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Teucrium Population

Teucrium Population

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Flora 201 (2006) 108–119
Morpho-anatomical differentiation of the balkan populations of the species
Teucrium flavum
L. (Lamiaceae)
$
Branislava Lakus ˇic ´
a,
Ã
, Dmitar Lakus ˇic ´
b
, Radis ˇa Janc ˇic ´
a
, Branka Stevanovic ´
b
a
Faculty of Pharmacy, Institute of Botany, University of Belgrade, Vojvode Stepe 450, 1100 Belgrade, Serbia and Montenegro
b
Faculty of Biology, Institute of Botany and Botanical Garden
Jevremovac
, University of Belgrade, Takovska 43,11000 Belgrade, Serbia and Montenegro
Received 3 January 2005; accepted 26 May 2005
Abstract
Ecological plasticity, i.e., inter-population differentiation of the species
Teucrium flavum
was analyzed on the basisof morpho-anatomical variability of its five populations from the maquis (Cisto-Ericetea and Cisto-Micromerietea),rocky grounds (Festuco-Brometea) and rocky crevices (Asplenietea rupestris) in the Eumediterranean and sub-Mediterranean region. Univariate statistic analysis included 22 quantitative characters related to the leaf and stemanatomy and morphology. In order to establish the variability and significance of morpho-anatomical differentiation,principal component analyses (PCA), multivariate analyses of variances (MANOVA), discriminant componentsanalysis (DCA) and clustering, according to the UPMGA method based on Mahalanobius’ distances, have been done.The morpho-anatomical analysis of plants from the five distant populations confirmed that the species
T. flavum
belongs to malacophyllous xeromorphic species. It was established that the plants from all the five populationsanalyzed are distinguished by stable conservative xeromorphic characteristics. There is a difference between thepronounced xeromorphic plants belonging to Eumediterranean populations and the subxeromorphic sub-Mediterranean ones.
r
2005 Elsevier GmbH. All rights reserved.
Keywords:
Teucrium flavum
; Evergreen shrub; Morphology; Anatomy; Leaf; Indumentum
Introduction
The Mediterranean species
Teucrium flavum
L. belongsto Sect.
Chamaedrys
(Miller) Schreber (Tutin and Wood,1972), which includes about 30 taxa, at the species andsubspecies rank, distributed mostly in the MediterraneanBasin. The species range extends over the entireMediterranean Basin, from the eastern coasts of Spainacross France, Italy, Croatia, Serbia and Montenegro,Albania and Greece, to Turkey in the east, and in thenorth of Africa in Algeria, Tunisia and Morocco (Meuselet al., 1978)(Fig. 1).
T. flavum
is an evergreen, branchy, semi-woody shrub,often over 50cm tall, that belongs to the life form of suffruticose chamaephytes. By such a habit
T. flavum
markedly differs from other representatives of the sect.
ARTICLE IN PRESS
www.elsevier.de/flora0367-2530/$-see front matter
r
2005 Elsevier GmbH. All rights reserved.doi:10.1016/j.flora.2005.05.001
$
This paper represents a part of the Ph.D. thesis of B. Lakus ˇic ´,entitled ‘‘Morphological Variability and Ecological Differentiation of Species of Genus
Teucrium
L. (Lamiaceae) in Yugoslavia’’ defended atthe Faculty of Biology, University of Belgrade, in the year 2000.
Ã
Corresponding author.
E-mail addresses:
blakusic@pharmacy.bg.ac.yu (B. Lakus ˇic ´),dlakusic@bfbot.bg.ac.yu (D. Lakus ˇic ´), bstev@bfbot.bg.ac.yu (B. Stevanovic ´).
 
