FLORA MALESIANA SERIES I- SPERMATOPHYTA Flowering Plants Vol. 8, part 2 Revisions INDEX TO REVISED FAMILIES Aceraceae. . . 4: 3, 592 Actinidiaceae s. str. . .4: 37 Aizoacese: 5 574. 5 sais 8; 207 Alismataceae Be Sh 62915 Amaranthaceae 4: 69, 593; 6: 915 Anacardiaceae . . In press Ancistrocladaceae . . .4: 8 Aponogetonaceae 4: 11; 7: 213 Balanophoraceae. . . . 7: 783 Basellaceae .... . .5: 300 Batidacede ... .-. .5: 414 Betulaceae Se 207 2.62917 Bignoniaceae Peart rope Bixaceae’sstrs | 222... ../45239 Burmanniaceae A 132592 Burseraceae 5: 209, 567; 6: 917; 7: 820 Butomaceae :43.5.°-' .. - 52118 Byplaaceaeioen es ds @allitrichaceae’. . . . . 4: 251 Campanulaceae 6: 107, 928 Cannabinaceae .. . .4: 223 Capparidaceae. .... 6: ‘61 Caprifoliaceae 4: 175, 598; 6: 928 Cardiopteridaceae . . .7: 93 Celastraceae 6: 227, 389, 930 Centrolepidaceae. . . .5: 421 Ceratophyllaceae. . . .4: 41 Chenopodiaceae 4: 99, 594: 6: 932 Glethracedes =. 4. +4. TRIBE) Cochlospermaceae . . .4: 6] Combretaceae 4: 533; 5: 564; 6: 932 Connaraceae 5: 495; 6: 933 Convolvulaceae 4: 388, 599; 5: 558; 6: 936; 7: 823 @omaceae:) of, 24.’ 8: 85 Corynocarpaceae 4: 262; 5: 557 Crassulaceae ..... 4: 197 Crypteroniaceae . . . .8:187 WRPETAGCAG Wh)... e 7: 435 Datiscaceaes ogc. 4: 382 Dichapetalaceae 5: 305; 6: 941 Dilleniaceae . . 4: 141; 7: 824 Dioscoreaceae. ... . 4: 293 Dipsacaceae.. t+ ane . 4: 290 Droseraceae. . 4:.3/72)52/557 Elatinaceae: 2 — See 4: 203 Epacridaceae .«.)... . 62422 Ericaceae . . . . 6: 469, 943 Erythroxylaceae . . . .5: 543 Fagaceae hs as taet = perce Ficoidaceae =. 2 -3/)..7)-: 4: 267 Flacourtiaceae 5: 1, 565; 6: 943; 7: 827 Flagellariaceae. . .. . 4: 245 Geraniaceae.. ... . 6: 445 Gnetaceae. . . 4: 336; 6: 944 Gonystylaceae. .... 4: 349 Goodeniaceae 5: 335, 567; 6: 949; 7: 827 Haemodoraceae . . . .5: 111 Haloragaceae . 2... 7: 239 Hamamelidaceae. . . .5: 363 Hippocrateaceae . . . . 6: 389 Hydrocaryaceae ... .4: 43 Hydrocharitaceae 5: 381; 6: 952 Hydrophyllaceae . . 4: 207 Hypericaceae aeons oe. $2 1 Icacinaceae 7 e414 eee Inidaceae wane wee es hae, Juglandaccae eas sae 6: 143 JUUCACEde 2. et eee 4: 210 Juncapinacedelie. + es). 4: 57 Labiatae Sho scott TD DYESS Recdceten ais eink DOES, emnacedey fy. "aise. 2; 2A9 Lentibulariaceae . SPA Loganiaceae. . 6: 293, 953 Lophopyxidaceae FCI OD Malpighiaceae. . ... Sen 25 Martyniaceae ..... 4: 216 Molluginaceae. ... . 4: 267 Moringacede 20 acute 4: 45 Myoporaceae ..... 4: 265 Mryricaceaer 2) 8225. 4: 277 INajadaceae eS 6: 157 Nyctaginaceae. .... 6: 450 INyssacede>- | a ack ee. 4: 29 Ochnacedey sare Me Onaeracedey ko st dee O8 Oxalidareal> hss 53 A. IAS Papaveraceae. 8 te 5: 114 LIBRARY Passifioraceae . .\..%, - 7: 405 heaanacede.. «36 a: 4: 216 Pentaphragmataceae . . 4: 517 Pentaphylacaceae 531s Philydraceae. . ... . ; 4: 5 Phytolaccaceae 4: 229 Pittosporacese’. 2 3. 5. | 5: 345 Plumbaginaceae . . . . 4: 107 Podostemaceae 4: 65; 6: 963 Polemoniaceae. . 4: 195 Pontederiaceae. . 4: 255 Portulacaceae ..... 72 YZ Primiulacedé=- . .. 2.) 6073 Protedceden! . Gis cei 7: 806 Thymelaeaceae 4: 349; 6: 1, 976; 7: 830 Mrapacese <3. a a 4: 43 Mnvonmiaceae | .7.-. ave 4: 59 AIRTHEACeEAG 26. ar een 4: 235 MiypHacede 4. . S25 tame 4: 243 Wimacede 253. ie owe S$: 34 Umbelliferae 4: 113, 595; 5: 555: 6: 983: 7: 830 Valenianaceae 0. 5c 4: 253 Violaceae . 7: 179, 831 Xyridaceae 4: 366, 598 Zygophyllaceae ie a NICAL GARDEN BRONX, NEW YORK 10458 —As - - « ULMACEAE (E. Soepadmo, Kuala Lumpur) . Monoecious or dioecious (?), evergreen, deciduous or semideciduous shrubs or trees, (in Mal.) unarmed and often buttressed. Growth habit (in Mal.) flush-wise, except for Trema and Parasponia. Indumentum of simple, bulbous-based unicellular hairs and/or multicellular glandular hairs. Stipules caducous or rarely rather long persistent and completely enclosing the bud, extrapetiolar or intrapetiolar, basally attached or rarely peltately attached to the nodes, free or connate. Leaves simple, (in Mal.) alternately arranged, petioled, pinnately nerved or triplinerved at base, often asymmetrical at base, entire or variously serrate. Inflorescences |-many- flowered, 3, 2, S2, or $Y, axillary, subterminal, or borne on leafless, older branch- lets or on short, lateral, leafless new shoots, paniculate, racemose, thyrsoid, cymoid, or rarely capitate, bracteate; bracts minute, caducous. Flowers functionally 3, 2, or 8. — 3 Flowers solitary or in condensed cymoid clusters along the rachis, sessile or short-pedicelled; perianth (4~)5(—7)-lobed, lobes free or variously connate, imbricate or induplicate-valvate in bud; stamens as many as tepals, attached to the base of and opposite the perianth lobes, straight or incurved in bud; anthers ovoid, ellipsoid or subreniform, apiculate or non-apiculate, initially tetrasporangiate, later becoming 2-celled, dehiscing lengthwise, introrse or extrorse; pistillode present or absent, if present either rather well developed or rudimentary, densely whitish to silvery, soft or hirsute pubescent. — 2 & & Flowers sessile or stalked, solitary in the axils of the upper new leaves or arranged in various types of in- florescences ; perianth herbaceous or thin-coriaceous, (4-)5(—7)-lobed, lobes always imbricate in bud and connate at base, (in Mal.) long persistent; staminodes or stamens as many as perianth lobes or absent; ovary superior, 2-carpellate, (in Mal.) 1-celled, sessile or stipitate; style 1, tubular, short or absent, stigmatic arms 2, slender, often bifid to deeply lobed at the tip, adaxially papillose-stigmatic for their entire length; ovule 1, anatropous to hemi-anatropous, subapical, pendulous, bitegmic. Fruit a drupe or a samara, faintly angular or flat and winged. Seed mostly exalbuminous; embryo large, straight or curved; cotyledons flat-convex, fleshy, straight or variously folded, often foliaceous. Germination mostly epigeal. Distribution. There are 15 genera, c. 200 spp., widely distributed in the tropics, subtropics, and temperate regions of Europe (as far north as 70°, Scandinavia), Africa (South of Sahara), continental Asia, Malesia, Australia (Queensland and New South Wales), Pacific Islands (as far as Tahiti; incl. also Hawaii and the Galapagos Is.), North, Central, and South America (as far south as 40°, Argentina). Fig. 1. Fossils. Various