Susy Paper Tweb_1175092 Webbia

7/25/2019 Susy Paper Tweb_1175092 Webbia

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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]


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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.

Webbia: Journal of Plant Taxonomy and Geography 5


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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

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