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Reduction of Blinkworthia (Convolvulaceae) based on multilocus phylogenetic reconstruction and resurrection of a species from synonymy revealed by phenetic analyses

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Summary

Blinkworthia was established as a tropical Asian genus in the morning glory family (Convolvulaceae) with a unique combination of habit, bract, corolla and fruit morphology. Recently, however, investigations of the morphology and anatomy have shown that none of these diagnostic characters can be used to delimit Blinkworthia as an independent genus. This study resolves the generic circumscription using phylogenetic analysis of molecular data from four loci: ITS, trnL-trnF, matK and rps16. The results showed that all members of Blinkworthia were nested within Argyreia, concordant with the current morphological concept of Argyreia. All information robustly supported that Blinkworthia should be subsumed under Argyreia prompting us to make the nomenclatural transfers from Blinkworthia to Argyreia and make the new combinations A. convolvuloides and A. lycioides. In addition, phenetic analyses revealed that a species endemic to Burma, A. campanuliflora, should be resurrected from the synonymy of B. convolvuloides. The three species are circumscribed, described, illustrated and their ecology, distribution and vernacular names are summarised, all documented with specimen citations. An identification key is provided for these three closely related species.

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Acknowledgements

The authors would like to sincerely thank all members of the Burma Flora Project team — Wattana Tanming, Yumiko Baba, Jonathan Peeraman, Aye Mya Mon, and colleagues, of the Kochi Prefectural Makino Botanical Garden (Japan), the Forest Research Institute (Burma), and the Queen Sirikit Botanic Garden (Thailand) — for allowing PR to join their collecting excursion in Burma in 2016. In addition, Dr Yotsawate Sirichamorn and Watchara Arthan instructed PR in the molecular phylogenetic approaches. We wish to acknowledge the curators and staff at the herbaria BCU, BK, BKF, BM, CAL, E, G, GXMI, HITBC, IBK, IBSC, K, KUN, L, NAS, NY, P, PE, and W for facilitating herbarium specimen study. PR would also like to thank the Science Achievement Scholarship of Thailand (SAST) for research funding and financial support for this study. PT thanks the Frontier Research Grant, and Mahidol University, for research funding. Gretchen Wade, Harvard Botany Libraries, located and obtained critical literature. Heimo Rainer (W) and Avishek Bhattacharjee (CAL) provided digital images by permissions of the Directors, Naturhisorisches Museum Wien of Austria (W) and Botanical Survey of India (CAL), respectively, for a type specimen critical to making a lectotype choice. GS would like to thank Faculty of Graduate Studies, Mahidol University, for funds that enabled a visit to Mahidol University to complete this manuscript. We thank the reviewers for helping improve our manuscript.

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  1. Department of Plant Science, Faculty of Science, Mahidol University, King Rama VI Road, Bangkok, 10400, Thailand

    Pantamith Rattanakrajang, Pirada Sumanon & Paweena Traiperm

  2. Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK

    Pantamith Rattanakrajang & Timothy Utteridge

  3. Harvard University Herbaria, 22 Divinity Ave, Cambridge, MA, 02138, USA

    George Staples

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  1. Pantamith Rattanakrajang
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Appendices

Appendix 1. Accession details of samples used in phylogenetic analyses in this study.

Taxon name, source or geographic region, voucher with collector number (acronym of herbarium), GenBank accession number for ITS, trnL-trnF, matK and rps16 respectively. An asterisk (*) indicates sequences newly performed in this study; an en-dash (–) indicates missing data.