macrofossils (woods, drupes, and leaf-impressions) and microfossils (pollen grains) attributed to Ulmaceae have been reported by different authors from various localities in Alaska, North America, Europe, continental Asia, and Japan. Amongst the older records are pollen grains of the Ulmus-Zelkova-type from Golden Valley Formation in North Dakota and Rocky Mountains, U.S.A. (Paleocene) and wood and leaf-impressions attributed to U/mus from Rocky Mountains and to Planera from Alaska (Late Paleocene). By the middle of the Eocene and throughout the Miocene and Pliocene fossils of Ul/maceae become more abundant (common) in the northern hemisphere, especially in Europe and North America. However, very little is known about the geological history of the family in Asia. Of the Malesian genera belonging to the tribe Ce/toideae, fossil records of Gironniera (identifi- cation very doubtful) go back to Middle Eocene (Alaska), that of Ce/tis to Miocene (Japan), and Trema to Upper Oligocene (Japan). — Literature: Berry, Tree Ancestor (1923) 146; WATARI, Jap. J. Bot. 11 (1941) 385; J. Fac. Sc. Un. Tokyo III, 6 (1952) 97; La Morte, Mem. Geol. Soc. Am. 51 (1952) 112, 260, 346, 360; PrakAsH & BARGHOORN, J. Arn. Arb. 42 (1961) 185, 347; Greouss, Tert. Angios. Hung., Ak. Kiado Budapest (1969) 83; Fercuson, Verh. Kon. Ned. (31) ZTIBL A , Rene: . . yD awevy YORE iy Ci AB EN , 32 FLORA MALESIANA [ser. I, vol. 8? mr XX/ °, Cd LEG Fig. 1. Approximate range of Ulmaceae. Ak. Wet. sect. II, 60 (1971) 103; LEopotp & MacGiniTiE in Graham, Flor. & Paleofl. Asia & N.E. America (1972) 147; WoLFE, /.c. 200; TANAI, /.c. 235; WOLFE, Brittonia 25 (1973) 334. Ecology. In Malesia species of the Ulmaceae may be found in both primary and secondary forests, from sea-level up to 2000 m; they are more common in the lowlands and hills. Apart from Ulmus lanceaefolia, Celtis rigescens, Gironniera nervosa and G. subaequalis which may attain up to 45 m in height and 100 cm in diameter, all species are understorey shrubs or trees. Of the 6 genera occurring in Malesia, species of Aphananthe, Celtis, Gironniera and Ulmus are basically primary forests inhabitants, though they may survive and thrive in secondary forests as well. Of these, Ce/tis and Gironniera species are the most widely spread and may be found growing on various types of soils, including those derived from limestone. In Malesia Ulmaceae are found under both everwet and more seasonal climatic conditions, but some show preference for one or the other. In Celtis two groups of species may be dis- tinguished, viz the group of C. tetrandra, timorensis and rubrovenia, and that of C. philippensis. The former is either semideciduous or deciduous and found mainly under a drier and more seasonal climate, while the latter is evergreen and grows better in more humid environmental conditions. Aphananthe and Ulmus species are mainly found in regions subject to a rather distinct seasonal climate, and they are either semideciduous or deciduous. Members of the genera Parasponia and Trema are pioneer plants preferring and thriving well in newly opened up habitats, e.g. forest clearings, thickets, roadsides, flood-plains, on volcanic ashes, efc. Fig. 10, 11, 17. They are usually short-lived (at least in Malaya, 5-7 years) and soon will die out, particularly when over-grown by the other more aggressive and long-lived pioneer plants, e.g. species of Macaranga, Mallotus, and Grewia, etc. For this reason species of Trema and perhaps also those of Parasponia are usually not or rarely found in old secondary forest. It also may be noted here that most Trema species grow, thrive, and are more widely spread in the western parts of Malesia, while Parasponias are more common in the eastern parts, especially in New Guinea. The structure and position of the inflorescence and flowers, particularly the amount of pollen grains produced and the structure of the stigmas, and also the absence of nectary, seem to suggest that pollination is most likely affected by wind, though insects may not be ruled out altogether as possible agents for pollination. Except for Ulmus, which produces a dry, flat, winged fruit, the other Malesian genera have various types of fleshy drupes which turn to bright yellow, orange, or deep-red in colour when ripe. These drupes are most probably dispersed by various species of frugivorous birds or arbo- reous mammals. Alternatively, at least in some species, e.g. Celtis philippensis var. wightii which is very common in coastal vegetation, fruit dispersal may be carried out by water currents. In Ulmus the winged fruits are easily dispersed by wind. 1977] ULMACEAE (Soepadmo) 33 It should be noted here that there is a very high percentage of seed abortion in Malesian genera for reasons unknown. This is made good by the production of a great number of flowers and fruits, produced regularly throughout the year or at least twice a year. Except for U/mus, the embryo is protected by a strong, hard and durable endocarp. In all genera endosperm is usually scanty or absent. Anatomy. For general surveys also covering the older literature see SOLEREDER, Syst. Anat. Dicot. Stuttgart (1899) 860-865 and ibid. (1908) 295; METCALFE & CHALK, Anat. Dicot. Oxford (1950) 1271-1278; Sweirzer, J. Arn. Arb. 52 (1971) 523-585. Additional selected references: Denay, l’Appareil conducteur foliaire des Urticacées, des Moracées et des Ulmacées (Ordre des Urticales), Arras (1934); JANssonius, Mikr. 6, Leiden (1934) 1-308 (wood anatomy, under Urti- caceae); LEROY, Bull. Mus. Nat. Hist. Nat. Paris sér. 2, 18 (1946) 118-123 & 180-184 (taxonomy, and anatomy of Aphananthe); DEN BERGER, Determinatietabel Malesié, Veenman, Wageningen (1949) (wood identification); RAo, GovINDU & THIRUMALACHAR, J. Indian Bot. Soc. 29 (1950) 224-226 (aerial roots, Trema) ; JANSSONIUS, Blumea 6 (1950) 407-464 (wood anatomical affinities) ; Descu, Mal. For. Rec. 15 (1954) 618-620 (wood); JutrE, Nova Guinea n.s. 10 (1959) 241-278 (wood); MosELEY, Brittonia 25 (1973) 356-379 (anatomy and relationships). SWEITZER’s study (/.c.) is the most up-to-date survey.of leaf and wood anatomy of the Ulma- ceae. Although his extensive research materials included very few Malesian species his general conclusions are probably