makes possible production of novel crops with new treats which in practice can never
be obtained by breeding. Very important feature of this method is possibility to predict
novel treats of transgenic crops. It enables to skip millions of years of evolution that
would be necessary for nature to produce plants with so different treats. This skip
made impossible accommodation between transgenic crops and environment, and
develops concerns regarding the application of transgenic plants. However, transgenic
plants can provide substantial benefits: better nutritional characteristics, increased
productivity, longer shelf life, environmental tolerance, pest and disease resistance,
etc.
Production Process of Transgenic Plants
Molecular biology methods enable gene transfer and production of transgenic
plants. These methods are commonly named genetic engineering. Actually, all
molecular biology methods that can be involved as tools for changing of genetic
constitution are named genetic engineering. Production of transgenic plants consists
of: isolation of a gene from donor organism, insertion of the gene into plant cell,
obtaining of whole plants from transformed cells grown in tissue culture, plant
breeding and testing.
Isolation of a Gene from a Donor Organism. This is the most difficult step in
production of transgenic plants. This step is mostly based on fundamental research in
determination of structure and function of donor organism genes. It is very important
to locate and determine the role of a gene, its function and treats that the gene is
responsible for. Identification of expression mechanisms of the gene and its influence
on other genes, metabolic pathways of gene products is very important. Procedure of
this step depends very much on results of such researches. In contrast to this step, all
other steps are processed under well determined procedures. Isolated gene that is to
be inserted into a plant is termed transgene.
Insertion of the Gene into Plant Cells. Currently, there are two types of
vehicles which can be used for gene insertion into plants. The most often used vehicle
is Agrobacterium tumefaciens (Figure 1). A. tumefaciens is a gram negative, rod
shaped, bacteria, which is the causal agent of Crown Gall disease (the formation of
tumors) of dicots. It has been used as gene transfer vehicle into dicots, and just
recently into monocots. A. tumefaciens contains chromosomal DNA and plasmid
(known as the Ti-plasmid for tumor-inducing plasmid). In order to be virulent the
bacterial Ti-plasmid has to contain a small segment of DNA termed T-DNA
(transferred DNA), and vir (virulence) genes that direct the infection process. A.

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tumefaciens are attracted to wound sites of a plant by chemotaxis. This is a response
to the release of some common root components (sugars and particularly phenolic
compounds such as acetosyringone). Acetosyringone activates vir genes on the Tiplasmid.
Activity of this gene leads to the production of opine permease, that is
inserted in the bacterial cell membrane for uptake of compounds (opines) that will be
produced by the tumors, and production of an enzyme restriction endonuclease that
excises part of the Ti-plasmid termed the T-DNA. Excised T-DNA released by the
bacterium enters the plant cells and in an unknown way integrates into the
chromosome. T-DNA dictates change in the functioning of those cells which cause
formation of tumors.
Figure 1. Simplified organization of a sequence constructed for insertion into a plant genome.
Insertion of a transgene into plant cell genome does not mean that the gene will
be expressed. Unexpressed gene will not code any product (protein), and thus, will
not provide any new treat to the plant. In order to provide gene expression a promoter
DNA sequence has to be inserted together with the gene, and as well as a termination
sequence (terminator). Terminator signals that gene coding sequences ends, and that
transcription of the DNA has to be ended. A given promoter and terminator can be
combined with various genes (Figure 2). The degree of gene expression, the region of
plant body where the gene will be expressed, and the plant life cycle stage depend on
which promoter is applied.
A marker gene has to be inserted together with the transgene. It is necessary for
selection of cells which received and expressed a transgene. It encodes a protein
which provides resistance to an agent (usually antibiotics or herbicides) which is toxic
to plant cells. Thus, cells that received transgene will be resistant to the toxic agent
(they will survive), and all other cells will die.
Figure 2. Bacterium Agrobacterium tumefaciens.

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Ti-plasmid used in gene transfer process is not a wild type plasmid, but plasmid
especially constructed for this purpose. T-DNA sequence which causes tumor growth
is deleted and just its border sequence is retained. A. tumefaciens with such Tiplasmid
is not a pathogen. It retains ability to insert the plasmid into the cell, but the
plasmid cannot cause tumor growth. Transgene is to be inserted into the Ti-plasmid,
instead of removed T-DNA sequence and transferred into a plant cell. The cell is
maintained in tissue culture and transgene is incorporated into the plant chromosome.
The cells in tissue culture are grown in media containing nutrients and hormones
necessary for cell growth.
Obtaining of Whole Plants from Transformed Cells Grown in Tissue
Culture. After the treatment with A. tumefaciens, it is necessary to select transformed
cells. The selection is carried out by replacement of standard media with selectable
media, which contain an agent (usually antibiotic or herbicide) toxic for the cells.
Marker gene inserted into plant chromosome together with the transgene provides
resistance to the toxic agent. Thus, the cells that received marker gene will survive in
the tissue culture, and all other will die. Survived cells will be maintained to produce
an embryo and eventually whole plant.
Plant Breeding and Testing. A limited number of plant lines is possible to use
in effective gene transfer. They are usually not elite lines. To obtain an elite line with a
transgene it is necessary to breed transgenic plant with an elite line. After the
breeding process, a transgenic plant containing at least 98% of elite genes is
produced. The next step is testing of the plant (expression of transgene, stability of
inheritance of the treats, unexpected features of the plant etc.).
Another method for insertion of a transgene is the gene gun. All steps in
production of transgenic plants applying gene gun are pretty much the same as with
A. tumefaciens, except gene insertion. Gene gun method applies microscopic gold
particles to deliver a transgene into the plant cell nucleus. Golden particles coated
with a large amount of transgenes together with a marker gene are accelerated with
air pressure and shoot at tissue culture cells. The golden particles will pass into the
cell nucleus. Coated DNA will be dissolved and inserted into the chromosome. The
method with A. tumefaciens is more effective than gene gun method, but it cannot be
applied to every plant species. A. tumefaciens is ussualy applied with dicots, and
gene gun with monocots.
Transgenic Plants Benefits
New treats introduced by insertion of transgenes into plants can provide a
number of benefits: pest and disease resistance, herbicide tolerance, better nutritional
characteristics, increased productivity, longer shelf life, environmental tolerance, etc.

