J. Jpn. Bot. 91 Suppl.: 99–111 (2016)

New or Noteworthy Collections from Myanmar (9) Agapetes () from Northwestern Myanmar

a b c Nobuyuki Tanaka , Tetsuo Ohi-Toma , Hiroko Murata , d b, Mu Mu Aung and Jin Murata *

aDepartment of Botany, National Museum of Nature and Science, 4-1-1, Amakubo, Tsukuba, 305-0005 JAPAN; bBotanical Gardens, Graduate School of Sciences, the University of Tokyo, 3-7-1, Hakusan, Bunkyo-ku, Tokyo, 112-0001 JAPAN; cFaculty of Pharmaceutical Sciences, Setsunan University, Nagaotoge-cho, Hirakata, Osaka, 573-0101 JAPAN; dForest Research Institute, Forest Department, Ministry of Natural Resources and Environmental Conservation, Yezin, Nay Pyi Taw, MYANMAR *Corresponding author: [email protected]

(Accepted on August 2, 2016)

As the result of field explorations to northwestern Myanmar, a total of five species of the Agapetes (Ericaceae) were collected. Of these, two species were hitherto undescribed taxa. They were recorded with citation of voucher specimens and a photograph of each species was provided. Two new species were described as Agapetes oxycoccoides and A. pentastigma. Agapetes oxycoccoides is similar to A. bracteata, but distinguished by the corymbose inflorescence without distinct bract, and more strongly reflexed longer corolla lobes. Agapetes pentastigma is morphologically similar to A. hillii, but differs in having a greenish-yellow collora with ladder-like red stripes, and a short peduncle. (Continued from Acta Phytotax. Geobot. 65: 89–97, 2014)

Key words: Agapetes oxycoccoides, Agapetes pentastigma, Myanmar, new species, phylogeny.

The genus Agapetes D. Don ex G. Don is 2005, Watthana 2012, Tong and Xia 2014). assignable to the tribe Rchb. of the In morphological and anatomical characters, subfamily Arn. (Ericaceae). The Agapetes is closely related to from SE past classification treated the species from New Asia (Stevens 1985). The molecular phylogeny Guinea and the SW Pacific as Agapetes (Airy indicated that Agapetes species nested among Shaw 1958). The species, however, except Vaccinium species, cf. V. gaultheriifolium A. scortechinii (King & Gamble) Sleumer, of (Griff.) Hook. f. and V. nummularia Hook. f. & which taxonomic status has been in question, Thomson, although only three Agapetes species, are currently placed in Paphia Seeman (Kron A. buxifolia Nutt. ex Hook. f., A. hosseana Diels, et al. 1999, Stevens 2004). Agapetes, holding and A. serpens (Wight) Sleumer, were analyzed about 80 species at present, is distributed (Kron et al. 2002). Currently it is thought that from the Himalayas to China, and Indo- many species of Agapetes are part of the same China to Southeast Asia (Fang and Stevens lineage as many SE Asian-Malesian Vaccinium

—99— 100 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue

(Fang and Stevens 2005). Myanmar, and A. pentastigma was close to A. Field expeditions to Kachin State, odontocera (Wight) Hook. f., distributed in E northwestern Myanmar to update the flora India. of Myanmar were carried out. Northwestern Myanmar is known for its wealth of plant Materials and Methods species diversity. Our previous works revealed Plant materials many new records and also new taxa based on Field excursions to the Hukaung Valley Tiger these northwestern excursions (Tanaka et al. Reserve, Kachin State, northwestern Myanmar 2006, Tanaka and Hughes 2007, Tanaka and have been conducted twice, in 2005 and 2007. Hayami 2011, Tanaka et al. 2016). The Hukaung Valley is one of the largest Differences in the amount of rainfall in conservation sites in the country, but it remains the dry and rainy seasons are remarkable in floristically poorly known. The valley is a flat Myanmar, and it is presumed that there alluvial plain measuring about 80 km north– are adapted to the environment by having a south by 50 km east–west, surrounded on all characteristic life cycle. The genus Agapetes sides by hills (Cruickshank and KoKo 2003). with its characteristic storage rootstock may be A total of five species of Agapetes were one of such a kind of plant, and it has apparently collected in the evergreen tropical lowland diversified in Myanmar. Including twenty new forest in Shinbweyang, northwest of Tanaing, species that were previously described from Hukaung Valley, and they were morphologically Myanmar by Airy Shaw (1935, 1948, 1959, investigated. Voucher specimens were deposited 1960a, 1960b), Kress et al. (2003) listed 54 in TI, TNS and RAF. species of Agapetes from Myanmar. Recently Tong and Xia (2014) added two new taxa, DNA sequencing and phylogenetic analysis A. putaoensis Y. H. Tong & N. H. Xia and A. For molecular phylogenetic analyses, wardii W. W. Smith var. heterotricha Y. H. 24 species and one variety of Agapetes, Tong & N. H. Xia to the flora of Myanmar, for including A. scortechinii, eight representative a total of 56 taxa to date. However, many of species of Vaccinium including species that them were known from fragmentary information are morphologically and phylogenetically based on the type and few additional herbarium related to Agapetes (V. gaultheriifolium and specimens. Observation on living plants in V. nummularia; Kron et al. 2002), and two the field is desirable for a taxonomic and species of Paphia were collected (Appendix phylogenetic revision of Agapetes and related 1). Considering the phylogenetic relationship species. The identifications of the specimens of Vaccinieae, Andromeda polifolia L. and collected by our fieldworks led us to recognize Dimorphanthera kempteriana Schltr. were used five species of Agapetes (Ericaceae) from as outgroups. Total genomic DNA was extracted the field of northwestern Myanmar. Based on from silica-gel-dried leaf tissue, using the morphological and phylogenetic investigations modified HEPES-CTAB method described by two of them were recognized as new to Ohi-Toma et al. (2010). science. We describe these two species as A. Nucleotide sequences of the chloroplast oxycoccoides and A. pentastigma in this study. trnK intron (including the matK gene) were Molecular phylogenetic analysis was also determined using polymerase chain reaction applied to understand the relationship of these (PCR) and direct bi-directional sequencing. species among the related taxa. It suggested PCR amplification was conducted using that A. oxycoccoides was related to A. setigera TaKaRa ExTaq (TaKaRa Bio, Shiga, Japan), D. Don ex G. Don, distributed in India and with the following cycling conditions: December 2016 Tanaka et al.: Agapetes from NE Myanmar 101

