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Goniothalamus (Blume) Hook.f. & Thomson, comprising over 130 species of trees and shrubs, is one of the largest palaeotropical genera in the Annonaceae, mainly distributed from India and Sri Lanka to tropical Australia and the South Pacific Islands (Saunders & Chalermglin, 2008; Nakkuntod et al., 2009; Turner, 2014; Thomas et al., 2017) . In China, 11 species are currently recognized (Li & Gilbert, 2011) .
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During monitoring the living collections of Annonaceae in Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences from 2018 to 2021, two Goniothalamus species with flowers and fruits were observed. The two species were introduced to Xishuangbanna Tropical Botanical Garden (XTBG) in 1997 and 2002. After a detailed examination of the morphological characters of our materials, related literatures, and herbarium specimens, we identified them as Goniothalamus sesquipedalis (Colebr. ex Wall.) Hook. f. & Thomson and G. peduncularis King & Prain, respectively. G. sesquipedalis was originally distributed in India, Bangladesh and Myanmar (Turner, 2015, 2018) , and G. peduncularis was only reported in Myanmar (Turner, 2015, 2018) . The two species therefore represent two new records for the flora of China, in which we update the descriptions and illustrate them herein.
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皱叶哥纳香  ;新拟,图版I;图版Ⅲ: A
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Goniothalamus sesquipedalis (Colebr. ex Wall.) Hook. f. & Thomson, Fl. Ind.: 108, 1855. (Plate I; Plate Ⅲ: A)
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Type: Guatteria sesquipedalis Colebr. ex Wall. in Wallich, Pl. Asiat. Rar.3: 42, t.266, 1832. Type: Wallich, Pl. Asiat. Rar.3: t.266, 1832. (lectotype, designated by Turner, Nordic J. Bot.33: 270, 2015) .
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Shrubs, to 2 m tall. Young branches glabrous. Leaf laminas oblong, 22-47 cm long, 5.7-10.5 cm wide, length/width ratio 3.4-7.2, apex acuminate to caudate, base cuneate, sometimes attenuate, coriaceous, glabrous abaxially and adaxially; midrib glabrous and (strong) prominent abaxially, glabrous and impressed adaxially; secondary veins 16-22 pairs, interarching5-10 mm from recurved margin, impressed adaxially, raised abaxially; tertiary veins reticulate, distinct; petioles 1.3-2 cm long, 2.5-5 mm in diameter, glabrous. Flowers greenish-yellow, solitary, axillary or extra-axillary, often on the main trunk (cauliflory) and on older branches (ramiflory) ; flowering pedicels 3-5 mm long; pedicel bracts 2-6, 2-4 × 1.5-3 mm. Sepals 3, 5-11 mm long, 5-7 mm wide, basally connate, apex acute to obtuse, ovate, puberulent abaxially, sparsely puberulent adaxially, greenish-yellow. Outer petals 3, 25-28 mm long, 6-8 mm wide, length/width ratio 3.1-4.6, lanceolate, puberulent abaxially and adaxially, greenish-yellow. Inner petals 3, shortly clawed and free in lower part, cohering and forming an ovate-caudate cone at apex, 15-18 mm long, 5-6.5 mm wide, length/width ratio 2.5-3, ovate-lanceolate, puberulous abaxially and adaxially, sometimes shortly lanate abaxially and adaxially in upper part, yellowish green, base obtuse to a4.5-5 mm claw. Stamens 70-80 per flower, 2-2.2 mm long, 0.8-0.9 mm wide; connectives apiculate. Carpels ca.12 per flower, ovary 2.5-3 mm long, light green, white pubescent; stigma and style2-2.2 mm long, puberulous. Immature fruits green, mature fruits orange-red; fruiting pedicels 7-8.5 mm long, 2.5-3 mm in diameter. Monocarps one seeded, fresh monocarps 21-24 mm long, 10-12 mm wide, length/width ratio 2-2.2, dry monocarps 16-18.8 mm long, 6.9-7.4 mm wide, length/width ratio 2.2-2.7, ovoid to ellipsoid-ovoid, base rounded, apex attenuate, sparsely puberulous, glossy, pericarp medium-thick, ca.2 mm thick, stipes 2-3 mm long, ca.3 mm in diameter. Seeds 13-14 mm long, 6.5-7 mm wide, ovoid, testa slightly with latitudinal stripes, dark brown to black brown, aril yellowish brown, endosperm ruminations lamellate.