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Pest and Disease Resistance. Insertion of a transgene (isolated from a soil
bacterium Bacillus thuringiensis) coding a crystalline (Cry) protein can introduce pest
resistance to transgenic plants. In the intestine gut, the protein is broken down to
release delta-endotoxin that creates pores in the intestinal lining of pets. This creates
ion imbalance, dysfunction of digestive system, and death of the insect in a few days.
Product of Cry gene is considered safe for human and birds, and less harmful for nontarget
insects than other insecticides. Insecticides based on Cry gene have been used
for long time, but in this case they are produced by plant in the plant body. Thus,
insects eating such plant will eventually die. Plants containing this transgene are
named Bt (Bacillus thuringiensis) plants. Bt plants available by now are: Bt cotton (to
control European corn borer, Southwestern corn borer, and corn earworm), Bt cotton
(to control cotton bollworm and tobacco budworm), and Bt potato (to control Colorado
potato beetle). Bt plants can decrease application of insecticides, and so, the
surrounding environment is no longer exposed to large amounts of harmful insecticide.
Time and labor saving is achieved too.
Transgenic papaya and squash carrying virus coat protein gene are resistant
to virus. This gene produces virus coat protein before an infection. After the infection,
plant cells will not produce this protein because of co-suppression (plant cell
mechanism which suppresses production of the viral protein), and virus cannot
replicate.
Herbicide Tolerance. Weeds can significantly decrease crop yield, and so
herbicides are widely applied for weed control. Sometimes farmers apply more than
one herbicide because they are specific for a particular weed, and they are applied at
particular growth stage. Herbicides are long lasting and can persist in the soil for
years. Transgenic herbicide tolerant plants have a transgene providing resistance to
new herbicide that can kill all kinds of plants (including weeds). These herbicides
break down in the soil quickly. So far, there are two herbicides that are used with
herbicide tolerant plants: Liberty (glufosinate) and Roundup (glyphosate). These
herbicides break down quickly in contrast to conventional herbicides that can remain
in the soil for more than a year, and thus prevent farmers to plant crops sensitive to
them in future. Fast degradation of these new herbicides protects environment of large
scale contamination, which can be common in the case of conventional herbicides. An
example of transgenic herbicide tolerant plant is Roundup Ready soybean.
Long-Lasting Products. Some plants (strawberries, tomatoes, pineapples,
sweet peppers and bananas) are genetically modified to produce less enzyme that
cause products to rot. These plants can remain firm and fresh for long time. The first
long-lasting plant modified plants was tomato, which came on the market in 1994.
Such tomato can tolerate a longer transport time and it can be allowed to ripen in the
sun before picking - resulting in a better tasting tomato.

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Improvements of nutritious characteristics of the plants. Nutritious
characteristics of a plant can be improved by inserting a transgene. An example is
golden rice. It contains beta-carotene which is converted into vitamin A in the body.
For the golden rice to make beta-carotene three new genes are implanted: two from
daffodils and the third from a bacterium.
Transgenic Plants Risks and Concerns
Transgenic plants can provide many benefits, but in the same time some risks
and concerns. Insertion of transgenes into plants, which introduces novel treats to the
plants, can change interactions among plants and animals. This can cause ecological
and other kinds of problems.
Gene Transfer from Transgenic Crops to Their Wild Relatives. Many crops
have wild relatives which can be crossed by pollination. Thus, there is real danger of
gene transfer from crops to wild population including weeds (K a i s e r , 2001; P o p p y
and W i l k i n s o n , 2005). This possibility is supported by researches. Wild population
would receive novel treats that can make them resistant to many factors. This is
especially critical for weeds. Weeds resistant for herbicides, plant diseases, climatic
factors etc. would be a nightmare for every farmer. Transgene transfer to wild
relatives that are not weeds is not desirable either, because this could change natural
balance in the wild nature and cause ecological problems.
Crop to Crop Gene Flow. Gene transfer is possible from transgenic to
conventional crops (H a l l et al., 2000; R i t a l a e t al., 2002).The transfer is processed
by pollination, and pollen can be carried by wind and insects. After this gene flow,
farmers that plant conventional crops will have transgenic even if they dislike them.
Long-term application of transgenic crops and gene flow can lead even to complete
loss of conventional crops. This is the most likely to occur in the case of highly
outcrossing plants in contrast to highly self-pollinating plants.
Antibiotic resistance. All transgenic crops in the process of their production
have to contain a marker gene which is inserted to insure that a transgene has been
inserted into the plant. It is usually an antibiotic resistance gene. There is a concern
that bacteria, which inhabit intestine of human and animals, could be transformed by
transfer of a DNA fragment carrying the antibiotic resistance gene from the intestine
into the bacterial cell. After such transformation, bacteria would become resistant to a
given antibiotic. This could lead to increase of amount of antibiotic resistant bacterial
strains in the nature, and would cause problems in human and animal medical
treatment. However, application of new markers that do not represent antibiotic
resistance genes would resolve this problem. An example of such markers is green
fluorescent protein and mannose (Joersbo et al., 1998). Another way to resolve this

