Indian J. Genet., 73(3): 278-286 (2013) DOI : 10.5958/j.0975-6906.73.3.041 Morphological relationship among the species of Indian occurrence

L. K. Bharathi1*, A. D. Munshi, T. K. Behera, K. Joseph John2, K. V. Bhat3 and A. S. Sidhu4

Indian Agricultural Research Institute, New Delhi 110 012; 2Regional Station, National Bureau of Genetic Resources, Thrissur 680 656; 3National Bureau of Plant Genetic Resources, New Delhi 110 012; 4Indian Institute of Horticultural Research, Bengaluru 560 089

(Received: April 2013; Revised : June 2013; Accepted : July 2013)

Abstract basic aspects of neglected and underutilized crops Analysis of morphological data for determining the genetic hinders their development and their sustainable variation within seven Momordica species lead to the conservation. Besides, the information available on recognition of three groups. First group containing M. germplasm is scattered and not readily accessible, charantia and M. balsamina, was characterized by n = 11, i.e., only found on regional floras. The genus annual, monoecious, non tuberous roots and muricate - tubercled fruit surface. The second group comprised of M. Momordica () comprises of 60 species dioica, M. sahyadrica, M. subangulata subsp. renigera and distributed in the warm tropics of both hemispheres M. cochinchinensis which was characterized by n = 14, [1], chiefly in Africa and with about 10 species in South perennial, dioecious, tuberous tap roots and echinate-soft East Asia [2]. papillate fruit surface. The third group contained a single species M. cymbalaria and was characterized by n = 9, In India, apart from bitter gourd (Momordica perennial, monoecious, tuberous tap roots, and ribbed fruit charantia L.), the genus Momordica comprises a surface. The species accommodated in group I and II were similar in nature of arils, male flowers, seed surface, number of small and often poorly known species but symmetrical anthers, ripe fruit colour and nature of epicarp bear edible fruits esteemed for their medicinal, while Group III could be differentiated from other two groups nutritional properties and taste. Taxonomic confusion in these characters. The range of intra-specific variation, exists in the genus because of lack of familiarity with particularly for fruit and seed characters, was high in all the species studied. At inter-specific level, wide variation was taxa and widespread use of common names and is observed for all the quantitative traits studied. Extensive region specific and poorly represented in herbaria. The collection, characterization, evaluation for resistance to similarity of the common characters, taken as key to biotic/abiotic stresses and conservation of species diversity distinguish between dioecious taxa has led to widely assumes great priority in crop improvement programmes of Momordica. conflicting taxonomic treatments of this genus in South and South East Asia. In the present investigation, Key words: Momordica, morphological variation, seven taxa viz., M. charantia, M. balsamina, M. dioica, species relationship M. sahyadrica, M. subangulata, M. cochinchinensis and M. cymbalaria were studied and presented Introduction evidence on species relationships within Momordica from morphological data analysed by phenetic Morphological studies provide information which can methods. The results presented here are therefore a be used for practical plant identification and fair attempt to establish the phylogenetic relationships hypothesizing phylogenetic relationships. The limited within the Momordica species of Indian occurrence. information available on many important and frequently

*Corresponding author’s e-mail: [email protected] 1Present address: Central Horticultural Experiment Station, Aiginia, Bhubaneswar, Orissa, India Published by Indian Society of Genetics & Plant Breeding, F2, First Floor, NASC Complex, PB#11312, IARI, New Delhi 110 012 Online management by indianjournals.com August, 2013] Morphological relationship among the Momordica species 279