Table. 1. List of primers for amplification and sequencing Primers Sequence from the 5’ end Note* Reference trnK-3914F TGGGTTGCTAACTCAATGG P, S Johnson and Soltis (1994) trnK-5-intron_Rv AGCTTTTCCCGCAACAGATAC S This study trnK-5-intron_Fw TTTCCAAAATCAAAAGAGCG S This study matK 10 ATGATTCTGTTGATACATTC P, S Kato et al. (1998) matK 800F CACTTGCTYATGATCGTGGT P, S This study matK 4R GCATCTTTTACCCARTAGCGAAG S Bremer et al. (2002) matK 1650R ACATTCCCAGAAATTKACAAGGT P, S This study matK 4F CTTCGCTAYTGGGTAAAAGATGC P, S Bremer et al. (2002) matK ERI_R GCACAAGAAAGTCGAAGTAT P, S Dunning and Savolainen (2010) matK 1660F TTTTCCATATGGGCTCAACC P, S This study matK AST_R CAAATAATATCCAAATACCAA P, S Dunning and Savolainen (2010) matK-3-intron Fw TACGGAGGAAGAGCAGGTTT S This study trnK-2R AACTAGTCGGATGGAGTAG P, S Johnson and Soltis (1994) *P. PCR amplification. S. Sequencing. denaturation at 96°C for 45 sec, followed by 33 coded gap-states were used as a fifth character; cycles at 96°C for 45 sec, annealing at 50°C for however, the length polymorphisms caused by 45 sec, extension at 72°C for 1 min, and a final mononucleotide repeats [poly(A) or poly(T)] extension at 72°C for 10 min. For amplifying and other unalignable portions were excluded and sequencing the trnK intron, we used newly from gap coding. In the data matrix, when designed primers in addition to published substitutions or overlapping gaps occurred at universal primers (Table 1). Amplification gap positions, the differences were indicated products were purified using illustra ExoProStar by adding ‘n’ to the position of the substitution (GE Healthcare UK Ltd., Buckinghamshire, and/or gap-state. In order to use coded gaps, England, UK). The purified PCR fragments were Maximum Parsimony analysis was applied using amplified using the Big Dye Terminator v3.1 PAUP* v. 4.0a149 (Swofford 2002). Nucleotide Cycle Sequencing Kit (Applied Biosystems, substitutions and coded gaps were given equal Foster City, California, USA). Cycle-sequencing weights, and the tree search was performed reactions were performed using the same primers using the branch-and-bound search option with for the PCR amplification and newly designed MulTrees in effect. The strict consensus tree primers for sequencing. Complementary strands of the most parsimonious trees was generated, of the sequenced regions were assembled and and changes of substitutions and gaps were edited using ATGC v. 4.2 (GENETXY Co., reconstructed on the tree using ACCTRAN Tokyo, Japan). All newly generated nucleotide character optimization. The MP bootstrap sequences were deposited in the DNA Data support (BS) values were estimated using a Bank of Japan (DDBJ) and their accession heuristic search of 10,000 replicates, with simple numbers are shown with sample information in sequence addition, TBR branch swapping, Appendix. Multrees in effect, and no limit of MaxTree. The nucleotide sequences were manually aligned and gap coding was employed as Results and Discussion described in Ohi-Toma et al. (2010). The Morphological characters 102 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue

Fig. 1. Strict consensus tree of 20 most parsimonious trees, based on chloroplast DNA sequences of trnK intron including the matK gene (tree length = 477, CI = 0.8532, RC = 0.7508). Branch lengths were estimated based on ACCTRAN character optimization and a scale bar (five site change) is shown. Numbers on branches are bootstrap values (over 50%). Generic names of Agapetes and Vaccinium are abbreviated as AGA and VAC, respectively. Two new species are indicated in bold. Thick branch is the AGA-VAC clade.

