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Volume 12 Number 4 2002 NOVON Volume 12 NOVON Number 4 2002 Diplopanax vietnamensis, a New Species of Nyssaceae from VietnamÐOne More Living Representative of the Tertiary Flora of Eurasia Leonid V. Averyanov Komarov Botanical Institute of the Russian Academy of Sciences, Prof. Popov str., 2, St.-Petersburg, 197376, Russia. [email protected] Nguyen Tien Hiep Institute of Ecology and Biological Resources of the National Center for Natural Sciences and Technology of Vietnam, Nghia Do, Cau Giay, Hanoi, Vietnam ABSTRACT. A new species of Diplopanax, D. viet- resentatives of this highland tropical ¯ora and its namensis (Nyssaceae, Cornaceae s.l.), was discovered corresponding climatic zone have disappeared al- in the mountains of southern Vietnam. It differs from most completely. Small relictual and depauperate the single known species, Diplopanax stachyanthus, elements of it can still be found in Portugal, the by short, simple, densely pubescent in¯orescences Colchis, and in a few other orographically favored and large yellow-white ¯owers with ®ve prominent continental regions. The largest remaining humid bosses on the ¯ower disc. This genus (with two spe- warm Tertiary ¯ora was shifted to Southeast Asia. cies, D. stachyanthus and D. vietnamensis) is now re- One of the richest assemblages of ancient, tropical garded as congeneric with Mastixicarpum, represen- Tertiary taxa may now be observed in highland trop- tatives of which were an integral component of the ical areas of Southeast Asia. Primary mountain for- Mastixia-like paratropical broad-leaved evergreen ests of this region include a surprisingly large pro- vegetation that covered much of the Northern Hemi- portion of archaic genera well represented in the sphere from the uppermost Cretaceous to the late Mio- fossil record of the Upper Cretaceous and Lower Ter- cene, about 65 to 7 million years ago. tiary recorded from Europe, East Asia, and North Key words: Diplopanax, Nyssaceae, Tertiary America (Kubitzki & Krutzsch, 1998; Martinetto, ¯ora of Eurasia, Vietnam. 1998). Among them are numerous Actinidiaceae, Annonaceae, Berberidaceae, Chloranthaceae, The montane ¯ora within the eastern part of the Daphniphyllaceae, Fagaceae, Hamamelidaceae, Jug- Indochinese Peninsula is still relatively unknown. landaceae, Lardizabalaceae, Lauraceae, Magnoli- Without a doubt it is diverse and conceals a great aceae, Menispermaceae, Nyssaceae, Pentaphylaca- number of exciting discoveries. Floristically, the ceae, Platanaceae, Ranunculaceae, Rhoipteleaceae, highland forests of this area represent relictual Sabiaceae, Saururaceae, Theaceae, as well as a num- remnants of a primitive, tropical ¯ora that occupied ber of gymnosperm genera such as Amentotaxus Pil- wide tropical and subtropical areas of Europe, Asia, ger (Cephalotaxaceae), Cephalotaxus Siebold & Zuc- and North America during the early Tertiary, ap- carini ex Endlicher (Cephalotaxaceae), Glyptostrobus proximately 40±70 million years ago (Axelrod et Endlicher (Taxodiaceae), Keteleeria CarrieÁre (Pina- al., 1998; Kubitzki & Krutzsch, 1998; Zhang & Lu, ceae), Pseudolarix Gordon (Pinaceae), and a great 1998). Due to subsequent climate cooling and in- number of others (Axelrod et al., 1998; Martinetto, creased aridity, the European and North Asian rep- 1998; Nguyen, 1998; Nguyen & Nguyen, 1998; Wu NOVON 12: 433±436. 2002. 434 Novon Figure 1. Diplopanax vietnamensis Averyanov & T. H. Nguyen. Ða. Flowering leafy branchlet. Ðb. Portion of in¯o- rescence with mature ¯ower buds just before opening. Ðc. Open ¯ower. Ðd. Adaxial surface of petal. Ðe. Calyx, style, and bosses on the disc. (All drawn from an isotype, L. Averyanov & T. H. Nguyen VH 596, LE.) Volume 12, Number 4 Averyanov & Nguyen 435 2002 Diplopanax vietnamensis (Nyssaceae) & Wu, 1998). These plants were widely distributed extant species comprises southern China and Viet- in the Northern Hemisphere during the Eocene and nam. Diplopanax stachyanthus is the type species of Miocene. Most of these plants from these areas ex- the genus. The following dichotomous key is provid- isted during the past 5±10 million years including ed for the identi®cation of Diplopanax species. such famous ``living fossils'' as Cathaya Chun & Kuang (Pinaceae), Metasequoia Miki ex Hu & W. C. KEY TO THE MODERN SPECIES OF DIPLOPANAX Cheng (Taxodiaceae), and Pseudolarix Gordon (Pin- aceae) (Axelrod et al., 1998). Species of the genus 1a. Tree, about 8±12(15) m tall; in¯orescence a ra- ceme or panicle branched at the base, 15±30 cm Diplopanax Handel-Mazzetti are typical representa- long; rachis of in¯orescence glabrous or sparsely tives of this ¯ora. pubescent in basal portion; mature ¯ower buds Diplopanax (with a single species, D. stachyan- before opening about 3 mm wide; ¯owers egg- thus Handel-Mazzetti) was described in 1933 from yellow, about 8 mm diam.; disc of the ¯ower without prominent bosses and 6 smooth; style southern China and was initially placed in the Ar- glabrous ................ 