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Leaf venation studies of 30 varieties ofMangifera indica L. (Anacardiaceae)
Bhavna Sharmaa, Susy Alberta*and Haresh Dhadukb
aDepartment of Botany, Faculty of Science, The M. S. University of Baroda, Vadodara, India;
bDepartment of Agriculture Botany,
B.A. College of Agriculture, Anand Agricultural University, Anand, India
(Received 22 February 2016; nal version received 2 April 2016)
Leaf morphological characters are important, especially for plant identication and classication. Leaf morphologicalcharacters are mainly used for classifying a particular genus, species and varieties, but leaf architectural patterns are notmuch studied. This study has been undertaken for identication of 30 Mangifera indica L. varieties based on venationcharacteristics. The results showed that pinnate type are common in the Kaju, Totapuri and Badshahpasand varieties,which are usually characterized by thick primary veins. Secondary veins were of camptodromous type. Minor venationwas measured up to ve orders and all varieties studies showed mbriate venation. Other qualitative characters of
secondary vein (angle of divergence, course, intersecondary vein), tertiary vein (pattern), higher order venation (course),veinlet branching, areole development, areole arrangement and areole shape were undertaken. Quantitative observationsincluded number of areoles per mm2, average size of areole per mm2, number of veinlets entering areole per mm2 andvein termination per mm2. Cluster analysis was performed to observe the relationship between the varieties. Closelyrelated varieties were found grouped in the dendrogram depending upon their hierarchy. Five clusters were formedrepresenting different vein pattern parameters.
Keywords:Cluster analysis; leaf; Mangifera indica; vein architecture
Background
Mangifera indica L. belongs to family Anacardiaceae,
which comprises about 600 species of plants that live in
temperate and subtropical countries. They are mainly
trees or shrubs with resiniferous secretory ducts in barkand foliage, plants are turpentine smelling, polygamo-
dioecious or dioecious. Fruit drupaceous or dry and
indehiscent containing one seed, epicarp thin, mesocarp
usually eshy, brous and resinous, endocarp crusta-
ceous to bony. Mango, the king of fruits, has been in
cultivation in India since prehistoric times. Evergreen
trees, andro-monoecious with functionally staminate and
bisexual owers in same inorescence.
Leaves of M. indica are simple, without stipules,
arranged alternately, petiolate and with pulvinus base.
Flowers small, 4- or 5-merous with imbricate aestivation.
The term
Leaf architecturedenotes the position and formof the elements constituting the outward expression of leaf
structure. In angiosperm families leaf architectural studies
have been conducted by many researchers, namely Hickey
(1973) and Melville (1976). These include venation pat-
tern, marginal conguration, leaf shape and gland position.
In palaeobotany, macrofossils showing leaf venation pat-
terns are extensively used to identify fossil taxa (Walther
1998). It is expected that the architectural properties of leaf
venation are related to functional aspects. The high inter-
specic variability of leaf venation patterns indicates
strong selective pressures acting on the architectural
arrangement of the conducting bundles of a leaf.
The angiosperm ora exhibit a wide range of leaf
architecture. Although foliar architecture has long been
used as a taxonomic tool, the coherent classication of
dicotyledon leaf architecture by Hickey (1973) stimu-
lated a wider interest in the subject. Leaf veins havetwo main functions; they provide physical support for
the lamina, allowing the leaf to maintain its three-
dimensional structure and orientation in space. In addi-
tion, leaf veins provide a transport system for the
movement of water and carbohydrates within the leaf
(Roth-Nebelsick et al. 2001). Veins are in continuity
with the vasculature of the stem or branch to which the
leaf is attached and form an interconnected reticulum
surrounded by ground tissues within the pane of the
leaf.
Venation patterns are strongly correlated with leaf
shapes (Dengler & Kang 2001), so must be consideredin that context. A useful summary of the terminology for
describing the leaf shape is given by Judd et al. (1999).
A typical leaf consists of a leaf blade (lamina), attached
by a petiole (stalk) to the stem. Simple leaves have a
single, connected blade. A simple leaf is called entire if
its margin (edge) forms a smooth arc, toothed if the
margin has small protrusions, and lobed if the margin is
signicantly indented, dividing the blade into distin-
guishable lobes.
This study was undertaken to categorize the relation-
ship between 30 varieties ofM. indica based upon the
vein architectural patterns.