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problem is to remove the marker after the development of transgenic plant, when its
presence is not required any more (Zuo, 2001). Transgenic plants with green
fluorescent protein as a marker and with removed markers are already submitted for
authorization and their large scale production can be expected soon.
Allergenicity. Insertion of a transgene into a plant can cause production of a
product which is an allergen. This allergen will cause allergic reaction, which can be
fatal to persons who consume such plants and are allergic. Some plants do not
contain any allergen, but insertion of a transgene which produces them can cause
health problem and even death of people who are allergic. Allergic people who do not
know that a transgenic plant contains a given allergen can be in great danger. An
example is transgenic soybean with inserted gene from Brazil nut (Nordlee et al.,
1996). The gene was inserted to improve nutritional characteristics of soybean, but
people who were allergic to Brazil nut were allergic to mentioned transgenic soybean
too. This soybean has never been approved for market application. Now, all
transgenic plants are tested for allergens to solve this problem.
Terminator Technology. In 1990s U.S. Department of Agriculture and Delta
and Pine Land Company developed a method for protection of technology. The goal of
this method is to force farmers to pay intellectual property for development of
transgenic plants. Actually, many farmers all over the world save some seeds to plant
next year. If they buy transgenic seeds from a company developer of the transgenic
plant once, they do not have to buy it anymore because they can produce transgenic
seeds by themselves, saving some of the seeds in next generation. In 1998,
developers of mentioned method were awarded a joint patent. The method is named
Technology Protection System (TPS). TPS makes possible production of seeds which
is sterile in the second generation. Seeds obtained after first generation is good for
consumption, but plant embryo is killed by TPS, making seed unsuitable for planting.
Thus, if farmers want to plant transgenic plants, they have to buy seeds from company
developer of the seed.
Advantage of this technology is that plant pollinated with transgenic plants will
be sterile as well, and it will prevent gene flow from transgenic plant to the other
plants. Another advantage is that this will encourage biotech companies for further
development of transgenic plants with novel treats, especially those that are not so
profitable for them.
Disadvantage is that some farmers, especially small and farmers in developing
countries, will not be able to buy seeds every season. That would increase cost of the
production, and it can lead to their bankruptcy in competition with large
manufacturers. Another disadvantage is that neighboring non-transgenic crops and
wild plants would be affected, because after possible pollination their seeds would be
sterile and it could cause lower yield. This impact would be higher on non-transgenic
crops than on wild plants.

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There is another system for protection of technology named Trait-specific
Genetic Use Restriction Technology or T-Gurt. It was developed by agro-chemical
companies including Monsanto and AstraZeneca. T-Gurt does not cause production of
sterile seeds. The seeds in next generation would not express transgenes. However, if
seeds are sprayed with a chemical, sold solely by the manufacturer of seeds, the
transgene would be expressed.
Benefits of this technology are that transgenes would not be expressed in the
wild plants, and farmers can plant seeds obtained in the next generation (but without
expression of transgenes if they do not buy the chemicals). Disadvantages are that
transgenes would be transferred by pollination to the neighboring population (crops
and wild plants) in spite the fact that it will not be expressed and spraying of the seed
with the chemicals can make negative impact on environment.
A new sophisticated system for protection of technology which produces sterile
seeds is under development. This seed becomes fertile if it is sprayed with a chemical
sold by company developer of the transgenic plant.
None of terminator technologies has been applied yet. A great resistance to its
application given by farmers all around the world is one of the reasons for delaying of
the application. However, it is hard to believe that large agro-chemical companies will
quit these projects which can enable them to make additional large scale profit.
Production of Transgenic Crops around the World
In 1996, the first transgenic crops were planted in the USA for commercial use.
Cultivation of transgenic crops has shown permanent increase (Figure 3) according to
ISAAA (ISAAA, 2006). In 2006, they were grown on 102 million hectares worldwide.
The increase has been observed in both, industrial and developing countries. The
USA is the country with the largest area planted with transgenic crops. European
countries do not plant large areas with transgenic plants, and their amounts are
modest in comparison to some other countries (table 1). In 2006, the most often
planted transgenic crops around the world were: soybean (58.6 million hectares),
maize (25.2 million hectares), cotton (13.4 million hectares), canola (4.8 million
hectares).

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Natura Montenegrina, 6/2007
Figure 3. Global area of transgenic plants in
million hectares (1996-2006).
COUNTRY
CULTIVATION
AREAS
TRANSGENIC PLANTS
USA 54,6 S,M,C,R, Squash, Papaya,
Alfalfa
Argentina 18,0 S,M,C
Brazil 11,5 S, C
Canada 6,1 R,M,S
India 3,8 C
China 3,5 C
Paraguay 2,0 S
South Africa 1,4 M,S,C
Uruguay 0,4 S,M
Philippines 0,2 M
Australia 0,2 C
Romania 0,1 S
Mexico 0,1 C,S
Spain 0,1 M
Colombia

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Obradović: INFLUENCE OF TRANSGENIC PLANTS ON ENVIRONMENT
CONCLUSION
There are many contradictory opinions in connection with transgenic plants.
Even experts in that field do not have common opinion. Some experts have opinion
that application of transgenic plants can be very dangerous for environment and
human health. The others have opinion that risks are overestimated, and that future is
in transgenic plants. Actually, probably both groups of experts are right. Application of
transgenic plants without appropriate regulations can be very risky. The new
transgenes inserted into them, and novel treats, can disturb harmony among
organisms that has been built by nature for many millions of years. This
disorganization of an ecosystem is not predictable and cannot be always prevented.
Restoring of the consequences could be very difficult and probably in majority of the
cases impossible. Transgenes that are accidentally transferred to the other plants,
including wild environment, will stay there forever. However, application of transgenic
plants can increase agricultural production, decrease cost of products and save
environment by decreasing amount of applied chemicals (or by application of safer
chemicals). Transgenic plant can be resistant to various plant diseases and
environmental conditions which can make them very suitable for agricultural
production. The problems caused by introduction of transgenes cannot be solved by
nature, or to be more accurate, nature would need very long time to do that with
dangerous consequences. Thus, only solution is to prevent all known problems and to
begin with application of transgenic plants with precautions. Regulations, which would
prevent large scale application without preliminary assays and approval, are
necessary at national and as well as international levels.
Contemporary obtained improvements obtained by transgenic technology are
very modest in comparison to benefits that will be in the future. Now, we enjoy y just
novice movements in this field. Benefits in the future will be so huge that agricultural
production without transgenic plants can be compared to transportation with animaldrawn
vehicles.
LITERATURE
BEVAN, M.W., FLAVELL, R.B. and C H I L T O N M.D. 1983: A Chimaeric
Antibiotic Resistance Gene as a Selectable Marker for Plant Cell
Transformation. - Nature 304: 184-187.
F R A L E Y , R.T., R O G E R S , S.B. and H O R S C H . R.B. 1983: Use of a Chimeric
Gene to Confer Antibiotic Resistance to Plant Cells. - Advances in Gene
Technology: Molecular Genetics of Plants and Animals. Miami Winter Symposia
Vol. 20: 211-221.
F R A L E Y , R.T., R O G E R S , S.G., H O R S C H , R.B., S A N D E R S , P.R., F L I C K ,
J.S., A D A M S , S.P., B I T T N E R , M.L., B R A N D , L.A., F I N K , C.L., F R Y ,