1. D. stachyanthus aliaceae (Handel-Mazzetti, 1933). Later, based on 1b. Tree, 15±25 m tall; in¯orescence not a branched the study of fruit structure, it was suggested that spike, 5±8 cm long; rachis of in¯orescence the genus has af®nities with Mastixia (Nyssaceae) densely pubescent in basal portion; mature ¯ow- and especially with some fossil Mastixiaceae (Hoo er buds before opening 4±5 mm wide; ¯owers yellow-white, about 10±12 mm diam.; disc of the & Tseng, 1978; Tseng, 1983). Comparison of extant ¯ower with 5 prominent bilobed bosses sur- Diplopanax fruit pyrenes and fossil fruit remnants rounding the base of the style; style sparsely pu- of Miocene species of Mastixicarpum M. E. J. bescent ................. 2. D. vietnamensis Chandler gave evidence that the two genera may be congeneric (Eyde & Xiang, 1990). It is remark- 1. Diplopanax stachyanthus Handel-Mazzetti, able that species of Mastixicarpum were wide- Sinensia 3(8): 197±198. 1933. TYPE: China. spread and integral components of Tertiary Euro- Guangxi: Kwangsi, N. Luchen, Miu-shan, Bin- pean ¯oras especially during the Miocene. Fruits long, 1200 m, 14 June 1928, R. C. Ching 5969 of these plants are the most common fossils in lig- (holotype, LU; isotype, NY). nite beds of Europe from the uppermost Cretaceous This species is distributed in southern China (Prov- to the late Miocene, roughly 65 to 7 million years inces Hunan, Guangdong, Yunnan, and Guangxi ago (Eyde & Xiang, 1990). Zhuang Autonomous Region) and northern Vietnam Woody fruits of Diplopanax and Mastixicarpum (Provinces Cao Bang and Vinh Phuc). Diplopanax have a certain similarity and obviously represent stachyanthus grows in wet evergreen broad-leaved the same woody-fruited evolutionary line in the montane primary and open secondary forests and family Nyssaceae (Cornaceae s.l.). It was long sup- shrubs, wood sides, at elevations of 900±1600 m on posed that this line of woody-fruited relatives had soils developed predominantly on silicate sands, clay, died out in the later Miocene about 5±10 million granite, and rhyolites. It ¯owers in April±June. years ago (Eyde & Xiang, 1990). With the descrip- tion of the new species, at least two modern species Specimens examined. CHINA. Kwangsi: Shap Man still exist in Southeast Asia. The second extant spe- Taai Shan, near Iu Shan village, SE Shang-sze, Kwangtung cies, Diplopanax vietnamensis, was recently discov- Border (Shang-sze District), 23 May 1933, W. T. Tsang 22361 (LE). VIETNAM. Cao Bang Prov.: Le A pass at ered in southern Vietnam (Kon Tum Province). It elevation 1600 m a.s.l., 24 Nov. 1976, Khoi, Nhan & Ve was observed as an occasional co-dominant of the 71 (LE). Vinh Phuc Prov.: Tam Dao 2, 29 May 1977, ®rst forest stratum in primary, wet, evergreen, T. B. Nguyen 121 (LE). broad-leaved tropical mountain forests. Other com- mon trees in these forests are also relictual genera 2. Diplopanax vietnamensis Averyanov & T. H. of Tertiary age from such families as Betulaceae, Nguyen, sp. nov. TYPE: Vietnam. Prov. Kontum: Fagaceae, Hamamelidaceae, Lauraceae, Magnoli- cloud evergreen primary forest on N slope of aceae, and Sabiaceae. This unique region repre- Ngoc Linh mountain system, 2380 m, 9 Mar. sents a model for the study of the ¯ora, vegetation, 1995, L. Averyanov, N. T. Hiep VH 5961 (holo- and climate reconstructions from now-extinct type, HN; isotypes, AUU, LE, MO, P). Figure 1. Northern Hemisphere forest ¯oras of Tertiary age (Martinetto, 1998). This area in Southeast Asia may be the birthplace and cradle of the Holarctic and 1 Index VH (abbreviation of words Vietnamese High- lands) is used in the numbering of herbarium collections even Paleotropic ¯ora (Wu & Wu, 1998). that were made during ®eldwork associated with the U.S. According to available data, Diplopanax includes National Geographic Society exploration program ``Flora two extant and four extinct species. Distribution of of Highlands of South Viet Nam.'' 436 Novon In¯orescentia brevis, simplex (eramosa), spiciformis, 5± component of the secondary forest stratum. This 8 cm; in¯orescentiae rhachis crassa, basi dense pilosa; tree ¯owers in March±May. ¯ores albido-¯avides, magni, 10±12 mm in diametro, dis- cus ¯orum cum 5 crassis excrescentiis; stylus disperse Paratypes. VIETNAM. Prov. Kontum: cloud ever- pilosusÐspecies nova a D. stachyantho (sola antea cog- green primary forest on N slope of Ngoc Linh mountain nita species generis) dignoscor. system at 2400±2450 m, 6 Mar. 1995, L. Averyanov & N. T. Hiep VH 561 (AUU, HN, LE, MO, P); Kon Plong Distr., Tree 15±25 m tall. Branchlets glabrous, apically Hieu Municipality,
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