largely applicable to the Malesian species as well. The wood anatomy is indicative of the mutual affinities of all Ulmaceous genera. Shared characters are: predominantly simple vessel perforations, short vessel members, alternate intervessel pits. Fibres with simple to slightly bordered slit-like pits. Parenchyma at least partly vasicentric. Genera of the tribe U/meae (in Malesia only represented by U/mus lanceaefolia) have exclusively homocellular rays. In Ce/teae (in Malesia all other genera) at least part of the ray tissue is heterocellular. The wood of U/mus lanceaefolia differs from all species described in literature in lacking the ring porosity and the typical ulmiform arrangement of vessel clusters (original observation). Instead, its vessel distribution resembles the diffuse porous group of tropical Celtis species. In Celtis the very striking differences in vessel distribution between tropical and extratropical species are well documented (cf. Sweitzer, /.c.). Although from SwEITZErR’s and other publica- tions some quantitative and qualitative differences between Malesian genera of Celteae can be deduced, our knowledge is still based on too limited materials to allow conclusions on diagnostic and systematic implications. The leaf anatomy of U/maceae at the same time supports its coherence as a family and provides an interesting diversity, of great potential diagnostic and systematic value. All U/maceae share the dorsiventral leaf architecture. The stomata are confined to the abaxial epidermis and are of the anomocytic type. The indumentum includes bulbous-based unicellular trichomes the walls of which are usually silicified. Mineral inclusions of calcium carbonate or silica in cystoliths (with or without pegs) are of common occurrence. The trichome-complement, presence or absence of mucilage cells, crystal complement, loose or compact structure of the spongy tissue, petiole and midrib vasculature show a considerable diversity. SWEITZER’s data and other reports from the literature do not yet allow a leaf anatomical characterization of the individual Malesian taxa, but preliminary studies are indicative that this will be possible if more material is studied. The entire evidence from vegetative anatomy supports the traditional placement of U/maceae in Urticales. — P. BAAs. Palynology. Based on size, sculpturing of exine and number of pores, pollen grains of Ulmaceae may be divided into two major morphological types, namely the U/mus-type and the Celtis-type. In the Ulmus-type the pollen are oblate to subspherical, amb convex or straight; (4-)5(-7)-porate, 20-30 by (26—)28-38(-—51) um, pore circular to elliptic, c. 2-3 by 3-4 um, slightly thickened around its margin; exine rugulate-reticulate. Genera with this type of pollen grains are: Ampelocera, Hemiptelea, Holoptelea, Phyllostylon, Planera, Ulmus, and Zelkova. In the Celtis-type the pollen is suboblate to spherical, amb convex; (2—)3—4(—5)-porate, pores circular or elliptic (elongated towards the poles), often annular and protruding, c. 2-3 by 3-4 um; 17-25 by 19-30 um; exine more or less smooth but for very fine (1-1.5 um) scabrae. This type is found in Aphananthe, Celtis, Chaetacme, Gironniera, Lozanella, Parasponia, Ptero- 34 FLORA MALESIANA [ser. I, vol. 82 celtis and Trema. It may be noted here that pollen grains of Gironniera, Parasponia and Trema are usually slightly smaller and have finer exine sculpturing than those of other genera with Celtis-type of pollen, while the pollen grains of the Malesian species of Trema and those of Parasponia parviflora MiQ. are predominantly diporate. As for pollen grains, Ulmaceae are very closely allied to Moraceae and Urticaceae, particularly to the former. According to NAIR (1967) the pollen type found in Ulmaceae, Moraceae and Urticaceae is derived from a tricolpate type of Ranalean stock. — Literature: ERDTMAN, Pollen Morph. & Taxon. 1 (1956) 442; IkusE, Pollen Grains of Japan (1956) 62; PRAGLOWsKI, Grana Palyn. 3 (1962) 45-65; KUPRIANOVA, Kom. Bot. Inst. Ac. Sc. USSR 1 (1965) 54-58; Nair & SHARMA, Bot. Notis. 118 (1965) 177- 186; STRAKA, Pollen et Spores 8 (1966) 241-264; Nair, Rev. Palaeobot. & Palyn. 3 (1967) 81-91; MALLIK & CHAUDHURI, Bull. Bot. Soc. Beng. 22 (1968) 105-108; TsuKADA, Bot. Mag. Tokyo 81 (1968) 385-395; Rao & Leer, Pacif. Sc. 24 (1970) 255-268; HUANG, Pollen Fl. Pl. Taiwan (1972) 235; SowuNnmi, Grana Palyn. 13 (1973) 145-186; ADAmMs & Morton, Atl. Pollen Trees & Shrubs Canad. & U.S. 9 (1974) pl. 17; STOCKMARR, Grana Palyn. 14 (1974) 103-107; Kepves & PARDUTZ, Acta Biol. Szeged. 20 (1974); HAMILTON, Pollen et Spores 18 (1976) 54-57. Embryology. Apart from several species of U/mus and Holoptelea very little is known about the sporogenesis and embryogenesis of the U/maceae. From a very limited information so far published it appears that the anthers are initially tetrasporangiate but become bisporangiate just before anthesis through the breakdown of the adjoining wall between the locules. The anther- wall development conforms with the so-called basic-type in which the parietal cells divide both anticlinally and periclinally to form the endothesium layer, two (Trema and Ulmus) or three to four (Holoptelea integrifolia) middle-layers and glandular tapetum. Simultaneous cytokinesis in the microspore mother-cells follows meiosis and as a result the pollen grains are initially arranged in either tetrahedral or decussate tetrads. At anthesis the pollen grains are either 2-celled (Holop- telea and Trema) or 3-celled (U/mus). In Celtis, Holoptelea and Trema up to 80% of the pollen grains produced are sterile or imperfectly developed. The ovule is anatropous to hemianatropous, bitegmic, crassinucellar or tenuinucellar (in a few species of U/mus) with the micropyle formed by both integuments (Celtis and Trema) or by the inner integument only (Holoptelea and Ulmus). In Holoptelea and Trema the megaspore mother-cell divides into 4 daughter cells arranged in a linear tetrad, and of these only the chalazal megaspore develops into Polygonum-type of embryo- sac. In U/mus, however, the embryo-sac is tetrasporic and either belongs to Adoxa- or Drusa-type or variation of these two types with 4-12 antipodal cells. The pollen tube enters the ovule either through the micropyle, the integuments or the chalaza. Endosperm formation is nuclear and the tissue is either diploid or triploid and later becomes cellular. Embryo development conforms with the Onagrad-type in Holoptelea and Solanad-type in U/mus. Polyembryony is a common phenomenon, especially in U/mus. The mature embryo is straight with broad, flat or plano- convex, equal or slightly unequal cotyledons in Holoptelea, Planera, Phyllostylon, Ulmus, and Zelkova, or curved with ascending hypocotyle and narrow, incurved or induplicate-plicate or variously folded cotyledons which are mostly unequal in length in Ampelocera, Aphananthe, Celtis, Gironniera, Parasponia, Pteroceltis and Trema. — Literature: SHATTUCK, Bot. Gaz. 40 (1905) 205-223; LELIVELD, Rec. Trav. Bot. Néerl. 