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J.S., G A L L U P P I , G.R., G O L D B E R G , S.B. H O F F M A N N , N.L. and
W O O , S.C. 1983: Expression of Bacterial Genes in Plant Cells. - Proceedings
of the National Academy of Sciences 80: 4803-4807.
FRAMOND, A.J., BEVAN, M.W., BARTON, K.A., FLAVELL, F. and
C H I L T O N , M.D. 1983: Mini-Ti Plasmid and a Chimeric Gene Construct: New
Approaches to Plant Gene Vector Construction. - Advances in Gene Technology:
Molecular Genetics of Plants and Animals. Miami Winter Symposia Vol 20: 159-
170.
H A L L , L., T O P I N K A , K., H U F F M A N , J., and D A V I S , L. 2000: Pollen Flow
between Herbicide-resistant Brassica napus Is the Cause of Multiple-resistant B.
napus Volunteers. - Weed Science 48: 688-694.
HERRERA-ESTRELLA, L., DEPICKER, A., Van MONTAGU, M. and
S C H E L L , J. 1983. Expression of Chimaeric Genes Transfered into Plant Cells
Using a Ti-plasmid-derived Vector. - Nature 303: 209-213.
ISAAA (The International Service for the Acquisition of Agri-biotech Applications)
Briefs (2006) 35.
JOERSBO, M., DONALDSON, I., KREIBERG, J., PETERSEN, S.G.,
B R U N S T E D T , J. and O K K E L S , F.T. (1998): Analysis of Mannose
Selection Used for Transformation of Sugar Beet. Molecular Breeding 4: 111-
117.
K A I S E R , J. 2001: Breeding a Hardier Weed. - Science 293: 1425-1426.
M U L L E R , H.J. 1926: The Problem of Genetic Modification. Fifth International
Congress of Genetics. Berlin.
M U R A I , N., S U T T O N , D.W., M U R R A Y , M.G., S L I G H T O M , J.L., M E R L O ,
D.J., REICHERT, N.A., SENGUPTA-GOPALAN, C., STOCK, C.A.,
B A R K E R , R.F., K E M P , J.D. and H A L L ., T.C. 1983: Phaseolin Gene from
Bean Is Expressed after Transfer to Sunflower via Tumor-inducing Plasmid
Vectors. - Science 222: 476-482.
NORDLEE, J.A., TAYLOR, S.L., TOWNSEND, J.A., THOMAS, L.A. and
B U S H , R.K. 1996: Identification of a Brazil-nut Allergen in Transgenic
Soybeans. - New England Journal of Medicine 334: 688-692.
POPPY, G.M. and W I L K I N S O N M.J. 2005: Gene Flow from GM Plants. -
Blackwell Publishing.
RITALA, A., NUUTILA, A.M., AIKASALO, R., KAUPPINEN, V. and
T A M M I S O L A , J. 2002: Measuring Gene Flow in the Cultivation of Transgenic
Barley. Crop Science 42: 278-285.
SCHELL, J., Van MONTAGU, M., HOLSTERS, M., ZAMBRYSKI, P.,
J O O S , H., I N Z E , D., H E R R E R A - E S T R E L L A , L., D E P I C K E R , A., D e

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BLOCK, M., CAPLAN, A., DHAESE, P., Van HAUTE, E.,
HERNALSTEENS, J-P., De GREVE, H., LEEMANS, J.,
DEBLAERE, R., WILLMITZER, L., SCHRODER, J. and
O T T E N , L . 1983: Ti Plasmids as Experimental Gene Vectors for Plants.
Advances in Gene Technology: Molecular Genetics of Plants and Animals. -
Miami Winter Symposia Vol. 20: 191-209.
Z U O , J., N I U , Q.W., M O L L E R , S.G. and C H U A , N.H. 2001: Chemical-regulated,
Site-specific DNA Excision in Transgenic Plants. - Nature Biotechnology 19:
157-161.
Received: 27.04.2007.

