Genet Resour Crop Evol (2011) 58:1263–1274
DOI 10.1007/s10722-011-9752-z
NOTES ON NEGLECTED AND UNDERUTILIZED CROPS
Morphological variability in 17 wild elephant foot yam
(Amorphophallus paeoniifolius) collections from southwest
India
Shirly Raichal Anil • E. A. Siril • S. Suhara Beevy
Received: 22 February 2011 / Accepted: 31 August 2011 / Published online: 15 September 2011
Ó Springer Science+Business Media B.V. 2011
Abstract Amorphophallus paeoniifolius (Dennst.)
Nicolson is a tuberous herb occurring in the wild and
cultivated state. Vegetative morphological characters
were studied at full foliage stage in 17 accessions of
A. paeoniifolius which include two cultivars, 15 wild
accessions and a related species, A. dubius Blume. The
first principal component (PC) accounted for 42.32%
of phenotypic variance followed by second for another
18.38%. Major traits that accounted for more vari-
ability in both PC1 and PC2 include offset shape,
cormel weight per corm, corm fresh weight, petiole
surface pattern and canopy spread. The unweighted
pair- group method with mathematical averaging
(UPGMA) clustering method revealed three principal
clusters which separated all the accessions between
Euclidean distances of 0.3–1.6. Both cluster analysis
and principal co-ordinate analysis revealed T10, a
morphotype of A. paeoniifolius var. campanulatus
(Decne.) Sivad. cultivated in Tamil Nadu and P19
accession of A. paeoniifolius var. paeoniifolius from
Karnataka as morphologically distinct, which needs
further validation on the basis of floral characters or
molecular markers and G3 from Gujarat as an
immediate ancestor of cultivated elephant foot yam.
The genotypic (GCV) and phenotypic (PCV) coeffi-
cients of variation, broad sense heritability (h2B) and
S. R. Anil
E. A. Siril (&)
S. S. Beevy
Department of Botany, University of Kerala, Kariavattom,
Thiruvananthapuram 695 581, India
e-mail: easiril@yahoo.com
genetic advance (GA) as a percent of the mean
assessed for 18 accessions revealed high heritability
estimates. A highly significant (P \ 0.01) correlation
coefficient between circumference of petiole base and
corm diameter, corm height, corm weight, east west
spread and north south spread suggests that circum-
ference of petiole and canopy spread are indicators to
corm weight and size.
Keywords Amorphophallus paeoniifolius

Genetic advance
Genotypic coefficient of variation

Multivariate analysis
Neglected and underutilized
crops
Phenotypic coefficient of variation