32 (1935) 543-573; Capoor, Beih. Bot. Centralbl. 57 (1937) 233-249; WALKER, Am. J. Bot. 37 (1950) 47-52; HJELMQvisT & GAZzZI, Bot. Notis. 118 (1965) 329-360; Davis, Syst. Embryol. Angiosp. (1967) 266-267. Chromosomes. From various published data it seems that the chromosome number in the Ulmeae (Holoptelea, Ulmus, and Zelkova) isn = 14 and 2n = 28, 42, and 56, though reports of n = 15 and 30 have been made on U/mus americana. In the Celtideae the number seems to be less constant varying from n = 10, 2n = 20, 28, 40 in Celtis (9 spp.); n = 30, 2n = 84 in Chaetacme (2 spp.); ton = 10, 10 + B, 18, 20, and 80 in Trema (3 spp.). It may be noted here that as for chromosome number, U/maceae seems to be closely related to Moraceae where n = 12-16, 2n = 24, 26, 28, 42, 56, and 84, and to Urticaceae of which n = 14, 28 and 2n = 22, 24, 28, 52, and 84. — Literature: KRAUSE, Ber. Deut. Bot. Ges. 48 (1930) 9-13; Planta 13 (1931) 29-84; WALKER, Science 75 (1932) 107; SAx, J. Arn. Arb. 14 (1933) 82-84; BowbDeEN, Am. J. Bot. 32 (1945) 195; DARLINGTON & WyLiE, Chromos. Atlas Fl. Pl. (1955) 182-183; MANGENOT & MANGENOT, Bull. Jard. Bot. Brux. 28 (1958) 315-329; ArorA, Bull. Bot. Surv. India 2 (1960) 1977] ULMACEAE (Soepadmo) 35 305; GAJAPATHY, Bull. Bot. Surv. India 3 (1961) 49-51; GRUDZINSKAJA & ZAKHARYEVA, Bot. Zhurn. 52 (1967) 641-651; Hsu, Taiwania 13 (1967) 117-129; MEHRA & GILL, Taxon 17 (1968) 574-576; J. Arn. Arb. 55 (1974) 663-677; Feporov (ed.), Chromos. Numb. FI. Pl. (1969) 710- 711; GADELLA c.s. Acta Bot. Neerl. 18 (1969) 74-83; MEHRA & Hans, Taxon 18 (1969) 310-315; TATAYUK & TURCHANINOVA, Tsitologia & Genetika 4 (1970) 397-401 ; HANs, Cytologia 36 (1971) 341-345; Niewaus, Taxon 20 (1971) 355; MeHRa, Nucleus 15 (1972) 64-83; SARKAR, Taxon 22 (1973) 652. Chemotaxonomy. SOLEREDER mentioned the more or less general occurrence of cystoliths and cystolith-like structures (SiO, + CaCO;) in Ulmaceae. The tendency to accumulate carbon- ate of lime seems to be very strong in this family ; CaCO, is deposited in wall structures (e.g. hairs, cystoliths) and in cell lumina (e.g. in heartwood of Ul/mus and Celtis; in seed coat cells of Celtis). Often oxalate of lime is also present in large amounts; solitary and clustered crystals occur in the family. Anatomically easily detectable internal excretion comprises also mucilage production. The mucilage is deposited in epidermal cells (many taxa) or in mucilage idioblasts in the meso- phyll of some genera and in barks and flowers of most species of U/mus. The bark of Ulmus rubra Muu. (‘Slippery Elm’) was used formerly as a mucilaginosum in official medicine. In mucilage-rich elm barks large mucilage idioblasts may develop to lysigenous mucilage cavities. Chemically elm bark mucilages are characterized by a high content of galacturonic acid, galac- tose, 3-0-methylgalactose and rhamnose. U/maceae are moderately strong accumulators of polyphenolic compounds. Derivatives of caffeic acid, catechins, pro-anthocyanidins (formerly leucoanthocyanidins), flavonols (especially glycosides of kaempferol and quercetin) and con- densed (= flavanoid) tannins seem to occur more or less ubiquiteous in leaves, fruits, barks and woods. According to LEBRETON flavonoid constituents with a trihydroxylated B-ring (in casu myricetin and prodelphinidin), an assumedly primitive feature, are restricted to Celtideae. (+)-Catechin was definitely identified in leaves, twigs and barks of European elms and its 7-xyloside was isolated from the stem-bark of U/mus americana L. C-Glycoflavons (tremasperin) occur in leaves of Trema aspera BL., and the wood of Zelkova serrata (THUNB.) MAKINO contains large amounts of the fungistatic 6-C-glucoflavonoids keyakinin and keyakinol. Tannin contents of woods, barks, leaves and fruits are moderate (mostly less than 10%). There is only one report in literature indicating a possible co-occurrence of galli- and ellagitannins with condensed tannins in Ulmaceae; bark and wood of Celtis australis L. contain gallic acid and derivatives of ellagic acid according to CHARI c.s. (1968). Much chemical work was performed with elm barks and especially elm woods in connection with ‘Dutch Elm Disease’. Cadinane-type oxigenated sesquiterpenes seem to be present in the young wood of every species. On aging (heartwood formation) or after fungal infection, synthesis and accumulation of fully aromatic (cadalenal, hydroxycadalenal) and (or) o-quinonoid (the mansonones) cadinane derivatives take place in American elm species belonging to the sections Trichoptelea, Microptelea and Chaetoptelea; they seem to be absent from the sections Blepharo- carpus and Madocarpus in which all European elms are included. It deserves mentioning that the antifungal cadalenals and mansonones represent phytoalexin-like stress compounds in U/mus, and occur at the same time as normal heartwood constituents in U/mus and Zelkova (but not in Celtis); they are chemically identical with, or biochemically closely related to the gossypol- mansonone-group of constituents of many Malvaceae, Bombacaceae and Sterculiaceae (man- sonones were first detected in the wood of Mansonia altissima A. CHEV.). It was recently shown that hemigossypol, the precursor of the long-known gossypol, is a phytoalexin in many malva- ceous plants and that p-quinonoid derivatives of hemigossypol are engaged in the plants resistance against attack by several phytophagous insects (J. R. GRAY c.s. J. C. S. Chem. Commun. 