IPOMOEEAE: *Argyreia barbigera Choisy: Burma, L. S. Mang 96014 (BKF), OM699030, OM759910, OM759935, OM759960. Argyreia campanuliflora (Gage) Thoth. *(1): Burma, K. Fujikawa et al. 90125 (MBK), OM699023, OM759903, OM759928, OM759953; *(2): Burma, P. Rattanakrajang et al. 113 (QBG), OM699024, OM759904, OM759929, OM759954. Argyreia capitiformis (Poir.) Ooststr. *(1): Thailand, Mae Hong Son, P. Rattanakrajang et al. 71 (BKF), OM699033, OM759913, OM759938, OM759963; *(2): Thailand, Chiang Mai, P. Rattanakrajang et al. 80 (BKF), OM699034, OM759914, OM759939, OM759964. Argyreia caudata Ooststr.: Malaysia, see Simões et al. (2015), SAN 152877 (SAN), KP261908, KP236602, KR024892, KR025035. *Argyreia confusa (Prain) Raizada: Thailand, Staples et al. 1432 (BKF), OM699032, OM759912, OM759937, OM759962; Argyreia (Blinkworthia) convolvuloides (Prain) Rattanakrajang & Traiperm *(1): China, Li Jianwu 1011 (QBG), OM699025, OM759905, OM759930, OM759955; *(2): Burma, Mandalay, P. Rattanakrajang et al. 116 (QBG), OM699027, OM759907, OM759932, OM759957; *(3): Burma, Pyin Oo Lwin, P. Rattanakrajang et al. 117 (QBG), OM699026, OM759906, OM759931, OM759956. Argyreia erinacea Ooststr.: Malaysia, see Simões et al. (2015), SAN 152883 (SAN), KP261909, KP236603, KR024893, KR025036. Argyreia henryi Craib: Thailand, see Simões et al. (2015), Staples 1324 (SING), KP261910, KP236605, KR024895, KR025038. Argyreia laotica Gagnep.: Thailand, see Simões et al. (2015), Staples 1390 (SING), KP261911, KP236606, KR024896, KR025039. Argyreia (Blinkworthia) lycioides (Choisy) Traiperm & Rattanakrajang *(1): Thailand, Kanchanaburi, P. Rattanakrajang & P. Traiperm 5 (BKF), OM699020, OM759900, OM759925, OM759950; *(2): Thailand, Chiang Mai, P. Rattanakrajang & P. Traiperm 6 (BKF), OM699017, OM759897, OM759922, OM759947; *(3): Thailand, Chiang Mai, P. Rattanakrajang & P. Traiperm 14 (BKF), OM699018, OM759898, OM759923, OM759948; *(4): Thailand, Ratchaburi, P. Rattanakrajang & P. Traiperm 49 (BKF), OM699021, OM759901, OM759926, OM759951; *(5): Burma, Shan State, Y. Baba et al. 104920 (MBK), OM699022, OM759902, OM759927, OM759952; *(6): Burma, Mandalay Region, P. Rattanakrajang & P. Traiperm 101 (QBG), OM699019, OM759899, OM759924, OM759949. *Argyreia melvillei (S.Moore) Staples: Thailand, Staples et al. 1419 (BKF), OM699031, OM759911, OM759936, OM759961. Argyreia mollis (Burm. f.) Bojer: Thailand, see Simões et al. (2015), Staples 1391 (SING), KP261912, KP236607, KR024897, KR025040. Argyreia nervosa (Burm. f.) Bojer (1): Singapore, Singapore Botanic Garden, Singapore (cultivation), see Simões et al. (2015), SBG 20031385 (SING), KP261913, KP236608, KR024898, KR025041; *(2): Thailand, Bangkok, P. Rattanakrajang & P. Traiperm 141 (BKF), OM699039, OM759919, OM759944, OM759969. *Argyreia roxburghii (Sweet) Choisy (1): Thailand, P. Traiperm 482 (BKF), OM699037, OM759917, OM759942, OM759967; (2): Burma, K. Fujikawa et al. 95334 (MBK), OM699038, OM759918, OM759943, OM759968. Argyreia siamensis (Kerr) Staples: Thailand, see Simões et al. (2015), Staples 1412 (SING), KP261914, KP236609, KR024899, KR025042. *Argyreia stenophylla (Kerr) Staples & Traiperm: Thailand, P. Traiperm 610 (BKF), OM699029, OM759909, OM759934, OM759959. *Argyreia variabilis Traiperm & Staples: Thailand, S. Suddee s.n. (BKF), OM699028, OM759908, OM759933, OM759958. Argyreia zeylanica (Gaertn.) Voigt *(1): Sri Lanka, P. Traiperm 419 (BKF), OM699035, OM759915, OM759940, OM759965; *(2): Sri Lanka, P. Traiperm 420 (BKF), OM699036, OM759916, OM759941, OM759966. Astripomoea grantii (Rendle) Verdc.: Tanzania, see Stefanović et al. (2002), Koyombo 1000 (MO), –, AY101073,–, –. Astripomoea malvacea (Klotzsch) A.Meeuse: Tanzania, see Stefanović et