Chamaedrys
which might indicate the relict character of the species and its belonging to the ancient Mediterra-nean flora (Tutin and Wood, 1972). On the basis of investigation of flowers and inflorescenses of the genus
Teucrium
,Ka ¨stner (1978)introduced a new classifica-tion of the genus, in which the species
T. flavum
isincluded in Sect.
Pollium
Schreber.In the Balkan Peninsula
T. flavum
is a strictlycalciphilous plant that inhabits limestone and dolomite,at altitudes between 0 and 200m. It is an importantelement of maquis and garrigue vegetation as well as of their degradation forms in the Mediterranean and sub-Mediterranean floristic region (Lakus ˇic ´, 2000).The aim of the present study was to establish whetherthere exists a morpho-anatomical differentiation be-tween Eumediterranean and sub-Mediterranean popula-tions of the xeromorphic species
T. flavum
, bearing inmind the environmental, particularly climatic differ-ences between their habitats. Three out of five popula-tions analyzed here developed under conditions of theEumediterranean climate, on the coasts of the Adriaticand Aegean Sea. Another two populations inhabit sub-Mediterranean locations (the Montenegro hinterland, inthe canyons of Cijevna and Morac ˇa rivers), character-ized by the perhumid-sub-Mediterranean Adriatic cli-mate (Table 1).
Material and methods
Plant material and morpho-anatomical analysis
A morpho-anatomical analysis was done on plantsamples from five populations of the species
T. flavum
growing in the Adriatic (Montenegro) and Aegean(Greece) part of the Mediterranean Basin. The collectedplant material was either placed in a herbarium or fixed
ARTICLE IN PRESS
Fig. 1.
Map of distribution of 
Teucrium flavum
L.
Table 1.
Ecological characteristics of the habitats of the analyzed populationsBudva(Montenegro)Lus ˇtica(Montenegro)Peloponnesus(Greece)Canyon of Cijevna(Montenegro)Canyon of Morac ˇa(Montenegro)Biogeography Adriatic provinceof EumediterraneanregionAdriatic provinceof EumediterraneanregionAegean province of EumediterraneanregionAdriatic provinceof SubmediterraneanregionAdriatic provinceof SubmediterraneanregionVegetation Rocky crevices(
Asplenietearupestris
)Maquis (
Cisto-Ericetea
)Maquis (
Cisto-Micromerietea
)Rocky grounds(
Festuco-Brometea
)Rocky grounds(
Festuco-Brometea
)Substratum Limestone Limestone Limestone Limestone LimestoneAltitude 40m 80m 80m 200mAverage annualtemperature15–16
1
C 1516
1
C 1718.5
1
C 1016
1
C 1016
1
CAverage Januarytemperature7–8
1
C 78
1
C 8.510.5
1
C 15
1
C 15
1
CAnnualprecipitation13002300mm 13002300mm 350600mm 15003100mm 15003100mm
B. Lakus ˇic ´et al. / Flora 201 (2006) 108119 109
 
