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Pollen Morphology and Taxonomy in Malesian Phyllanthus (Phyllanthaceae)

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Pollen Morphology and Taxonomy in Malesian Phyllanthus (Phyllanthaceae) J. Jpn. Bot. 91 Suppl.: 257–292 (2016) Pollen Morphology and Taxonomy in Malesian Phyllanthus (Phyllanthaceae) a, b c a Ming-Jou WU *, Tseng-Chieng HUAng , Chia-Chin LIU , Yeng-Juei CHEN , a a a a Yung-Sheng CHAng , Chia-Lun HSU , Shin-Yi WU , Ai-Yi TSEng , a a a Yin-Chen CHAng , Cheng-Chuan LIU and Anusara KAEWMUAN aDepartment of Natural Resources and Environmental Studies, National Dong Hwa University, Shoufeng, Hualien 974, TAIWAN; bInstitute of Plant Biology, National Taiwan University, Taipei, 106 TAIWAN; cDepartment of Life Sciences, Tzu Chi University, Hualien, 970 TAIWAN *Corresponding author: [email protected] (Accepted on January 9, 2016) Pollen morphology in 89 of 127 Malesian Phyllanthus species (Phyllanthaceae) was investigated. Thirty-four pollen types are recognized. They are grouped into five pollen classes. These pollen classes are I. Colporate pollen, II. Incomplete synaperturate pollen, III. Synaperturate pollen, IV. Clypeate pollen, and V. Porate pollen. A key for the pollen classes and types is provided. Evolutionary trends in pollen morphology and infrageneric classification inPhyllanthus are discussed. Key words: Euphorbiaceae, Malesia, Phyllanthaceae, Phyllanthus, pollen class, pollen type. A total number of 110 Phyllanthus species Phyllanthus. Pollen morphology of 89 out of 127 in Malesia was estimated in 2001 (Malesian species was investigated. Previous palynological Euphorbiaceae Newsletter 11). Since studies on Malesian Phyllanthus species were then, several regional taxonomic revisions reported mainly by Punt (1980) and Chen et accompanying pollen studies of Malesian al. (2009). Punt examined 37 New Guinea Phyllanthus have been done by the authors Phyllanthus and reported nine pollen types, and including Peninsula Malaysia (Chang 2012), Chen et al. examined 21 Philippine Phyllanthus Borneo (Hsu 2011), Central Malesia (Wu and recognized eight pollen types. Many authors 2012), the Philippines (Chen 2009, Chen et al. have discussed pollen morphology, phylogeny 2009) and New Guinea (Chang 2010, Tseng and infrageneric classification inPhyllanthus and 2012). Based on these studies, 127 Malesian concordance with the value of palynological data Phyllanthus species are recognized. In this paper, in Phyllanthus taxonomy. Punt (1967) proposed palynological data of these studies and some evolutionary trends of seven pollen characters new data collected from Java (Liu 2015) and to discuss evolutionary trends in Phyllanthus Sumatra (Kaewmuan 2016) are compared and pollen types. Webster (1957) and Webster integrated for grouping pollen types and pollen and Carpenter (2002, 2008) discussed pollen classes to discuss evolutionary trends in pollen morphology and systematics of neotropical and morphology and infragenus classification in paleotropical Phyllanthus, and grouping sections —257— 258 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue Table 1. Voucher specimens of Malesian Phyllanthus pollen class and pollen type. The numbers in circle correspond to those in Map 1 Pollen class Pollen type Taxa Voucher specimens Locality Iα 1. P. amarus ① P. amarus Schumach. & Thonn. Idenburg 17 (L) Sumatra N. F. Stein 10 (SING) Peninsular Malaysia E. Kuswata Kartawinata 1063 (L); C. G. C. Reinwardt s.n. Java (L 0135108) F. H. Endert 2098 (L) Borneo D. R. Pleyte 172 (L) Moluccas M.