*Corresponding author. Email:[email protected]
2016 Dipartimento di Biologia, Universit di Firenze
Webbia: Journal of Plant Taxonomy and Geography, 2016
http://dx.doi.org/10.1080/00837792.2016.1175092
mailto:[email protected]://dx.doi.org/10.1080/00837792.2016.1175092http://dx.doi.org/10.1080/00837792.2016.1175092http://www.tandfonline.com/http://www.tandfonline.com/http://www.tandfonline.com/mailto:[email protected]7/25/2019 Susy Paper Tweb_1175092 Webbia
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Materials and methods
Thirty cultivars of M. indica were collected from
Junagadh Agricultural University, Junagadh for the pre-
sent study. Mature leaves from the terminal part of the
branch were collected from 10 representative plants. The
leaves were immersed in 80% ethanol for 48
72 h withseveral changes of solvent in order to remove chloro-
phyll pigments. The leaf samples were then washed and
treated with 35% NaOH at 60C for 2436 h. The
digested leaf tissue was carefully brushed apart to obtain
the leaf skeleton. These were further hardened by treat-
ing with a saturated chloral hydrate solution for several
days, washed, dehydrated and preserved. The major
venation pattern was studied with the help of a dissect-
ing microscope. The venation pattern of minor veins was
studied by cutting small bits from the central parts of the
leaf skeletons. The terminology of Hickey (1973) is fol-
lowed for the description of leaf architecture. For statisti-
cal analysis, hierarchical cluster analysis was performed
using Minitab. Clustering analysis was performed using
average linkage between groups.
Results
In the present study, cluster analysis of 30 varieties in
M.indica was performed and the different pattern of
venation has been evaluated microscopically.
Cluster analysis
The dendrogram resulting from hierarchial cluster analy-
sis grouped the 30 varieties of M.indica in different clus-
ters (Figure 1). Five clusters were visible in the
dendrogram, each cluster with closely related varieties
depending upon the vein architecture.
Venation pattern
The major venation pattern in all varieties was the pin-
nate type with a single primary vein (midvein) serving
as the origin for the higher-order venation. The size of
the primary vein was determined by calculating the ratio
between vein widths to leaf width 100%. It was calledmassive if more than 4%; Stout, if between 2% and 4%;
Moderate, if between 1.25% and 2%; and Weak, if
< 1.25% (Hickey 1973). The highest value of primary
vein size was seen in Badshahpasand (Figure 4C), fol-
lowed by Totapuri (Figure 5B) and Kaju (Figure 5D).
They were described as moderate whereas in all the
other varieties it was weak. Here the secondary veins did
not terminate at the margin but were found upturned and
gradually diminishing apically inside the margin, con-
nected to the superjacent secondaries by a series of cross
veins without forming prominent marginal loops and ter-
med as camptodromous. In minor venation, the highest-order veins were identied up to ve orders. Marginal
ultimate venation was mbriate in all the varieties.
The course of the primary vein in all the varieties
was straight, lacking noticeable curvature or change in
course and unbranched, lacking ramications of primary
rank. The next veins were the secondaries, for which the
angle of divergence was measured between the branch
and the continuation of the source vein above the pointof branching. The varieties showed acute, obtuse and
right angles. An acute angle can be narrow, moderate or
wide. The classication (Hickey 1973) is as follows: (i)
Acute (angles < 80): Narrow < 45, Moderate 4565,
Wide 6580; (ii) Right angle or approximately so
(80100); and (iii) Obtuse (angles > 100).
