Uncovering five new species of Polyalthia (Annonaceae, Miliuseae) from Thailand: molecular and morphological congruence
Abstract
Based on molecular phylogenetic analyses using seven plastid DNA markers (matK, rbcL, ndhF and ycf1 exons; trnL intron; psbA-trnH and trnL-trnF intergenic spacers) and detailed morphological investigations, five new species of Polyalthia Blume (Annonaceae Juss.) from Thailand are recognized, described and illustrated. These species are: P. acuminatissima Wiya, Damth. & Chaowasku sp. nov. (from Nakhon Phanom Province, northeastern region), P. esanensis Wiya, Damth. & Chaowasku sp. nov. (from eastern and northeastern regions), P. chiangmaica Wiya, Damth. & Chaowasku sp. nov. (from Chiang Mai Province, northern region), P. maxwelliana Wiya, Chanthamrong & Chaowasku sp. nov. and P. miliusifolia Wiya, Chanthamrong & Chaowasku sp. nov. (both from Kanchanaburi Province, southwestern region). All five species belong to one of the three major clades within the genus. Three of them (P. acuminatissima, P. esanensis and P. chiangmaica) were previously misidentified as P. evecta (Pierre) Finet & Gagnep., which is polyphyletic. Comparisons with their phylogenetically related or morphologically similar species are provided and a key to the species of this major clade occurring in Thailand is included. The conservation status of all five new species is provisionally assessed as Data Deficient.
References
Akaike H. 1974. A new look at the statistical model identification. IEEE Transactions on Automatic Control 19 (6): 716–723. https://doi.org/10.1109/TAC.1974.1100705
Ast S. 1938. Anonacées. In: Humbert H. (ed.) Flore générale de l’Indo-Chine. Suppl. 1: 59–123. Muséum national d’Histoire naturelle, Paris.
Bân N.T. 2000. Thực Vật Chí Việt Nam [Flora of Vietnam]. Vol 1: Họ Na [Annonaceae]. Science and Technics Publishing House, Hanoi.
Blume C.L. von 1825. Bijdragen tot de Flora van Nederlandsch Indië. Part 1. Ter Lands Drukkerij, Batavia [Jakarta]. https://doi.org/10.5962/bhl.title.115427
Blume C.L. von 1830. Flora Javae nec non insularum adjacentium. Parts 28–29. Frank, Brussels. https://doi.org/10.5962/bhl.title.48445
Bunchalee P. 2022. Polyalthia. In: Newman M.F., Barfod A.S., Esser H.J., Simpson D.A. & Parnell J.A.N. (eds) Flora of Thailand. Vol. 16 (part 1: Annonaceae): 238–259. Prachachon Printing, Bangkok.
Bunchalee P., Chantaranothai P., Johnson D.M. & Murray N.A. 2019. Polyalthia khaoyaiensis (Annonaceae), a new species from Thailand. Phytotaxa 405 (3): 171–179. https://doi.org/10.11646/phytotaxa.405.3.6
Bunchalee P., Leeratiwong C. & Johnson D.M. 2021a. Two new species and a new record of the genus Polyalthia (Annonaceae) from Peninsular Thailand. Phytotaxa 510 (3): 239–250. https://doi.org/10.11646/phytotaxa.510.3.4
Bunchalee P., Johnson D.M., Murray N.A. & Leeratiwong C. 2021b. Three new species of Polyalthia (Annonaceae) from Thailand and Laos. Phytotaxa 512 (4): 272–282. https://doi.org/10.11646/phytotaxa.512.4.3
Bunchalee P., Leeratiwong C. & Sinbumroong A. 2025. Polyalthia khaosokense (Annonaceae), a new species from Peninsular Thailand with a key to the Polyalthia evecta species complex. Phytotaxa 683 (2): 184–188. https://doi.org/10.11646/phytotaxa.683.2.10
Candolle A.L.P.P. de 1832. Mémoire sur la famille des Anonacées, et en particulier sur les espèces du pays des Birmans. Mémoires de la Société de Physique et d’Histoire naturelle de Genève 5: 177–221.