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NATURA MONTENEGRINA, PODGORICA, 6:151-152
Caldesia parnassifolia (L.) P a r l NEW SPECIES IN MONTENEGRIN FLORA
Vera BIBERDŽIĆ 1
1 Natural History Museum of Montenegro, Trg Vojvode Bećir Bega Osmanagića 16,
81000 Podgorica, Montenegro, E-mail: prmuzej@cg.yu
During macrophyta research of Skadar Lake and its surroundings, the species
Caldesia parnassifolia (L.) Parl was found, representing the first finding of this taxon
in Montenegro. A very small population of floating leaves was found at the locality
called Pančeva oka.
Caldesia parnassifolia (L.) Parl represents the remainder of the tertiary
hydrophyte flora in our country and in the Balkan Peninsula. In phylogenetic respect, it
is most similar to Alisma genus, which is characterized by a cosmopolitan distribution.
Pollination is entomophilic, and the dispersal of seeds is hydrochory i.e. by water. It is
vegetatively propagated via adventitious tree buds, blossoms in July and August. It
belongs to the subtropical flora component.
Genus Caldesia Parl. is represented in the European Flora by one species:
Caldesia parnassifolia (L.) Parl. (T u t i n et al. 1964-1980) and it is generally
distributed in tropical and subtropical regions of Africa, Asia and Australia. In the
temperate zone of Eurasia it is also distributed disjointly: one part of the areal can be
found in Central and Southern Europe (to France to the west, to Ukraine to the east,
to Lithuania to the north and to central Italy to the south) (T u t i n et al 1964-1980,
Vukojič i ć , S., J a n k o v i ć , M. 1999), while the other one involves eastern Asia
(Manchuria, China, Japan).
Prodromus Flore penisulae Balcanicae, Hayek 1932-1933, records that Caldesia
parnassifolia (Bassi) Parl is present in Bosnia and Herzegovina?, Serbia and Bulgaria.
The species has also been recorded in Croatia (N i k o l i ć , T. 2005)
The taxon has been included in the European Red List for Flora. It has also
been included in the List of the Convention on the Conservation of European Wildlife
and Natural Habitats (Bern Convention) and in the List of Species of the Habitats
Directive (Annex IIb, IVb.)
In future research, we will try to find this taxon at other localities as well, to
determine the size of the population and to identify the degree to which it is
endangered in Montenegrin flora.

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LITERATURE
H A Y E K , A. 1932-1933: Prodromus Florae Penisulae Balcanicae. - 3 Fedes Repert.
(Beih) 30.
NIKOLIĆ , T., T O P I Ć J. ur. 2005: Crvena knjiga vaskularne flore Republike
Hrvatske. Kategorije EX,RE,CR,EN i VU. - Ministarstvo kulture, Državni zavod
za zaštitu prirode, Zagreb, 4-695.
TUTIN, T. G., HEYWOOD, V. H., BURGES, N. A. MOORE., VALENTINE,
D. H., WALTERS, S. M., WEBB, D. A. (eds) 1964 - 1980: Flora Europaea 1-
5. - Cambridge University Press.
VUKOJIČ I Ć , S., J A N K O V I Ć , M. 1999: Caldesia parnassifolia (L.) Parl u
Stevanović, V.: Crvena knjiga flore Srbije, 1, Taksoni iščezli iz Srbije. -
Ministarstvo za životnu sredinu Republike Srbije, 57-136.
Received: 08.11.2007.

NATURA MONTENEGRINA, PODGORICA, 6:153-160
ADDITIONS TO THE FLORA OF MONTENEGRO: Setaria verticilliformis Dumort.,
Setaria viridis (L.) PB. subsp. pycnocoma (Steud) Tzvel., Impatiens balsamina L.
AND Catalpa bignoniodes Walt.
Danijela STEŠEVIĆ 1) Nejc J O G A N 2)
1)
Faculty of Science, University of Montenegro, Cetinjski put bb, 81 000 Podgorica, denist@cg.yu
2) Department of Biology BF, University in Ljubljana, Večna pot 111, SI-1000 Ljubljana, Slovenia,
nejc.jogan@bf.uni-lj.si
Setaria verticilliformis Dumort.
According to compilation by R o h l e n a 1942 later amended by P u l e v i ć
2005 in the flora of Montenegro genus Setaria PB. is represented with only 3 species:
Setaria glauca (= Setaria pumila (Poir.) Roem. & Schult.), S. verticillata (L.) PB. and
S. viridis (L.) PB.
Recently, two additional taxa of Setaria have been recorded: S. verticilliformis
Dum. and S. viridis (L.) PB. subsp. pycnocoma (Steud) Tzvel.
The first mentioned species has been recorded in the city area of Podgorica (city
lawns, waste places and roadsides). Using keys in Pignati (1982a) species can be
determined as S. ambigua Guss. Checking additional literature sources (H i t c h o c k
1950, C l a y t o n 1980, W i s s k i r c h e n & H a e u p l e r 1998, M a r t i n č i ć
1999, A e s c h i m a n n et al 2004, F i s c h e r et al 2005,
http://herbarium.usu.edu/treatments/Setaria.htm), we noticed that opinion about
taxonomic status of this taxon differs between authors. According to ITIC (Integrated
Taxonomic Information System - http://itis.gov/) its accepted scientific name is S.
verticilliformis Dumort. Some more important synonyms are listed below:
= Setaria verticillata (L.) PB. var. ambigua (Guss.) Parl.
= Setaria verticillata × viridis
= Setaria ambigua (Guss.) Guss., non Schraeder
= Setaria decipiens Schimp. ex Nyman
= Setaria gussonei Kerg.
S. verticilliformis is thermocosmopolitan species (P i g n a t i 1982a), so we
presume that the reason why it was not reported for the flora of Montenegro up to
now, might be its confusion with similar S. verticillata or S. viridis.
Photos of panicles of these 3 Setaria species, macrodetails of rachis and
bristles, and some distinguishing characters are given below:

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Natura Montenegrina, SHORT COMMUNICATION, 6/2007
bristles are retroversly scabrous, rachis is retroversly scabrous
bristles are mostly antroversly scabrous, rachis is antroversly scabrous
Photos by: Danijela Stešević
bristles are antroversly scabrous, rachis is hispid and villous