Category of secondary vein in all 30 varieties was
cladodromous type, in which the secondaries freely
branch towards the margin. Course of secondary vein
was either straight without noticeable deviation;
recurved, arching basally for a portion of its course;
curved uniformly, arc smooth or gradually increasing in
the degree of curvature; or sinuous, repeated smoothchanges in the direction of curvature. In varieties Batli
(Figure 2B), Jhumakhiya 1 (Figure 2C), Sindoria
(Figure 2D), Pairi (Figure 2E), Jamadar (Figure 2G),
Rucchado (Figure 3A), Ladoo (Figure 3B), Kesar (Fig-
ure 3D), Jhumakhiya 2 (Figure 3E), Sopari (Figure 3F),
Langdo (Figure 3G), Aamabdi (Figure 3H), Neelam
(Figure 4B), Badshahpasand (Figure 4C), Rajapuri
(Figure 4G), Jahangir (Figure 5A), Jhamrukhiyo
(Figure 5C), Kaju (Figure5D), Aamirpasand (Figure 5E)
and Gajariyo (Figure 5F), the course was curved
uniformly. It was sinuous in Asadiyo (Figure 4A) and
Totapuri (Figure5B), whereas in Dudhpendo (Figure4E),Alphonso (Figure 4F) and Fazli (Figure 4F) it was
straight. Recurved course was found in Cowasji
(Figure 2A), Goto (Figure 2F), Khodi (Figure 3C) and
Desi (Figure 4D); whereas in Mulgoa (Figure 2H) it was
curved abruptly. All of the secondary veins were
unbranched except Totapuri, in which branches had one
or more secondary ramications. Secondary vein spacing
was either uniform as seen in Cowasji (Figure 2A), Batli
(Figure2B), Jhumakhiya 1 (Figure2C), Pairi (Figure2E),
Goto (Figure 2F), Rucchado (Figure 3A), Ladoo
(Figure 3B), Jhumakhiya 2 (Figure 3E), Sopari
(Figure 3F), Asadiyo (Figure 4A), Neelam (Figure 4B),
Badshahpasand (Figure 4C), Desi (Figure 4D),
Dudhpendo (Figure 4E), Alphonso (Figure 4F), Rajapuri
(Figure 4G), Jahangir (Figure 5A), Jhamrukhiyo
(Figure 5C), Aamirpasand (Figure 5E) and Gajariyo
(Figure 5F), or it can be seen increasing towards the
base, as observed in Sindoria (Figure 2D), Jamadar
(Figure 2G), Mulgoa (Figure 2H), Khodi (Figure 3C),
Kesar (Figure 3D), Langdo (Figure 3G), Fazli
(Figure 4H) and Kaju (Figure 5D). Vein spacing was
seen to decrease towards the base in Aambadi
(Figure 3H). Intermediate veins between second-order
and third-order veins, generally originating from the
medial primary vein, interspersed among the secondaryveins known as intersecondary veins. All varieties
showed composite intersecondary veins, which were
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made up of coalesced tertiary vein segments for over
50% of their length. Intersecondary veins had a width
and course similar to the secondaries, but they were usu-
ally thinner than the costal secondaries and did not reach
the margin.
The present variety had either strong intersecon-
daries, which were prominently seen in varieties Cowasji
(Figure 2A), Khodi (Figure 3C), Kesar (Figure 3D),
Sopari (Figure 3F), Aambadi (Figure 3H), Desi (Fig-
ure 4D), Dudhpendo (Figure 4E), Alphonso (Figure 4F),
Rajapuri (Figure 4G); or had weak intersecondaries, as
found in Batli (Figure 2B), Jhumakhiya 1 (Figure 2C),
Sindoria (Figure2D), Pairi (Figure2E), Goto (Figure2F),
Jamadar (Figure 2G), Mulgoa (Figure 2H), Rucchado
(Figure 3A), Ladoo (Figure 3B), Jhumakiya 2
(Figure 3E), Langdo (Figure 3G), Asadiyo (Figure 4A),
Neelam (Figure 4B), Badshahpasand (Figure 4C), Fazli
(Figure 4H), Jahangir (Figure 5A), Totapuri (Figure 5B),
Jhamrukhiyo (Figure 5C), Kaju (Figure 5D), Aamir-
pasand (Figure5E) and, Gajariyo (Figure5F).
The pattern of tertiary veins was either ramied or
reticulate. (i) Ramiedtertiary veins branch into higher
orders without rejoining the secondary veins, which
include exmedial (i.e. branching oriented towards the leaf
margin) or transverse, where branching was orientedacross the intercostal area. (ii) Reticulatetertiary veins
anastomose with other tertiary veins or with the sec-
ondary veins. They include random reticulate, in which
the angles of anastomoses vary, or orthogonal reticulate,
where the angles of anastomoses are predominantly at
right angles. Ramied pattern was seen in Cowasji
(Figure 2A), Batli (Figure 2B), Asadiyo (Figure 4A),
Badshahpasand (Figure4C), Desi (Figure 4D), Alphonso
(Figure 4F) and Jhamrukhiyo (Figure 5C). A random
reticulate pattern was seen in Jhumakhiya 1 (Figure 2C),
Jamadar (Figure 2G), Mulgoa (Figure 2H), Ladoo (Fig-
ure3B), Khodi (Figure 3C), Kesar (Figure3D), Aambadi
(Figure 3H), Neelam (Figure 4B) and Dudhpendo (Fig-
ure 4E), while it was found orthogonal reticulate in Pairi
(Figure 2E), Goto (Figure 2F), Rucchado (Figure 3A),
Jhumakhiya 2 (Figure 3E), Sopari (Figure 3F), Langdo
(Figure 3G), Jahangir (Figure5A), Kaju (Figure 5D) and
Gajariyo (Figure5F).