-J. Wu 2951 (TAI) Palawan L. J. Brass 26029 (L) New Guinea ② P. debilis J. G. Klein ex Willd. W. J. C. Kooper 555 (L) Java J. W. Purseglove 4034 (L) Peninsular Malaysia H. Wiriadinata 1362a (L) Borneo J. Elbert 2903 (L) Sulawesi E. E. Henty NGF49173 (L) New Guinea 2. P. erythrotrichus ③ P. erythrotrichus C. B. Rob. C. B. Robinson BS29333 (L) Luzon, Rizal 3. P. maderaspatensis ④ P. maderaspatensis L. J. Jeswiet s.n. (L 0159955) Java C. W. Kooy 413 (L) Lesser Sunda Islands 4. P. microcarpus ⑤ P. microcarpus Müll. Arg. Hardial 486 (L) Peninsular Malaysia A. Hoogerwerf s.n. (L0135559) Java ⑥ P. novae-hollandiae Müll. Arg. R. Schodde 2425 (L) New Guinea 5. P. acinacifolius ⑦ P. acinacifolius Airy Shaw & G. L. Webster C. D. Sayers NGF19967 (L) New Guinea ⑧ P. flaviflorus (Lauterb. & K. Schum.) Airy Shaw R. D. Hoogland 5422 (L) New Guinea ⑨ P. kostermansii Airy Shaw A. J. G. H. Kostermans & W. Soegeng-Reksodihardjo New Guinea 656 (L) ⑩ P. poli-borealis Airy Shaw Sleumer BW14087 (LAE) New Guinea ⑪ P. stultitiae Airy Shaw L. J. Brass 11534 (L) New Guinea Iβ 6. P. urinaria ⑫ P. benguetensis C. B. Rob. E. D. Merrill 9684 (L) Luzon ⑬ P. chamaepeuce Ridl. R. Kiew RK2408 (SING) Peninsular Malaysia H. Winkler 3156 (L) Borneo ⑭ P. nozeranii Rossignol & Haicour J. Sinclari SF39526 (SING) Peninsular Malaysia K. Abdul A2889 (L) Borneo ⑮ P. urinaria L. subsp. urinaria C. G. G. J. van Steenis 24290 (L) Sumatra J. W. Purseglove P4034A (SING) Peninsular Malaysia J. P. Mousset 117 (L) Java P. Buwalda 5973 (L) Moluccas M.-J. Wu 2954 (TAI) Palawan J. W. Purseglove BSIP9864 (LAE) New Guinea December 2016 Wu et al.: Pollen in Malesian Phyllanthus 259 Table 1. Continued Pollen class Pollen type Taxa Voucher specimens Locality ⑯ P. urinaria L. subsp. nudicarpus Rossignol & Haicour Bakar s.n. (SING 0036094) Peninsular Malaysia J. W. F. Slik 2004-13 (L) Borneo A. Zippelius s.n. (L0136094) Lesser Sunda Islands M.-J. Wu 2952 (TAI) Palawan ⑰ P. submarginalis Airy Shaw E. Schmutz 2975 (L) Lesser Sunda Islands ⑱ P. aff. urinaria 1 E. Schmutz 3057 (L) Lesser Sunda Islands ⑲ P. aff. urinaria 2 L. L. Co 5993 (TAI) Luzon 7. P. roseus ⑳ P. ridleyanus Airy Shaw T. C. Whitmor KEP FRI4273 (L) Peninsular Malaysia ㉑ P. roseus (Craib & Hutch.) Beille C. Davidson 1344 (L) Peninsular Malaysia 8. P. tenellus ㉒ P. tenellus Roxb. V. F. Schiffner 2153 (L) Java 9. P. securinegioides ㉓ P. securinegioides Merr. M. D. Sulit PNH16268 (L) Palawan 10. P. glochidioides ㉔ P. glochidioides Elmer G. E. Edaño PNH40177 (L) Luzon ㉕ P. aff. glochidioides M. Q. Lagrimas PNH39954 (L) Luzon ㉖ P. pectinatus Hook. f. H. N. Ridley 5030 (SING) Peninsular Malaysia 11. P. collinsae ㉗ P. collinsae Craib E. Schmutz 850 (L) Lesser Sunda Islands J. A. J. Verheijen 3467 (L) Lesser Sunda Islands 12. P. triphlebius ㉘ P. triphlebius C. B. Rob. M.