Diversity and Conservation of Plants and Traditional Knowledge 233-240, 2014

Edited by: S. Panda & C. Ghosh

Published by: M/s Bishen Singh Mahendra Pal Singh, Dehra Dun

DIVERSITY OF CYPSELAR FEATURES AND THEIR

TAXONOMIC SIGNIFICANCE IN THREE SPECIES OF
THE GENUS CENTAUREA, TRIBE CARDUEAE OF
ASTERACEAE
Bidyut Kr. Jana and Sobhan Kr. Mukherjee
Department of Botany, University of Kalyani, Kalyani-741235,
West Bengal, India.
E-mail: janabidyutkumar@yahoo.com, sobhankr@gmail.com
Detailed studies on mature cypselas of 3 species belonging to genus Centaurea
Linnaeus such as C. aspera Linnaeus, C. scabiosa Linnaeus and C. stoebe
Linnaeus in the tribe Cardueae have been studied to show the difference of
morphological and anatomical features at the species level. Morphological
features of the apical part, surface hairs, location of vascular trace, structure
of carpopodium and pappus bristles of cypselas are valuable taxonomically.
Anatomically, testal features are more important than the pericarp in the
tribe Cardueae and have potential value for characterization of taxa. On the
other hand, the structure of endosperm cannot be treated as significant
taxonomic parameter. An artificial key has been provided on morphological
and anatomical features of cypselas for the identification of species.
Key words: Asteraceae; Cypselar morpho-anatomy; Centaurea.

INTRODUCTION
From the systematic viewpoint the tribe Cardueae is very much disputive and
its tribal delimitation is also controversial. The tribe Cardueae is one of the largest
in Asteracee, contains 73 genera and over 2,360 species and the tribe shows great
variations in habit, floral morphology and cypselar anatomy but all show some
common features including the key characters of style morphology, which is
thickened below the branches of the upper region and usually provided with a
collar of hairs. According to Bentham (1873) the broadly defined tribe Cardueae
consists of four subtribes i.e. Echinopsidinae, Carlininae, Carduinae and
Centaureinae, which are followed for simplicity. Apical caruncle is present in many
genera of subtribe Carduinae. Subtribe Carduinae and mainly Centaureinae show
higher structural differentiation than the remaining subtribes, according to Wagner
(1977). A limited cladistic-morphological analysis, by Bremer (1994) supported the
monophyly of the group. Molecular analyses also proved the monophyletic nature
of Cardueae. In the classical delimitation of the tribe, Staehelina and the
Xeranthemum group were placed in Carlininae by Bremer (1994). Geographical
distribution of this tribe is mainly in Mediterranean in its widest sense (including

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Diversity of cypselar features and their taxonomic significance in three species ...

the Irano-Turanian region), with a major centre of diversification in central Asia.

The mountains of central Asia (Tian-Shan, Pamir and the Himalayas) constitute
the eastern boundary for most of the genera. The importance of cypselar
morphological or / and anatomical features has been recognized in the tribe
Cardueae by Lavialle(1912), Carlquist(1958), Chauhan (1972), Dittrich (1968) and
Singh & Pandey (1984). In this context, Roth (1977) has expressed his idea regarding
the usefulness of cypselar features "Not only is the external morphology of the
achenium very characteristic, but also its internal structure shows certain qualities
which can be used taxonomically". The present paper deals with detailed
morphological and anatomical features of mature cypselas in three species
belonging to the genus Centaurea under the tribe Cardueae with the aid of light
compound microscope. The main objective of the study is to show that the diacritical
features of cypselas can be used as significant taxonomic parameter for isolation
of each taxon and also to provide additional data for upgradation of existing
system of classification.
Mature cypselas of 3 species were collected from the two herbaria of the
world (Table 1).
Table 1. Sources of the studied materials
Names of taxa
Centaurea aspera Linnaeus

Centaurea stoebe Linnaeus

Centaurea scabiosa Linnaeus

Sources of specimens
Botanischer Garten der Universitat
Zurich,
Zollikerstrasse 107, CH-8008 Zurich/ Switzerland.
Specimen no.- XX0BRISS-20093677
Humboldt- Universitat Zu Berlin, Institut fur
Biologie, Spezielle Botanik u. Arboretum, D 12437 Berlin, Germany. Specimen no.- 74
Humboldt- UniversitatZu Berlin, Institutfur
Biologie, Spezielle Botanik u. Arboretum, D 12437 Berlin, Germany. Specimen no.- 72

Different parts of cypselas such as structure of epicarpic cells in cypselar

surface, distribution of vascular trace through the ribs, relative thickness of ribs
and furrows, structure of pappus bristle and scales were observed and were drawn
in both compound research microscope as well as stereo dissecting binocular
microscope. Cypselas were processed following Jana & Mukherjee (2012).