As the result of the morphological species known from Myanmar and neighboring investigation, five species were recognized. regions according to studies of related previous Three were identified as A. burmanica W. literatures (Airy Shaw 1935, 1948, 1959, 1960a E. Evans, A. hillii Brandis (Fig. 4) and A. and 1960b, Stevens 1985, Fang and Stevens setigera D. Don ex G. Don (Fig. 5), however, 2005, Watthana 2012). These species are newly the other two taxa did not match any other described as A. oxycoccoides and A. pentastigma December 2016 Tanaka et al.: Agapetes from NE Myanmar 103

Fig. 2. Agapetes oxycoccoides J. Murata, Nob. Tanaka & Ohi-Toma. A. Habit. B. Vertical section of flower showing style, stigma, and ovary. C. Side view of flower. D. Front view of flower. E. Stamens with pubescent flat filaments and connate anther tubules. Scale bar: 2 cm for A, 5 mm for B–E. 104 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue in the taxonomic treatment. analysis. Thus, the species most closely related Morphology of the stigma may be one of the to A. oxycoccoides is still not clear. To clarify important characters for Agapetes identification the rigorous systematic position for these based on our extensive observations on the two new species further phylogenetic study Myanmar Agapetes specimens. As previously with additional taxa including A. bracteata is recognized (Fang and Stevens 2005), the required. stigma shape of Agapetes are divided into two types: 1) small, punctate, inconspicuous Taxonomic treatment stigmas, and 2) thickened capitate, conspicuous Description of new taxa stigmas, and they could be a useful character for Agapetes oxycoccoides J. Murata, Nob. identification. In addition to our new species, Tanaka & Ohi-Toma, sp. nov. [Fig. 2] Agapetes pentastigma, A. sikkimensis Airy Shaw, Type: MYANMAR. Kachin State: 1–4 miles A. setigera, A. hillii, and A. odontocera also have from Shinbweyan toward Pansaung, Hukaung the latter type of stigma. Valley Tiger Reserve, 26°42ʹ26ʺ–43ʹ56ʺN, 96°11ʹ25ʺ–12ʹ33ʺE, cultivated in the greenhouse Phylogenetic relationship of Koishikawa Botanical Garden, J. Murata & The aligned sequence length of trnK intron al. 1401 (TI–holotype, RAF–isotype). (including the matK gene) for all samples Agapetes oxycoccoides is morphologically was 2,564 bp, including 11 coded gaps. The most similar to A. bracteata Hook. f. ex C. B. maximum parsimony analysis resulted in a strict Clarke but differs in the corymbose inflorescence consensus tree from 20 most-parsimonious without distinct bracts, a much more strongly trees with tree length of 477 (CI = 0.8532, RC = reflexed corolla, and pseudoverticillate leaves. 0.7508). Their strict consensus tree with branch Epiphytic shrub up to ca. 1 m tall with sub- lengths was generated (Fig. 1). cylindrical woody tubers, adnate to the tree The phylogenetic tree indicated that Agapetes trunk; branches terete, slightly reticulately is not monophyletic because a part of Vaccinium fissured when old, pubescent. Leaves species were nested in the same clade. Most spirally aggregated at the top of branches species of Agapetes belonged to the clade with (pseudoverticillate); petiole 3–4 mm long, several Vaccinium (the AGA-VAC clade), but A. glabrous; blades ovate-elliptic, 4–6 cm long, 2.0– scortechinii, the only species of Agapetes section 2.7 cm wide, glabrous, veins brochidodromous, Pseudoagapetes (Airy Shaw 1958), is outside midveins raised on lower surface, secondary of the clade. As it has already been pointed out veins 3–5 pairs, with fine veins conspicuous (Stevens 2004), this species should be treated on both surfaces, base attenuate, margin entire, under different genus. revolute, apex obtuse. Inflorescences axillary or In the AGA-VAC clade, A. pentastigma is on leafless stem, 1–6-flowered; peduncle 3.5–4.5 most closely related to A. odontocera and the mm long; bracts small, inconspicuous; pedicels two species formed a clade with A. sikkimensis. spreading, slender, 2–2.5 cm long, pubescent. These species share thickened capitate, Flowers nodding, 1–1.2 cm long, 1 cm wide. conspicuous stigmas. Agapetes oxycoccoides Calyx tube reddish, widely campanulate, 3.5–4.0 forms a clade with A. setigera, but the nucleotide mm across; lobes whitish, triangular, ca. 1.5 mm variations between them are large. Agapetes long, apex acute. Corolla white, lobed more than oxycoccoides is morphologically similar to A. half its length; lobes triangular-lanceolate, 6–7.5 bracteata Hook. f. ex C. B. Clarke rather than mm long, strongly reflexed at anthesis. Stamens A. setigera. However, unfortunately, we could 10, 1–1.2 cm long; filaments flat, ca. 1 mm long, not include A. bracteata in this phylogenetic white, pubescent; thecae maroon, ca. 3 mm long, December 2016 Tanaka et al.: Agapetes from NE Myanmar 105