Higher-order venation included quaternaries and
quinternaries. The course of quaternary veins was either
relatively randomly oriented or orthogonal. Varieties
Cowasji (Figure 2A), Jhumakhiya 1 (Figure 2C), Sindo-
ria (Figure 2D), Goto (Figure 2F), Khodi (Figure 3C),
Jhumakhiya 2 (Figure 3E), Sopari (Figure 3F), Aambadi
(Figure 3H), Neelam (Figure 3B), Alphonso (Figure 4F),
Rajapuri (Figure 4G), Fazli (Figure 4H), Totapuri
(Figure 5B), Kaju (Figure5D), Aamirpasand (Figure 5E)and Gajariyo (Figure 5F) showed an orthogonal course.
Randomly oriented condition was seen in Batli
Table 2. Vein architecture features in Mangifera indica L.
Sr.no. Variety
No. of areoles permm2
Average size of areoles permm2
Veinlet entering areole permm2
Vein termination permm2
1 Cowasji 36 12 20 542 Batli 15 26.5 12 61
3 Jhumakhiya 1 13 25 15 624 Sindoria 9 22 15 845 Pairi 11 23.3 11 666 Goto 11 22.6 18 557 Jamadar 24 18 21.5 708 Mulgoa 31 14 24 649 Rucchado 8 28.6 13 8610 Ladoo 5 48 12 9811 Khodi 20 11 28 10612 Kesar 28 21 16.5 10213 Jhumakhiya 2 10 31 18 8414 Sopari 17 28 26 9015 Langdo 16 25.3 24 6316 Aambadi 9 22 29 7817 Asadiyo 12 23.6 32 81
18 Neelam 17 23.3 48 9019 Basdshahpasand 18 21 41 9220 Desi 13 27.6 36 11021 Dudhpendo 17 20.3 42 11322 Alphonso 19 18.6 56 9523 Rajapuri 15 21.5 43 9224 Fazli 18 26 58 9825 Jahangir 14 22.5 43 10226 Totapuri 19 18 56 9627 Jhamrukhiyo 22 16.5 47 10628 Kaju 18 22.5 60 9829 Aamir pasand 21 20 55 8930 Gajariyo 17 24 63 104
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(Figure 2B), Pairi (Figure 2E), Jamadar (Figure 2G),
Mulgoa (Figure 2H), Rucchado (Figure 3A), Ladoo
(Figure 3B), Kesar (Figure 3D), Langdo (Figure 3G),
Asadiyo (Figure 4A), Badshahpasand (Figure 4C), Desi
(Figure 4D), Dudhpendo (Figure 4E), Jahangir
(Figure5A) and Jhamrukhiyo (Figure 5C).
Areoles are the smallest areas of the leaf tissues sur-
rounded by veins, which taken together form a contigu-
ous eld over most of the area of the leaf. The
appearance and characteristics of these areoles are termed
as areolation (Hickey 1973). The areoles were formed by
all types of veins and the veins contributed to one or
more sides of the areole. The shape may be triangular,
quadrangular, 5 or polygonal (Table 1). The development
of areole was imperfect in Batli (Figure 2B), Jhumakhiya
1 (Figure 2C), Pairi (Figure 2E), Jamadar (Figure 2G),
Mulgoa (Figure 2H), Rucchado (Figure 3A), Ladoo
(Figure 3B), Khodi (Figure 3C), Kesar (Figure 3D),
Aambadi (Figure 3H), Neelam (Figure 4B), Badshah-
pasand (Figure 4C), Desi (Figure 4D), Rajapuri
(Figure 4G), Fazli (Figure 4H), Jahangir (Figure 5A) and
Gajariyo (Figure 5F). Well-developed areoles were seen
in Cowasji (Figure 2A), Sindoria (Figure 2D),
Jhaumakhiya 2 (Figure 3E), Sopari (Figure 3F), Langdo
(Figure 3G), Asadiyo (Figure 4A), Dudhpendo
(Figure 4E), Alphonso (Figure 4F), Totapuri (Figure5B),
Jhamrukhiyo (Figure 5C) and Aamirpasand (Figure 5E)
whereas they were incomplete in Goto (Figure 2F).