RESULTS
Centaurea aspera Linnaeus
Morphology (Fig. 1A-D; 3 F-I)
Cypsela homomorphic, 4 mm x 1 mm with pappus, 3 mm x 1 mm without pappus,
white brown, stripped, obovate, slightly curved, upper portion truncate whereas
basal portion slightly tapered, spheroidal in cross sectional configuration. Surface
pubescent, ribs absent. Surface hairs ascending in orientation with the surface,
made up of body cells and basal cells. At the upper portion of cypsela stylopodium
present, inconspicuously arranged, fully immersed in the nectar. At the basal
region of cypsela carpopodium present, diameter of carpopodium same as the
base of the cypsela; cells outline visible, distinguishable from the other cells of

Bidyut Kr. Jana and Sobhan Kr. Mukherjee

235

cypsela, symmetric, ring like. Carpopodial cells thin-walled, not-pitted, horizontally

arranged in 1 row. At the upper portion of cypsela pappus present, ecoronate,
homomorphic, yellowish in colour, unequally arranged, represented by 85 - 90
barbellate-setose type of pappus bristles, arranged in (4-5) rings, persistent.
Anatomy (Fig. 2B)
Cypsela spheroidal in cross section. Ribs absent. Cypselar wall 0.05 mm and
0.04 mm wide at dorsal and lateral lobe region respectively. Pericarp thick, on an
average 0.04 mm wide, differentiated into two zones- epicarp and mesocarp.
Endocarp absent. Epicarp uni-seriate, made up of thin-walled, rectangular,compactly
arranged, parenchyma cells. Mesocarp consists of only thick-walled, compactly
arranged, irregular to ovoid parenchymatous tissue. Testa attached with pericarp,
approximately 0.06 mm thick, uniseriate, made up of horizontally placed, thickwalled, compactly arranged, pitted, palisade parenchyma cells. Endosperm persists
in mature cypsela, uniseriate, made up of horizontally placed, thick-walled,
compactly arranged, parenchyma cells. Mature embryo occupies a major part of
the cypsela; cotyledons 2 in number, arranged at right angle to the axis of cypsela,
containing 6 resin ducts (3 ducts in each cotyledon).
Centaurea scabiosa Linnaeus
Morphology (Fig 1E-I; 3 J-L)
Cypsela homomorphic, 3 mm x 1 mm with pappus, 2 mm x 1 mm without pappus,
yellow brown, stripped, obovate, upper portion truncate whereas basal region
slightly tapered. Ellipsoidal in cross section. Surface sparsely pubescent. Surface
hair made up of body and basal cell, ascending in orientation with the surface. At
the upper portion of cypsela, stylopodium present, inconspicuously arranged,
fully immersed in the nectar. At the basal region of cypsela, carpopodium present;
diameter of it same as the base of the cypsela, cells outline visible, distinguishable
from other cells of the cypsela. Carpopodial cells thin-walled, more or less
rectangular, not pitted, uniseriately arranged. At the upper portion of cypsela pappus
present, ecoronate, homomorphic, arranged in 2-3 circle, light yellow in colour,
unequally arranged, barbellate-setose,35-45 in number, persistent.
Anatomy (Fig. 2A)
Cypsela elliptic in cross section. Ribs absent. Cypselar wall 0.04 mm and 0.05
mm wide at dorsal and lateral lobe region respectively. Pericarp thin, on an average
0.04 mm wide, differentiated into two zones- epicarp and mesocarp. Endocarp absent.
Epicarp uni-seriate, made up of thick-walled, hexagonal, compactly arranged,
parenchymatous cells. Mesocarp made up of compactly arranged, irregular-ovoid,
thick-walled parenchyma cells. Secretary duct present in mesocarpic region below
the dorsal and lateral lobe region. Vascular trace also present in mesocarpic region.
Testa attached with pericarp, approximately 0.07 mm thick, uniseriate, made up of
radially arranged, thick-walled, compactly arranged palisade parenchyma cells.
Endosperm persists in mature cypsela, uniseriate, made up of thick-walled,
hexagonal, parenchymatous cells. Mature embryo occupies a major part of the
cypsela; cotyledons 2 in number, arranged at right angle to the axis of cypsela,
containing 10 resin ducts (5 ducts in each cotyledon).