Plant genetic resources
Wild accessions
Introduction
The genus Amorphophallus Blume ex Decne., a
tuberous herb belonging to the family Araceae and a
paleotropical aroid comprising of more than 200
species (Grob et al. 2002; Van der Ham et al. 2005;
Sedayu et al. 2010; Jaleel Abdul et al. 2011) occurring
in Africa, Madagascar, India, continental South East
Asia, Malesia and North East Australia (Mayo et al.
1997) is one of the true marvels of nature and a true
treasure of variation (Hetterscheid and Ittenbach
1996). Striking variations can be accounted in the
spathe characters, appendix, tuber characters, petiole
and individual flowers (Hetterscheid and Ittenbach
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1996). In India, sixteen species were reported
(Sivadasan and Jaleel 2000). In the present study,
A. paeoniifolius (Dennst.) Nicolson, alone was con-
sidered for a detailed morphological characterization
mainly due to its economic importance. A. paeoniifo-
lius is chiefly valued for its edible tuber and profound
medicinal properties (Hanelt and IPK 2001; Ochse and
Van der Brink 1980). A. paeoniifolius has its distri-
bution in Madagascar eastwards via India to Malesia,
southern China, Indo-China, Polynesia, northern Aus-
tralia (Hetterscheid and Ittenbach 1996; Lebot 2009).
Nicolson (1987) has identified the cultivated form as
A. paeoniifolius Nicolson var. campanulatus (Decne.)
Sivad., commonly called elephant foot yam, with
smooth bright green petiole with white blotches and
leaflet bases not decurrent to junction of the three main
petiolules and a round corm with or without very small
cormels. The corms, young petiole (Dey 1896; Raghu
et al. 1999) and unopened inflorescence of this
cultivated form are used as vegetable. Elephant foot
yam is a rich source of carbohydrate, protein, minerals
like calcium, iron, phosphorous, vitamin A, B, C,
flavanoids, and fibre (Kay 1987; Shilpi et al. 2005).
Important medicinal attributes of Amorphophallus
include hepatoprotective, antioxidant, uterus stimu-
lating agent (Singh et al. 2011; Laderman1983). In
India, A. paeoniifolius var. campanulatus is exten-
sively cultivated in the states of Gujarat, Maharashtra,
Kerala, Tamil Nadu and Andhra Pradesh (Anonymous
1998). Wild congenerics of the cultivated A. pae-
oniifolius known as A. paeoniifolius Nicolson var.
paeoniifolius (Nicolson 1987) is characterized by
strongly muricate dark green to purplish petiole,
mottled and leaflet bases strongly decurrent to the
junction of the three main petiolules. The wild form,
A. paeoniifolius var. paeoniifolius possess powerful
therapeutic action against piles and gastro-intestinal
disorders (Raghu et al. 1999). The corms are aperient,
carminative and expectorant. The corm extract applied
externally as an irritant to treat acute rheumatism,
administered internally in the treatment of dysentery,
diarrhoea, piles, hemorrhoids and in the formulation of
indigenous medicines to cure inflammatory conditions
and ophthalmia (Drury 1873; Watt 1889; Bala et al.
2007; Purwal et al. 2011). The wild form is having
emmenagogue, antibacterial properties and the roots
extracts are used to cure abdominal colic, elephanti-
asis, skin and blood diseases, fistula, glandular swell-
ings in the neck, urinary diseases and dropsy and has
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Genet Resour Crop Evol (2011) 58:1263–1274
depressant action on central nervous system (Jayawe-
era 1981; Dey et al. 2011). The crushed seed relieves
tooth ache and the seed paste applied to relieve
rheumatic swellings (Anonymous 1998). It is used in
the folk medicine to cease tumor growth, lung
swelling asthma, vomiting, abdominal pain, piles,
hemophilic conditions, skin diseases, obesity, dyspep-
sia, debility and control intestinal worms (Kirthikar
and Basu 1987; Siraj and Balachandran 1994; Nadk-
arni and Nadkarni 2000; Prajapathi et al. 2004).
Elsewhere this taxon was referred as A. paeoniifolius f.
sylvestris Backer with rough petiole (Hanelt and IPK
2001).
Wild relatives of crop plants are considered as
valuable resources in the crop improvement pro-
grammes. Traditionally morphological and phenolog-
ical characteristics are used to develop quantitative
estimates of genetic similarities and relationships
between the wild and cultivated relatives (Mac Key
1988). Despite the custom use of molecular markers in
the recent years, morphological descriptor in genetic
diversity analysis is still worthwhile. The multivariate
analysis and in particular principal component and
cluster analysis have been an important strategy for
characterization, evaluation and classification of plant
genetic resources, especially when large numbers of
accessions are to be assessed for several characters
(Peeters and Martineelli 1989).
Earlier classifications of family Araceae was based
on striking floral characters (Nicolson 1982). How-
ever, other workers (Mondal 2005; Sharma 2009)
argued for equal emphasis to both vegetative and
reproductive characters as both of them are subjected
to different types of selection pressures. Nevertheless,
no serious attempts have been made till now in
designing a valid key for identifying various species
of Amorphophallus based on vegetative morphological
characters with few exceptions (Hetterscheid and