Materials and methods Results and discussion

Plants analyzed in this study (Table 1) were grown Qualitative assessment of accessions both under field conditions and in National Phytotron Morphological diversity of seven species of Momordica Facility (NPF) of Indian Agricultural Research Institute was studied in this investigation (Fig. 1). The findings (IARI), New Delhi. The sets of morphological of the qualitative assessment of these seven species descriptors (40 qualitative and 15 quantitative and key morphological differences observed for the characters) were selected (Tables 2 and 4) on the basis vegetative and reproductive traits have been presented of Monograph of Indian Cucurbitaceae [3] and species in Table 2. Species relationship through multivariate descriptions in the taxonomic treatment of the genus analysis on qualitative traits was established. (2). Data on qualitative and quantitative characters were analysed using NTSYS-pc (Ver. 1.80) statistical The UPGMA dendrogram revealed two main package [4]. The multivariate statistic was used for clusters (Fig. 2). Cluster I consists of M. charantia, M. establishing relationship among the taxa. The data balsamina, M. dioica, M. sahyadrica, M. subangulata matrix was used to calculate the correlation among subsp. renigera and M. cochinchinensis which had various character states scored. The correlation solitary flowers, prominent bract in male flowers, coefficient matrix was subject to eigenvectors analysis. symmetrical anthers and fleshy fruits with slimy wet The eigenvectors derived were used to extract the red arils. Cluster I was found divided into two clearly first three most informative principal components. separated sub clusters i.e. sub cluster Ia including M. Those three components were plotted in both three charantia and M. balsamina which are characterized dimentional and biplot mode in various combinations. by epigeal germination, monoecious sex expression Only the biplots of the first two most informative and annual habit with 11 bivalents of meiotic components were presented. Similarly, principal chromosomes. Although both the species (M. component analysis (PCA) was performed for data on balsamina and M. charantia) share >80% similarity 15 quantitative traits. they could be easily distinguished from each other.

Table 1. Materials of Momordica used in the study

Species Accessions used Distribution Source/collector for morphological study M. charantia 14 Tamil Nadu, Kerala, Andhra Pradesh, Karnataka, IARI, New Delhi Orissa, Madya Pradesh, Uttar Pradesh, Chattisgarh, West Bengal, Assam, Rajasthan M. balsamina 1 Rajasthan, Gujarath NBPGR (Regional station) Thrissur M. dioica 30 Western and Eastern Ghats (Tamil Nadu, Kerala, CHES, Bhubaneswar Andhra Pradesh, Karnataka, Maharashtra, Goa), Orissa, Madya Pradesh, Uttar Pradesh, Chattisgarh, West Bengal, Assam, Rajasthan M. sahyadrica 3 Western Ghats of Kerala, Karnataka, Goa, Tamil Nadu NBPGR (Regional station), and Maharashtra Thrissur M. subangulata 3 West Bengal, Assam (Kokrajar and Kamrup districts), subsp. renigera Meghalaya and Mizoram (Naga-Khasi-Jainthia and CHES, Bhubaneswar and Garo hills), Arunachal Pradesh, Nagaland, NBPGR (Regional station), Tripura and Sikkim Thrissur M. cochin- 2 Andaman islands, Orissa, West Bengal, Assam, CHES, Bhubaneswar chinensis Tripura M. cymbalaria 4 Tamil Nadu, Andhra Pradesh, Karnataka, Maharashtra S. Anbu, (Former Dean), Horticultural College and Research Institute, Periyakulam, Tamil Nadu 280 L. K. Bharathi et al. [Vol. 73, No. 3

Fig. 1. Morphological diversity of Momordica spp. A-Seeds, B-Roots, C-Leaves, D-Position of flower bract, E- Flowers showing absence/presence of petal spot, F-Stigma, G-Filament and thecae, H-Sepal, I-Petal, J-Fruit

Fig. 2. Phenogram showing relationships of Momordica species using 40 qualitative characters August, 2013] Morphological relationship among the Momordica species 281

Table 2. Qualitative characters, character states and resultant data matrix of Momordica species