-J. Wu 3374, 3375 (TAI) Luzon 13. Pilate ㉙ P. aff. triphlebius M. Ramos BS43203 (L) Bohol Iγ 14. P. hookeri ㉚ P. hookeri Müll. Arg. J. H. Coert 521 (L) Java A. Hoogerwerf 252 (L) New Guinea 15. P. emblica ㉛ P. emblica L. F. W. Junghuhn s.n. (L 0159502) Sumatra S. Kiah SK613 (SING) Peninsular Malaysia F. W. Junghuh s.n. (L 0159536) Java P. W. Korthals s.n. (L 0159572) Borneo E. Demandt s.n. (L0159581) Lesser Sunda Islands 16. P. oxyphyllus ㉜ P. columnaris Müll. Arg. R. E. Holttum 15134 (SING) Peninsular Malaysia ㉝ P. oxyphyllus Miq. E. F. de Vogel 2986 (L) Sumatra J. W. Purseglove 4135 (L) Peninsular Malaysia C. Ploem s.n. (L 0230439) Java P. Hommel 111ZII (L) Java II 17. P. casticum ㉞ P. acidus (L.) Skeels S. M. Popta 613/97 (L) Java J. Elbert 2035 (L) Lesser Sunda Islands F. J. M. Gaerlan et al. PPI10249 (L) Luzon 260 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue Table 1. Continued Pollen class Pollen type Taxa Voucher specimens Locality ㉟ P. ciccoides Müll. Arg. F. J. M. Gaerlan et al. PPI10249 (L) New Guinea ㊱ P. reticulatus Poir. T. S. Suppiah KEP FRI11438 (L) Peninsular Malaysia Koorders, S.H. 20576 (L) Java W. Giesen 19A (L) Borneo J. G. Hallier 1301 (L) Borneo H. Hallier 4117 (L) Luzon J. E. Teysmann HB11622 (L) Lesser Sunda Islands J. Elbert 5538 (L) Sulawesi 18. P. maritimus ㊲ P. caesius Airy Shaw & G. L. Webster C. Versteegh BW(Ind.) 4701. (L) New Guinea ㊳ P. maritimus J. J. Sm. L. J. Toxopeus 368 (L) Moluccas J. J. F. E. de Wilde 1205 (L) New Guinea ㊴ P. pullenii Airy Shaw & G. L. Webster P. van Royen 6288 (L) New Guinea ㊵ P. ramosii Quisumb. & Merr. C. Mabesa 853 (L) Luzon 19. P. aeneus ㊶ P. ardisianthus Airy Shaw & G. L. Webster L. J. Brass 11357 (LAE) New Guinea ㊷ P. cuscutiflorus S. Moore A. K. Nura NK261 (KEP) Peninsular Malaysia ㊸ P. finschii K. Schum. V. Novotny 9 (L) New Guinea ㊹ P. merinthopodus Diels P. F. Stevens LAE58638 (L) New Guinea ㊺ P. paniculatus Oliv. P. Katik LAE 56277 (L) New Guinea ㊻ P. praelongipes Airy Shaw & G. L. Webster R. D. Hoogland 4737 (L) New Guinea ㊼ P. tagulae Airy Shaw & G. L. Webster L. J. Brass 27986 (LAE) New Guinea 20. P. papuanus ㊽ P. bourgeoisii Baill. C. L. Ledermann 9740 (L) New Guinea ㊾ P. rheophilus Airy Shaw E. E. Henty NGF12403 (L) New Guinea ㊿ P. rosselensis Airy Shaw & G. L. Webster A. N. Gillison NGF25404 (L) New Guinea P. tenuirhachis J. J. Sm. W. H. de Vriese s.n. (L0135901) Sulawesi D. R. Pleyte 503 (L) New Guinea 21. P. ovatifolius P. clamboides (F. Muell.) Diels R. D. Hoogland 9044 (L) New Guinea P. effuses S. Moore R. D. Hoogland 3917 (L) New Guinea P. gjellerupii J. J. Sm. H. A. van der Sijde BW (Ind.) 4188 (L) New Guinea P. ovatifolius J. J. Sm. Jaheri s.n. (L0135238) Moluccas December 2016 Wu et al.: Pollen in Malesian Phyllanthus 261 Table 1. Continued Pollen class Pollen type Taxa Voucher specimens Locality P. papuanus Gage L. J. Brass 6960 (L) New Guinea III 22. P. aphanostylus P. aphanostylus Airy Shaw & G. L. Webster L. J. Brass 32430 (L) New Guinea 23. P. tabularis P. tabularis Airy Shaw J. F. Veldkamp 8359A (L), 11220 (LAE) New Guinea P. warburgii K.Schum. Atasrip 117 (L) New Guinea 24. P. rubriflorus P. rubriflorus J. J. Sm. H. J. Lam 7758 (L) New Guinea IV 25. P. aff. samarensis P. aff. samarensis C. E. Ridsdale 1479 (L) Luzon 26. P. tenuipes P. tenuipes C. B. Rob. C. E. Ridsdale ISU276 (L) Luzon 27. P. virgatus P. virgatus G. Forst. Asdat 197 (L) Sumatra Unknown (L0136230) Java P.
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