Ittenbach 1996). The present study aimed to analyze
the vegetative morphological variation in A. paeoniifo-
lius and to derive their evolutionary relationships.
Materials and methods
A comprehensive collection of A. paeoniifolius germ-
plasm was made due to the continuous field explora-
tion in Western Ghats and also through National
Bureau of Plant Genetic Resources (NBPGR, New
Genet Resour Crop Evol (2011) 58:1263–1274
Delhi, India). The collections were maintained in the
Botanic Garden, Department of Botany, Kariavattom,
Thiruvananthapuram, Kerala, India. The collections
consisted of 15 accessions of A. paeoniifolius var.
paeoniifolius, two accessions of A. paeoniifolius var.
campanulatus and one accession of A. dubius Blume.
(Table 1). An important common feature noticed in
two A. paeoniifolius varieties and two different species
selected in the present study are long styled flowers
(Nicolson 1987). On the basis of features like petiole
surface pattern, habit and presence of large number of
globose cormels, A. dubius is regarded as closely
allied to A. paeoniifolius var. campanulatus (Hooker
1894; Gamble 1967) thus, included in the present
study. Further A. dubius was elevated as synonym of
A. paeoniifolius elsewhere (Hetterscheid and Itten-
bach 1996). Among cultivated form (A. paeoniifolius
var. campanulatus), ‘Gajendra’—a popular high
yielding non-acrid variety (Shirly and Palaniswami
2008) released for the cultivation in India and another
local cultivar namely ‘karunaikizhangu’ were consid-
ered for the present study. Specific codes were
allocated to each accession (Table 1).
Observations on 30 vegetative qualitative traits
(Table 2) and 14 quantitative traits were scored at full
foliage stage of all the accessions mainly based on
descriptors developed by National Bureau of Plant
Genetic Resources for elephant foot yam (Abraham
et al. 2006) with few additional characters as appro-
priate to wild collections. The quantitative traits were
recorded on the following; breadth of largest leaflet
(BLLFT), breadth of smallest leaflet (BSLFT), cormel
weight per corm (CRLWTC), corm diameter (CDM),
cormel length (CRLL), cormel number (CRLN), corm
fresh weight (CFW), corm height (CHT), east west
spread of canopy (EWSPD), north south spread
(NSSPD), length of petiole (LOP), circumference of
petiole base(GOP), length of largest leaflet (LLLFT)
and length of smallest leaflet (LSLFT).
The data on both qualitative and quantitative traits
were recorded on ten plants in each accession. The
observations in a complied form encompasses mean of
3 years data from 2008–2010. Multivariate analysis
was performed by numeric taxonomic techniques
using the procedure of principal component analysis
(Sneath and Sokal 1973). To bring out the patterns of
similarity and dissimilarity, data was subjected to
cluster analysis based on UPGMA method to group the
18 accessions. Principal coordinate analysis (PCoA)
1265
followed by construction of scatter plots based on
ordination of accessions was performed. Morphomet-
ric analyses were performed using Multivariate Sta-
tistical Package MVSP Version 3.1 (Kovach
computing Services, Wales, UK).
In addition to the above 14 quantitative traits,
lengths of three main petioles (LP1, LP2 and LP3)
were considered for analysis of variance (ANOVA)
using SPSS 7.5 (SPSS Inc. Chicago, IL, USA). In
order to determine relatedness of various traits con-
sidered in the present investigation, Pearson’s multiple
correlation matrix was plotted. In addition to these by
using data on quantitative traits, variance components,
coefficient of variation, heritability and genetic
advance (GA) were determined (Johnson et al. 1955).
Results and discussion
Principal component analysis
The first principal component (PC) accounted maxi-
mum variation (Table 3). The first PC had cormel
development, cormel shape, petiole surface pattern,
leaflet margin, cormel number, cormel weight per corm,
cormel length, corm fresh weight, East–West spread and
petiole length as traits with the highest loadings.
However, cormel development, cormel shape, cormel
weight per corm, cormel length and cormel number had
positive values. Therefore PC1 distinguished accessions
based on distinct corm and cormel characters.
The second component accounted for 18.38% of the
variability with dominance of corm characters such as
corm fresh weight, corm diameter and corm height as
well as foliar characters like petiole surface pattern,
east–west spread, north–south spread, circumference
of petiole and length of petiole. The third principal
component accounted for 11.84% and contributed by
cormel development, corm colour (abaxial and adax-
ial), corm flesh colour, petiole surface pattern, corm
fresh weight. Major traits that accounted for more
variability in PC1, PC2 and PC3 include cormel shape,
corm fresh weight and petiole surface pattern. The
highest loaded variables in PC1, PC2 and PC3 were
cormel weight per corm (0.66), corm fresh weight
(0.73) petiole surface pattern (0.36), respectively.
Hence corm and cormel characters and foliar charac-
ters especially petiole surface pattern are important in
distinguishing various accessions of A. paeoniifolius.
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1266 Genet Resour Crop Evol (2011) 58:1263–1274