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Distribution: China (Yunnan) (new record) ; India (Sikkim, Assam, Nagaland, Manipur, Meghalaya) (Grierson, 1984; Karthikeyan et al., 2009) ; Bangladesh (Khanam & Rahman, 2002) ; Myanmar (Kress et al., 2003; Kang et al., 2017) .
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Additional specimens examined: China(中国). Yunnan(云南): Yingjiang County(盈江县),Xima Township(昔马乡),Nabangba(那邦坝),24°44′ N,97°33′ E,400 m,5 November 1974,Tao Guoda(陶国达)013190(specimen No. 001927 [barcode,HITBC0040552!],specimen No. 001928 [barcode,HITBC0040553!]);the same location,April1979,s. coll.(specimen No. 066463 [barcode,HITBC0040554!]);Yingjiang County(盈江县),Xima Township(昔马乡),24°45′ N,97°42′ E,260 m,3 December 1981,Tao Guoda(陶国达)12774(specimen No. 001929 [barcode,HITBC0040551!]);Yingjiang County(盈江县),Nabangba(那邦坝),Tongbiguan Natural Reserve(铜壁关自然保护区),10 October 2011,Zhou Shishun(周仕顺)11197(HITBC 0040504);Yingjiang County(盈江县),Labang(拉邦),voucher from a cultivated plant at the Xishuangbanna Tropical Botanical Garden,Chinese Academy of Sciences,21 July 2019,Yang Bin(杨斌),XTBG0077(HITBC);15 February 2020,Yang Bin(杨斌),XTBG0079(HITBC);9 May 2020,Yang Bin(杨斌)&;Xiao Chunfen(肖春芬),XTBG0114(HITBC);24 May 2021,Yang Bin(杨斌),XTBG0243(HITBC)。
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Notes:The specimens of Goniothalamus sesquipedalis deposited in Herbarium of Xishuangbanna Tropical Botanical Garden (HITBC specimen No. 001927, 001928, 001929, 066463) was previously misidentified as Goniothalamus multiovulatus Ast, which is native to Vietnam (Turner, 2018) . G. multiovulatus can be differentiated from G. sesquipedalis by having 4.5-5.5 cm long, 2-2.5 cm wide, cylindrical monocarps with 2-4 hairy seeds per monocarp (Ast, 1938) . Hou (2003) misidentified them (HITBC specimen No. 001927, 001928, 001929) as Goniothalamus macrophyllus (Bl.) Hook. f. et Thoms, which is distributed in Malaysia, Singapore, Indonesia, Thailand (Saunders, 2002; Saunders & Chalermglin, 2008; Turner, 2018) . G. macrophyllus differs from G. sesquipedalis by having the leaves with a distinctive and diagnostically important ‘granular’ appearance abaxially because of immersion of tertiary and lower order veins (Saunders, 2002; Saunders & Chalermglin, 2008) . G. sesquipedalis differs from G. macrophyllus in several important respects: the leaves of G. sesquipedalis with secondary venation very prominent adaxially and forming intramarginal vein, tertiary venation prominent and reticulate, whereas the leaves of G. macrophyllus with secondary venation ± plane to impressed adaxially and not forming intramarginal vein, tertiary venation unconspicuous (Saunders, 2002) ; G. sesquipedalis can also be distinguished from G. macrophyllus by having3-5 mm (vs.5-11.5 mm) long flowering pedicels; lanceolate, greenish-yellow (vs. ovate, orange-pink) outer petals when mature; ovate-lanceolate (vs. ovate) , 15-18 mm (vs.7-15 mm) long inner petals (Saunders, 2002; Saunders & Chalermglin, 2008) .