Table. 2. Comparison of Agapetes oxycoccoides with its related species Characters A. oxycoccoides A. bracteata A. setigera A. lobbii Leaves ovate-elliptic, ovate-lanceolate, ovate-lanceolate, narrowly elliptic, pseudoverticillate sparsely alternate pseudoverticillate sparsely alternate Inflorescence corymbose, racemose, axillary and racemose, racemose, on leafless stem terminal on leafy stem on leafless stem on leafless stem Peduncle short, glabrous, elongated, pubescent, elongated, glandular hairy elongated, glabrous without distinct bracts with distinct bracts Pedicel glabrous glandular hairy glandular hairy glabrous Calyx crimson green green reddish Collora widely campanulate, campanulate, white, sparsely cylindrical, red, campanulate, pale pink, white, pubescent, pubescent near the base, 2.5–3 cm long 2.2–2.9 cm long 1–1.2 cm long 1.2–1.5 cm long Corolla lobes strongly reflexed, white, strongly reflexed, white, short spreading, greenish upper 2/5 of collora 6–7.5 mm long 7–9 mm long strongly reflexed, white Stigma punctate punctate capitate punctate papillate; tubules brownish yellow, 6–6.5 mm Region, Hukaung Valley Tiger Reserve, long; spurs undeveloped. Style white, glabrous, 26°42ʹ32ʺN, 96°11ʹ55ʺ–13ʹ01ʺE, cultivated ca. 7.5 mm long, base greenish, slightly exerted in the greenhouse of Medicinal Plant Garden, from connate anther tubules; stigma punctate. Setsunan University, H. Murata & J. Murata Fruit unknown. 1402 (TI–holotype, RAF, TNS–isotype). Distribution: Thus far known only from the Agapetes pentastigma is similar to A. hillii type locality. Brandis distributed in India and Myanmar but Etymology: The specific epithet is derived differs in having a greenish-yellow collora with from the resemblance of flower shape to ladder-like red stripes (vs. V-shaped pattern), Vaccinium oxycoccos L. in having a strongly and 1/2 divided calyx lobes (vs. deeply divided reflexed corolla. nearly to the base), shorter peduncle and longer Note: The phylogenetic analysis (Fig. 1) slender pedicels (vs. thickend at apex end). indicated that this species was related to A. Epiphytic shrub up to 1.5 m high, with setigera D. Don ex G. Don, but A. setigera sub-cylindrical woody tubers, adnate to the is very much distinct in having racemose tree trunk; branches subterete. Leaves spirally inflorescences with glandular hairs, a tubular arranged on top of branches; leaf blade oblong- and orange-red corolla, and a capitate stigma. lanceolate, 15–18 cm long, 4–6.2 cm wide, dull Other similar species are compared in Table yellow green on upper surface, whitish green 2, although they were not available for our beneath, apex acuminate, base attenuate, margin molecular phylogenetic analysis. inconspicuously and distantly sinuous, glabrous on both surfaces, leathery, sessile; midrib Agapetes pentastigma J. Murata, Nob. prominent above, raised on lower surface; Tanaka & H. Murata, sp. nov. [Fig. 3] secondary veins 20–25 pairs. Inflorescence Type: MYANMAR. Kachin State: along corymbose, 4–7 flowered, leaf-axillary; the Ledo Road, between Namyung and peduncle very short, glabrous; pedicel 2.5–3 Shinbweyang, 5–7 miles from Shinbweyang cm long, articulate at both ends, grooved; bracts toward Namyung, in border area of Sagaing attached at the base of the pedicel, lanceolate, 106 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue

Fig. 3. Agapetes pentastigma J. Murata, Nob. Tanaka & H. Murata. A. Habit. B. Close view and vertical section (right) of flowers showing stamens and stigma. C. Front view of flowers. D. Close view of thickened capitate stigma with pentagonal appearance at the top. Scale bar: 5 cm for A, 2 cm for B–C, 3 mm for D. December 2016 Tanaka et al.: Agapetes from NE Myanmar 107