Table 1 Details of collection of A. paeoniifolius from various localities

Sl. no Accession Place of Genetic group/species Status wild
code collection*/origin or cultivated

1. V1 Vittal, DK A. paeoniifolius var. paeoniifolius Wild

2. V4-P Vittal, DK A. paeoniifolius var. paeoniifolius Wild
3. P1 Puttur, DK A. paeoniifolius var. paeoniifolius Wild
4. P5 Puttur, DK A. paeoniifolius var. paeoniifolius Wild
5. K4 Thrissur, KL A. paeoniifolius var. paeoniifolius Wild
6. P17 Puttur, DK A. paeoniifolius var. paeoniifolius Wild
7. P18 Puttur, DK A. paeoniifolius var. paeoniifolius Wild
8. P19 Puttur, DK A. paeoniifolius var. paeoniifolius Wild
9. T1 Vithura, TVM, KL A. paeoniifolius var. paeoniifolius Wild
10. T10 Nagercoil, TN A. paeoniifolius var. campanulatus Cultivated
11. A.d NBPGR A. dubius Wild
12. K3-1 Thrissur, KL A. paeoniifolius var. paeoniifolius Wild
13. K3-2 Thrissur, KL A. paeoniifolius var. paeoniifolius Wild
14. K3-3 Thrissur, KL A. paeoniifolius var. paeoniifolius Wild
15. K5 Thrissur, KL A. paeoniifolius var. paeoniifolius Wild
16. G3 Navasari, GUJ A. paeoniifolius var. paeoniifolius wild
17. GJ (Gajendra) Kovvur, AP A. paeoniifolius var. campanulatus (National variety) cultivated
18. M1 Moozhiyar, PTA, KL A. paeoniifolius var. paeoniifolius Wild
* DK Dakshina Kannada, Karnataka, PTA Pathanamthitta, Kerala, GUJ Gujarat, AP Andhra Pradesh, TVM Thiruvananthapuram, TN
Tamilnadu

Cluster analysis distinct cormels and smooth petioles with identical

UPGMA cluster analysis with 44 variables revealed
three principal clusters which separated all the acces-
sions at a Euclidean distance of 1.6 (Fig. 1). In the first
principal cluster a morphotype of A. paeoniifolius var.
paeoniifolius from Navasari, Gujarat (G3)—a wild
accession, clustered with A. paeoniifolius var. cam-
panulatus cv. ‘Gajendra’ at a distance of 1.5. These
were the most distantly placed among all the acces-
sions. Clustering of these accessions was due to large
corm size and canopy spread. The cultivar ‘Gajendra’
has smooth petiole and massive habit with large corm
with few cormels where as G3 accession had highly
muricate or scabrous petiole, massive habit, and large
corm with many cormels (5–20) which indicates that
G3 may be an escape from cultivated type or the
former may be a wild ancestor. Both had light green
petiole with white blotches. G3 may be an immediate
ancestor of cultivated elephant foot yam and the large
distance between GJ and G3 may be due to the long
period of evolution through selection .
In the second principal cluster A. dubius clustered
with T10 (‘karunaikizhangu’) due to the presence of
petiole surface pattern (light green with white
blotches), but are distant by a Euclidean distance of
1.19. The local cultivar T10 had long cormels on a
small corm. This might have evolved from A. dubius
due to homology in corm and cormel pattern. A. dubius
has distinct globose cormels. However, accession T10
with many long cormels having medicinal properties
was thus selected for cultivation. G3 and A. dubius in
the above two clusters may be the primitive ones as
they are found in the wild state and cultivated ones
might have evolved from that after many cycles of
selection.
All the remaining wild accessions constituted the
third cluster which confirms the varietal status of
A. paeoniiflius var. paeoniifolius. The third principal
cluster may be divided into two groups, the first group
being T1, K3-1, K3-2 and M1 with K3-2 and T1
standing well apart. They clustered together due to
rare occurrence of cormels in these accessions, but K3-
2 has a distinct petiole pattern (Table 2) and T1 having
another pattern (Table 2). Accession T1 remained
apart within this cluster at a Euclidean distance of 0.9.
T1 is distinguished from the other accessions of this

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Genet Resour Crop Evol (2011) 58:1263–1274 1267

Table 2 List of qualitative variables evaluated for the systematics of A. paeoniifolius