Chaowasku T. & Kessler P.J.A. 2013. Seven new species of Miliusa (Annonaceae) from Thailand. Nordic Journal of Botany 31 (6): 680–699. https://doi.org/10.1111/j.1756-1051.2012.01785.x
Chaowasku T. & Kessler P.J.A. 2014. Miliusa cambodgensis sp. nov. (Annonaceae) from Cambodia and M. astiana, M. ninhbinhensis spp. nov. from Vietnam. Nordic Journal of Botany 32 (3): 298–307. https://doi.org/10.1111/j.1756-1051.2013.00219.x
Chaowasku T., Johnson D.M., van der Ham R.W.J.M. & Chatrou L.W. 2012. Characterization of Hubera (Annonaceae), a new genus segregated from Polyalthia and allied to Miliusa. Phytotaxa 69: 33–56. https://doi.org/10.11646/phytotaxa.69.1.6
Chaowasku T., Thomas D.C., van der Ham R.W.J.M., Smets E.F., Mols J.B. & Chatrou L.W. 2014. A plastid DNA phylogeny of tribe Miliuseae: insights into relationships and character evolution in one of the most recalcitrant major clades of Annonaceae. American Journal of Botany 101 (4): 691–709. https://doi.org/10.3732/ajb.1300403
Chaowasku T., Johnson D.M., van der Ham R.W.J.M. & Chatrou L.W. 2015. Huberantha, a replacement name for Hubera (Annonaceae: Malmeoideae: Miliuseae). Kew Bulletin 70: 23. https://doi.org/10.1007/s12225-015-9571-z
Chaowasku T., Damthongdee A., Jongsook H., Ngo D.T., Le H.T., Tran D.M. & Suddee S. 2018a. Enlarging the monotypic Monocarpieae (Annonaceae, Malmeoideae): recognition of a second genus from Vietnam informed by morphology and molecular phylogenetics. Candollea 73: 261–275. https://doi.org/10.15553/c2018v732a11
Chaowasku T., Damthongdee A., Jongsook H., Nuraliev M.S., Ngo D.T., Le H.T., Lithanatudom P., Osathanunkul M., Deroin T., Xue B. & Wipasa J. 2018b. Genus Huberantha (Annonaceae) revisited: erection of Polyalthiopsis, a new genus for H. floribunda, with a new combination H. luensis. Annales Botanici Fennici 55 (1–3): 121–137. https://doi.org/10.5735/085.055.0114
Chaowasku T., Aongyong K., Damthongdee A., Jongsook H. & Johnson D.M. 2020. Generic status of Winitia (Annonaceae, Miliuseae) reaffirmed by molecular phylogenetic analysis, including a new species and a new combination from Thailand. European Journal of Taxonomy 659: 1–23. https://doi.org/10.5852/ejt.2020.659
Chernomor O., von Haeseler A. & Minh B.Q. 2016. Terrace aware data structure for phylogenomic inference from supermatrices. Systematic Biology 65 (6): 997–1008. https://doi.org/10.1093/sysbio/syw037
Couvreur T.L.P., Dagallier L.P.M.J., Crozier F., Ghogue J.P., Hoekstra P.H., Kamdem N.G., Johnson D.M., Murray N.A. & Sonké B. 2022. Flora of Cameroon 45 – Annonaceae. PhytoKeys 207: 1–532. https://doi.org/10.3897/phytokeys.207.61432
Damthongdee A. & Chaowasku T. 2022. Miliusa parvifolia comb. nova based on Popowia parvifolia (Annonaceae, Miliuseae). Annales Botanici Fennici 59 (1): 313–315. https://doi.org/10.5735/085.059.0144
Damthongdee A., Saengpho C., Wiya C., Baka A., Chanthamrong K., Ue-aree P. & Chaowasku T. 2024. Drepananthus khaosoi sp. nov. (Annonaceae) from southern Thailand, with molecular phylogenetic reconstructions. Anales del Jardín Botánico de Madrid 81 (1): e144. https://doi.org/10.3989/ajbm.596