Fig. 4. A, B. Agapetes hillii Brandis. A. Close view of a branch with flowers in natural habitat. B. Natural habitat. C, D. Agapetes burmanica W. E. Evans. C. Close view and vertical section (right) of flowers showing stamens and stigma. D. Close view of inflorescence. Scale bar: 4 cm for A, 2 cm for C–D. apex attenuate, 1.5–2 mm long, glandular hairs bands, 4.5–5 cm long, 1–1.5 cm wide, glabrous; at margin. Calyx tube yellowish-green, tubular, lobes olive green, triangular-lanceolate, 7–8 mm 4.5–5 mm long, 5–7 mm wide, glabrous; calyx long, reflexed at anthesis. Stamens 10, 4–4.5 lobes 5, lobes 1/2 of the length, triangular, cm long; filaments flat, ca. 5.5 mm long, white, 5–6 mm long, reddish, apex attenuate. Corolla pubescent; thecae reddish brown, 6.5–7 mm tubular, olive green with red ladder-like pattern long, papillate; tubules brownish yellow, 4–4.5 consisting of longitudinal stripes and transverse cm long. Style reddish-white, glabrous, 4–5.5 108 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue

Table. 3. Comparison of Agapetes pentastigma with its related species Characters A. pentastigma A. odontocera A. burmanica A. hillii Leaf oblong-lanceolate lanceolate oblong-lanceolate lanceolate Inflorescence corymbose, axillary on small corymbose, corymbose, corymbose, terminal young leafy stem on leafless old stem on leafless old stem Peduncle very short, glabrous almost lacking almost lacking, glabrous glabrous Pedicel glabrous puberulous glabrous glabrous Calyx divided to 1/2, lobes divided to 1/2, shortly lobed, deeply divided nearly to triangular, reddish green lobes triangular lobes triangular, red the base, lobes narrowly triangular Collora about 4 cm long, greenish about 1.5 cm long when dry, about 6 cm long, pink about 5 cm long, yellow with red longitudinal (reddish yellow with red with red transverse zig- greenish-yellow with stripes and red transverse transverse zig-zag bands) zag bands red V-shaped patterns (non zig-zag) bands Corolla lobes green green green green Stigma capitate, green punctate, brownish yellow punctate, green capitate, green cm long, exerted ca. 4 mm from the connate Hukaung Valley Tiger Reserve, J. Murata & al. 041355 anther tubules; stigma capitate, 3–3.5 mm wide, (MBK), and H. Murata 1601 (TI, TNS, RAF). green. Fruit unknown. Distribution: China and Myanmar. Distribution: Thus far known only from the Note: This species was described from type locality. N.E. Burma based on the Forrest specimen Etymology: The specific epithet is derived (Htawgaw, G. Forrest 24981, E). The plant from its representative character, a large collected in northwestern Myanmar is slightly thickened capitate stigma with pentagonal different from the typical A. burmanica in having appearance at the top. very short corolla lobes with rounded apex. To clarify the range of variation of calyx shape in A. Note: The phylogenetic analysis (Fig. 1) burmanica further observations in the field are indicated that, among the species examined, this required. species was closest to A. odontocera Benth. & Hook. f. distributed in India. Morphologically, Agapetes hillii Brandis, Indian Trees: 404 however, this species is easily distinguished (1906); Kress & al., Contr. U.S. Natl. Herb. 45: from the latter by having much larger flowers 213 (2003). [Fig. 4A, B] and a distinctly capitate stigma. Other similar Specimen examined: MYANMAR. along the Ledo species are compared in Table 3, although some Road, between Namyung and Shinbwiyang, 5–7 miles of them were not available for our molecular from Shinbwiyang toward Namyung, in border area of phylogenetic analysis. Sagaing Region, J. Murata & al. 041479 (MBK, TI). Distribution: India and Myanmar. Other collected taxa from northwestern Myanmar Agapetes burmanica W. E. Evans in Notes Agapetes setigera D. Don ex G. Don, Gen. Roy. Bot. Gard. Edinburgh 15(74): 199, pl. 219 Hist. 3: 862 (1832); C. B. Clarke, in Hook. f., Fl. (1927); Kress & al., Contr. U.S. Natl. Herb. 45: Brit. Ind. 3: 443 (1881); Brandis, Indian Trees: 213 (2003); Fang & Stevens in Wu & Raven, Fl. 404 (1906); Nath D. M., Bot. Survey S. Shan China 14: 507 (2005). [Fig. 4C, D] State: 132 (1960); Kress & al., Contr. U.S. Natl. Specimen examined: MYANMAR. Kachin State. Herb. 45: 214 (2003). [Fig. 5] East of Shinbweyang, in border area of Sagaing Region, Specimen examined: MYANMAR. Kachin State. December 2016 Tanaka et al.: Agapetes from NE Myanmar 109

Fig. 5. Agapetes setigera D. Don ex G. Don. A. Habit. B. Close view of flower showing stamens and stigma by removing collora. C. Front view of flower showing capitate stigma. Scale bar: 2 cm for A, 1 cm for B–C.