(A)
(B)
Figure 1. Dendrogram showing ve clusters and linkages between the varieties. (A) Number of clusters. (B) Distance betweencluster centroids.
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Arrangement of areole was well oriented or random type.
Veinlets are the ultimate veins of the leaf, which occa-
sionally cross the areoles to become connected distally,
they were either simple or branched. Simple veinlets
may be linear or curved. The branched ones may divide
once or twice dichotomously. The veinlets were linear inall the studied varieties except for Sindoria (Figure 2D),
Goto (Figure 2F), Jhumakhiya 2 (Figure 3E), Asadiyo
(Figure 4A), Neelam (Figure 4B), Desi (Figure 4D),
Rajapuri (Figure4G), Jahangir (Figure5A), Jhamrukhiyo
(Figure 5C) and Gajariyo (Figure 5F) in which it was
curved.
Number and size of areole, veinlets entering areoles,and vein termination are given in Table 2. Variation in
Figure 2. Cleared leaf section showing vein patterns. (A) Cowasji, well-developed areole with dichotomous branched veinlets. (B)Batli, imperfect areole with dichotomous branched veinlets. (C) Jhuamakhiya 1, imperfect areole with dichotomous branched veinlets.(D) Sindoria, well-developed areole with dichotomous branched veinlets. (E) Pairi, imperfect areole with dichotomous branchedveinlets. (F) Goto, incomplete areole with dichotomous branched veinlets. (G) Jamadar, imperfect areole with dichotomous branchedveinlets. (H) Mulgoa, imperfect areole with dichotomous branched veinlets.
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the number of areoles was noticed, ranging from 10 to
30 the highest number was observed in Cowasji (36)
and the lowest in Ladoo (5). A large number of areoles
in the variety indicates smaller areole size per mm2.
This can be seen in Cowasji (Figure 2A) and Mulgoa(Figure 2H). Highest numbers of veinlets entering
areoles were seen in Gajariyo (63) and the lowest was
found in Pairi (11). Number of vein terminations was
highest in Dudhpendo (113) and lowest in Cowasji
(54). Marginal ultimate venation was found to be
incomplete in Gajariyo but all the other varieties had ambrial vein.
Figure 3. Cleared leaf section showing vein patterns. (A) Rucchado, imperfect areole with dichotomous branched veinlets. (B)Ladoo, imperfect areole with dichotomous branched veinlets. (C) Khodi, imperfect areole with dichotomous branched veinlets. (D)Kesar, imperfect areole with dichotomous branched vein lets. (E) Jhumakhiya 2, well-developed areole with dichotomous branchedveinlets. (F) Sopari, well-developed areole with dichotomous branched veinlets. (G) Langdo, well-developed areole with dichotomousbranched vein lets. (H) Aambadi, imperfect areole with dichotomous branched veinlets.
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Cluster analysis
The dendrogram resulting from hierarchical cluster anal-
ysis grouped the 30 varieties of M. indica in different
clusters (Figure 1). Five clusters were visible in the den-
drogram, each cluster with closely related varietiesdepending upon the vein architecture.
Discussion
Leaf veins are an important aspect of leaf structure and
are responsible for both the mechanical support of
leaves and the long-distance transport of water,
nutrients and photoassimilates (Onoda et al. 2011;Malinowski 2013).
Figure 4. Cleared leaf section showing vein patterns. (A) Asadiyo. well-developed areole with dichotomous branched veinlets. (B)Neelam, imperfect areole with dichotomous branched veinlets. (C) Badshahpasand, imperfect areole with dichotomous branched vein-lets. (D) Desi, imperfect areole with dichotomous branched veinlets. (E) Dudhpendo, well-developed areole with dichotomousbranched veinlets. (F) Alphonso, well-developed areole with dichotomous branched veinlets. (G) Rajapuri, imperfect areole withdichotomous branched vein lets.(H) Fazli, imperfect areole with dichotomous branched veinlets.