Darriba D., Taboada G.L., Doallo R. & Posada D. 2012. jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9: 772. https://doi.org/10.1038/nmeth.2109
Edgar R.C. 2004. MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32 (5): 1792–1797. https://doi.org/10.1093/nar/gkh340
Endlicher S. 1839. Genera plantarum secundum ordines naturalis disposita. Part 11. Beck, Vienna. https://doi.org/10.5962/bhl.title.728
Finet A. & Gagnepain F. 1906. Contributions à l’étude de la flore de l’Asie orientale [Anonacées]. Bulletin de la Société botanique de France 53 (Mémoires 4): 55–170. https://doi.org/10.1080/00378941.1905.10833364
Felsenstein J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39 (4): 783–791. https://doi.org/10.1111/j.1558-5646.1985.tb00420.x
Goloboff P.A. & Catalano S.A. 2016. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics 32 (3): 221–238. https://doi.org/10.1111/cla.12160
Goloboff P.A., Farris J.S., Källersjö M., Oxelman B., Ramírez M.J. & Szumik C.A. 2003. Improvements to resampling measures of group support. Cladistics 19 (4): 324–332. https://doi.org/10.1111/j.1096-0031.2003.tb00376.x
Hasegawa M., Kishino H. & Yano T. 1985. Dating of the human-ape splitting by a molecular clock of mitochondrial DNA. Journal of Molecular Evolution 22: 160–174. https://doi.org/10.1007/BF02101694
Hewson H.J. 1988. Plant Indumentum: A Handbook of Terminology. Australian flora and fauna series 9. Australian Government Publishing Service, Canberra.
Hooker J.D. & Thomson T. 1855. Flora Indica: Being a Systematic Account of the Plants of British India, Together with Observations on the Structure and Affinities of Their Natural Orders and Genera. Pamplin, London. https://doi.org/10.5962/bhl.title.57706
Hooker J.D. & Thomson T. 1872. Anonaceae. In: Hooker J.D. (ed.) The Flora of British India. Vol. 1 (part 1): 45–94. Reeve & Co., London. https://doi.org/10.5962/bhl.title.678
IUCN Standards and Petitions Committee 2024. Guidelines for using the IUCN Red List categories and criteria. Version 16. Prepared by the Standards and Petitions Committee. Available from https://nc.iucnredlist.org/redlist/content/attachment_files/RedListGuidelines.pdf [accessed 3 Jul. 2025].
Jessup L.W. 2007. Annonaceae. In: Wilson A.J.G. (ed.) Flora of Australia. Vol. 2: 18–57. CSIRO, Melbourne.
Jovet-Ast S.1940. Anonacées nouvelles d’Indochine. Notulae Systematicae 9: 73–88.
Khumchompoo S. & Thongpukdee A. 2005. Polyalthia kanchanaburiana (Annonaceae): a new species from Thailand. Thai Forest Bulletin (Botany) 33: 35‒41.
Kurz W.S. 1875. Descriptions of new plants from the Nicobar Islands (including a few from the Andaman Islands). Journal of Botany, British and Foreign 13: 321–333.
Miller M.A., Pfeiffer W. & Schwartz T. 2010. Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop (GCE): 1–8. IEEE, Piscataway. https://doi.org/10.1109/GCE.2010.5676129
Minh B.Q., Schmidt H.A., Chernomor O., Schrempf D., Woodhams M.D., von Haeseler A. & Lanfear R. 2020. IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution 37 (5): 1530–1534. https://doi.org/10.1093/molbev/msaa015
Miquel F.A.W. 1865. Anonaceae Archipelagi Indici. Annales Musei Botanici Lugduno-Batavi 2: 1–45.