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Stešević, Jogan: Additions to the flora of Montenegro . . .
Setaria viridis (L.) PB. subsp. pycnocoma (Steud) Tzvelev
Taxonomy of the group of closely related taxa between wild S. viridis on one
side and cultivated S. italica, on the other, is quite blurred. Most probably the reason
for that is, that the wild mentioned species is supposed to be a progenitior of the
cultivated one, so they are morphologically linked by an array of intermediates. Those,
more resembling typical S. viridis are often recognized as S. viridis subsp. pycnocoma
and the ones closer to typical S. italica but still retaining some “wild” character states
as presence of bristles, disarticulating ripe spikelets etc. are called S. italica subsp.
moharia (Alef.) Koern. Main distinguishing characters delimiting mentioned subspecies
of S. viridis are (compiled by Jogan in Martinčič et al. 2007):
- Stem more than 80 cm high (less than 50 (80) cm in type subsp.)
- Panicle lobed and interrupt towards the base, more than (0.8) 1.2 cm wide (vs.
not lobed and up to 0.6 (0.8) cm wide)
- Spikelets (2.2) 2.4-2.6 (2.8) mm long (and only (1.8) 2-2.4 (2.5) mm in subsp.
viridis)
The most important synonyms of S. viridis
subsp. pycnocoma are:
- Setaria viridis (L.) PB. var. major
(Gaudin) Peterm.
- Setaria gigantea (Franch. & Sav.)
Makino
- Setaria pycnocoma (Steud.) Nakai
- Panicum comosum Steud.

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As an obligate segetal weed, S. viridis subsp. pycnocoma, is supposed to be of
a hybrid origin between cultivated and wild growing relatives mentioned above.
Darmency (2005) compiled a “dynamic” model of possible relationship among wild,
weedy, crop and feral forms in the Setaria italica/S. viridis group:
Species Process Resulting Type Conclusion Likelihood
S. viridis Selection by man S. italica Domestication High
S. italica Seed lost at harvest S. italica Volunteerism Very low
S. italica Back mutation S. viridis ssp.
pycnocoma
S. viridis Genetic variability S. viridis ssp.
pycnocoma
S. italica and
S.viridis
Interspecific gene
flow
S. viridis ssp.
pycnocoma
Endoferality
Mimicry
Exoferality
Hyper
rare
Possible
High
In Montenegro, up to now, S. viridis subsp. pycnocoma has been recorded in
Gornji Kokoti (roadside Podgorica – Cetinje), but we can expect its scattered
occurrence also in other lowland parts of the country. Its recent discovery in the
discussed territory (together with the first discovered S. decipiens) showed a need for
taxonomic revision of this genus in the flora of Montenegro.
Impatiens balsamina L. (fam. Balsaminaceae)
In the flora of Montenegro, genus Impatiens L. was present with only one
species - Impatiens noli tangere (R o h l e n a 1942, P u l e v i ć 2005), which is
according to M o o r e (1968) the only Impatiens species native to Europe. All others
are neophytes from Asia and North America (M o o r e 1968, S l a v n i k 1995).
During our field survey undertaken on June 22, 2007, at waste place on Tološko
polje, we have found a few indviduals of unfamiliar Imaptiens, later identified as
Impatiens balsamina (M o o r e 1968, P i g n a t i 1982b). It is annual, up to 1m tall,
with alternate, eliptical to lanceolate-obovate leaves, with serrate margins and 15-20
teeth on each side. Flowers are solitary or in groups up to 3 in leaf axils, pinkish to
purplish to white. Capsula pubescent.

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Impatiens noli tangere
Impatiens balsamina
Photos by: Danijela Stešević
Impatiens balsamina originates from South East Asia and just after its
introduction to Europe (1542) it stared to spread spontaneously (http://www.bbg.org/
gar2/topics/plants/handbooks/flowers/5b.html).
According to available sources I. balsamina is reported in several European
countries as ephemeral escape from the gardens (Italy, UK, Czech Republic, Austria,
Hungary; P i g n a t i 1982b, C l e m e n t s & F o s t e r 1994, P y š e k et al. 2002, E
s s l & R a b i t s c h 2002, B o t o n d & Z o l t a n 2004), occasionally naturalized
(Austria, France, Italy, Czech Republic, (?)Yugoslavia; M o o r e 1968, G r e t u e r et
al. 1984) or maybe even an invasive alien species in E Europe (http://www.sevin.ru/
invasive/dbases/plants/species.html).
In some other regions of the world, such as in Austral-Pacific region (http://www.
gisp.org/downloadpubs/AP_NATIO.PDF), Pacific islands (http://www.hear.org/pier/reports/
kiribati_appendix1.htm), Micronesia (http://www.hear.org/pier/pdf/kosrae_report.pdf),
Peru (http://i3n.iabin.net/documents/progress_peru_cbd_informe.doc), etc... I. balsamina
has status of invasive species.
Considering the appearance of I. balsamina at Tološko polje, we presume its
seeds were brought here with the waste from nearby gardens, in which it is grown as
an ornamental plant, so in Montenegro it can be only an ephemeral garden escape.

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Catalpa bignonioides Walt. (fam. Bignoniaceae)
Catalpa bignonioides (photo by Danijela Stešević)
Catalpa bignonioides originates from South East part of North America. It is very
decorative deciduous tree, up to 15m high, with rounded to broad crown. Leaves are
opposite, with simple, broadly egg- shaped leaflet up to 30cm long, sometimes with
one or two lateral lobes. Petiole is up to 16cm long. Flowers are big (3-4 cm), white
and two lipped, with two yellow strips in inner part as well as purple spots.
Inflorescence is a panicle. It is flowering in July. It reproduces by seeds and
vegetatively (Š i l i ć 1990).
Common catalpa is deliberately introduced in Europe as an ornamental plant
and as growing wild it is reported for several countries but mostly only as an
ephemeral escape (e.g. Italy, UK, Austria, Czech Republic, Hungary; P i g n a t i
1982b, G r e u t e r et al. 1984, C l e m e n t s & F o s t e r 1994, E s s l & R a b i t s c
h 2002, P y š e k et al. 2002, B o t o n d & Z o l t a n 2004). It may be more
problematic only in Spain (D a n a et al. 2005 reports its weedy occurrence in
Andalucia) and possibly also an invader in some parts of E Europe
(http://www.sevin.ru/invasive/dbases /plants/species.html), but the definition of “invader” in
the sense of that source is not clear-cut.