Characters/species* 1 2 3 4 5 6 7 Chromosome number (n) (0= 9, 1=11, 2=14) 1 1 2 2 2 2 0 Ploidy (0. diploid, 1.tetraploid) 0 0 0 0 1 0 0 Germination (0. epigeal, 1. hypogeal) 0 0 1 1 1 1 1 Habit (0. herbaceous, 1. suffrutescent, 2. lianas) 0 0 1 1 1 2 0 Duration (0. annual, 1. perennial) 0 0 1 1 1 1 1 Breeding system (0. monoecious, 1. dioecious) 0 0 1 1 1 1 0 Anthesis (0=4-8 AM, 1=11-12 AM, 2=7-9 PM) 0 0 2 0 0 0 1 Reproduction (0. sexual, 1. Sexual & asexual) 0 0 1 1 1 1 1 Nature of roots (0. fibrous, 1. tuberous) 0 0 1 1 1 1 1 Tuberous adventitious roots (0. absent, 1. present) 0 0 0 0 1 0 0 Nature of tap root tubers (0. absent, 1. soft fleshy, 2. woody) 0 0 1 1 1 2 1 Leaf lobing (0. absent, 1. angled, 2. deeply lobed, 3. mixed occurrence) 2 2 3 3 0 2 1 Petiole umbilical glands (0. absent, 1. present) 0 0 0 0 0 1 0 Male inflorescence-morphological type (0. solitary/pseudo raceme, 1. racemose) 0 0 0 0 0 0 1 Nature of male flower bract (0. foliaceous, 1. absent/scar like) 0 0 0 0 0 0 1 Male flower bract shape (0. flat, 1. hooded, 2. absent) 0 0 1 1 1 1 2 Male flower bract position in stalk (0. below middle, 1. above middle, 2. 0 1 2 2 2 2 3 Peduncle tip, 3. absent) Floral nectarines (0. naked, 1. closed) 0 0 1 1 1 1 0 Petal spot (0. absent, 1. present) 0 0 0 0 1 1 0 Shape of receptacle tube (0. cup, 1. saucer) 0 0 0 0 1 1 0 Calyx coloration (0. non pigmented, 1. light cream yellow, 2. purple 3. black) 0 0 1 2 2 3 3 Sepal tip (0. acute, 1. obtuse) 0 0 0 0 1 0 0 Petal color (0. yellow, 1. greenish white) 0 0 0 0 0 0 1 Petal tip (0. broad obovate, 1. acute/subacute, 2. acuminate) 0 0 1 0 2 1 0 Shape of corolla leaves (0. obovate, 1. sub elliptic, 2. oblong lanceolate) 0 0 2 0 0 1 0 Corolla venation (0. less prominent, 1. embossed) 0 0 0 0 1 1 0 Filament &thecae color (0. white, 1. black) 0 0 0 0 1 1 0 Anthers (0. symmetrical, 1. asymmetrical) 0 0 0 0 0 0 1 Stigma color (0. green, 1. yellow) 0 0 1 1 1 1 0 Fruit surface (0. tubercled, 1. spiny, 2. ribbed) 0 0 1 1 1 1 2 Texture of pericarp (0. fleshy, 1. papery) 0 0 0 0 0 0 1 Nature of epicarp (0. delicate, 1. shell like) 0 0 0 0 0 1 0 Fruit shape (0. ovoid ellipsoid/oblong, 1. pyriform) 0 0 0 0 0 0 1 Fruit color at ripening (0. orange/red, 1. green) 0 0 0 0 0 0 1 Placenta color on ripening (0. red, 1. white) 0 0 0 0 0 0 1 Aril on ripening (0. slimy wet, 1. dry granular) 0 0 0 0 0 0 1 Aril color (0. orange red, 1. white) 0 0 0 0 0 0 1 Seed shape (0. sub tridentate, 1. round oval, 2. subglobose, 3. cog wheel) 0 1 2 3 3 3 1 Seed sculpture (0. present, 1. absent) 0 0 0 0 0 0 1 Seed margin (0. dented, 1. smooth) 0 0 0 0 0 0 1 *Part of the data was used by the authors in another research paper; 1 - M. charantia, 2 - M. balsamina, 3 - M. dioica, 4 - M. sahyadrica, 5 - M. s. subsp. renigera; 6 - M. cochinchinensis, 7- M. cymbalaria 282 L. K. Bharathi et al. [Vol. 73, No. 3