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Diversity of cypselar features and their taxonomic significance in three species ...

Centaurea stoebe Linnaeus

Morphology (Fig. 1J-L;3A-E)
Cypsela homomorphic, 4 mm x 1 mm with pappus, 3 mm x 1 mm without pappus,
yellow brown, oblanceolate, straight, upper portion truncate, whereas basal portion
slightly tapered, striated with entire margin, ellipsoidal in cross section. Surface
rough and glabrous, containing 2 lateral lobes. On the surface ribs absent. Surface
hairs absent. At the upper portion of cypsela, stylopodium present, fully immersed
in the nectary. At the basal region of cypsela, carpopodium present, diameter same
as the base of the cypsela, asymmetric, biconvex. Carpopodial cells distinguishable
from other cells of the cypsela, thin-walled, not pitted, more or less hexagonal,
arranged in 2-3 layers. At the upper surface of cypsela, pappus present,
homomorphic, light yellow, unequal, 40 -65 in number, barbellate- setose type,
arranged in 2-3 circles, persistent.

L
Fig. 1. Morphology of cypselas: A - D. Centaurea aspera. A. Cypsela; B. Upper part of
Cypsela; C. Carpopodial cella; D. Upper part of pappus bristle. E -1. Centaurea scabiosa:
E. Cypsela; F. Basal part of Cypsela; G Upper part of Cypsela; H. Upper part of pappus
bristle; I Carpopodial cells. J - L. Centaurea stoebe: E. Cypsela; F. Basal part of Cypsela; G
Upper part of Cypsela; H. Upper part of pappus bristle; I Carpopodial cells.

Anatomy (Fig. 2C)

Cypsela elliptic in cross sectiona. Ribs absent. Cypselar wall 0.1 mm and 0.08
mm wide at dorsal and lateral lobes region respectively. Pericarp thin, on an average
0.09 mm wide, differentiated into 3 zones - epicarp, mesocarp and endocarp. Epicarp

Bidyut Kr. Jana and Sobhan Kr. Mukherjee

237

uni-seriate, made up of thick-walled, quadrangular, compactly arranged, parenchyma

cells. Mesocarp Consist of thick-walled, compactly arranged, round-ovoid
parenchyma cells with small cell lumen. Endocarp made up of uniseriately arranged,
hexagonally placed, parenchyma cells.
Testa attached with pericarp, approximately 0.05 mm thick, uniseriate, made up
of radially elongated, thick walled, palisade parenchyma cells. Endosperm persists
in mature cypsela, bi-seriate. Outer layer made up of crusted layer of cells whereas
inner layer made up of uniseriately arranged, thick-walled, horizontally placed,
parenchyma cells. Discrete patches of vascular trace present in between testa and
endosperm. Mature embryo occupies a major part of the cypsela; cotyledons two
in number, arranged at right angle to the axis of cypsela containing 6 resin ducts (3
ducts in each cotyledon)

DISCUSSION
All the studied species of Centaurea Linnaeus (C. aspera Linnaeus, C.
scabiosa Linnaeus, C stoebe Linnaeus) exhibit homomorphism.
Morphologically, the colour and size are less diagnostic than shape of cypselas.
Similarly surface ornamentation, pappus structure and carpopodium are
valuable than stylopodium. Colour of cypsela is also variable and depends on
the maturity of cypsela. In Centaurea aspera, cypsela is white brown in colour;
whereas in C. stoebe, C. scabiosa, cypselas are yellow brown in colour. The

Fig. 2. Cross section of cypselas: A. Centaurea scabiosa; B. C. aspera; C. C.

stoebe [Abbreviations: Ep - Epicarp; Me - Mesocarp; Pa - Parenchyma; T Testa; E - Endosperm; V.T. - Vascular trace]

238

Diversity of cypselar features and their taxonomic significance in three species ...