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Plate I Goniothalamus sesquipedalis (Colebr. ex Wall.) Hook. f. & Thomson
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长梗哥纳香  ;新拟,图版Ⅱ;图版Ⅲ: B
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Goniothalamus peduncularis King & Prain, J. Asiat. Soc. Bengal, Pt.2, Nat. Hist.67: 284, 1898. (Plate Ⅱ; Plate Ⅲ: B)
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Type: Burma, Upper Burma, Kachin Hills, Sima Road, 1 000 feet, 1897, Shaik Mokim s.n. (lectotype CAL [CAL000004500], designated by Turner, Gard. Bull. Singapore70 (1) : 523, 2018; isolectotype K [K000380823], Plate Ⅲ: B) .
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Goniothalamus lii X. L. Hou &;Y. M. Shui,Acta Bot. Yunnan.,25(3): 258,2003. Syn. nov. Type: China(中国). Yunnan(云南): Yingjiang County(盈江县),Xima Township(昔马乡),Nabangba(那邦坝),300 m,7 November 1974,Tao Guoda(陶国达)013254(holotype,KUN1263140,Plate Ⅲ: C;isotype,HITBC [specimen No. 001921,barcode 0040594!])
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Plate Ⅱ Goniothalamus peduncularis King & Prain
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Shrubs, to 4 m tall. Young branches glabrous. Leaf laminas oblong, elliptic-oblong to oblanceolate-oblong, 15-23 cm long, 4.4-7 cm wide, apex acuminate, base cuneate, thinly papery, glabrous abaxially and adaxially; midrib raised abaxially, impressed adaxially; secondary veins 10-16 pairs, interarching3-7 mm from recurved margin, impressed adaxially, raised abaxially; tertiary veins reticulate, raised abaxially, unconspicuous adaxially; petioles 8-13 mm long, with shallow furrows adaxially, glabrous, black when dry. Inflorescence often solitary, sometimes 2-flowered, axillary, often on the main trunk (cauliflory) and rarely on older branches (ramiflory) ; peduncle degenerate, 1-2 mm long; flowering pedicels 9-15 mm long; pedicel bracts 2-4, 1-2 mm long, ca.1 mm wide. Sepals 3, ovate, 5-7 mm long, 3-6 mm wide, basally connate, apex acute to obtuse, sparsely puberulent abaxially, conspicuous puberulent along margin, puberulent adaxially, yellowish-green. Outer petals 3, 25-37 mm long, 12-16 mm wide when fresh, 24-34 mm long, 9-12 mm wide when dry, ovate to ovate-lanceolate, puberulent abaxially and adaxially, often yellowish-green, grey white when mature. Inner petals 3, shortly clawed and free in lower part, cohering and forming an ovate-attenuate to ovate-acuminate cone at apex, 13-15 mm long, 5-6 mm wide when fresh, 12-14 mm long, 4-5 mm wide when dry, ovate-lanceolate to oblong-lanceolate, puberulous abaxially and adaxially, densely in upper part, greenish yellow, base obtuse to a2-3 mm claw, apex attenuate to acuminate. Stamens ca.100 per flower, 2.5 mm long; connectives slightly apiculate, sometimes hemispherical when dry, yellowish brown pubescent. Carpels 20-30 per flower, ovary ca.2 mm long, greenish-yellow, white pubescent; stigma and style2-3 mm long. Immature fruits green, mature fruits deep blackish blue; fruiting pedicels 1.2-1.5 (-4.8) cm long. Monocarps 4-25, 1-2 seeded per monocarp, fresh monocarps 17-25 mm long, 8-9 mm wide, ellipsoid to ellipsoid-oblong, base rounded, apex rounded with a small tip, sub-glabrous to sparsely puberulent, pericarp ca.1 mm thick when fresh, stipes 5-12 mm long. Seeds 10-13 mm long, 6.5-8 mm wide, ellipsoid, testa with white puberulous hairs outside, yellowish brown when immature, endosperm ruminations lamellate.