Ref. Character description Descriptor states
no

1. Habit 1-small, 2-massive

2. Size of corm and cormel 1-cormels very small, 2-cormels more prominent
3. Corm shape 1-sub globose, 2-flat globose, 3-depressed globose, 4-saucer-shaped,
5-vertically elongate, 6-irregularly subglobose
4. Offset development 1-absent, 2-seasonal, 3-gradual
5. Offset shape 1-spindle shaped, 2-globose, 3-conical, 4-rhizomatous, 5-elongate,
6-spindleshaped elongate
6. Corm surface 1-rough, 2-smooth
7. Colour of corm surface (upper) 1-black, 2-dark brown, 3-white
8. Colour of corm surface (lower) 1-black, 2-dark brown, 3-white
9. Hairiness of corm 1-pubescent 2-glabrous
10. Colour of corm flesh 1-pale greenish yellow, 2-light yellowish green, 3-light yellow, 4-yellowish
white
11. Foliar phenophase 1-lasting one growing season, 2-rarely long lasting
12. Petiole shape 1-terete, 2-angular
13. CS of the upper portion of petiole near the 1-trigonal, 2-triangular, 3-terete
junction of primary partition
14. Petiole nature 1-smooth, 2-scaberulous, 3-scabrid, 4-scabrous, 5-hairy
15. Petiole colour 1-unicolorous, 2-variously blotched
16. Petiole surface pattern 1-light green with white blotches, 2-dark green with white blotches, 3-Blackish
or blackish green (unicolorous), 4-grey coloured or flesh coloured with dark
green or black small patches and white dots or various combinations of these,
5-Green with dark green long dots which are fused or separate to give a
blackish green appearance, 6-Green with dark green long dots which are
fused or separate to give a blackish green appearance and white irregular
small spots which are not fused, 7-Flesh coloured with dark green long dots,
8-Green with brown spots sometimes fused and separate, 9-Light brownish
with dark brownish patches, 10-Blackish with grayish white tinch or irregular
mixing in between giving a blackish overall appearance, 11-Light green with
white dots, 12-Ivy green with white stripes and/or clear cut white patches.
17. Rachis nature 1-naked, 2-narrowly winged with supernumerary leaflets, 3-broadly winged
with supernumerary leaflets
18. Extent of decurrence 1-decurrent to halfway to the junction of rachises, 2-decurrent to the junction of
rachises, 3-not decurrent
19 Leaflet shape 1-obovate, 2-elliptic, 3-mostly elliptic-few obovate, 4-orbicular, 5-oblong-
elliptic-lanceolate, 6-lanceolate, 7-elliptic-ovate, 8-elliptic-obovate,
9-lanceolate-elliptic-ovate, 10-elliptic-oblong-obovate
20. Leaflet apex 1-acute, 2-acuminate, 3-acute-acuminate, 4-cuspidate, 5-acuminate-cuspidate
21. Leaflet margin 1-entire, 2-erose, 3-undulate (wavy), 4-slightly wavy
22. Leaflet appearance (abaxial) 1-shiny, 2-not shiny (glabrous)
23. Leaflet appearance (adaxial) 1-shiny, 2-not shiny
24. Leaflet colour (abaxial) 0-vivid yellowish green, 1-strong olive green, 2-deep bluish green, 3-strong
olive green with reddish margin, 4-deep bluish green dark green with reddish
margin, 5-brilliant yellowish green
25. Leaflet texture 1-chartaceous, 2-choreaceous
26. Corm emergence 1-regular yearly emergence, 2-shy yearly emergence or delayed
27. Petiole partition 1-primary partition alone, 2-primary and secondary partition, 3-up to tertiary
partition, 4-quarternary partition

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Table 2 continued
Ref. Character description Descriptor states
no

28. Canopy spread 0-sparse, 1-medium, 1-thick

29. Cormel number grouping 0-No cormel or rare occurrence of cormel;1- \ 4 cormels; 2-[ 4 cormels
30. Phenology-vegetative phase 1-normal season (April–Sep), 2-late (Aug–Nov)

Table 3 Principal component analysis in 18 accessions of to the presence of identical petiole pattern. The
A. paeoniifolius, Eigen values, Eigen vectors and percent of accessions P17 and K4 are least distant among all
variation accounted by the first two principal components accessions (0.3), though they are from different
(highly loaded variables in combined analysis given and sig-
localities. Both are similar in their petiole characters
nificant PC values indicated in boldface) (log10 transformed)
(Table 2), and canopy spread. The accessions V1, V4-
Variables PC1 PC2 PC3 P, K5 and K3-3 with thick canopy grouped together
Cormel development 0.260 0.040 0.087 with significant intra-clusteral distance. V4-P, K5 and
Cormel shape 0.250 0.125 20.164 K3-3 are having identical petiole pattern and thick
Colour of corm (abaxial) 0.010 -0.032 0.101 canopy whereas V1 has a separate pattern. The intra
Colour of corm (adaxial) 0.018 -0.092 0.225 clusteral distance observed within this cluster may be
Corm flesh colour -0.051 0.051 -0.160
indicative of the different stages in the evolution of
Petiole surface pattern -0.117 -0.223 0.363
A. paeoniifolius.
In the above analysis, accession T10, (‘karunaikizh-
Extent of decurrence 0.019 -0.108 -0.010
angu’) a popular cultivar in Tamil Nadu, is placed in a
Leaflet margin -0.151 -0.040 -0.010
different cluster from that of ‘Gajendra’. Both T10 and
Cormel no 0.230 0.018 -0.006
GJ are cultivated forms of A. paeoniifolius var.
CRLWTC 0.668 0.287 0.029
campanulatus and show similarity in petiole surface
CDM -0.060 0.239 0.035
pattern but T10 stands distinct from all the remaining
CRLL 0.184 0.084 -0.087
accessions due to the presence of long cormels and
CRLN 0.388 -0.007 0.233
small corm. T1 and P19 were the outliers in the third
CFW -0.209 0.735 0.261
principal cluster. The voucher specimens of these
CHT -0.049 0.212 0.050
accessions are being deposited in the Herbarium of the
EWSPD -0.134 0.166 -0.012
Department of Botany, University of Kerala .
GOP -0.103 0.203 -0.072
LLLFT -0.112 0.003 -0.016
NSSPD -0.136 0.167 -0.028 Principal co-ordinate analysis (PCoA)
LOP -0.112 0.186 -0.061
Eigen values 0.432 0.188 0.121 PCoA involving both qualitative and quantitative
Percent variation 42.32 18.38 11.84 characters considered all together (Fig. 2) gave true
Cumulative percentage 42.32 60.70 72.54 species coordination for A. dubius and cultivated
elephant foot yam ‘Gajendra’ (GJ) indicating them as
cluster due to the presence of light yellowish green separate species. The accession from Navasari, Gujarat
corm flesh colour. (G3) also stood apart as it is distinguishable from all
The second group within this principal cluster was accessions in the study due to the presence of the
the largest with 10 accessions with P19 placed well highest number of small globose cormels (16–20) in a
apart within this group at a distance of 1.1. P19 is fully matured corm. Accession T10 stood far apart
distinct from all accessions in having a massive habit due to the presence of more number of long cormels
and delayed or shy or inconsistent emergence of and the findings are in tune with the plotted cluster
vegetative phase. P1, P17, K4, P18 and P5 clustered diagram.
together because of their medium canopy with P1 The cluster and principal component analysis on
standing apart. P18 and P5 are close to each other due vegetative morphological characters revealed the