Along the Ledo Road, near Shinbweyang, in border area of Botanical Garden, University of California Sagaing Region, Hukaung Valley Tiger Reserve, cultivated Botanical Garden, and Kyoto Botanical in the greenhouse of Koishikawa Botanical Garden, J. Gardens. This study was partially supported by Murata & T. Ohi-Toma 1602 (TI, RAF). Distribution: India and Myanmar. JSPS KAKENHI Grant Numbers 13375003 (to J. Murata). The authors are grateful to the Forest References Department, Ministry of Natural Resources Airy-Shaw H. K. 1935. Studies in the : I. New and and Environmental Conservation of Myanmar less known species of Agapetes. Bull. Misc. Inform. for permission to conduct this study in the Kew. 1935: 24–53. Hukaung Valley Tiger Reserve, and for their Airy-Shaw H. K. 1948. Studies in the Ericales: V. Further notes on Agapetes. Kew Bull. 3: 77–104. support and collaboration. Thanks are due to Airy-Shaw H. K. 1958. Studies in the Ericales: XI. Further Prof. M. Kanzaki and Dr. A. Nakanishi and new species and notes on the Agapetes of continental the following institutions for providing us the Asia. Kew Bull. 13: 468–512. materials for molecular phylogenetic analysis; Airy-Shaw H. K. 1960a. Studies in the Ericales: XII. A Royal Botanical Garden Edinburgh, Ofuna few new or noteworthy Agapetes from the Calcutta herbarium. Kew Bull. 14: 110–113. Botanical Garden, Sakuya Konohana Kan, Airy-Shaw H. K. 1960b. Studies in the Ericales: XIII. A San Francisco Botanical Garden, South China new Agapetes from the Shan States of Burma. Kew 110 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue

Bull. 14: 229–230. Principal Climbers, etc. Recorded from Burma. Forest Banik D. and Sanjappa M. 2007. Five new varieties of Department, CCF, Rangoon. Agapetes (Ericaceae) from India. J. Bot. Inst. Texas 1: Ohi-Toma T., Wu S. G., Yadav S. R., Murata H. and 903–912. Murata J. 2010. Molecular phylogeny of Typhonium Bremer B., Bremer K., Heidari N., Erixon P., Olmstead sensu lato and its allied genera in the tribe Areae of the R.G., Anderberg A. A., Källersjö M. and Barkhordarian subfamily Aroideae (Araceae) based on sequences of E. 2002. Phylogenetics of based on 3 coding six chloroplast regions. Syst. Bot. 35: 244–251. and 3 non-coding chloroplast DNA markers and the Rodger A. 1922. List of Trees, Shrubs, Herbs and Principal utility of non-coding DNA at higher taxonomic levels. Climbers, etc. Recorded from Burma with Vernacular Molec. Phylogen. Evol. 24: 274–301. Names. (ed. 2). Forest Department, Rangoon. Conlon T. 2015. Agapetes D. Don ex G. Don – Jewels Stevens P. F. 1985. Notes on Vaccinium and Agapetes of the east at the Royal Botanic Garden Edinburgh. (Ericaceae) in Southeast Asia. J. Arn. Arb. 66: 471– Sibbaldia no. 13: 61–81. 490. Cruickshank R. D. and Ko Ko 2003. Geology of an amber Stevens P. F. 2004. New taxa in Paphia and locality in the Hukawng Valley, Northern Myanmar. J. Dimorphanthera (Ericaceae) in Papuasia and the Asian Earth Sciences 21: 441–455. problem of generic limits in Vaccinieae. Edinb. J. Bot. Dunning L. T. and Savolainen V. 2010. Broad-scale 60: 267–298. amplification ofmatK for DNA barcoding. Bot. J. Linn. Swofford D. L. 2002. PAUP*: Phylogenetic analysis using Soc. 164: 1–9. parsimony (and other methods), version 4.0 beta 10. Fang R. Z. and Stevens P. F. 2005. Agapetes. In: Wu. Z. Y. Sinauer, Sunderland. and Raven P. H. (eds.), Flora of China. 14: 504–517. Tanaka N. and Hayami Y. 2011. Begonia kachinensis (Sect. Science Press, Beijing and Missouri Botanical Garden Sphenanthera, Begoniaceae), a new species from Press, St. Louis. Myanmar. Acta Phtotax. Geobot. 61: 151–154. Hundley H. G. 1987. List of Trees, Shrubs, Herbs and Tanaka N. and Hughes M. 2007. Begonia (Sect. Principal Climbers, etc. Recorded from Burma with Sphenenthera) hayamiana (Begoniaceae), a new Vernacular Names. 4th revised ed. Forest Department, species from in Northern Myanmar. Acta Phytotax. Swe Daw Oo Press, Rangoon. Geobot. 58: 83–86. Hundley H. G. and Chit K. K. 1961. List of Trees, Shrubs, Tanaka N., Kobayashi S., Ohi-Toma T. and Murata J. Herbs and Principal Climbers, etc. Recorded from 2006. New or noteworthy plant collections from Burma with Vernacular Names. 3rd ed. Superintendent, Myanmar (1): Hydrobryum japonicum, Balanophora Goverment Printing and Staty., Rangoon. subcupularis, Rhopalocnemis phalloides and Sonerila Johnson L. A. and Soltis D. E. 1994. MatK DNA sequences laeta. J. Jpn. Bot. 81(6): 324–331. and phylogenetic reconstruction in Saxifragaceae s.str. Tanaka N., Ohi-Toma T., Aung M. M. and Murata J. Syst. Bot. 19: 143–156. 2016. Systematic Notes on the genus Hedychium Kato H., Oginuma K., Gu Z., Hammel B. and Tobe H. (Zingiberaceae) in Myanmar. Bull. Natl. Mus. Nat. 1998. Phylogenetic relationships of Betulaceae based Sci., Ser. B, 42: 57–66. on matK sequences with particular reference to the Tong Y. H. and Xia N. H. 2015. New taxa of Agapetes position of Ostryopsis. Acta Phytotax. Geobot. 49: (Ericaceae) from Myanmar. Phytotaxa 184: 39–45 89–97. (DOI: http://dx.doi.org/10.11646/). Kress W. J., DeFilipps R. A., Farr E. and Kyi D. Y. Y. 2003. Watthana S. 2012. The genus Agapetes D. Don ex G. Don A Checklist of the Trees, Shrubs, Herbs, and Climbers (Ericaceae) in Thailand. Thai J. Bot. 4: 99–116. of Myanmar. Cont. U.S. Natl. Herb. 45: 1–590. Kron K. A., Fuller R., Crayn D. M., Gadek P. A. and Quinn Appendix C. J. 1999. Phylogenetic relationships of epacrids and List of samples used in the phylogenetic analysis (the vaccinioids (Ericaceae s.l.) based on matK sequence species, source, voucher specimen, and DDBJ/GenBank data. Pl. Syst. Evol. 218: 55–65. accession number of trnK intron). Kron K. A., Powell E. A. and Luteyn J. L. 2002. Agapetes brandisiana W. E. Evans: Cultivated in South Phylogentic relationships within the bluberry tribe China Botanical Garden, Liao s.n. (IBSC), LC168848. A. (Vaccinieae, Ericaceae) based on sequence data burmanica W. E. Evans, Myanmar: Kachin, J. Murata 1601 from matK and nuclear ribosomal ITS regions, with (TI), LC168849. A. buxifolia Nutt. ex Hook. f., Cultivated comments on the placement of Satyria. Amer. J. Bot. in Royal Botanic Garden Edinburgh (19696067), Conlon 89: 327–336. C505 (E), LC168850. A. flava (Hook. f.) Sleumer, Buhtan: Kurz S. 1877. Forest Flora of British Burma.Vol. 2. p. 87. Cultivated in Royal Botanic Garden Edinburgh (19822403), The Superintendent of Government Printing, Calcutta. Conlon C329 (E), LC168851. A. hosseana Diels, Thailand: Lace J. H. 1912. List of Trees, Shrubs, Herbs and Cultivated in Royal Botanic Garden Edinburgh (19672592), December 2016 Tanaka et al.: Agapetes from NE Myanmar 111