1976, 109; J. A. VeEcu c.s. l.c. 144). As far as ecological chemistry (defensive substances) is concerned, Ulmaceae much resemble members of the order Malvales. Leaf, bark and wood waxes were investigated by several authors in recent time. They seem to consist mainly of alkanes, long-chain fatty acids, wax alcohols and phytosterins. Additionally pentacyclic triterpenes are often present; 6-amyrin (i), lupeol (ii), betulin (iii), friedelin (iv), friedelanol (v), moretenol (vi), simiarenol (vii) and simiarenon (viii) were reported from leaves and (or) barks of Celtis australis L. (iii), C. laevigata WiLLD. (vi), Holoptelea integrifolia PLANCH. (iv, Vv), Trema guineensis FICALHO (reported as T. orientalis BL.; vii, viii), Ulmus americana L. (ii, esterified with cerotinic acid) and Zelkova 36 FLORA MALESIANA [ser. I, vol. 82 serrata MAKINO (iv). The heartwood of Holoptelea integrifolia PLANCH. yielded 2«-hydroxy-3- epioleanolic acid (G. MisrA c.s. Planta Medica 27, 1975, 290); this is the only triterpenic acid isolated hitherto from U/maceae. Seeds of Ulmaceae seem to store predominantly proteins and fatty oils. The oils have linolic (Celtis, Chaetacme, Trema), oleic (Holoptelea) or capric (Ulmus, Zelkova) acids as main fatty acid. Species of Ce/tis and Pteroceltis accumulate small amounts of quebrachitol in leaves; this cyclitol could not be detected in leaves of species of Ulmus and Zelkova (Hemiptelea included). Alkaloid-like compounds are recorded in literature from members of Ampelocera, Aphananthe, Celtis, Gironniera, Trema and Ulmus, but only in the case of Ampelocera ruizii KLOTZSCH an alkaloid-like compound isolated from leaves was chemically identified; it proved to be an «-pyridone derivative related to trigonelline (R. H. BURNELL c.s. Lloydia 38, 1975, 444). The foetid smell of some Celtis woods of India, Indonesia (‘kaju tai’) and Africa is caused by skatol. Several species of U/maceae are reported to be toxic in literature. GRESHOFF isolated a toxic bitter principle from the leaves of Aphananthe aspera (THUNB.) PLANCH. (= Homoioceltis aspera BL.) which he compared with his streblide (from Streblus asper Lour.; strebloside is now known to be a cardenolide). Leaves of Trema cannabina Lour. (= Sponia virgata PLANCH.) and of T. aspera BL. (= T. cannabina) were reported to be cyano- phoric; both species, however, are polymorphic with regard to cyanogenesis if the botanical identification of all plant samples investigated hitherto was correct. Leaves of T. aspera (= T. cannabina) contain another toxic principle called trematoxin; its chemical structure is not yet known. From the taxonomic point of view three facts deserve special mentioning: (1) U/maceae are generally included in Urticales; their chemistry agrees rather well with such a classification as is indicated by patterns of mineralisation and phenolic compounds. (2) The chemistry of Ulmaceae resembles members of Malvales in several respects: chemistry of stress compounds; mucilages with high contents of galactose, rhamnose and galacturonic acid; some features of the poly- phenolic and triterpenic patterns. (3) The classification of Ulmaceae in Ulmoideae and Celti- doideae (ENGLER’s Syllabus 2, 1964) or Ulmeae and Celtideae (HUTCHINSON, General of Flowering Plants 2, 1967) is not very satisfactory from the chemical point of view (see cadinane-type sesquiterpenes including mansonones and capric acid as main fatty acid in seed oils in U/mus and Zelkova, but not in Celtis). For more phytochemical details and references see my ‘Chemotaxonomie der Pflanzen’ 6 (1973) 545-554, 762-763, 791, 796. — R. HEGNAUER. Taxonomy. The family name Ul/maceae was first introduced and defined by MirBEL in 1815, at which time it included only Ce/tis and U/mus. Link (1831) proposed splitting Ulmaceae into two separate families, i.e. U/maceae to include U/mus and related genera, and Celtidaceae com- prising Celtis and its allies, an opinion which was supported by GRUDZINSKAYA (1967). However, all contemporary taxonomists generally agree to regard Ulmaceae as a natural taxon closely related to Moraceae and Urticaceae and to include these families in the order Urticales. Any difference of opinion is usually restricted to the inclusion or exclusion of a few genera in the family. In the most recent treatise, HUTCHINSON (1967) divided the family into two tribes, namely the Ulmeae (fiowers bisexual, fruit not drupaceous, embryo straight, cotyledons flat or longitudi- nally folded) to include: Holoptelea, Planera, Phyllostylon, and Ulmus, and the Celtideae (flowers unisexual or sometimes bisexual, fruit drupaceous, embryo curved, cotyledons mostly variously folded) comprising Ampelocera, Aphananthe, Celtis, Chaetacme, Gironniera, Hemiptelea, Lozanella, Mirandaceltis, Parasponia, Pteroceltis, Trema and Zelkova. This subdivision was supported by SwWEITZER (1971) who studied the anatomy of leaf and wood. However, as has been mentioned under Embryology and Palynology, the embryo of Ze/kova is straight, and the pollen (also of Ampelocera and Hemiptelea) belongs to the Ulmus-type (see also ERDTMAN, 1956). Furthermore in many species of Celtis the flowers are bisexual, and in U/mus lanceaefolia and U. parvifolia the flowers are either functionally male or female. This seems to indicate that the tribal subdivision as proposed by HUTCHINSON is not a clear cut case, but that Ulmaceae is a natural taxon. It should be noted further that the Mexican genus Mirandaceltis is in the present study regarded as congeneric with Aphananthe. As for phylogenetic relationship, there seems to be two different opinions. Bessey (1915) and THORNE (1968, 1973) placed Ulmaceae along with Moraceae and Urticaceae in the superorder 1977] ULMACEAE (Soepadmo) 37 PI ae < ES \ | Fig. 2. Ulmus lanceaefolia Roxs. ex WALL. a. Habit, nat. size, b. fruit, x 2, c. persistent cup-shaped perianth, x 2, d-e. embryo, nat. size, g. flowering twig, x 2/,, h. cluster of ¢ flowers, x 8, i. ¢ flower, x 14, j. cluster of 3 flowers, x 2, k-l. 3 flower, x " ad HANSEN c.s. 11265, g-i ScuMuTz 3024, j—/ LisTeR 38 FLORA MALESIANA [ser. I, vol. 8? Malviiflorae, and considered them as families having a very close affinity to or derived from the Malvales. On the other hand, authors such as HUTCHINSON (1967), CRONQUIST (1968), TAKHTAJAN (1969), SweITzER (1971), efc., are of the opinion that Ulmaceae, Moraceae, and Urticaceae are closely allied to or have been derived from the Hamamelidales. — Literature: MIRBEL, Elém. Phys. Veg. Bot. (1815) 905; Link, Handb. 