These species differed in the position of floral bract in of M. cochinchinensis was reported much earlier [14]. the peduncle (near to the axis/lower half in M. charantia However, M. subangulata subsp. renigera which does and upper half in M. balsamina) and anther filaments not have the glands was quoted as M. cochinchinensis (fused to give globose appearance in M. charantia while in various literatures [15-22]. it was split into lobes in M. balsamina). However, in few reports the small wild variety of bitter gourd (M. The close relationship between M. dioica and M. charantia var. muricata) has been mistakenly sahyadrica as well as M. subangulata subsp. renigera mentioned as M. balsamina [5]. Both the species and M. cochinchinensis could be easily recognized. showed greater homology in plant morphology [6] and Close affinity among these species have been reported karyomorphology [7]. based on molecular (1, 9-10); morphological [23-24] and karyomorphological [7] similarity. The specimens M. balsamina was found to be closer to M. of M. sahyadrica were placed under M. dioica at the charantia among the available Indian Momordica Central National Herbarium at Kolkata due to strong species as reported earlier [6]. However, M. charantia morphological similarity between these two species was reported to be closer to M. angolensis; M. [23]. M. subangulata subsp. renigera which is supposed balsamina, M. welwitschii [8] and M. foetida [9]. The to be an allotetraploid clustered together with M. claim might be true as M. angolensis, M. welwitschii cochinchinensis. It is in the expected line as M. and M. foetida (African species) were not included in cochinchinensis is considered as one of the putative the present study. The results clearly revealed three parents of M. subangulata subsp. renigera [24]. It was groups as was deduced earlier through molecular also evident from the PCA that (Fig. 3) the relationship markers [10] and reproductive compatibility [11], between M. charantia and M. balsamina; M. dioica and corresponded well to the recent classification [1]. The M. sahyadrica and M. subangulata subsp. renigera and clear separation of the monoecious from the dioecious M. cochinchinensis were close in PCA ordination species and close similarities within the monoecious forming distinct phylogenetic groups. species suggested that both the monoecious species (M. charantia and M. balsamina) might have evolved The second major group (cluster II) contains M. from a common ancestor and has diverged cymbalaria; which is distinguished from other morphologically from the dioecious species. Momordica species of Indian occurrence in characters like male flowers borne in short raceme, asymmetrical Sub cluster Ib includes M. dioica, M. sahyadrica, anthers, white coloured dry arils, papery epicarp and M. subangulata subsp. renigera and M. cochinchinensis smooth and shiny round non bitten seeds. M. with 14 bivalents of meiotic chromosome numbers, cymbalaria showed hypogeal germination, monoecious hypogeal germination, dioecious sex expression and sex expression, perennial habit with n = x = 9 bivalents perennial habit. M. subangulata subsp. renigera differed of chromosomes in the male gametes. M. cymbalaria from other species of cluster Ib (M. dioica, M. sahyadrica, M. cochinchinensis) with respect to its ploidy level (tetraploid), tuberous adventitious roots, extremely long fruit stalk and unlobed leaves. Petal spot was present in M. subangulata subsp. renigera and M. cochinchinensis and was absent in M. dioica and M. sahyadrica. M. cochinchinensis was remarkably distinct from other species in its growth habit (>25m long), petiolar umbilical glands, non dehiscent leathery epicarp, flat surfaced seeds and fruit size (300-600g). The flowers of M. dioica had smaller petals and did not have petal spot in their petals which are main distinguishing character from M. cochinchinensis and M. subangulata subsp. renigera [12]. The similarity of the common characters taken as key to distinguish between dioecious taxa of Momordica lead to widely conflicting treatment of this group in South and South Fig. 3. Principal component analysis ordination based East Asia [13]. Presence of umbilical glands in petiole on 40 qualitative characters of Momordica species of Indian occurrence August, 2013] Morphological relationship among the Momordica species 283 accessions were qualitatively and quantitatively M. subangulata subsp. renigera exhibited equal different from the rest of the Indian Momordica species. similarity values (67.5%) with M. dioica and M. Bharathi et al. [7, 10] highlighted its distinctness from cochinchinensis. The PCA was performed and the other Momordica species of Indian occurrence. It is scatterplot is presented in Fig. 3. It was evident from possible that M. cymbalaria originated along with other the figure that the relationship between M. charantia African species from a progenitor species different and M. balsamina, M. dioica and M. sahyadrica, and from the dioecious Momordica species of Indian M. subangulata subsp. renigera and M. cochinchinensis occurrence [10]. were close in PCA ordination forming distinct groups. PCA separated M. cymbalaria as a distinct species. Maximum similarities in qualitative traits (Table 3) were exhibited by M. charantia Vs M. balsamina Quantitative assessment of accessions (95%) followed by M. sahyadrica Vs M. dioica (87.5%), The mean, range and standard deviation for 15 M. subangulata subsp. renigera Vs M. sahyadrica quantitative traits in different Momordica species are (77.5%) and was the least in M. cymbalaria Vs M. presented in Table 4. It was evident from table 4 that cochinchinensis (17.5%). It was further observed that

Table 3. Similarity matrix values based on 40 qualitative traits between Momordica species

M. M. M. M. M. s. subsp. M. M. charantia balasamina dioica sahyadrica renigera cochinchinensis cymbalaria M. charantia 1 M. balasamina 9.500 M. dioica 5.250 5.250 1 M. sahyadrica 6.000 6.000 8.750 1 M. s.subsp. renigera 4.000 4.000 6.750 7.750 1 M. cochinchinensis 4.250 4.250 6.750 7.000 7.250 1 M. cymbalaria 3.750 3.750 3.750 4.250 2.250 1.750 1