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During the field survey undertaken on June 7, 2007 over the city area of
Podgorica, at the left bank of the River Morača - near Botun, we have found a single
individual of common Catalpa. Due to the fact that this species reproduces by seeds,
we presume that the seed is brought by birds or wind from the nearby gardens where
it is planted as ornamental.
LITERATURE
A E S C H I M A N, D., L A U B E R, K., M O E S E R, D.M., T H E Y R I L L A T, J.P. 2004:
Flora Alpina 1, Haupt. 1159 pp.
B O T O N D, M. & Z O L T Á N, B.D. eds. 2004: Biological Invasions in Hungary. Invasive
Plants. - Természet BUVAR Alapítvány Kiadó
C L A Y T O N, (1980): Setaria Beauv, in T u t i n at al. eds. Flora Europaea vol. 5.,
Cambridge, p. 263-264
C L E M E N T, E.J. & F O S T E R, M.C. 1994: Alien Plants of the British Isles. - Botanical
Society of the British Isles, London. 603 pp.
D A N A, E.D., S A N Z, M., V I V A S, S. and S O B R I N O, E. 2005: Especies Vegetales
Invasoras en Andalucía. Junta de Andalucía
D A R M E N C Y H. 2005: Incestuous Relations of Foxtail Millet (Setaria italica) with
Its Parents and Cousins. In: Gressel J (ed) Crop Ferality and Volunteerism. CRC
Press, Boca Raton, pp 81–96
E S S L, F. & R A B I T S C H, W. eds. 2002: Neobiota in Österreich. - Umweltbundesamt
GmbH, Wien. 432 pp.
F I S C H E R, M. A., W. A D L E R & K. O S W A L D, 2005: Exkursionsflora. Österreich,
Liechtenstein, Südtirol. - Biologiezentrum der Oberösterreichischen Landesmuseen,
Linz, 1373 pp.
G R E U T E R, W., B U R D E T, H.M., L O N G, G. eds. 1984: Med-Checklist- Pteridophyta
(ed. 2) Gymnospermae Dicotiledones (Acanthaceae-Cneoraceae), Conservatoire
et jardin botaniques de la Ville de Geneve. 330 pp.
H I T C H C O C K, A. S. 1950: Manual of the Grasses of the United States, 2nd ed.
(Revised by A. Chase.) Washington, D.C.: Government Printing Office. 1051 pp.
M A R T I N Č I Č, A., T. W R A B E R, N. J O G A N, A. P O D O B N I K, B. T U R K,
B. V R E Š, V. R A V N I K, B. F R A J M A N, S. S T R G U L C K R A J Š E K,
B. T R Č A K, T. B A Č I Č. M. A. F I S C H E R, K. E L E R & B. S U R I N A,
2007: Mala flora Slovenije. 4. izd. Tehniška založba Slovenije, Ljubljana. 845 pp.
M O O R E, D.M. 1968: Impatiens L, in T u t i n at al. eds. Flora Europaea vol. 2.,
Cambridge, p. 240-241
R O H L E N A, J. 1941-1942: Conspectus Florae Montenegrinae, Preslia 20-21, pp. 506
P I G N A T I, S. 1982a: Flora d’ Italia, vol 3, Edagricole. Bologna. p. 612-613
P I G N A T I, S. 1982b: Flora d’ Italia, vol 2, Edagricole. Bologna. p. 72-73, 618
P U L E V I Ć, V. 2005: Materials for the Vascular Flora of Montenegro, Special edition
of The Republic Institute for Nature Protection of Montenegro, Podgorica. pp.
218

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S L A V N I K, B. 1995: Rod Impatiens v České republice, Preslia 67: 193–211.
P Y Š E K, P., S A L D O, J. & M A N D A K, B. 2002: Catalogue of Alien Plants of the
Check Republic, Preslia 74: 97-186
Š I L I Ć, Č. 1990: Ukrasno drveće i grmlje, IP “Svjetlost”, Zavod za udžbenike i
nastavna sredstva, Sarajevo - Zavod za udžbenike i nastavna sredstva,
Beograd, 221 pp.
W I S S K I R C H E N, R. & H A E U P L E R, H. 1998: Standardliste der Farn- und
Blütenpflanzen Deutschlands, Stuttgart: Ulmer, 765 pp.
Received: 14.11.2007

NATURA MONTENEGRINA, PODGORICA, 6:161-163
Duchesnea indica (Andr.) Focke, NEW ALIEN SPECIES IN THE FLORA OF
MONTENEGRO
Igor T O M O V I Ć 1) & Danijela STEŠEVIĆ 1) ∗
1) Faculty of Sciences, University of Montenegro, Cetinjski put bb, 81 000 Podgorica, Montenegro
∗ Corresponding author denist@cg.yu
According to two capital papers about the flora of Montenegro R o h l e n a (1942)
& P u l e v i ć (2005) and recently published contributions to the flora of Montenegro
(Stešević 2005, 2006a, 2006b, H a dž i a b l a h o v i ć 2006, etc.), we can conclude
that record of Duchesnea indica (Andr.) Focke in Berane presents its first record for
the flora of Montenegro.
This South East Asian ornamental plant belogs to family Rosaceae. It is
characterized by perennial herbaceous form, with epigeal rooting stolons. Stem is up
to 50cm; leaves are trifoliate, rather long petiolate, with obovate, crenate leaflets and
lanceolate stipules; flowers are yellow, solitary nor or slightly exceeding the leaves;
sepals are 10mm, epicalyx segments broadly ovate, exceeding the sepals; petals 8
mm; receptacle strongly, bright red, tasteless (V a l e n t i n e 1968).
Due to the its ecological preferences (J a c k o w i a k 1992; L a u b e r & W a g n e r
2001) species mostly inhabits relatively moist, nitrophilous and shaded habitats.
At the first sight it looks likes native strawberry. Characters that clearly
distinguish these species are given in table below:
Native strawberry (Fragaria)
Sepals are nearly equal
Flowers are white
Fruits are juicy, sweet and with pleasant
aroma
Indian strawberry (Duchesnea)
Outer sepals are bigger that inner
Flowers are yellow
Fruits are juicy, sour and without aroma
Photos by: Danijela Stešević