2 (1831) 445; Bessey, Ann. Mo. Bot. Gard. 2 (1915) 109-164; ERDTMAN, Pollen Morph. & Pl. Tax. (1956) 442-443; GruDZINsKAyYA, Bot. Zhurn. 52 (1967) 144-150; HUTCHINSON, Genera of Flowering Plants 2 (1967); CRoNquist, Evol. & Class. Fl. Pl. (1968) 166-167; THORNE, Aliso 6 (1968) 57-66; Brittonia 25 (1973) 395-405; TAKHTAJAN, FI. PI. Orig. & Disp. (1969) 210-212; Sweitzer, J. Arn. Arb. 52 (1971). Uses. 1. Timber. Throughout the north temperate regions the tough, strong and durable wood with attractive appearance and excellent bending quality of many species of Celtis and Ulmus is extensively used for various purposes including shipbuilding, panelling, furniture, boxes, crates, veneers, etc. and that of Zelkova and Phyllostylon for making weaver’s shuttles, scales, piano-keys, efc. In Central America timber of Chaetoptelea (= ?Ulmus) is used for railway sleepers, frames and wheels of vehicles. In Africa and India wood of Holoptelea is utilized for various building purposes. In Malesia and neighbouring countries except Aphananthe cuspidata, Celtis rigescens, C. hildebrandii, C. tetrandra, Gironniera nervosa, Ulmus lanceaefolia and a few others, the trees seldom reach timber size, and as a consequence very little is known about their usage. Of these species the timber is locally used for making planks in house-building and other light constructions. The soft wood of Trema and other species of Gironniera is used locally for making tea-chests and match-sticks, for firewood and charcoal. 2. Bark. Due to the high content of mucilagenous substances, decoction of barks of Holop- telea, Parasponia, Trema and Ulmus mixed with some other ingredients is used in local folk medicines to cure ailments such as inflammation of mucous membrane, rheumatism, etc. The tough fiber is known to be used locally for making ropes. 3. Root. Decoction of roots of Gironniera and Trema species mixed with other substances is used to cure sore mouth, diarrhoea, and also applied as protective medicine after child-birth. 4. Leaves. Especially of Trema species leaves are used as fodder, though due to the presence of glucocides they could be poisonous if consumed in a large quantity. 5. Fruits. In India fruits of Celtis and Holoptelea are known to be eaten. 6. Shade trees. Trema has been used for shade in coffee and cocoa plantations in various parts of Asia. 7. Soil conservation. In South Africa Trema has been planted to protect soils against erosion (SCHEEPERS c.s.). As both Trema and Parasponia species come up in dense seral stands on eruptiva, on fresh volcanic ash, are sometimes pioneers on lavastreams, and are almost invariably an important constituent of thickets, seral regrowths, and secondary forest, I would emphasize that they may represent an untapped cheap source for soil conservation for poor, eroded soils and old mining lands. They have all the favourable qualities of pioneer plants, indifference to soil, pro- ducing abundant seed, and that already at a very early age, and furthermore they are available almost throughout the year. Curiously I do not know of experiments by the Indonesian Forestry Service in this respect. — Literature: BURKILL, Dict. Econ. Prod. Mal. Pen. (1935) 513-514, 1088-1089, 2213-2214; MeTCALFE & CHALK, Anat. Dicot. (1950) 1277; SCHEEPERS c.s. Tijd. Natuurwet. S. Afrika Akad. Wet. & Kunst. 8 (1968) 105-120; Sweitzer, J. Arn. Arb. 52(1971) 52a. KEY TO THE GENERA 1. Flowers always borne on bare older branches, and organized in a condensed cluster on short leafless lateral shoots; perianth cup-shaped, 5-7-lobed; ovary (fruit) stipitate. Fruit a dry, flat, winged samara. Embryo straight es wee ie dl ee eee APS eee 1. Ulmus 1. Flowers axillary, or rarely borne in a condensed capitate thyrse on older branches (Gironniera celtidi- folia); perianth 4~5-lobed, with the lobes free from one another except for their base; ovary sessile. Fruit a fleshy drupe. Embryo variously curved. 2. Leaves triplinerved at base, or if pinnately nerved the stipules do not leave a circular scar around the node; lateral nerves less than 5 pairs. 3: Stipules intrapetiolar, conate-s° +): "0G Oe ae ee 6 Lee ee 2. Parasponia 3. Stipules extrapetiolar, free. 1977] ULMACEAE (Soepadmo) 39 4. Female flowers borne in condensed, multiflowered raceme. Perianth of male flower induplicate- valvate. Fruit compressed, elliptic lens-shaped in cross-section .......... 3. Trema 4. Female flowers solitary in the axils of leaves or borne in a cymoid cluster of 2-3. Perianth lobes of male flowers imbricate. Fruit faintly 3—5-angular in cross-section. 5. Male flowers borne in a 2-3-flowered cymoid inflorescence or in a much-branched paniculate, subterminal inflorescence. Female flowers borne in a racemose cluster of 2-10 or in a $8, much- branched racemose inflorescence; staminodes mostly present. Cotyledons broad, variously folded SES Ir et eS , k ounies oe aoe ue eS. Bey bdo ee 2 4. Celtis 5. Male flowers organized in a condensed, multiflowered raceme. Female flowers always solitary or rarely borne in a 2-3-flowered racemose, $2 inflorescence; staminodes always absent. Cotyledons 2 LOT OTe Tay 27 8 RS Ge ee, ee 5. Aphananthe 2. Leaves pinnately nerved; lateral nerves more than 5 pairs; stipules free but overlapping each other, m falling leaving circular scar around the node. . ......4.+..4+ +56. 6. Gironniera 1.ULMUS LINNE, Gen. Pl. ed. 5 (1754) 106; ENDL. Gen. Pl. (1837) 276, Suppl. 2 (1842) 29; PLANCH. Ann. Sc. Nat. III, 10 (1848) 259; in DC. Prod. 17 (1873) 154; BAILL. Hist. Pl. 6 (1877) 137; B. & H. Gen. Pl. 3 (1880) 351; Hook. f Fl. Br. Ind. 5 (1888) 480; ENGL. in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 62; BERNARD, Bull. Herb. Boiss. II, 5 (1905) 1097; ibid. 6 (1906) 23; SCHNEIDER, Oest. Bot. Z. 66 (1916) 21, 65; in Sargent, Pl. Wils. 3 (1917) 238; GaGnep. Fl. Gén. I.-C. 5 (1927) 674; TuTin, FI. Europ. 