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Carlquist (1961) has stated that leaves have many
features of potential taxonomic signicance. One of these
is the venation pattern (Foster & Arnot 1960). Hickey
(1973) has provided a classication of vein architecture,
in which importance of major and minor venation pat-
terns along with other features like shape, base, apex,
texture, margin etc. are specied. There is a distinct
venation pattern in each plant. Venation can be differen-
tiated depending upon the number of size classes.
According to Plymale & Wylie (1944), the rst, second
and third categories from major and the ultimate veinlets
constitute the minor venation pattern. Ummu Hani
Badron et al. (2014) reported variable patterns of leaf
venation as signicant because they can be used as
additional data, especially in the group identication of
species and also to differentiate Ficus species.
The present study reveals that the leaves in all theselected varieties ofM. indica L. were simple with the
bronchidodromous type of venation. The course of
primary veins in all the varieties was straight and
unbranched. In leaves with bronchidodromous venation,
usually there is a single primary vein serving as the ori-
gin for the higher order venation, secondary veins not
terminating at the margins, and secondaries joined
together in a series of prominent arches (Hickey 1973).
Studies on Brassicaceae showed craspedodromous or
pinnate-festooned bronchidodromous type of major
venation pattern (Rao & Inamdar 1983). In the family
Acanthaceae, the major venation patterns observed were
pinnate craspedodromous in Acanthus ilicifolius and
acrodromous in Lepidagathis trinervis (Chaudhari &
Inamdar 1984). In Coffea arabica, simple to campto-
brochidodromous type of venation was found with com-
posite intersecondary veins (Mishra et al. 2010).
Intersecondary veins are observed in some varieties.They are intermediate in thickness between those of
Figure 5. Cleared leaf section showing vein pattern. (A) Jahangir, imperfect areole with dichotomous branched veinlets. (B)Totapuri, well-developed areole with dichotomous branched veinlets. (C) Jhamrukhiyo, well-developed with dichotomous branchedveinlets. (D) Kaju, imperfect areole with dichotomous branched veinlets. (E) Amirpasand, well-developed with dichotomous branchedveinlets. (F) Gajariyo, imperfect areole with dichotomous branched veinlets.
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secondary and third-order veins. They may be either sim-
ple (as in Sindoria, Sopari, Badshahpasand and Totapuri)
or composite, as in other varieties (Table 2).
According to Levin (1929), Gupta (1962) and
Verghese (1969), the vein islet number is constant for a
species while Banerjee & Das 1972), Sehgal & Paliwal(1975) and Jain (1978) suggested that the vein ending
and vein islet termination are highly variable.
In the present study, the course of higher-order venation
in some varieties was orthogonal or relatively randomly ori-
ented. Tertiary veins arise from the secondaries, and were
relatively thinner than the secondary veins. The predomi-
nant angles of tertiary veins were acute. The pattern was
randomly reticulate, orthogonal reticulate or ramied. The
relationship of the tertiary veins to midvein was oblique
with three angles remaining approximately constant or at
right angles or longitudinal approximately parallel. The
nextner order of the veins was known as quaternary vein.
The course of orientation was either orthogonal or relativelyrandomly oriented. The veins originating from these and
those of equal size form lower orders the quinternaries.
Marginal ultimate venationthe veins at the margins
form two different patterns in different varieties. The major
portion of the marginal ultimate venation was mbriate,
where higher veins orders fused into a vein running just
inside of the margin (mbrial vein).
Minor venation patterns are considered as an impor-
tant aspect in taxonomic studies. The main function of
marginal venation in a leaf is to avoid desiccation,
therefore in coffee cultivars it has been observed that
the leaves with incompletely looped margins have anadaptive advantage in drought tolerance (Mishra et al.
2010).
According to Nelson & Dengler (1997), procambium
of the midvein (primary vein) is the rst morphologically
recognizable vein order, the secondary veins are formed in
continuity with the primary veins and appear in basipetal,
acropetal or divergent patterns, depending upon the spe-
cies. Minor vein orders are established in continuity with
the previous procambium. Studies conducted by Scarpell
et al. (2006) and Donner et al. (2009) found that auxin
ow is required to achieve normal vein patterning and vein
density.
Cluster analysis was performed using the average
linkage method. In this method, the distance between
two groups is dened as the average of the distances
between all pairs of individuals of the two groups. Five
clusters are seen in the Figure 1. Variety Jamadar and
Kesar, Rucchado and Pairi, Aamirpasand and Aambadi,
Neelam and Jamadar, Langdo and Sopari and many
others were found linked in joining clusters (Figure 1).