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PLATE - 1 . Photographs of different parts of cypselas in Centaurea. Figs. A - E. C. stoebe:

A. Cypsela; B. Apical part with pappus; C. Part of surface; D. Basal carpopodium; E. Part
of pappus. Figs. F - I. C. aspera: F. Cypsela; G. Apical part with pappus; H. Part of
surface; I. Basal carpopodium. Figs. J - L . C. scabiosa: J. Basal carpopodium; K. Apical
part with pappus; L. Part of surface.

shape of cypsela is also variable but it has not been considered as an important
taxonomic parameter. Among the studied species, Centaurea aspera and C.
scabiosa shapes of cypsela are obovate, whereas in C. stoebe, cypsela is
oblanceolate. In all the studied species, phytomelanin layer is absent. In C.
scabiosa and Centaurea aspera, surface is pubescent, whereas in C. stoebe
surface is rough and glabrous. In all the studied species, stylopodium is
inconspicuous in size. All the studied cypsela, at the basal region carpopodium
exists, which is a very important taxonomic character. In Centaurea aspera
and C. scabiosa, carpopodial cells are arranged in single row, whereas in C.
stoebe, carpopodial cells are arranged in 2-3 rows. Presence of carpopodium is
an important taxonomic character to separate different taxa. Information about
the different types of abscission zone of cypselas in Asteraceae has been
presented in the work of John (1921). This zone has been recognized as 'callus
or podocarp' by Robinson (1913), 'carpopod' by Blake (1918), 'separation

Bidyut Kr. Jana and Sobhan Kr. Mukherjee

239

tissue' by Roth (1977), 'mechanical tissue' by Jefferey (1987), or as

'carpopodium' by many authors after Mattfeld i.e., Robinson & Brettell (1973),
Robinson & King (1977), Wetter (1983), Jeffrey' (1992) and Short etal (1989).
So carpopodial features have definite systematic value for characterization of
taxa. At the upper portion cypsela possesses pappus represented by barbellatesetose type of bristles. Pappus structure is also a very important character
from taxonomic view point. Mukherjee & Nordenstam (2008) have mentioned
the diversity of pappus structure in some tribes of Asteraceae.
In all the studied species, anatomical structures are also important. In all the
studied species of Centaurea (C. aspera, C. stoebe and C. scabiosa) epicarp zone
is made up of uniseriately arranged parenchyma cells. In all the studied species,
mesocarp is made up of compactly arranged, thick-walled, parenchyma cells. In C.
scabiosa, resin duct is found in mesocarpic region. In other studied species, resin
ducts are absent in mesocarpic zone. Presence of resin cavity, within the pericarp in
many members of the tribe Cardueae has been reported by Lavialle(1912). Testal
features are also an important taxonomic parameter. In all the studied species, testal
layer is represented by thick-walled, palisade parenchyma like cells. It is an important
feature in the tribe Cardueae. Number of resin ducts in each cotyledon is also variable
from 6-10. From the cypselar viewpoint it has been indicated that C. scabiosa is to
some extent advanced than the other two species of Centaurea.
From the above observations it can be concluded that cypselar morphoanatomical features have significant value for separation of taxa and also to show
the relationship among them.
Key to the studied species of Centaurea
la. Resin duct present in pericarp: number of resin duct in each cotyledon five

C. scabiosa
lb. Resin duct absent in pericarp: number of resin duct in each cotyledon three
2
2a. Cypselar surface rough and glabrous; pappus bristles arranged in 2-3 rows... C. stoebe
2b. Cypselar surface pubescent; pappus bristle arranged in 4-5 rows
C. aspera

Acknowledgements
Authors are grateful to the Curators and Directors of Botanischer Garten der
Universitat Zurich, Zollikerstrasse 107, CH-8008 Zurich / Switzerland and HumboldtUniversitat Zu Berlin, Institut fur Biologie, Spezielle Botanik u. Arboretum, D 12437 Berlin, Germany for providing authenticated identified research specimens
for our work.

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