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Genet Resour Crop Evol (2011) 58:1263–1274
Fig. 1 UPGMA
phenogram based on
qualitative and quantitative
characters
Fig. 2 PCoA based on qualitative and quantitative characters
existence of variability among the investigated acces-
sions. The multivariate analysis recorded considerable
variation and similarities among the accessions,
principally due to corm or cormel characters and
petiole surface pattern. The greater importance of such
characteristics was revealed in cluster and PCoA.
1269
Cormel shape, number, length, cormel weight per
corm had positive values in PC1. At the same time
petiole surface pattern and other leaf characters were
highlighted in the PC2. However, they had negative
values. In sweet potato Veasey et al. (2007) had
reported the shape of storage root and secondary flesh
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1270
colour as important characters contributing to diver-
sity. In Smallanthus sonchifolius (Poepp. et Endl.) H.
Robins (yacon) also tuber shape is the most important
character contributing to diversity and it was proved to
be genetically determined Valentovaä et al. (2006). In
the present study corm flesh colour was an important
trait which accounted for variability in PC3. Tuber
flesh colour is an important character contributing to
diversity in genus Dioscorea (Dansi et al. 1999). In
potato, Andean folk classification system is based on
tuber characteristics along with other characters.
Petiole pigmentation has been an important aerial
vegetative descriptor in sweet potato (Veasey et al.
2007). As per PCoA, petiole surface pattern was an
important character in distinguishing the various
accessions as evident due to PC1, PC2 and PC3.
Nicolson (1987) has grouped long styled species
such as A. dubius, A. campanulatus, A. paeoniifolius
under Amorphophallus sect. Candarum. Several char-
acters have been used to segregate the species in
various parts of Asia. Unfortunately these characters
segregate separately rather than in groups making it
difficult to define the taxa (Nicolson 1987) and
suggests that muricate petiole and strongly decurrent
leaflet bases are prioritized vegetative characters
distinguishing wild A. paeoniifolius from the culti-
vated ones (Hanelt and IPK 2001). In the present study
two varieties viz., paeoniifolius and campanulatus
were clustered in two different principal clusters
except T10 which clustered with A. dubius in a
different principal cluster. In a previous systematic
analysis (Nicolson 1987) also these two varieties were
placed into different groups. Major characters attrib-
uted to this grouping in the present study was
variations in petiole surface pattern, corm and cormel
characters, corm flesh colour, leaflet characters and
canopy spread. An intra-specific variability based on
morphological characters was highlighted and they
clustered at distance between 0.26 and 1.2 which
indicated intermediate stages in the evolution of
cultivated elephant foot yam. But the morphotypes
P19, T10 may be considered as distinct taxonomic
units at least at varietal level if this morphological
characterization remains constant (Pissard et al. 2008)
and those morphological marker systems allowed
classifying the accessions and morphotypes (Arbizu
et al. 1997) but needs validation with floral characters
and molecular markers.
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Genet Resour Crop Evol (2011) 58:1263–1274
Analysis of variance
Analysis of variance performed on various quantita-
tive data showed significant (P \ 0.05) variation
among the 18 accessions studied (Table 4). Breadth
of largest leaflet was significantly (P \ 0.05) high in
K5. Cormel length was significantly high in V4
followed by K3-3. Cormel number per corm was
significantly (P \ 0.05) high in G3 followed by T10.
Corm diameter, corm fresh weight, corm height,
circumference of petiole base, EW spread, NS spread
and petiole length was significantly high (P \ 0.05) in
cultivar ‘Gajendra’, indicating the large size of the
plant. Length of largest leaflet was maximum in K3-1,
M1 and K3-2 and was significant (P \ 0.05).
The highest coefficient of variation (Table 4)
estimated for cormel weight per corm (213.3), cormel
length (112.0), cormel number (178.6) and corm fresh
weight (104.6) signify the existence of high degree of
variability with regard to these traits and the same can
be used for distinguishing the accessions. The coef-
ficient of variation was lowest for breadth and length
of largest and smallest leaflet and petiole length
indicating that leaf characters are least important in
distinguishing the accessions. However, qualitative
foliar characters have some role in distinguishing the
accessions as evident by PCA where petiole surface
pattern was one of the highly loaded variables in the
first three principal components.
The phenotypic coefficient of variation (PCV) was
slightly higher than the genotypic coefficient of
variation (GCV) for all the characters studied signi-
fying that genotypic factors exerted a reasonable effect
in estimating the variation. Though environment
influenced the expression of these characters to some
extent (Table 5) as is the case in most of the crops e.g.
West African rice (Okelola et al. 2007), the magnitude
of error variance was relatively lower than the
genotypic variance for all traits. Lower level of error
variance indicates that the genotypic component
drives the total variance for these characters. It can
be concluded that most of the variability observed in
the phenotype for different characters has more of a
genetic than a non-genetic basis. The variability due to
genotypic variance indicates considerable scope for
selection. The high heritability estimates for most of
the characters indicates that environmental factors did
not greatly affect phenotypic variation for the traits;
Genet Resour Crop Evol (2011) 58:1263–1274 1271