Conclusion
This study has signicant taxonomic importance because
the studied varieties can be separated based on veinarchitecture and vein patterns. The result can be used as
additional and supportive data for group clustering and
identication, especially for Mangifera varieties. Indeed,
similar studies are found useful in differentiated species
of Gossypium, Camellia and Coffea. It is suggested that
many other plant taxa can be extensively studied and
classied using vein architecture as supporting analysis
in species classi
cation.
Key for the identication of variety based on vein
architecture
1. Primary vein size - weak...........................................2
Primary vein size - moderate..................................28
2. Angle of divergence - right angle............................3
Angle of divergence - acute......................................9
3. Course of secondary veins - straight........................4
Course of secondary veins - curved..........................5
4. Pattern of tertiary vein - random reticulate.................
................................................................ Dudhpendo
Pattern of tertiary vein - ramied................Alphonso5. Course of secondary vein - curved abruptly...............
.........................................................................Mulgoa
Course of secondary vein - curved uniformly............6
6. Pattern of tertiary vein - reticulate orthogonal............
.......................................................................Gajariyo
Patterrn of tertiary vien - random reticulate..............7
7. Course of higher order venation - orthogonal.............
.........................................................................Neelam
Course of higher order venation - relatively
randomly oriented......................................................8
8. Veinlet branching - 4 to 5 times.................. Jamadar
Veinlet branching - 2 to 3 times.......................Kesar9. Angle of divergence - acute narrow........................10
Angle of divergence - either acute wide or acute
moderate...................................................................16
10. Course of secondary vein - sinuous..............Asadiyo
Course of secondary vein - curved or recurved......11
11. Course of secondary vein - recurved...................Desi
Course of secondary vein - curved..........................12
12. Course of secondary vein curved abruptly...Jahangir
Course of secondary vein - curved uniformly.........13
13. Pattern of tertiary vein - orthogonal reticulate........14
Patterns of tertiary vein - random reticulate............15
14. Course of higher order venation - orthogonal.............
..........................................................................Sopari
Course of higher order venation - relatively
randomly oriented...........................................Langdo
15. Areole development - imperfect .................Aambadi
Areole development - well developed.........................
...............................................................Aamirpasand
16. Angle of divergence of secondary vein - acute
wide..........................................................................17
Angle of divergence of secondary vein - acute
moderate...................................................................21
17. Course of secondary vein - straight....................Fazli
Course of secondary vein - either curved or
recurved....................................................................1818. Course of secondary vein - recurved......................19
Course of secondary vein - curved uniformly.........20
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19. Patterns of tertiary vein - ramied.................Cowasji
Patterns of tertiary vein - reticulate orthogona...Goto
20. Higher order venation - orthogonal.............Rajapuri
Higher order venation - relatively randomly
oriented..................................................Jhamrukhiyo
21. Course of secondary vein - recurved................KhodiCourse of secondary vein - curved uniformly.........22
22. Patterns of tertiary vein - ramied..........................23
Patterns of tertiary vein - reticulate.........................24
23. Patterns of tertiary vein - ramied apical............Batli
Patterns of tertiary vein - ramied exmedial...............
.......................................................................Sindoria
24. Patterns of tertiary vein - random reticulate............25
Patterns of tertiary vein - reticulate orthogonal.......26
25. Higher order venation - orthogonal....Jhumakhiya 1
Higher order venation - relatively randomly oriented
...........................................................................Ladoo
26. Higher order venation - orthogonal....Jhumakhiya 2
Higher order venation - relatively randomly oriented..................................................................................27
27. Vein termination per mm2 - 66...........................Pairi
Vein termination per mm2
- 86...................Rucchado
28. Angle of divergence of secondary vein - right angle
.......................................................................Totapuri
Angle of divergence of secondary vein - acute .....29
29. Angle of divergence of secondary vein - acute
narrow................................................Badshahpasand
Angle of divergence of secondary vein - acute
moderate .............................................................Kaju
References
Banerjee ML, Das S.1972. Minor venation pattern in the Indianacres. In: YS Murthy et al., editors, Advances in plantmorphology. Meerut, India: Sarita Prakasan; p. 5157.
