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Phylogenetic Implications of Hemipenial Morphology in Sri Lankan Agamid Lizards

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Phylogenetic Implications of Hemipenial Morphology in Sri Lankan Agamid Lizards SCIENTIFIC CORRESPONDENCE diseases was also made (Table 2). Only tion of legendary variety Kosi of M. ar- 8. Margina, A. and Zheljazkov, V., J. Es- Kosi was found resistant in different vensis. This variety can be utilized for sential Oil Res., 1994, 6, 607–615. combinations of diseases (Figure 1 c). breeding programmes to develop multi- 9. Shukla, R. S., Singh, H. B., Kalra, A., Two varieties, namely Himalaya and ple disease-resistant varieties against five Singh, V. P. and Kumar, S., J. Med. Aromat. Plant Sci., 1999, 21, 311–315. Kalka, showed resistance to rust and leaf major fungal diseases of menthol mint in 10. Singh, A. K. and Kanuja, S. P. S., Spice spot diseases. Variety Damroo showed India. India, 2007, 20, 14–17. moderately resistant reaction to powdery mildew, leaf spot, leaf blight, and stem 1. Singh, K., Singh, S., Chand, S., Anwar, ACKNOWLEDGEMENTS. We are grateful blackening and rot disease combinations. M. and Patra, D. D., Maps dew 2007, 3, to the Director, CIMAP, Lucknow for provid- Two genotypes, S-10-11-45 and S-13-2- 35–37. ing the necessary facilities and Dr N. K. Patra, 125, were found moderately resistant to 2. Kalra, A., Singh, H. B., Patra, N. K., CIMAP Resource Centre, Udham Singh Na- gar for help with experiments and for critical rust and moderately susceptible to pow- Pandey, R., Shukla, R. S. and Kumar, S., J. Hortic. Sci. Biotechnol., 2001, 76, reading of the manuscript. We also thank Dr dery mildew and leaf spot disease. SS-15 546–548. H. P. Singh for doing statistical analysis. genotype showed susceptible reaction 3. Shukla, R. S., Chauhan, S. S., Gupta, M. against all the diseases. L., Singh, V. P., Naqvi, A. A. and Patra, Received 18 February 2008; revised accepted Based on the present study, variety N. K., J. Med. Aromat. Plant Sci., 2000, 1 August 2008 Kosi of menthol mint was identified as 22, 453–455. resistant, showing multiple disease resis- 4. Shukla, R. S., Singh, H. B., Kalra, A. and R. S. SHUKLA tance. This variety has been granted US Kumar, S., In International Conference M. ALAM* Patent (no. PP 12,426) on 26 February on Integrated Plant Disease Management BIRENDRA KUMAR for Sustainable Agriculture, IARI, New 2002. Additionally, this genotype is V. P. SINGH highly productive covering about 80% of Delhi, 10–15 November 1997, p. 448. 5. Shukla, R. S., Gupta, M. L. and Singh, the menthol mint-growing areas and is Central Institute of Medicinal and H. N., J. Mycol. Plant Pathol., 2001, 31, quantitatively commercially acceptable Aromatic Plants, 10 395–396. by the farmers and industry . The result 6. Kumar, B., Singh, H. P. and Patra, N. K., P.O. CIMAP, presented here has been also validated by Crop Improv., 2007, 34, 95–99. Lucknow 226 015, India achieving the EOAI-SOM Award 2003– 7. Patra, D. D. et al., J. Med. Aromat. Plant *For correspondence. 05 for the development and dissemina- Sci., 1998, 20, 364–367. e-mail: [email protected] Phylogenetic implications of hemipenial morphology in Sri Lankan agamid lizards While useful in differentiating some of the species). Although recent molecular the ventral sulcus; (6) lateral and medial taxa at the species level1, the morpho- analyses7,8 have shown all these genera sulcus distinct throughout the length of logy of reptile hemipenes has often been to belong to the subfamily Draconinae, each lobe; (7) length of entire organ considered to be of limited phylogenetic some forms exhibit highly derived char- greater than its width; (8) minute den- value2,3 because of remarkable diver- acters such as the rostral horns of Cera- ticulation present on calyces; (9) sulcus gence even within individual genera4–6. tophora, ovoviviparity of Cophotis and traverses apex; (10) each lobe with more Here we test for similarity between a mo- the highly developed extension of the than 11 flounces; (11) ventral sulcus with lecular phylogeny and a phylogeny based canthus rostralis in Lyriocephalus. transverse ridges; (12) transverse ridges on 15 morphological character states in Because there is no consensus on what present along more than half of length of the hemipenes of 17 of the 18 species (in constitutes plesiomorphic and apomor- the ventral sulcus; (13) calyces subequal six genera) of dragon-lizards in Sri Lanka phic states in the reptile hemipenis3, and along the entire length of the organ; (14) (Squamata: Agamidae) and show that (a) on the relative phylogenetic importance entire length of the lateral and medial molecular and morphological phylogenies of these characters, we gave equal weights sulcus with calyces; (15) calyces present show remarkable congruence, and (b) to all 15 character states used in the mor- only on the lower half of both lateral and hemipenial morphology is informative in phological analysis as follows (score ‘1’ medial sulci. resolving supra-specific relationships for ‘yes’, and ‘0’ for ‘no’, using standard We used PAUP (version 4.0 b10)10 