Fig. 4. Principal component analysis ordination based on 15 quantitative characters of Momordica species of Indian occurrence 284 L. K. Bharathi et al. [Vol. 73, No. 3 SD M cymbalaria 0.77 0.07 5.69 0.37 0.52 0.02 (5.1-8.4) (2.0-2.5) (5.3-6.1) (2.7-3.6) (0.7-0.9) (2.5-3.0) (2.9-3.2) 0 SD 0.81 6.08 1.55 1.12 2.21 0.20 1.03 3.05 0.43 5.58 3.04 0.10 13.63 M. M. M 24.67 12.78 11.01 25.53 454.57 194.47 100.14 cochinchinensis (8.2-8.7) (1.1-1.5) (7.0-7.8) (1.4-2.0) (2.4-2.7) (1.3-2.2) (12.0-13.6) (10.3-11.7) (13.4-15.1) (90.5-109.8) (317.1-592.1) SD 3.81 1.14 8.42 0.36 1.35 0.17 6.96 1.65 7.38 0.54 1.61 0.35 2.34 0.71 2.53 0.17 1.79 0.47 1.05 renigera M M.subangulata 27.27 11.01 50.58 10.20 22.61 11.51 13.08 subsp. (7.0-1.7) (9.0-12.1) (7.0-11.1) (19.9-24.3) (10.2-12.3) (10.9-12.3) SD 0.39 0.51 9.83 2.48 0.94 5.79 1.88 4.22 0.27 3.75 0.35 2.21 4.04 M. sahyadrica M 5.3 16.47 11.13 29.94 35.05 (6.8-7.2) (7.8-9.1) (8.1-9.6) (16.0-16.8) (24.1-31.5) (24.1-25.2) (10.6-11.6) (27.4-31.3) (42.9-56.4) (30.5-38.1) (11.9-13.7) (21.6-29.5) SD 1.98 4.12 5.54 8.80 0.78 5.54 dioica species 13.55 10.94 16.54 (2.8-5.2) (6.7-7.2) (9.8-14.9) (5.0-10.0) MM Momordica 4.59 10.96 12.02 balsamina SD 2.68 3.79 8.56 M M. M. M. 5.36 0.76 4.23 6.53 1.59 7.22 0.46 8.54 0.64 6.96 0.81 4.18 6.65 1.08 0.87 0.10 0.56 0.49 0.10 0.48 0.09 0.61 0.21 0.34 0.05 1.72 0.30 6.12 1.18 2.33 3.32 0.96 2.35 0.38 2.05 4.02 0.81 7.38 0.27 2.26 0.39 1.25 3.62 0.51 7.02 0.36 5.97 3.15 3.42 3.86 0.79 7.02 0.32 8.84 2.09 14.2 1.15 2.81 0.32 2.98 0.41 2.18 2.31 0.33 3.41 0.45 4.53 0.08 7.57 2.58 0.96 0.20 3.77 1.13 1.37 2.54 0.70 3.39 0.25 8.40 5.90 3.92 1.22 0.22 0.99 0.64 0.05 0.74 0.04 1.27 0.25 2.03 0.20 0.71 0.07 0.68 0.06 0.66 0.57 0.08 0.63 0.03 0.67 0.07 1.62 17.17 18.82 15.08 14.47 Values in parantheses indicates range, M = Mean and SD Standard deviation (3.7-7.9) (2.3-5.5) (4.2-6.0) (4.0-7.8) (4.0-4.4) (3.4-4.1) (0.6-1.0) (0.4-0.8) (0.4-0.5) (0.4-0.8) (0.3-0.4) (1.4-2.0) (5.6-8.6) (5.1-7.6) (4.1-6.3) (1.7-2.9) (3.1-5.8) (7.1-7.7) (1.8-2.5) (3.0-4.7) (6.6-7.3) (2.2-3.4) (1.7-2.8) (3.1-3.9) (4.5-4.6) (5.8-9.4) (0.8-1.3) (0.6-0.8) (0.4-0.7) (0.5-0.7) (0.6-0.7) (1.6-1.7) (0.5-0.6) (2.5-6.4) (2.3-3.4) (3.2-3.7) (1.0-1.8) (0.5-0.7) (0.7-0.8) (1.1-1.5) (1.9-2.2) (0.6-0.8) (8.8-17.3) (7.6-28.7) (2.6-12.9) (6.1-60.3) (6.4-21.9) (4.0-23.0) (8.2-23.6) (13.2-20.3) charantia * Variation for quantitative characters in le 4. ab Internode length (cm) Leaf length (cm) Tendril length (cm) Petiole length (cm) Petiole index *-petiole length/leaf length; T Character Male flower diameter (cm) Female flower diameter (cm) Fruit length (cm) Fruit weight Fruit diameter (cm) Fruit stalk length (cm) 100-seed weight Number of seeds/fruit Seed length (cm) Seed width (cm) August, 2013] Morphological relationship among the Momordica species 285 the range of intraspecific variation, particularly for fruit study leave. The authors thank Director, NBPGR, New and seed characters, was high for all the species. At Delhi for providing the seeds of M. balsamina, M. interspecific level, wide variation was observed for all sahyadrica and Dr. S. Anbu, Professor, Horticulture the quantitative traits studied. The highest interspecific College and Research Institute, Periyakulam, diversity was observed between M. cymbalaria and TamilNadu for providing us the tubers/seeds of M. M. cochinchinensis in leaf length (2.21 to 12.78 cm), cymbalaria genotypes. internode length (3.05 to 11.01 cm), fruit weight (0.77 to 454.57 g), fruit length (2.81 to 14.2 cm) and fruit Reference diameter (0.96 to 7.57 cm). 