A review of dipterocarps - Center for International Forestry Research

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Biogeography and Evolutionary Systematics of Dipterocarpaceae Table 5. Distribution of living and fossil dipterocarp genera or section. Taxa S.Am Afri Mada Seyc India Sri-L Chin Burm InCh Thai Mal Born Indo Phil N.Gu Pakaraimaea O Marquesia O Monotes O O Vateriopsis O Vateria O* O Stemonoporus O Doona O Balanocarpus K. O O Vatica Kosterm. O* O O O O O O O O* O O Dipterocarpus * (?) O* O O* O* O O O O* O Anisoptera * O O O O O O O O Anthoshorea O O O? O O O O O O O s.Shorea (*°) O*° O O? O*° O*° O O O O*° O s.Hopea O* O O O O O O O O O O s.Dryobalanoides O O O O O O O O Parashorea O O O O O O O O Pentacme *(?) O O O (O) O O Sunaptea O O O O O O O O O Cotylelobium O O O O O Neobalanocarpus (O) O Dryobalanops * * O O* O* s.Richetioides (O) O O O O s.Rubroshoreae (O) O O O O s.Brachypterae O O O O s.Pachycarpae O O s.Rubellae O O s.Neohopea O Upuna O S.Am: South America; Afri: Africa; Mada: Madagascar; Seyc: Seychelles; Sri-L: Sri-Lanka; Burm: Burma; Chin: China; InCh: Indo-China; Thai: Thailand; Mal: Peninsular Malaysia; Born: Borneo; Indo: Sumatra, Java and other Indonesian islands but Borneo; Phil: Philippines; N.Gu: New Guinea; (O): extreme geographic position (Langkawi island for Malaysia, extreme S-W Thailand); O: living species; *: fossils; O*: living species and fossils; s.Shorea(*°): both section Shorea, and Shorea sensu lato for fossils; O*°: both section Shorea and Shorea sensu lato when precise taxonomic level not specified, particularly for fossils. Potential Taxa for Differentiation The preceding facts suggest that Dipterocarpus, Vatica, Hopea section Hopeae and Shorea (sections Anthoshorea and Shorea) could be the main Dipterocarpoideae taxa from which new forms could arise by diversification during periods of isolation of Indian and East Asian lands. This is supported by certain highly variable species which, in a single species, may contain much of the whole set of variations of the other species in their own genus, or even that of different other genera for example, Shorea roxburghii and Vatica pauciflora (respectively S. talura and V. wallichii: in 15 Maury 1978, Maury-Lechon 1979 a, b, Maury-Lechon and Ponge 1979). The new taxa should probably correspond to groups of species such as Hopea s. Dryobalanoides and all the Shorea of the ‘red-meranti’ group in the Malesian area, and perhaps also Balanocarpus Kosterm. in the Indo-Sri Lankan part. The limited taxa Vateriopsis, Vateria in the west and Upuna in the east, are residual genus with limited potential for differentiation. Anisoptera, with the fossil and present distribution area, perhaps partly shares this lack of evolutionary potential and could be a regressing group. With more limited areas, Parashorea and Pentacme
Biogeography and Evolutionary Systematics of Dipterocarpaceae from one part, and Dryobalanops from the other, could also enter this evolutionary-limited-potential group. Sunaptea, having been several times merged into Vatica, requires fossil and living distributions. The Sunaptea morphological and anatomical characters in embryos, fruit-seeds and seedlings could have had a much larger distribution. The same remarks concern Cotylelobium. Kostermans (1992) changed Cotylelobium scabriusculum (Thw.) Brandis into Sunaptea scabriuscula (Thw.) Brandis. Perhaps information on living and fossil dipterocarps from China, Indo-China and Burma could modify the present perception of their extension. It is also unknown whether the Vaticoxylon were of the Vaticae, Pachynocarpus or Sunaptea types. The absence of Cotylelobium among fossil forms results from a lack of detailed criteria in wood anatomy that prevents assessment of its presence. Close cooperative works between paleobotanists, wood anatomists of living forms and systematicists are needed to consolidate present conclusions on the mixed taxa of Vatica and Cotylelobium. Future work should particularly consider separately the Sunaptea, Vaticae and Pachynocarpus. The same treatment is necessary for the Shorea and Hopea sections. These remarks are especially pertinent for the new molecular approaches being rapidly developed, and which have been applied in a few instances to dipterocarps (e.g. Chase et al. 1993, Wickneswari 1993). Some of the diverse opinions, in all disciplines, are due to the studies being limited to a restricted number of species. It is therefore necessary to examine the whole set of species instead, with particular attention to intermediate ones such as Vatica heteroptera and V. umbonata group. The V. pauciflora (ex V. wallichii) case has already been mentioned above, together with Shorea roxburghii (ex S. talura). Maury- Lechon’s previous conclusions (Maury 1978, Maury- Lechon 1979a, b: Fig. 16, p.100) based on cotyledonary shapes and structures have been vindicated by Wickneswari’s results (1993: Fig. 1). These conclusions concern affinities between the Sunaptea group and Cotylelobium and their joint affinities with Upuna, as well as the connection of these three taxa with the closely related group of Anisoptera first, and then Dipterocarpus and Dryobalanops. Shorea bracteolata, the only Anthoshorea in Wickneswari’s study, has cotyledonary characters that are distinct from species such as S. roxburghii and S. resinosa (Maury 1978, Maury-Lechon 1979 a, b, Maury-Lechon and Ponge 16 1979). Consequently, the position of S. bracteolata reflects perhaps only partially the position of the whole group of species presently included within the Anthoshorea. Shorea resinosa and S. roxburghii cotyledonary shapes locate the Anthoshorea close to Doona and to Dryobalanops, Dipterocarpus and Cotylelobium. The present heterogeneity of Anthoshorea suggests the need for a re-examination both by DNA analysis and other approaches. The Dipterocarpus genus should also be examined for eventual correspondence between chemotaxonomic groups (Ourisson 1979) and biological characters such as seed sensitivity to desiccation and cold temperatures, seed and seedling resistance to pathogens by defensive secretions, and chemical type of root exudates for mycorrhizal fungi association. The statement that there is no relation between chemical groups and morphological features should be re-examined in considering flower characters, particularly stamen shapes, pollen and pollination, sexual and non-sexual reproduction, the fruit-seed-embryo-seedling sequence and the habitat. Phytogeographical Regions of Extinct Dipterocarps or Related Taxa No living or extinct monotoid (Tables 5 and 6) has been reported in Asia while diverse supposed dipterocarpoid fossils are described from Europe: Woburnia porosa wood from Bedfordshire Lower Cretaceous, U.K. (Stopes 1912, Kraüsel 1922, Schweitzer 1958, all in Aubréville 1976), flowers of Monotes oeningensis (Heer) Weyland from Upper Eocene in Hungary (Boureau 1957, Boureau and Tardieu-Blot 1955), and Tertiary fruits of west Germany, Switzerland and Austria (Gothan and Weyland 1964, in Aubréville 1976). Doubts were cast on these identifications (Bancroft 1933, Harris 1956 and Hughes 1961 in Aubréville 1976, Gottwald and Parameswaran 1966, 1968). Other doubtful fruits of Monotes type have even been reported from New York (USA) and from the Alaska Eocene putative but unlikely tropical forest (Wolffe 1969, 1977). Boureau (in Boureau and Tardieu-Blot 1955) doubted this sequence but he remained convinced of the real presence of Monotes in the European Cretaceous and Tertiary. Huge distances would then separate the living Monotoideae from the extinct ones, and the Asiatic- Malesian Dipterocarpaceae from the European fossils, without any fossils in between, notably in North Africa.