al. (2002), Koyombo 6971 (A), AF309166, AY101074, –, –. Ipomoea aquatica Forssk.: Malaysia, see Simões et al. (2015), FRI 70037 (FRIM), KP261915, KP236616, KR024912, KR025049. Ipomoea cairica (L.) Sweet: USA, Hawaii, see Simões et al. (2015), Staples 1399 (SING), KP261916, KP236618, –, KR025051. Ipomoea carnea Jacq.: Singapore (cultivation), see Simões et al. (2015), SGB 19971190 (SING), KP261917, KP236619, KR024914, KR025052. Ipomoea cordatotriloba Dennst.: USA, Florida, see Simões et al. (2015), Staples 1465 (SING), KP261919, KP236621, KR024916, KR025054. Ipomoea indica Merr: USA, Hawaii, see Simões et al. (2015), Staples 1397 (SING), KP261921, KP236625, KR024921, KR025058. Ipomoea intrapilosa Rose: CNC (cultivation, Mexico), see Simões et al. (2015), Simões 52 (BM), KP261922, KP236626, KR024922, KR025059. Ipomoea littoralis Blume: Malaysia, see Simões et al. (2015), SAN 152892 (SAN), KP261924, KP236628, KR024924, KR025061. Ipomoea marmorata Britten & Rendle subsp. marmorata: Ethiopia, see Simões et al. (2015), Sebsebe D. et al. 6939 (BM), KP261925, –, KR024925, KR025062. Ipomoea mauritiana Jacq.: CNC (cultivation, France), see Simões et al. (2015), Simões 48 (BM), KP261926, KP236629, KR024926, KR025063. Ipomoea pes-tigridis L.: Singapore (cultivation), see Simões et al. (2015), SBG 20100015 (SING), KP261928, KP236632, KR024929, KR025066. Ipomoea pogonantha Thulin: Ethiopia, see Simões et al. (2015), Sebsebe D. et al. 6937 (BM), KP261930, KP236634, KR024931, KR025068. Ipomoea polymorpha Schult.: Ethiopia, see Simões et al. (2015), Sebsebe D. et al. 6961 (BM), KP261931, KP236635, KR024932, KR025069. Ipomoea purpurea (L.) Roth: CNC (cultivation, Madagascar), see Simões et al. (2015), Simões 50 (BM), KP261932, KP236636, KR024933, KR025070. Ipomoea quamoclit L.: Singapore (cultivation), see Simões et al. (2015), SBG 20080990 (SBG), KP261933, KP236637, KR024934, KR025071. Ipomoea sagittifolia Burm. f.: Malaysia, see Simões et al. (2015), FRI 7036 (FRIM), KP261934, KP236638, KR024935, KR025072. Ipomoea sinensis (Desr.) Choisy: Ethiopia, see Simões et al. (2015), Sebsebe D. et al. 6920 (BM), KP261935, KP236639, KR024936, KR025073. Ipomoea tricolor Cav.: Singapore, see Simões et al. (2015), Simões 19 (BM), KP261936, KP236640, KR024937, KR025074. Ipomoea violacea L.: Malaysia, see Simões et al. (2015), FRI 70039 (FRIM), KP261937, KP236641, KR024938, KR025075. Lepistemon binectarifer (Wall.) Kuntze (1): Malaysia, see Simões et al. (2015), FRI 66675 (FRIM), KP261938, –, KR024940, KR025077; *(2): Thailand, Sumanon s.n. (BKF), OM699041, OM759921, OM759946, OM759971. Rivea clarkeana Craib [= Rivea ornata (Roxb.) Choisy] (1): see Manos et al. (2001), Larsen & Larsen 34275 (K), AF309148, –, –, –; *(2): Thailand, Mae Hong Son, P. Rattanakrajang et al. 70 (BKF), OM699040, OM759920, OM759945, OM759970. Stictocardia beraviensis Hallier f.: CNC (cultivation, Madagascar), see Simões et al. (2015), Simões 62 (BM), KP261939, KP236717, KR0245019, KR025136. Stictocardia laxiflora (Baker) Hallier f.: CNC (cultivation, Uganda), see Simões et al. (2015), Simões 49 (BM), KP261940, KP236718, KR0245020, KR025137. Stictocardia tiliifolia (Desr.) Hallier f. (1): USA, Hawaii, see Simões et al. (2015), Staples 1400 (SING), KP261941, KP236719, KR0245021, KR025138; (2): see Manos et al. (2001), REM 106 (DUKE), AF309155, –, –, –. MERREMIEAE: Merremia emarginata Hallier f.: Thailand, see Simões et al. (2015), Staples 1335 (BM), KP261993, KP236659, KR024958, KR025092. Merremia hederacea Hallier f.: Thailand, see Simões et al. (2015), Simões 26 (BM), KP261995, KP236664, KR024964, KR025098. Merremia hirta Merr.: Cambodia, see Simões et al. (2015), Simões 44 (BM), KP261996, KP236665, KR024965, KR025099.