1. Schaefer H. and Renner S. S. 2011. Phylogenetic relationships in the order and a new The PCA performed on the quantitative traits classification of the gourd family (Cucurbitaceae). revealed that the first three components accounted Taxon, 60: 122-138. for 52.27, 69.67 and 82.85% cumulative variation 2. De Wilde W. J. J. O. and Duyfjes B. E. E. 2002. progressively. The total proportion of variation explained Synopsis of Momordica (Cucurbitaceae) in South by the first three PCs, namely, 82.85% indicated high East Asia and Malaysia. Bot. Zhurn., 87: 132-148. reliability for the groupings obtained in the 3-D plot. 3. Chakravarty H. L. 1959. Monograph of Indian Figure 4 illustrates the accession distribution in the Cucurbitaceae. Rec. Bot. Surv. India, 17: 81. first three principal components and clearly showed 4. Rohlf F. J. 1992. NTSYS-PC – Numerical the separation of 57 accessions. The analysis and Multivariate Analysis System. Version 1.80. separated the accessions into four distinct groupings, Exeter Software, New York. the first one containing members of M. 5. Maurya I. B., Arvindakshan K., Sharma S. K. and cochinchinensis, second containing members of M. Jalwania R. 2007. Status of indigenous vegetables sahyadrica, third containing members of M. in southern part of Rajasthan. Acta Hort. (ISHS), 752: subangulata subsp. renigera and the fourth, grouping 193-196. Available at http://www.actahort.org/books/ of all other species accessions. The pattern of 752/752 30.htm clustering obtained with PCA is comparable to the 6. Pandey A. K., Niraj K. and Aishwarya K. 2007. UPGMA based analysis. The results further Taxonomy of the genus Momordica (Cucurbitaceae) substantiate the traits used in this analysis as relevant in India. Abstracts of Botany and Plant Biology, Joint Congress, Chicago, Illinois, 7-11 July 2007. for taxonomic classifications. Despite a number of studies including monograph, taxonomic reviews of 7. Bharathi L. K., Munshi A. D., Vinod, Behera T. K., the genus Momordica from South East Asia [2, 3, 6, Shanti C., Das A.B., Joseph J. K. and Vishalnath. 2011a. Cyto-taxonomical analysis of Momordica L. 14, 25, 26] there are no previously reported quantitative (Cucurbitaceae) species of Indian occurrence. J. studies comparing all seven Momordica species of Genet., 90: 21-30. Indian occurrence. The present study highlighted clear 8. Schaefer H. and Renner S. S. 2010. A three-genome quantitative morphological differentiation between the phylogeny of Momordica (Cucurbitaceae) suggests species. seven returns from dioecy to monoecy and recent long-distance dispersal to Asia. Mol. Phylogenet. A good representation of diversity has been Evol., 54: 553-560. assembled in M. dioica, M. subangulata subsp. 9. Ali A. M., Karuppasamy S. and Al-Hemaid F. M. renigera and M. charantia. However, other species need 2010. Molecular phylogenetic study of Luffa tuberosa extensive coverage. In the absence of any earlier (Roxb.) (Cucurbitaceae) based on internal attempt to collect and conserve this diversity, transcribed spacer (ITS) sequences of nuclear immediate steps need to be taken in this direction. ribosomal DNA and its systematic implication. Int. J. This perhaps also serves as introspection to the poor Bioinformatics Res., 2: 42-60. state of wild Momordica genepool collection and 10. Bharathi L. K., Parida S. K., Munshi A. D., Behera T. conservation in the national Agricultural Research K., Raman K. V. and Mohapatra T. 2011. Molecular System (NARS). diversity and phylogeny of Momordica spp. of Indian occurrence. Genet. Resour. Crop Evol., 59: 937-948. Acknowledgement 11. Bharathi L. K., Munshi A. D., Behera T. K., Vinod, The first author thanks Dr. S. D. Shikamany (Former Joseph J. K., Bhat K. V., Das A. B. and Sidhu A. S. 2012. Production and preliminary characterization Director, Indian Institute of Horticultural Research) and of novel inter-specific hybrids derived from Indian Council of Agricultural Research for granting Momordica species. Curr. Sci., 103: 178-186 286 L. K. Bharathi et al. [Vol. 73, No. 3