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Plate Ⅲ Type specimens and voucher specimen in the protologue of four species of Goniothalamus
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Distribution:China (Yunnan: Yingjiang, Cangyuan, Menghai) (new record) ; Myanmar (Kress et al., 2003; Turner, 2015, 2018) .
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Additional specimens examined: China(中国). Yunnan(云南): Yingjiang County(盈江县),Xima Township(昔马乡),Nabangba(那邦坝),300 m,10 December 1978,Tao Guoda(陶国达)17895(specimen No. 002077 [barcode,HITBC0040544!]);Menghai County(勐海县),Mengman Town(勐满镇),995 m,22°9′25.23″ N,100°5′34.29″ E,28 June2021,Zhou Shishun(周仕顺)19431(HITBC);909 m,22°10′50.79″ N,100°4′32.98″ E,29 June2021,Zhou Shishun(周仕顺)19474(HITBC);Cangyuan County(沧源县),Banhong Township(班洪乡),Fabaomangkuhe(法保芒库河)700 m,30 May 1974,Li Yanhui(李延辉)11770(specimen No. 001923 [barcode,HITBC0040547!],specimen No. 001925 [barcode,HITBC0040545!]);780-800 m,2 June1974,Li Yanhui(李延辉)11873(specimen No. 001926 [barcode,HITBC0040546!],Plate Ⅲ: C1);Banlao Township(班老乡),Shangbanlao(上班老)900 m,21 May 1975,Li Yanhui(李延辉)20975(specimen No. 001922 [barcode,HITBC0040548!]);Yingjiang County(盈江县),Longmen(陇门),voucher from a cultivated plant at the Xishuangbanna Tropical Botanical Garden,21 April2021,Xiao Wenqiang(肖文强),C400663(HITBC0031632!);6 May 2019,Sheng Caiyu(盛才余),C400846(HITBC0031129!);26 May 2020,Yang Bin(杨斌)&;Xiao Chunfen(肖春芬),XTBG0128(HITBC).
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Notes: Goniothalamus lii X. L. Hou & Y. M. Shui was characterized by its oblong or oblong-lanceolate leaf blades with (10-) 13-21 pairs of lateral veins, pedicels 9-13 mm long, sepals ca.5 × 4 mm, outer petals 2.5 × 1.3 cm, the stamens with hemispherical connectives, sometimes with densely brownish puberulent (Hou & Shui, 2003) , all these characters are consistent with G. peduncularis. Moreover, examination of relevant type materials of G. lii and observations on living plants of G. peduncularis introduced from Yingjiang County show that both species share the liner style and ellipsoid monocarps. Therefore, G. lii is considered as conspecific with G. peduncularis, and we propose it as a synonym with the latter in this study. In Flora of China, Li & Gilbert (2011) synonymized G. lii with G. yunnanenis W. T. Wang (Plate Ⅲ: D) (Wu & Wang, 1957) without further explanation. We don not agree with that treatment, however. As Hou & Shui (2003) mentioned in the protologue of G. lii, it is morphologically similar to G. yunnanenis, but clearly differs by having10-21 (vs.7-9) pairs of lateral veins and pedicels 9-13 mm (vs. ca.4 mm) long. Moreover, based on our observations of living plants, the distinction between these two species is obvious on many other characters. G.lii (here as G. peduncularis) has grey white outer petals when mature (Plate Ⅱ: d3) , styles linear (Plate Ⅱ: d7) , monocarp stipes 5-12 mm long (Plate Ⅱ: E, F, G) , and monocarps deep blackish blue when mature (Plate Ⅱ: F) ; whereas G. yunnanenis has redidish brown outer petals when mature, styles stick shape (Jiang & Li, 1979) , and monocarp stipes very short, less than 2 mm, and monocarps orange red to dark red when mature.