Appendix 2. Accession details of samples used in phenetic analyses in this study.

Accession number in this study, species name, source or geographic region, voucher with collector number (acronym of herbarium) respectively.

Argyreia campanuliflora (Gage) Thoth.: (1) Burma, Chin State, J. Murata et al. 24643 (MBK); (2) Burma, Magway Region, L. S. Man 88122 (MBK); (3) Burma, Chin State, L. S. Man 91786 (MBK); (4) Burma, Chin State, K. Fujikawa et al. 90187 (MBK); (5) Burma, Chin State, K. Fujikawa et al. 90125 (MBK); (6) Burma, Chin State, M. Matsumoto 80982 (BKF); (7-9) Burma, Chin State, P. Rattanakrajang et al. 113/1-3 (QBG); (10-12) Burma, Magway Region, Minbu, S. Mokim 418/1-3 (CAL). Argyreia (Blinkworthia) lycioides (Choisy) Traiperm & Rattanakr.: (13) Thailand, Chiang Mai, Mae Rim, P. Rattanakrajang & P. Traiperm 14 (BKF); (14) Thailand, Chiang Mai, Mae Rim, P. Rattanakrajang & P. Traiperm 20 (BKF); (15) Thailand, Chiang Mai, Mae Rim, P. Rattanakrajang & P. Traiperm 21 (BKF); (16) Thailand, Chiang Mai, Mae Rim, P. Rattanakrajang & P. Traiperm 22 (BKF); (17) Thailand, Kanchanaburi, Sri Sawat, P. Rattanakrajang & P. Traiperm 4 (BKF); (18) Thailand, Kanchanaburi, Sri Sawat, P. Rattanakrajang & P. Traiperm 5 (BKF); (19) Burma, Timmona, A. Huk. s.n. (P); (20) Thailand, Nakhon Sawan, Put 4075 (P); (21) Thailand, Chiang Mai, Muang, K. Larsen & B. Hansen 4287 (L); (22) Thailand, Nakhon Sawan, Put 4075 (L); (23) Thailand, Chiang Mai, Muang, A. F. G. Kerr 643 (L); (24) Burma, Meiktila, C. B. Collett 867 (L). Argyreia (Blinkworthia) convolvuloides (Prain) Rattanakr. & Traiperm: (25) Burma, Sagaing Region, J. Murata et al. 32311 (MBK); (26) China, Yunnan, L. J. Wu 1011 (QBG); (27) Burma, Mandalay Region, Pyin Oo Lwin, A. Rodges 859 (RAF); (28) Burma, Mandalay Region, Pyin Oo Lwin, Ba Pe 9606 (RAF); (29) China, Guangxi, L. K. De 3-15033 (GXMI); (30) China, Yunnan, L. C. Dui 1310 (HITBC); (31) China, Yunnan, Y. X. Di & Y. J. Suo s.n. (IBSC); (32) China, Guangxi, Q. H. Xing 3744 (IBSC); (33) China, Guangxi, Z. C. Quan A62361 (PE); (34) Burma, Mandalay Region, Pyin Oo Lwin, P. Rattanakrajang et al. 117 (QBG); (35) China, Yunnan, W. S. Gong 742 (KUN); (36) China, Yunnan, P. S. Ji 59-13003 (KUN); (37) China, Guangxi, L. Z. Ti 601026 (KUN); (38) China, Yunnan, T. D. Ding 1338 (KUN); (39) China, Guangxi, R. C. Ching 7353 (NAS); (40) China, Guangxi, R. C. Ching 7353 (NY); (41-42) Burma, Magway Region, Myingyan, J. C. Prazer s.n./1-2 (CAL).