Table 4 Ranges, means,

Characters Range Mean SE (±) CV F values
and standard error for 17
for accessions
characters in 15 accessions
of A. paeoniifolius var. BL leaflet (cm) 3.2–8.4 5.7 0.08 20.95 45.86*
paeoniifolius, two
accessions of Leaflet (cm) 0.9–3.5 2.2 0.04 27.55 11.48*
A. paeoniifolius var. Cormel wt per corm (g) 0–90 6.0 0.94 213.3 7.99*
campanulatus and one Corm dm (cm) 3.4–22 8.4 0.23 36.79 32.67*
accession of A. dubius
Cormel length (cm) 0–10 0.8 0.09 112.0 16.01*
Cormel no. 0–20 1.9 0.24 178.6 15.76*
Corm fw (g) 15–2,000* 297.5 23.17 104.6 33.78*
Corm ht (cm) 2–12.5 5.4 0.13 32.93 32.65*
Ewspread (cm) 30–135 75.9 1.85 32.71 27.95*
GOP (cm) 3.5–17 8.6 0.22 34.86 54.42*
Llleaflet (cm) 9.9–26 15.7 0.33 28.69 50.25*
Lsleaflet (cm) 1.9–7.3 4.2 0.07 25.45 13.19*
NSSpread (cm) 30–135 75.9 1.82 32.33 31.05*
Petiole length (cm) 22–127 54.3 1.52 37.55 43.75*
LP1 (cm) 4–17 8.8 0.23 36.35 33.89*
LP2 (cm) 4–119.5 9.9 0.66 90.05 3.99*
* Significant at P \ 0.05
LP3 (cm) 5–28 12.1 0.34 38.06 28.82*
level

Table 5 Summary statistics and estimates of phenotypic coefficient of variation (PCV) and genotypic coefficient of variation
(GCV), broad sense heritability and genetic advance in 18 accessions of A. paeoniifolius for 17 quantitative traits
Characters Mean Vp Ve Vg PCV (%) GCV (%) H2B GA GA (% of mean)