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Acknowledgement We are grateful to Mr. WANG Yuan for providing the specimens image and Prof. ZHU Xiangyun for useful discussion.
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参考文献
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AST MS, 1938. Identité de deux espèces d′Anonacées: Oxymitra gabriaciana Baillon et Goniothalamus saïgonensis Pierre mss ex Finet et Gagnepain. Espèces nouvelles d′Indo-Chine [J]. Bull Soc Bot France, 85(1): 50-53.
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摘要
番荔枝科(Annonaceae)是基部被子植物木兰目(Magnoliales)中较进化且物种数最多的科。目前的系统发育研究将番荔枝科划分为4个亚科,即蒙蒿子亚科(Anaxagoreoideae)、澄光木亚科(Ambavioideae)、番荔枝亚科(Annonoideae)和排石木亚科(Malmeoideae),有107属,2400多种,中国原产21 属约110 种。番荔枝科泛热带分布,是热带植物区系的优势类群,中国云南盈江位于云南省最西部边境,与缅甸东北部接壤,并与印度的东阿萨姆较近,植物区系处于东南亚(印度—马来西亚)热带生物区系向东亚亚热带-温带生物区系的过渡地带,属典型热带北缘性质,在植被地理和生物地理上十分重要,成为生物多样性保护的关键和热点地区。该区的热带雨林是印度阿萨姆和缅甸北部的热带雨林向东和向北扩散分布的边缘类型,是东南亚热带雨林在纬度和海拔分布上的极限类型。该文报道了采自中国云南省盈江县,引种保存于中国科学院西双版纳热带植物园的番荔枝科哥纳香属2个中国新记录种,即皱叶哥纳香 [Goniothalamus sesquipedalis (Colebr. ex Wall.) Hook. f. & Thomson]和长梗哥纳香(G. peduncularis King & Prain)。Flora of China将盈江哥纳香(G. lii X. L. Hou & Y. M. Shui)处理为云南哥纳香(G. yunnanensis W. T. Wang)的异名,基于活植物观察、馆藏标本和文献研究,该文对盈江哥纳香的分类地位进行了澄清,将其处理为长梗哥纳香的异名。皱叶哥纳香原记载产于印度、孟加拉国和缅甸等地,长梗哥纳香仅产于缅甸,该文对它们进行了补充描述,并提供彩色图版以便于鉴别。凭证标本存放于中国科学院西双版纳热带植物园标本馆(HITBC)。哥纳香属2个新记录的发现,丰富了中国番荔枝科植物多样性的认识,为中国云南热带植物区系属于热带亚洲(印度—马来西亚)植物区系,以及与缅甸北部、印度东北部植物区系的关系增加了例证。
Abstract
Goniothalamus sesquipedalis (Colebr. ex Wall.) Hook. f. & Thomson and G. peduncularis King & Prain are reported as two new records for the flora of China, the two species were found in Yingjiang County, Yunnan Province, China and conserved in Xishuangbanna Tropical Botanical Garden. G. lii X.L. Hou & Y.M. Shui has been treated as synonym with G. yunnanensis W. T. Wang in Flora of China. Here, we clarified its taxonomical status and treated as a new synonym with G. peduncularis based on living plants observation, the type specimens and the literatures revision. G. sesquipedalis was previously known from India, Bangladesh and Myanmar, and G. peduncularis was only in Myanmar. In this paper, we update the description and illustrate them based on the herbarium specimens and living collections. The voucher specimens are deposited in the Herbarium of Xishuangbanna Tropical Botanical Garden (HITBC).
Keywords
Goniothalamus ; Annonaceae ; new record ; Yingjiang County ; taxonomy