Appendix 3. List of morphological characters used in phenetic analyses in this study.

P1. Habit: (0) Shrub; (1) Climber (based on specimen notes). P2. Internode length (a): (0) a < 2 cm; (1) 2 cm ≤ a < 3 cm; (2) 3 cm ≤ a < 4 cm; (3) a ≥ 4 cm. P3. Petiole length (b): (0) b < 0.4 cm; (1) 0.4 cm ≤ b < 0.6 cm; (2) 0.6 cm ≤ b < 0.8 cm; (3) b ≥ 0.8 cm. P4. Leaf blade shape: (0) Elliptic; (1) Lanceolate; (2) Lanceolate (Lindley); (3) Lanceolate (Linnaeus); (4) Ovate. P5. Leaf blade width (c): (0) c < 1 cm; (1) 1 cm ≤ c < 2 cm; (2) 2 cm ≤ c < 3 cm; (3) 3 cm ≤ c < 4 cm; (4) c ≥ 4 cm. P6. Leaf blade length (d): (0) d < 2 cm; (1) 2 cm ≤ d < 3 cm; (2) 3 cm ≤ d < 4 cm; (3) 4 cm ≤ d < 5 cm; (4) 5 cm ≤ d < 6 cm; (5) c ≥ 6 cm. P7. Leaf blade apex: (0) Acute; (1) Emarginate; (2) Mucronate apiculate; (3) Obtuse. P8. Leaf blade base: (0) Cuneate; (1) Obtuse; (2) Rounded. P9. Peduncle length (e): (0) e < 0.5 cm; (1) 0.5 cm ≤ e < 1 cm; (2) 1 cm ≤ e < 1.5 cm; (3) e ≥ 1.5 cm. P10. Pedicel length (f): (0) f < 0.2 cm; (1) 0.2 cm ≤ f < 0.4 cm; (2) f ≥ 0.4 cm. P11. Ratio of peduncle length to pedicel length (g): (0) g < 5; (1) 5 ≤ g < 15; (2) g ≥ 15. P12. Bract shape: (0) Boat-shaped; (1) Circular or elliptic; (2) Spatulate. P13. Outer bract length (h): (0) h < 0.5 cm; (1) 0.5 cm ≤ h < 0.75 cm; (2) h ≥ 0.75 cm. P14. Outer sepal shape: (0) Elliptic; (1) Obovate; (2) Orbicular or sub-orbicular; (3) Ovate. P15. Outer sepal width (i): (0) i < 0.6 cm; (1) 0.6 cm ≤ i < 0.8 cm; (2) 0.8 cm ≤ i < 1 cm; (3) i ≥ 1 cm. P16. Outer sepal length (j): (0) j < 0.75 cm; (1) 0.75 cm ≤ j < 1 cm; (2) 1 cm ≤ j < 1.5 cm; (3) j ≥ 1.5 cm. P17. Ratio of outer sepal length to outer bract length (k): (0) k < 1.5; (1) 1.5 ≤ k < 2; (2) 2 ≤ k < 2.5; (3) k ≥ 2.5.

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Rattanakrajang, P., Sumanon, P., Traiperm, P. et al. Reduction of Blinkworthia (Convolvulaceae) based on multilocus phylogenetic reconstruction and resurrection of a species from synonymy revealed by phenetic analyses. Kew Bull 77, 859–883 (2022). https://doi.org/10.1007/s12225-022-10052-1

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