Page 1 and 2: A Review of Dipterocarps Taxonomy,
Page 3 and 4: ã 1998 by Center for International
Page 5 and 6: Authors S. Appanah Forest Research
Page 7 and 8: SPINs Species Improvement Network T
Page 9 and 10: Foreword The Center for Internation
Page 11 and 12: Introduction As a consequence, much
Page 13 and 14: Introduction each project that is m
Page 15 and 16: Biogeography and Evolutionary Syste
Page 23: Biogeography and Evolutionary Syste
Page 55 and 56: Conservation of Genetic Resources i
Page 65 and 66: Seed Physiology P.B. Tompsett Seed
Page 67 and 68: Seed Physiology 59 (Sasaki 1980, Ya
Page 69 and 70: Seed Physiology 61 § orthodox; and
Page 71 and 72: Seed Physiology 63 Table 4. Lowest-
Page 73 and 74: Seed Physiology 65 Table 5. (contin
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Seed Physiology 67 Table 6. Viabili
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Seed Physiology 69 Dipterocarp seed
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Seed Physiology 71 Roberts, E.H. 19
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Seed Handling 74 viability of the s
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Seed Handling 76 the basic activiti
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Seed Handling 78 Table 2. Seed (fru
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Seed Handling 80 Table 3. Mean inse
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Seed Handling 82 Seedling storage u
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Seed Handling 84 air will ensure ra
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Seed Handling 86 ASEAN Forest Tree
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Seed Handling 88 Tompsett, P.B. and
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Seedling Ecology of Mixed-Dipteroca
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Root Symbiosis and Nutrition Table
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Root Symbiosis and Nutrition Table
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Root Symbiosis and Nutrition specie
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Root Symbiosis and Nutrition in Sab
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Root Symbiosis and Nutrition where
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Root Symbiosis and Nutrition 5. Lim
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Root Symbiosis and Nutrition Group
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Root Symbiosis and Nutrition Takaha
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Pests and Diseases of Dipterocarpac
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Management of Natural Forests S. Ap
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Management of Natural Forests about
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Management of Natural Forests 4. Cl
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Management of Natural Forests were
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Management of Natural Forests incre
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Management of Natural Forests 1. He
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Management of Natural Forests which
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Management of Natural Forests Cheng
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Management of Natural Forests Uebel
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Plantations 152 macrocarpa, V. indi
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Plantations 154 are: Dipterocarpus
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Plantations 156 and no longer in ne
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Plantations 158 able to store them
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Plantations 160 ‘Predictive Test
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Plantations 162 Qureshi et al. (196
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Plantations 164 Tomboc and Basada (
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Plantations 166 are removed. Anothe
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Plantations 168 intervention. Sange
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Plantations 170 mono-layered. The r
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Plantations 172 diameter of 46.3 m
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Plantations 174 Ang, L.H., Maruyama
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Plantations 176 Cheah, L.C. 1971. A
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Plantations 178 Kadambi, K. 1957. B
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Plantations 180 Moura-Costa, P. 199
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Plantations 182 Regional Workshop o
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Plantations 184 Conference on Dipte
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Non-Timber Forest Products from Dip
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Scientific Index BIXALES, 25 BOLETA
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Scientific Index EPHEMEROPTERA, 119
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Scientific Index Ovalis section, 8,
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Scientific Index Shorea leptoderma,
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Scientific Index VATERINAE sub-trib
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General Index barus kapur, 192-3 ba
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General Index endangered species, i
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General Index distribution, 15 enda
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General Index nurse crops, 161, 165
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General Index seed material, cryopr
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General Index TPI, 139 tractors, 15
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