12. Bharathi L. K., Vishalnath, Naik G., Joseph J. K. 20. Ram D., Kalloo G. and Banerjee M. K. 2002. and Anbu S. 2009. Bitterless bittergourds from Popularising kakrol and kartoli: the indigenous nature’s bounty. Indian Hort., March-April: 24-25. nutritious vegetables. Indian Hort., Oct-Dec, 6-9. 13. Joseph J. K, Antony V. T. and Roy Y. C. 2007. On the 21. Rasul M. G., Hiramatzu M. and Okubo H. 2004. occurrence, distribution and taxonomy of Momordica Morphological and physiological variation in kakrol subangulata Blume ssp. renigera (G. Don) de Wilde (Momordica dioica Roxb.). J. Fac. Agr., Kyushu Univ., in India. Genet. Resour. Crop Evol., 54: 1327-1332. 49: 1-11. 14. Chakravarty H. L. 1946. Studies on Indian 22. Sanwal S. K., Kozak M., Kumar S., Singh B. and Cucurbitaceae. Indian J. agric. Sci., 16: 49. Deka B. C. 2011. Yield improvement through female 15. Patnaik B. P. and Patnaik A. 1976. Studies on homosexual hybrids and sex genetics of sweet gourd flowering, pollination, fruit set, fruit development and (Momordica cochinchinensis Spreng.) Acta Physiol. yield in Momordica cochinchinensis Spreng. Plant., 33: 1991-1996. Abstracts of Recent Advances in Plant Sciences. 23. Joseph J. K. and Antony V. T. 2007. Momordica Session 13. Vegetable crops pp. 123. sahyadrica sp.nov. (Cucucrbitaceae), an endemic 16. Shadeque A. and Baruah G. K. S. 1984. Sweet gourd: species of Western Ghats of India. Nord. J. Bot., 24: A popular vegetable of Assam. Indian Farming, 34: 539-542. 25-35. 24. Bharathi L. K., Vinod, Munshi A. D., Behera T. K., 17. Handique A. K. 1988. Hormonal induction of Shanti C., Kattukunnel J. J., Das A. B. and parthenocarpy in Momordica cochinchinensis Vishalnath. 2010. Cyto-morphological evidence for Spreng. Curr. Sci., 57: 896-898. segmental allopolyploid origin of Teasle gourd (Momordica subangulata subsp. renigera). 18. Vijay O. P. and Jalikop S. H. 1980. Production of Euphytica, 176: 79-85. parthenocarpic fruit with growth regulators in kakrol (Momordica cochinchinensis Spreng). Indian J. Hort., 25. Chakravarthy H. L. 1982. Fascicles of Flora of India 37: 167-169. II. Cucurbitaceae (Taxonomy and distribution). Rec. Bot. Surv. India, 17: 1-234. 19. Mohanty C. R., Maharana T., Tripathy P. and Senapati N. 1994. Interspecific hybridization in 26. Joseph J. K. 2005. Studies on ecogeography and Momordica species. Mysore J. Agric. Sci., 28: 151- genetic diversity of the genus Momordica L. in India. 156. Ph.D. Thesis, Mahatma Gandhi University, Kottayam, Kerala.