1 (1964) 65; Hutcu. Gen. FI. Pl. 2 (1967) 147; Touw & STEEN. Blumea 16 (1968) 84. — Fig. 2, 4-6. Deciduous or semideciduous trees or shrubs. /nnovations densely set with greyish to brownish simple hairs, glabrescent. Buds ovoid-conical or obovoid-globose, scales imbricate, hard and tough, glabrous. Stipules extrapetiolar, caducous. Leaves pinnately nerved, variously serrate to crenate, thin- to thick-coriaceous and rigid, glabrous or variously sparsely hairy at least beneath. Flowers % but of two kinds, one functionally g and the other functionally 2, variously stalked and spirally arranged in fascicles of 3-15 on short lateral shoots. Perianth mostly campanulate, variously 4-8-lobed. Anthers glabrous, reniform, extrorse. Ovary compressed, sometimes stipitate; style short. Ovule 1, anatropous to amphitropous. Fruit a dry and compressed nutlet surrounded by a membranous reticulate-venose wing. Seed: endosperm absent, embryo straight with planoconvex cotyledons. Germina- tion epigeal. Distr. About 20-25 spp., distributed in Europe (as far north as 68°), W. & SW. Russia, N. & NE. India, Burma, China, Korea, Japan, Formosa, Indo-China, N. Thailand, and in North America from N. Mexico to the U.S.A. east of the Rocky Mts as far north as 60°. In Malesia: 1 sp. so far known from a few localities in N. Sumatra, the Lesser Sunda Is. (Flores), and Central & S. Celebes. As has been indicated by SCHNEIDER, /.c., there seem to be three centres of distribution, i.e. the Euro- em crmtre (5-6 spp.), the Indian-E. Asian centre (10-15 spp.), and the North American centre (4-5 spp.). ig. 3. Fossils. Numerous fossils (pollen grains, leaf-impressions, and wood fragments) have been reported from various late Cretaceous and Tertiary deposits in Europe, Russia, China, Japan, North America, and Greenland. Fig. 3. Ecol. In Malesia the genus is so far known only from areas more or less subject to a seasonal climate at 200-1450 m. Taxon. Currently there is not a single worldwide monograph of the genus available for reference. The latest and perhaps the most comprehensive revision since PLANCHON’s work (1873) is that by SCHNEIDER (1916). He distinguished 26 spp. and recognized 5 distinct sections in the genus based on morphological characters derived from inflorescence, flowers, and fruits. When more specimens from China become available for further studies, | believe the number of species occurring in the Indian-E. Asian centre will have to be reduced considerably. 40 FLORA MALESIANA [ser. I, vol. 8? Fig. 3. Approximate range of U/mus L. with number of spp. in each of the three centres, Malesian localities belonging to a species of the Asian centre. Fossil localities outside the present range indicated by dots; adopted from BERNARD, /.c.; GREGUss (Tert. Angios. Hung., Ak. Kiado Budapest, 1969, 83), and La Motte (Mem. Geol. Soc. Am. 51, 1952, 346). Fig. 4. Peeling bark of Ulmus lanceaefolia RoxB. ex WALL., x 1/, (Photogr. ScHMuTz, 5 Nov. 1972, Flores, Nunang). 1. Ulmus lanceaefolia Roxs. ex WALL. Pl. As. Rar. 2 (1831) 86, t. 200; Roxs. Fl. Ind. ed. Carey 2 (1832) 66 (‘lancifolia’); PLANCH. Ann. Sc. Nat. III, 10 (1848) 281; in DC. Prod. 17 (1873) 162; Kurz, For. Fl. Burma 2 (1877) 473; GAMBLE, Man. Ind. Timb. ed. 1 (1881) 342; Hook. f. Fl. Br. Ind. 5 (1888) 480; Hems-. J. Linn. Soc. Bot. 26 (1894) 447; PRAIN, Beng. Pl. (1903) 718; BRANDIS, Ind. Trees (1906) 594; SCHNEIDER, Oest. Bot. Z. 66 (1916) 32; in Sargent, Pl. Wils. 3 (1917) 263; Merr. Contr. Arn. Arb. 8 (1934) 44; Touw & STEEN. Blumea 16 (1968) 84; MELVILLE & HEYBROEK, Kew Bull. 26 (1971) 24 (‘lanceifolia’). — U. hookeriana PLANCH. in DC. Prod. 17 (1873) 162; ENGL. in E. & P. Nat. Pfi. Fam. 3, 1 (1888) 62. — U. tonkinensis GAGNEP. Fl. Gén. I.-C. 5 (1927) 674. — Fig. 2, 4-6. Small to large tree up to 48 m, 70cm @, often with fluted trunk. Bark rough, pustulate, with large warty lenticels. Branchlets initially densely set with greyish to brownish curly simple hairs, later glab- rous and sparsely warty lenticellate. Buds obovoid- globose, c. 2-3 mm @; bracts dark brown. Stipules linear-lanceolate acute, c. 4-5 by 1-11/, mm, soon caducous. Leaves thin- to thick-coriaceous, lanceo- late to ovate-lanceolate, (2-)4-6(-9) by (1-)2-3 (—31/,) cm (index 2-21/,), broadest at or slightly below the middle, more or less glabrous, glossy; base rounded to attenuate-acute, unequal; margin serrulate to serrulate-crenulate; apex acute with blunt tip; midrib raised beneath and flattish to impressed above, as the petiole initially densely greyish, curly hairy on both surfaces, glabrescent; nerves (6~)10-12(-14) pairs, subparallel, often rather irregularly spaced, slightly raised beneath, flattish to impressed above, often forked near and towards the leaf-margin; reticulations fine, areo- late; petiole (2~-)3—-4(—6) by '/,-1 mm. Flowers in fascicles of 3-10. — Functionally 3 flowers globose before anthesis, 1!/,-2 mm @, subglabrous; lobes 5-6, obovate-lanceolate, c. 2 by 1mm; ts 1977] Fig. 5. Ulmus lanceaefolia Rox. ex WALL. with old leaves at Nunang (Photogr. ScHmMutTz, 15 Oct. 1972, Flores). glabrous, slender, c. 1 mm; anthers c. 1 by '/, mm, glabrous; pistillode compressed obovate-elliptic, glabrous. — Functionally 2 flowers (as seen under a very young fruit): perianth campanulate, lobes 5-6, rounded-elliptic, hairy along the margin; filaments slender, glabrous, 3-5 mm, anthers as in 3 flower; ovary stipitate, glabrous, + obovate- elliptic. Fruit obovate-elliptic, glabrous, including the wing 2-3'/, by 1'/,-2cm, stalk 5-10 mm, articulate, lower part hairy. Distr. China (?), India (E. Himalaya, Sikkim, Bhutan, Khasia Hills, Manipur, Assam), Bangla- desh, Burma (Hukong Valley, Chittagong Hills), Thailand (northern parts), Laos, Vietnam (Mt Bavi); in Malesia: N. Sumatra (Gajo- & Karo- Batak Lands), Lesser Sunda Is. (Flores), and Cele- bes (Poso; Bonthain). Fig. 7. Ecol. Scattered tree in lowland to submontane forest, 200-1450 m. In Thailand it is confined to forests along streams and in Flores it has been found on limestone. Fl. fr. in the northern hemis- phere Febr.