in 50% alcohol and deposited, respectively in theHerbarium of the Institute of Botany and BotanicalGarden ‘‘Jevremovac’’, Faculty of Biology, Universityof Belgrade (BEOU) and Herbarium of the Institute of Botany, Faculty of Pharmacy, University of Belgrade(HFF).
V
oucher specimens:1. Budva (Montenegro Eumediterranean): rockyvegetation (class.
Asplenietea rupestris
), at the alti-tude of 40m (
Lakusˇ ic´ , D
.,
25 May 1995, HFF – 
Fix.no. Bu 01-02
).2. Lus ˇtica – Stari Kras ˇic ´i (Montenegro – Eumediterra-nean): maquis (class.
Cisto-Ericetea
), limestone, atthe altitude of 80m (
Lakusˇ ic´ , D. & B. 2225/96,
27July 1996, BEOU, HFF – 
Fix.no. TFS
).3. Peloponnesus (Greece, Peloponnesus – Eumeditera-nean: maquis (
Stevanovic´ , V.
, May 1995, BEOU – 
Fix.no. TF
).4. Canyon Cijevna, village Dinos ˇa (Montenegro – sub-Mediterranean): rocky vegetation (class.
Festuco-Brometea
), limestone (
Lakusˇ ic´ , B., Jancˇ ic´ , R., Slav-kovska, V.,
09 July 1997, HFF – 
Fix.no. 11
).5. Canyon of Morac ˇa, Dromir (Montenegro sub-Mediterranean): rocks above the river Morac ˇa (class.
Festuco-Brometea
) (
Lakusˇ ic´ , B., Slavkovska, V.,Jancˇ ic´ , R.
12 July 1997, HFF – 
Fix.no. 67
).Anatomical analyses of leaves and stems were doneon permanent slides, prepared by the standard methodfor light microscopy. Cross-sections of the leaves (150samples) and stems (50 samples) were cut on a Reichertsliding microtome (up to 10
m
m thick). The sections werecleared in Parazone and thoroughly washed beforestaining in safranin (1% w/v in 50% ethanol) and alcianblue (1% w/v, aqueous).Epidermal peels (150 samples), for surface structuresand stomata analyses, were prepared using Jeffrey’ssolution (10% nitric acid and 10% chromic acid, 1: 1)and stained in safranin and alcian blue. All slides weremounted in Canada balsam after dehydration.Density and type of the leaf and stem hairs, as well asthe paradermal aspect of epidermal cells, were alsostudied with SEM (JOEL JSM-6460), for which thesamples were covered by gold.All morpho-anatomical measurements were donewith the Image Analyzer System Ozaria 2001 and thedata processed in the statistical package Statistica 4.5for Windows. For each of the quantitative characters, 30leaf samples and 10 stem samples were obtained fromdifferent individuals belonging to each of the fivepopulations analyzed.Twenty-two quantitative characters of the statisticalanalysis were grouped in three categories: I, Leaf anatomy characters (13); II, Leaf shape characters (4)and III, Stem anatomy characters (5).I.
Leaf anatomy characters
: (1) Height of adaxialepidermal cells; (2) thickness of palisade tissue; (3)thickness of spongy tissue; (4) height of abaxialepidermal cells; (5) number of palisade layers; (6)surface area of adaxial epidermal cells; (7) surfacearea of abaxial epidermal cells; (8) surface area of abaxial stomata; (9) number of abaxial stomata;(10) number of adaxial glandular hairs; (11)number of abaxial glandular hairs; (12) number of adaxial non-glandular hairs; (13) number of abaxialnon-glandular hair.II.
Leaf shape characters
: (14) Leaf length; (15)distance between the largest leaf width point andthe leaf top; (16) the largest width of the leaf; (17)leaf surface area.III.
Stem anatomy characters
: (18) Stem diameter; (19)stem diagonal; (20) stem cortex thickness; (21)thickness of the stem vascular cylinder; (22) stempith diameter.
Statistical analysis
For each of the quantitative characters a univariatestatistic analysis was done on the basis of the followingparameters: average value, minimum, maximum, stan-dard deviation and standard error. The significance of differences between the populations studied was estab-lished by multivariate analyses of variances (MANO-VA). The general structure of the sample variabilitywere established by Principal Component Analysis(PCA). For checking the hypothesis that the analyzedsample was composed of discrete groups, which aremorphologically differentiated one from the other, aDiscriminant Component Analysis (DCA) was done.Overall differences between the compared groups arepresented by Mahalanobius’ distances, which are usedfor clustering on the basis of UPGMA method.
Results
Leaf shape and anatomy
The leaves of 
T. flavum
are elliptically oval, being thewidest at the basal part, and rounded at the tip. The leaf margin is obtusely dentate. The leaf stalk is long. Itshould be pointed out that the leaf shape was always thesame in all the populations studied.In general, in all the populations studied the leaf length was between 12 and 27mm, whereas the leaf width ranged between 11 and 21mm. The leaf surfacearea varied between 80 and 370mm
2
.The leaf indumentum of all the plants studiedwas composed of glandular and non-glandular hairs
ARTICLE IN PRESS
B. Lakus ˇic ´et al. / Flora 201 (2006) 108–119110

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