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Duchesnea has worldwide distribution. According to W e b e r (2003)
geographical regions where it grows as native are: Northern Europe (Scandinavia:
Finland, Norway, Sweden); British Isles; Central Europe (Austria, Benelux states,
Denmark, France without the Mediterranean border, Germany, Switzerland); Southern
Europe (Southern France, Greece, Italy, Spain); Eastern Europe (all European states
east of Austria, Germany, Italy); European part of Russia, European part of Turkey,
Mediterranean Islands; Northern Africa (Algeria, Egypt, Libya, Morocco); Tropical
Africa (All African states between northern and southern Africa); Southern Africa
(Lesotho, Namibia, South Africa, Swaziland); Temperate Asia (Middle East, China,
Japan, Asian part of Russia); Tropical Asia (India, Sri Lanka, all states east of India
and bordering the south of China, Malaysia, Indonesia, Philippines); Australia, New
Zealand, Canada + Alaska; South-eastern USA (States of continental USA bordering
the Gulf of Mexico: Alabama, Florida, Louisiana, Mississippi, eastern Texas); Western
USA (States of continental USA bordering the Pacific Ocean); Remaining USA (All
states of continental USA between western and south-eastern USA); Mexico; Tropical
South America (all states of continental South America beyond Mexico and except
Argentina, Chile, Uruguay); Chile; Argentina; Cape Verde; Canary + Madeira; Azores;
South Atlantic Islands; Madagascar and Hawaii.
It is presumed that its introduction in Europe dates from the beginning of 19 th
Century (http://uk.encarta.msn.com/media_1481569451_781532557-1_1/Introduced_
Species_in_Europe.html).
First record of Duchesnea indica for the territory of former Yugoslavia (village
Turčin, near Varaždin in Croatia) was reported by T r i n a j s t i ć (1973). In his opinion
species started to spread spontaneously from Graz (Austria), via valley of river Drava.
For current distribution of Duchesnea in Croatia see Flora Croatica Database
(http://hirc.botanic.hr/fcd/).
Data about distribution of Indian strawberry in Slovenia are available in J o g a n
(2001).
In the region species is also reported in Serbia (J o v a n o v i ć , 1994).
At the territory of Montenegro spontaneously growing population of Duchesnea
indica is found in periurban area of Berane, at shady ruderal habitat type. Pathway of
its arrival remains unknown. Considering the fact that in Berane and in surrounding
area, Indian strawberry is not planted as ornamental plant, we exclude the possibility
that it escaped from cultivation.
We believe that further data about distribution of Duchesnea indica in the region
will help us to create precise hypothesis about the pathway of its arrival and spread
over the territory of Montenegro.
Acknowledgement: We would like to thank Prof. Vlado Matevski and dr. Ljiljana Topalić for
valuable communications about distribution of Duchesnea indica in the region.

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LITERATURE
H A DŽ I A B L A H O V I Ć, S. 2006: Floristic and chorological additions to the vascular
flora of Montenegro, Proceedings of the II International Symposium of Ecologists
of the Republic of Montenegro, p. 93-101
J A C K O W I A K, B. 1992: On the distribution of Duchesnea indica (Rosaceae) in
Vienna. Fragmenta Floristica et Geobotanica 37: 593-547.
J O G A N, N. ed. 2001: Materials for the Atlas of Flora of Slovenia, Centre for
Cartography of fauna and flora, Ljubljana, pp. 443
J O V A N O V I Ć, S. 1994: Ecological study of ruderal flora and vegetation of Belgrade,
Faculty of Biology, University in Belgrade, 222 pp.
L A U B E R, K. & W A G N E R, G. 2001: Flora Helvetica, Paul Haupt ed., Bern-
Stuttgart-Wien, 1615 pp.
R O H L E N A, J. 1941-1942: Conspectus Florae Montenegrinae, Preslia 20-21
P U L E V I Ć, V. 2005: Materials for the Vascular flora of Montenegro. - Special edition
of Republic institute for Nature Protetion of Montenegro, Podgorica, 218 pp.
S T E Š E V I Ć, D. & J O V A N O V I Ć S. 2005b: Contribution to the knowledge of non
indigenous flora of Montenegro, Proceedings of the Workshop devoted to 25 th
Anniversary of the Faculty of Sciences nad Mathematics, University of
Montenegro, Podgorica 8-9 September 2005, p. 65-78
S T E Š E V I Ć, D. 2006a: Gagea chrysantha (Jan) Schultes & Schultes fil. and Linaria
genistifolia (L.) Miller subsp. genistifolia two new taxon in the flora of
Montenegro. - Proceedings of II Interanational Symposium of Ecologist of the
Republic of Montenegro, Kotor 20-24. September 2006, p. 69-72
S T E Š E V I Ć, D. & J O G A N, N. 2006a: Two new neophytes in the flora of
Montenegro: Artemisia verlotiorum and Sporobolus vaginiflorus. - Natura
Montenegrina, 5: 173-175
T R I N A J S T I Ć, I. 1973: Duchesnea indica (Andr.) Focke (Rosaceae), nova
adventivna vrsta u flori Jugoslavije.- Acta Botanica Croatica 32: 261-266, Zagreb
V A L E N T I N E, D. H. 1968: Duchesnea Sm., in Tutin at al. eds. Flora Europaea vol.
2., Cambridge, p. 48
W E B E R, E. 2003: Invasive plant species of the world. A reference guide to
environmental weeds. - CABI Publishing, Wallingford, UK, 560 pp.
Received: 14.11.2007

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