to among these lizards. hemipenial nomenclature9; see Table 1): construct a cladogram based on these The Sri Lankan dragon-lizards include (1) hemipenis divided for more than half character states under a maximum parsi- three endemic genera, Lyriocephalus (one its length; (2) flounces present; (3) apex mony criterion (character state transfor- species), Cophotis (two species) and of each lobe divided symmetrically both mation–accelerated) and a heuristic search Ceratophora (five species), and three non- laterally and medially by sulcus; (4) sul- with stepwise addition starting tree op- endemic genera Otocryptis (two species), cus spermaticus bifurcated; (5) a fleshy tion, random stepwise addition and TBR Calotes (seven species) and Sitana (one cardioid structure present at the base of branch-swapping option. The tree was 838 CURRENT SCIENCE, VOL. 95, NO. 7, 10 OCTOBER 2008 SCIENTIFIC CORRESPONDENCE FIGURE 1 152 left unrooted. We used bootstrap analysis 91 Draco blandfordii a 115 b to determine support for the nodes (full 67 Japalura tricarinata heuristic, retaining groups with a fre- 154 Japalura variegata quency greater than 50%, 100 replicates). A single maximum parsimony tree was Otocryptis wiegmanni 155 recovered (tree length = 15). The tree 104 (Figure 1) clearly shows distinct clades Otocryptis nigristigma representing the genera recognized by both morphological and molecular phy- 143 7 Sitana ponticeriana logenies , suggesting that hemipenial 98 characters are indeed useful in diagnosing Calotes desilvai between genera. For instance, the two species of Otocryptis (bootstrap = 90), 100 five species of Ceratophora (bootstrap = Calotes ceylonensis 62) and seven species of Calotes (boot- 22 strap = 90) form monophyletic clades. The 102 90 Calotes liocephalus endemic genera Lyriocephalus and Co- (I) photis also, form a tight clade, recover- 35 94 53 Calotes liolepis 86 ing the same relationships derived in 29 7 (II) recent molecular analyses . 53 48 Calotes nigrilabris At the species level, however, hemip- enial characters were not always diagnos- 81 90 Calotes calotes tically informative. For example, within 69 the genus Ceratophora, C. stoddartii and 116 Calotes versicolor C. tennentii are indistinguishable by their 45 hemipenial morphology. Likewise, among 88 67 Calotes emma the island’s seven species of Calotes, 80 Calotes mystaceus only three (C. calotes, C. ceylonensis and 30 140 Salea horsfieldi C. versicolor) have mutually distinctive hemipenes. For example, the hemipenes 121 43 Ceratophora aspera of C. calotes and C. versicolor are indis- tinguishable from each other, but clearly Ceratophora karu 69 distinguishable from all other insular congeners by having calyces present only 44 on the lower half of both lateral and Ceratophora erdeleni 62 medial sulci; likewise C. liolepis, C. (III) liocephalus and C. desilvai are also dis- 60 Ceratophora tennenti 35 tinguishable by their hemipenial morpho- logy, while C. ceylonensis differs from 89 Ceratophora stoddartii the other six species by having the entire 51 length of lateral and medial sulcus with 90 Cophotis ceylanica calyces. 37 69 At the supra-species level, however, 104 Lyriocephalus scutatus (IV) several genera among the Sri Lankan Agamidae show synapomorphic hemip- 139 57 Aphaniotis fusca enial character states. For example, the 73 157 Gonocephalus grandis genera Otocryptis and Sitana have the 132 hemipenis divided for more than half its Bronchocela cristatella 182 length and flounces present on the Acanthosaura capra 44 hemipenial lobe. Otocryptis itself has the 140 53 Japalura flaviceps apex of each hemipenial lobe divided 163 Pseudocalotes brevipes symmetrically both laterally and medi- ally by a sulcus, and the sulcus spermati- Figure 1. Relationships of Sri Lankan agamid lizards based on phylogenies derived from (a) 7 cus bifurcated. These two character states 1041 (informative) bases of aligned mitochondrial DNA positions for 12 species, with relative branch-length indicated above each branch, and (b) 15 character states determined by morphol- are absent in its sister genus Sitana. The ogy of the hemipenes for 17 species. Drawings of hemipenes are not to scale, depict ventral state of having the sulcus traverse the view, and are centred vertically to the branches of the cladogram on right (b). Colour codes for hemipenial apex is shared by all mem- distribution of species are as follows: green, Endemic to Sri Lanka; red, Occurs in India; black, bers of the genus Ceratophora, while the Common to India and Sri Lanka; blue, Southeast Asia. Key generic synapomorphies include: (I) Sulcus spermaticus not bifurcated; (II) A fleshy cardioid structure present at the base of the ven- presence of a fleshy cardioid structure at tral sulcus, lateral and medial sulci distinct throughout length of each lobe; (III) Sulcus traverses the base of the ventral sulcus, and having hemipenial apex and (IV) Width of hemipenial organ greater than its length. Scale bar = 1 mm. the lateral and medial sulci distinct CURRENT SCIENCE, VOL. 95, NO. 7, 10 OCTOBER 2008 839 SCIENTIFIC CORRESPONDENCE Table 1.
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