BL leaflet 5.7 1.2 0.0 1.2 0.47 0.46 98 2.2 39.4

Bsleaflet 2.2 0.2 0.0 0.2 0.31 0.30 91 0.9 39.3
Cormel wt per corm 6.0 77.7 9.7 68.0 3.61 3.38 87 15.9 266.6
Corm dm 8.4 7.8 0.2 7.6 0.96 0.95 97 5.6 66.3
Cormel length 0.8 0.5 0.0 0.4 0.78 0.75 93 1.3 169.2
Cormel no. 1.9 7.3 0.5 6.8 1.97 1.91 94 5.2 279.2
Corm fw 297.5 79,528.0 2,350.4 77,177.5 16.35 16.11 97 563.8 189.5
Corm ht 5.4 2.6 0.1 2.5 0.69 0.68 97 3.2 59.4
Ewspread 75.9 484.6 17.3 467.3 2.53 2.48 96 43.7 57.6
GOP 8.6 8.0 0.1 7.8 0.97 0.96 98 5.7 66.7
Llleaflet 15.7 17.9 0.4 17.6 1.07 1.06 98 8.6 54.5
Lsleaflet 4.2 0.7 0.1 0.6 0.41 0.39 92 1.6 37.9
NSSpread 75.9 484.7 15.6 469.1 2.53 2.49 97 43.9 57.9
Petiole length 54.3 359.0 8.2 350.8 2.57 2.54 98 38.1 70.3
LP1 8.8 8.3 0.2 8.1 0.98 0.96 97 5.8 65.9
LP2 9.9 24.6 6.2 18.4 1.58 1.37 75 7.7 77.5
LP3 12.1 16.8 0.6 16.2 1.18 1.16 97 8.1 67.3

rather genetic constitution of the accessions was by cormel weight/corm, east–west spread, north -south
responsible for the variation. Maximum genotypic spread and petiole length and minimum for breadth of
variation was recorded in corm fresh weight followed smallest leaflet. Phenotypic variation also showed a

123
1272
similar trend. GCV and PCV were maximum for corm
fresh weight and cormel weight per corm and mini-
mum for breadth of smallest leaflet. However, GA as
percent of mean was high for corm fresh weight,
cormel length, cormel number and cormel weight/
corm. The high value for GCV and heritability along
with high GA% suggest that improvement is effective
through phenotypic selection and indicative of the
effect of additive genes (Pansey and Sukhatme 1985)
for these characters. High heritability associated with
moderate or low GA% observed for the remaining
characters may be attributed to the non-additive gene
action and improvement is possible through hybrid-
ization (Kamruzzahan et al. 2000).
Pearson’s correlation coefficient indicate a signif-
icant (P \ 0.01) correlation between circumference of
petiole base (GOP) and corm diameter (r = 0.793),
corm height (r = 0.753), corm weight (r = 0.773),
East–West spread (r = 0.750) and North–South
spread (r = 0.752). Similarly East–West spread
(r = 0.579) and North–South spread (r = 0.577) had
a highly significant correlation to corm weight. This
indicates that GOP and canopy spread are indicators
for the corm weight and size. Abraham et al. (2008)
has also reported thickness of petiole, corm diameter
and number of leaflets as yield attributing characters in
cultivated form of A. paeoniifolius var. campanulatus
based on GCV, PCV, heritability, genetic advance and
association studies.
In several occasions taxon identification is hard or
even impossible when only vegetative plant parts are
present Jaleel Abdul et al. (2011), which is often the
case during collection. Thus, present study suggests
that morphological characters such as cormel shape,
cormel weight per corm, corm fresh weight, corm flesh
colour and petiole surface pattern can be used for the
identification of various morphotypes of A. paeoniifo-
lius. The multivariate analysis separates two acces-
sions T10 and P19 as distinct morphotypes with at
least a varietal status. Morphological characterization
is an easily manageable tool that can be used for
delineation of wild germplasm. On the other hand
additional information about the genotype of the plant
based on molecular markers and breeding experiments
can supplement to resolve taxonomical problems in
this genus. Further, additional traits like medicinal
properties, acridity, resistance to collar rot disease
needs to be evaluated in wild germplasm. The wild
congenerics of the cultivated species harbour genetic
123
Genet Resour Crop Evol (2011) 58:1263–1274
resources of potential value, especially as the culti-
vated species are losing their ability to reproduce
sexually because of selective pressures to preferen-
tially allocate more photosynthate towards tuber
production (Singh and Gadgil 1995) and extreme
protogyny (Arakeri 1956; Sreekumari 2000). Hence
conservation of this rich diversity of wild relatives of
cultivated plants is important.
Acknowledgments The authors are thankful to Dr. (Mrs.)
Ashalatha S Nair, Professor and Head, Department of Botany,
University of Kerala for providing facilities. Thanks are due to
NBPGR, New Delhi for providing A. dubius germplasm. SRA is
thankful to the Director, Central Tuber Crops Research Institute
(Indian Council of Agricultural Research), Thiruvananthapuram,
Kerala for granting study leave.
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