Carlquist S. 1961. Comparative plant anatomy. Holt, Rinehartand Winston: New York, NY.
Chaudhari GS, Inamdar JA. 1984. Leaf architecture of someacanthaceae. Bot Mag Tokyo. 97:469481.
Dengler N, Kang J. 2001. Vascular patterning and leaf shape.Curr Opin Pl Biol. 4:5056.
Donner TJ, Sherr I, Scarpella E. 2009. Regulation of prepro-cambial cell state acquisition by auxin signaling inArabidopsis leaves. Development. 136:32353246.
Foster AS. 1972. Venation patterns in the leaves of Ephedra. J
Arn Arbor Harv Univ. 53:364
378.Foster AS, Arnot HJ. 1960. Morphology and dichotomous vas-
culature of the leaf of Kingdonia uniora. Am J Bot.47:684698.
Gupta R. 1962. Correlation of tissues in leaves. I. Absolutevein islet numbers and absolute veinlet termination num-bers. Ann Bot. 25:6570.
Hickey LJ. 1973. Classication of the architecture of dicotyle-donous leaves. Amer J Bot. 60:1733.
Inamdar JA, Mubthy GSR. 1978. Leaf architecture in someSolanaceae. Flora. 167:265272.
Jain DK.1978. Studies in Bignoniaceae III. Leaf architecture. JInd Bot Soc. 57:369387.
Judd WW, Campbell CS, Kellogg EA, Stevens PF. 1999. Plantsystematics: A phylogenetic approach. Sundrland, MA:Sinauer Associates.
Levin FA. 1929. The taxonomic value of vein islet areas basedupon a study of the genera Berosma, Cassia, Erythroxylonand Digitalis. J Pharm Pharmacol. 2:1743.
Malinowski R. 2013. Understanding of leaf developmentthescience of complexity. Plants. 2:396415.
Melville R. 1976. The terminology of leaf architecture. Taxon.25:549561.
Merrill EK. 1978. Comparison of mature leaf architecture ofthree types in Sorbus l. (Bosaceae). Bot Gaz. 139:447453.
Mishra MK, Padmajyothi D, Prakash NS, Ram AS, SrinivasanCS, Sreenivasan MS. 2010. Leaf architecture in Indian
Coffee (Coffea Arabica L.) cultivars and their adaptivesignicance. World J Fungal Plant Biol. 1:3741.Nelson T, Dengler N. 1997. Leaf vascular pattern formation.
Plant Cell. 9:1211135Onoda Y, Westoby M, Adler PB, Choong AM, Clissold FJ,
Cornelissen JH, Daz S, Dominy NJ, Elgart A, Enrico L,et al. 2011. Global patterns of leaf mechanical properties.Ecol Lett. 14:301312.
Plymale EC, Wylie RB. 1944. The major veins of mesomor-phic Leaves. Amer J Bot. 42:1827.
Rao NV, Inamdar JA. 1983. Leaf architectural studies in thebrassicaceae. Bot Mag Tokyo. 96:1528.
Roth-Nebelsick A, Uhl D, Mosbrugger V, Kerp H. 2001. Evo-lution and function of leaf venation architecture. Rev AnnBot. 87:553566.
Scarpell AE, Marcos D, Friml J, Berleth T. 2006. Control ofleaf vascular patterning by polar auxin transport. GenesDev. 20:10151027.
Sehgal L, Paliwal GS. 1975. Studies on the leaf anatomy ofEuphorbia II. Venation patterns. Bot J Linn Soc.68:173208.
Ummu HB, Talip N, Mohamad AL, Affenddi AEA and JuhariAAA.2014. Studies of leaf venation in selected taxa of thegenus Ficus L. (Moraceae) in Peninsular Malaysia. TropLife Sci Res. 25(2), 111125.
Verghese TM. 1969. A contribution to the failure venation ofScrophulariaceae. In: Choudhary KA, editor. Recentadvances in the anatomy of tropical seed plants. India:Hindustan publishing corporation, Delhi; p. 253266.
Walther H. 1998. Die tetraora von Hammerunterwiesenthal
(Freestaat Sachsen). Abhandungen des Staatiechen Muse-ums fur Mineralogie uns Geologie zu Dresden. 43(44):239264.
Yapp RH. 1912. Spiraea ulmaria L. and its bearing on theproblem of xeromorphy in marsh plants. Ann Bot. 26:815870.
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