Conlon C340 (E), LC168852. A. incurvata (Griff.) A. sikkimensis Airy Shaw, Bhutan: Cultivated in Royal Sleumer var. incurvata, Nepal: Cultivated in Koishikawa Botanic Garden Edinburgh (19842032), Sinclair & Long Botanical Garden, University of Tokyo (2016-002), T. Ohi- 5778 (E), LC168869. A. smithiana Sleumer, Cultivated Toma s.n. (TI), LC168853. A. incurvata (Griff.) Sleumer in Sakuya Konohana Kan, J. Murata s.n. (TI), LC168870. var. hookeri (C. B. Clarke) Airy Shaw, Nepal: Cultivated A. thailandica Watthana, Thailand: Cultivated in Royal in Royal Botanic Garden Edinburgh (19892150), Kew/ Botanic Garden Edinburgh (19991631), Argent 13 (E), Edinburgh Kanchenjunga Expedition (1989) 142 (E), LC168871. A. variegata D. Don ex G. Don, cultivated in LC168854. A. lacei Craib, Cultivated in Ofuna Botanical Royal Botanic Garden Edinburgh (19696068), Conlon Garden, T. Ohi-Toma s.n. (TI), LC168855. A. lobbii C. B. C124 (E), LC168872. Andromeda polifolia L., Japan: Clarke, Thailand: Doi Inthanon, Nakanishi s.n. (QBG), Cultivated in Koishikawa Botanical Garden, University of LC168856. A. mannii Hemsl., Myanmar: Chin, Kuroiwa Tokyo, T. Ohi-Toma s.n. (TI), LC168873. Dimorphanthera & al. 030270 (MBK), LC168857. A. aff. miniata (Griff.) kempteriana Schltr., cultivated in Kyoto Botanical Garden, Hook. f., Myanmar: Chin, Kuroiwa & al. 030448 (MBK), J. Murata s.n. (TI), LC168874. Paphia meiniana (F. LC168858. A. moorei Hemsl., Myanmar: Chin, Kuroiwa Muell.) Schltr., Australia: Cultivated in San Francisco & al. 030333 (MBK), LC168859. A. oblonga Craib, Botanical Garden (2001-0421), Bourell s.n. (CAS), cultivated in South China Botanical Garden, Liao s.n. LC168875. P. stenantha Schltr., Papua New Guinea: (IBSC), LC168860. A. odontocera (Wight) Hook. f., India: Milne Bay, cultivated in Royal Botanic Garden Edinburgh Cultivated in Royal Botanic Garden Edinburgh (19751313), (19762042), Woods 2521 (E), LC168876. Vaccinium Chamberlain 106 (E), LC168861. A. oxicoccoides J. amamianum Hatus., Japan: Amami, cultivated in Murata, Nob. Tanaka & Ohi-Toma, Myanmar: Kachin. Koishikawa Botanical Garden, University of Tokyo, T. Ohi- J. Murata & al. 041215 (TI), LC168862. A. aff. pensilis Toma s.n. (TI), LC168877. V. bulleyanum (Diels) Sleumer, Airy Shaw, China: Cultivated in Royal Botanic Garden cultivated in South China Botanical Garden, Liao s.n. Edinburgh (19962294), Gaoligong Shan Biotic Survey (IBSC), LC168878. V. fragile Franch., China: Cultivated in Expedition (1996) 8147 (E), LC168863. A. pentastigma Koishikawa Botanical Garden, University of Tokyo, T. Ohi- J. Murata, Nob. Tanaka & H. Murata, Myanmar: Kachin, Toma s.n. (TI), LC168879. V. gaultheriifolium Hook. f. ex J. Murata & al. 041479 (TI), LC168864. A. saligna C.B.Clarke, Nepal: Cultivated in University of California (Hook. f.) Hook. f., Bhutan: Cultivated in Royal Botanic Botanical Garden, Edinburgh Makalu Expedition (1991) Garden Edinburgh (19842035), Sinclair & Long 5785 (E), 778 (E), LC168880. V. leucobotrys (Nutt.) G. Nicholson, LC168865. A. scortechinii (King & Gamble) Sleumer, cultivated in South China Botanical Garden. Liao s.n. Malaysia: Cultivated in San Francisco Botanical Garden (IBSC), LC168881. V. nummularia Hook. f. & Thomson (1997-0424), Bourell s.n. (CAS), LC168866. A. serpens ex C. B. Clarke, cultivated in Sakuya Konohana Kan. T. (Wight) Sleumer, India: Cultivated in Koishikawa Ohi-Toma s.n. (TI), LC168882. V. oxycoccos L., Japan: Botanical Garden, University of Tokyo (89-2364), T. Ohi- Nagano, J. Murata s.n. (TI), LC168883. V. scopulorum W. Toma s.n. (TI), LC168867. A. setigera D. Don ex G. W. Sm., cultivated in South China Botanical Garden, Liao Don, Myanmar: Kachin. J. Murata 1602 (TI), LC168868. s.n. (IBSC), LC168884.

田中伸幸a，大井・東馬哲雄b，邑田裕子c，M. M. Aungd， b 邑田 仁 ：ミャンマー植物についての新知見 (9) 著者らが実施してきたミャンマー西北部の植物調査 Tanaka & H. Murata は柱頭の先が 5 角形に膨らむ特徴を によりツツジ科 Agapetes 属植物 5 種が採集された．形 持っており，よく似た A. hillii から花冠上の脈のパター 態比較および分子系統解析による近縁種の検討の結 ンや花序柄の長さで区別される． 果，このうち 2 種は新種であることが明らかとなった （a国立科学博物館植物研究部， ので記載・発表する．Agapetes oxycoccoides J. Murata, b東京大学大学院理学系研究科附属植物園， Nob. Tanaka & Ohi-Toma は ツ ル コ ケ モ モ Vaccinium c摂南大学薬学部薬用植物園， oxycoccos に似た花を苞が発達しない散房花序につける dミャンマー・林業省野生生物保護局） ことが特徴である．Agapetes pentastigma J. Murata, Nob.