Mols J.B., Kessler P.J.A., Rogstad S.H. & Saunders R.M.K. 2008. Reassignment of six Polyalthia species to the new genus Maasia (Annonaceae): molecular and morphological congruence. Systematic Botany 33 (3): 490–494. https://doi.org/10.1600/036364408785679752
Nge F.J., Chaowasku T., Damthongdee A., Wiya C., Soulé V.R.C., Rodrigues-Vaz C., Bruy D., Mariac C., Chatrou L.W., Chen J., Choo L.M., Dagallier L.-P.M.J., Erkens R.H.J., Johnson D.M., Leeratiwong C., Lobão A.Q., Lopes J.C., Martínez-Velarde M.F., Munzinger J., Murray N.A., Neo W.L., Rakotoarinivo M., Ortiz-Rodriguez A.E., Sonké B., Thomas D.C., Wieringa J.J. & Couvreur T.L.P. 2024. Complete genus-level phylogenomics and new subtribal classification of the pantropical plant family Annonaceae. Taxon 73 (6): 1341–1369. https://doi.org/10.1002/tax.13260
Pierre L. 1881. Flore forestière de la Cochinchine. Fasc. 2. Doin, Paris. https://doi.org/10.5962/bhl.title.61558
Pirie M.D., Chatrou L.W., Mols J.B., Erkens R.H.J. & Oosterhof J. 2006. ‘Andean-centred’ genera in the short-branch clade of Annonaceae: testing biogeographical hypotheses using phylogeny reconstruction and molecular dating. Journal of Biogeography 33 (1): 31–46. https://doi.org/10.1111/j.1365-2699.2005.01388.x
Rambaut A., Drummond A.J., Xie D., Baele G. & Suchard M.A. 2018. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systems Biology 67 (5): 901–904. https://doi.org/10.1093/sysbio/syy032
Ridley H.N. 1912. A botanical excursion to Pulau Adang. Journal of the Straits Branch of the Royal Asiatic Society 61: 45–65.
Ridley H.N. 1915. The plants of Koh Samui and Koh Pennan. Journal of the Federated Malay States Museums 5: 158–168.
Ridley H.N. 1920. Two new Siamese plants. Journal of the Federated Malay States Museums 10: 127.
Ronquist F., Teslenko M., van der Mark P., Ayres D.L., Darling A., Höhna S., Larget B., Liu L., Suchard M.A. & Huelsenbeck J.P. 2012. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systems Biology 61 (3): 539–542. https://doi.org/10.1093/sysbio/sys029
Roxburgh W. 1795. Plants of the Coast of Coromandel. Vol. 1. Bulmer, London. https://doi.org/10.5962/bhl.title.467
Saunders R.M.K., Su Y.C.F. & Xue B. 2011. Phylogenetic affinities of Polyalthia species (Annonaceae) with columellar-sulcate pollen: enlarging the Madagascan endemic genus Fenerivia. Taxon 60 (5): 1407–1416. https://doi.org/10.1002/tax.605016
Shailajakumari S., Sreekala A.K., Parthipan B. & Kumar E.S.S. 2019. Note on the status of Polyalthia malabarica var. longipedicellata M.Alister et al. (Annonaceae) – an endemic species of the southern Western Ghats. Global Journal of Bio-Science and Biotechnology 8: 117–118.
Simmons M.P. 2004. Independence of alignment and tree search. Molecular Phylogenetics and Evolution 31 (3): 874–879. https://doi.org/10.1016/j.ympev.2003.10.008
Simmons M.P. & Ochoterena H. 2000. Gaps as characters in sequence-based phylogenetic analyses. Systems Biology 49 (2): 369–381. https://doi.org/10.1093/sysbio/49.2.369
Staden R., Beal K.F. & Bonfield J.K. 2000. The Staden package, 1998. In: Misener S. & Krawetz S.A. (eds) Bioinformatics Methods and Protocols. Methods in Molecular Biology™: 115–130. Humana Press, Totowa. https://doi.org/10.1385/1-59259-192-2:115
Tamura K., Stecher G. & Kumar S. 2021. MEGA11: molecular evolutionary genetics analysis version 11. Molecular Biology and Evolution 38 (7): 3022–3027. https://doi.org/10.1093/molbev/msab120
Tang C.C., Xue B. & Saunders R.M.K. 2013. A new species of Goniothalamus (Annonaceae) from Palawan, and a new nomenclatural combination in the genus from Fiji. PhytoKeys 32: 27–35. https://doi.org/10.3897/phytokeys.32.6663
Tavaré S. 1986. Some probabilistic and statistical problems in the analysis of DNA sequences. Lectures on Mathematics in the Life Sciences 17: 57–86.
Thiers B. continuously updated. Index Herbariorum: A Global Directory of Public Herbaria and Associated Staff. New York Botanical Garden’s Virtual Herbarium. Available from https://sweetgum.nybg.org/science/ih/ [accessed 3 Jul. 2025].
Thwaites G.H.K. 1864. Enumeratio Plantarum Zeylaniae: an Enumeration of Ceylon Plants, with Descriptions of the New and Little Known Genera and Species, Observations on Their Habitats, Uses, Native Names, etc. Part 5. Dulau & Co., London. https://doi.org/10.5962/bhl.title.574
Turner I.M. 2012a. A new combination in Monocarpia (Annonaceae). Edinburgh Journal of Botany 69 (1): 95–98. https://doi.org/10.1017/S0960428611000424
Turner I.M. 2012b. The plant taxa of H.N. Ridley, 4. The primitive angiosperms (Austrobaileyales, Canellales, Chloranthales, Laurales, Magnoliales, Nymphaeales and Piperales). Gardens’ Bulletin Singapore 64 (1): 221–256. Available from https://ia902809.us.archive.org/13/items/biostor-146686/biostor-146686.pdf [accessed 3 Jul. 2025].
Turner I.M. 2018. Annonaceae of the Asia-Pacific region: names, types and distributions. Gardens’ Bulletin Singapore 70 (1): 409–744. https://doi.org/10.26492/gbs70(2).2018-11
van Heusden E.C.H. 1994. Revision of Haplostichanthus (Annonaceae). Blumea 39: 215–234.
Wiens J.J. 1998. Combining data sets with different phylogenetic histories. Systematic Biology 47 (4): 568–581. https://doi.org/10.1080/106351598260581
Wiya C. & Chaowasku T. 2021. Disintegration of Polyalthia debilis (Annonaceae): P. cambodica comb. nov., P. canaensis comb. et stat. nov., and P. suthepensis nom. nov. for Unona dubia. Phytotaxa 487 (3): 273–282. https://doi.org/10.11646/phytotaxa.487.3.7
Xue B., Su Y.C.F., Mols J.B., Kessler P.J.A. & Saunders R.M.K. 2011. Further fragmentation of the polyphyletic genus Polyalthia (Annonaceae): molecular phylogenetic support for a broader delimitation of Marsypopetalum. Systematics and Biodiversity 9 (1): 17–26. https://doi.org/10.1080/14772000.2010.542497
Xue B., Su Y.C.F., Thomas D.C. & Saunders R.M.K. 2012. Pruning the polyphyletic genus Polyalthia (Annonaceae) and resurrecting the genus Monoon. Taxon 61 (5): 1021–1039. https://doi.org/10.1002/tax.615009
Xue B., Thomas D.C., Chaowasku T., Johnson D.M. & Saunders R.M.K. 2014. Molecular phylogenetic support for the taxonomic merger of Fitzalania and Meiogyne (Annonaceae): new nomenclatural combinations under the conserved name Meiogyne. Systematic Botany 39 (2): 396–404. https://doi.org/10.1600/036364414X680825
Xue B., Tan Y.H. & Ye X.E. 2016. The identity of Polyalthia florulenta (Annonaceae): a second species of Wangia in China. Phytotaxa 283 (2): 163–171. https://doi.org/10.11646/phytotaxa.283.2.5
Xue B., Ding H.B., Yao G., Shao Y.Y., Fan X.J. & Tan Y.H. 2020. From Polyalthia to Polyalthiopsis (Annonaceae): transfer of species enlarges a previously monotypic genus. PhytoKeys 148: 71–91. https://doi.org/10.3897/phytokeys.148.50929
Yang Z. & Rannala B. 1997. Bayesian phylogenetic inference using DNA sequences: a Markov Chain Monte Carlo method. Molecular Biology and Evolution 14 (7): 717–724. https://doi.org/10.1093/oxfordjournals.molbev.a025811
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