—April; in Flores Nov. Taxon. U. lanceaefolia is very closely allied to U. parvifolia Jacq. from China and Japan. It differs from the latter by its narrow leaves with a shorter petiole, serrulate to serrulate-crenulate margin, and fewer lateral nerves, obovoid-globose buds, the campanulate perianth of the functionally ? os and the reticulate venation of the fruit; see 2 ULMACEAE (Soepadmo) 4] Uses. Very little is known about the usage of this species, but judging from the enormous size it can attain it must have been a useful timber in house-building, construction, efc., at least to the local inhabitants. Vern. Sumatra: péngki(h), poki, Karo-Batak, pongki, Toba-Batak; Lesser Sunda Is.: ngguling, nggulung, Flores; Celebes: mota, Bonthain. € ny Fig. 6. Ulmus lanceaefolia Roxp. ex WALL., leafless, in flower, + x 2/3, at Nunang (Photogr. ScHMUTZ, 5 Nov. 1972, Flores). Fig. 7. Range of Ulmus lanceaefolia Roxs. ex WALL. 42 FLORA MALESIANA [ser. I, vol. 8? | i Y y | Fig. 8. Parasponia rigida MeRR. & PERRY. a. Habit, x 2/,, b. twig-tip with stipules, c. connate stipules, from inside, d. detail of leaf undersurface, all x 3, e. ¢ flower, f. ditto in LS. g. young 9 flower, h. mature 2 flower, i. ditto in LS, all x 12, j. fruit, k. ditto in LS, both x 12 (af ANU 6463, g—k SCHODDE 4828). 1977] ULMACEAE (Soepadmo) 43 2. PARASPONIA Mia. Pl. Jungh. (1851) 68; Fl. Ind. Bat. 1, 2 (1859) 218; BL. Mus. Bot. 2 (1856) 65; PLANCH. in DC. Prod. 17 (1873) 194; ENGL. in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 65; J. J. SMITH in K. & V. Bijdr. 12 (1910) 662; BAcK. & BAKH. f. FI. Java 2 (1965) 12; Hutcu. Gen. FI. Pl. 2 (1967) 149. — Fig. 8, 10-11. Shrubs to medium-sized trees. Bark grey-brown, smooth to finely fissured ; inner bark fibrous and tough. /nnovations with dense appressed, silvery to greyish hairs. Stipules intrapetiolar, connate into a bifurcate unit and together enclosing the terminal bud, caducous. Leaves (in Mal.) triplinerved at base, concolorous, above non-scabrous to variously scabrous, mostly glabrous except for the midrib and lateral nerves, lower surface variously pubescent. Inflorescences axillary, 3, 9, or 32, much-branched, many-flowered, paniculate or thyrsoid, including the bracts densely short greyish appressed-pubescent. Flowers 5-merous. — ¢ Flower + globose, perianth lobes imbricate in bud; stamens glabrous, introrse; filaments subulate, glabrous; anthers reniform to subglobose, sub-basifixed, glabrous; pistil- lode obovoid-conical, compressed, surrounded by hirsute hairs at its base. — 9 Flower ovoid-conical; staminodes absent; ovary ovoid, slightly compressed; stig- matic arms short, simple; ovule anatropous. Drupe ovoid, slightly compressed pericarp fleshy and fibrous, endocarp hard and stony. Seed: endosperm scanty or copious; embryo curved, cotyledons equal, hypocotyle ascending. Distr. 5 spp., in Polynesia (Tahiti) and Melanesia (Fiji, New Hebrides, Solomons); in Malesia: New Guinea (incl. New Britain), Moluccas (Ternate, Banda), Philippines, Celebes, Lesser Sunda Is. (Lombok, Bali), Java, and S. Sumatra (Palembang). Fig. 9, 12. Fig. 9. Approximate range of the genus Parasponia Mia. (line); localities of P. andersonii (PLANCH.) PLANCH. (dots) and P. melastomatifolia J. J. S. (triangles). Ecol. In New Guinea and the Pacific islands the genus is found as a pioneer plant invading and occupy- ing newly available habitats from the lowland up to 2000 m. In Java the same situation was described b E. W. CLAson from natural regeneration on volcanic ash of Mt Kelud, together with Trema (Bull. Jard. Bot. Btzg III, 13, 1935, 509). Fig. 10, 11. Recorded as a ery on lavastreams of Mt Batur in Bali by De Voocp (Trop. Natuur 29, 1940, 48, f. 12). Grows well in all types of soils, including volcanic ash and limestone, very often gregariously together with Trema spp. and forming a dense thicket on ridges, hills and along river-banks. 44 FLORA MALESIANA [ser. I, vol. 8? TAXON. Parasponia is morphologically very similar to Trema but can easily be distinguished from the latter by its imbricate perianth lobes of the male flowers and intrapetiolar, connate stipules enclosing the terminal bud. KEY TO THE SPECIES 1. Basal nerves running up throughout the length of the leaf or nearly so. 2. Leaf thick-coriaceous, lower surface densely set with soft erect hairs; margin distinctly serrate; reticulations prominent beneath. Inflorescence 3 or 2 petiole... oh , at anthesis condensed and shorter than the 1. P. rigida 2. Leaf chartaceous to thin- -coriaceous, lower surface glabrous or sparsely appressed- hairy; margin finely serrulate to subentire; reticulations obscure. Inflorescence 3 or $9, at anthesis lax and longer than the petiole. Basal nerves running up to ee . 2. P. melastomatifolia 2/, the length of the leaf. "5. Leaf thick-coriaceous, upper surface strongly rugose and scabrous, lower surface densely pubescent; midrib, nerves, and reticulations prominent beneath . P. rugosa 3. Leaf chartaceous to thin-coriaceous, upper surface not or hardly rugose nor ‘scabrous, lower surface sparsely appressed pubescent or glabrous; midrib, nerves, and reticulations only slightly raised beneath. 4. Leaf elliptic-lanceolate, more or less glabrous; nerves more than 4 on each side, straight and ascend- ing at a narrow angle (less than 40°) from the midrib. Inflorescences mostly 32, very rarely g or 2 4. P. parviflora 4. Leaves ovate to ovate-elliptic, underneath sparsely appressed pubescent; nerves less than 4, usually 3 on each side, arcuating at a wider angle (c. 45—60°) from the midrib. Inflorescences 3 or 9, "rarely 32 = 1. Parasponia rigida MeRR. & PERRY, J. Arn. Arb. 22 (1941) 254. — Fig. 8. Small tree, up to 10m, 10cm @. Branchlets initially densely silver-appressed-hairy, glabrescent and sparsely warty lenticellate. Stipules ovate- lanceolate, 8-10 by 2-3mm. Leaves elliptic to ovate-lanceolate, thick-coriaceous, (5—)8—10(—12) by (11/,—)2-31/,(—4!/.) cm (index 2!/,-3'/,), broadest at or below the middle; base rounded to subcordate, mostly symmetrical; margin serrate; apex acute to acuminate; above more or less glabrous, rugulose and scabrous, beneath densely soft-hairy; midrib and nerves strongly raised beneath, flattish to im- pressed above; reticulations subscalariform, dis- tinct beneath; petiole 10-12 by 2-3 mm, terete. Inflorescences 3 or 2, very rarely $2, 10—S0-flowered, at anthesis condensed, c. +/,-1 cm long, usually shorter than the petiole. — ¢ Flowers 1-2 mm @, perianth lobes narrow-ovate, concave, c.2 by 1 mm, appressed-pubescent outside; filaments c. 1 mm; anthers ellipsoid, c. 1 by */, mm, pistillode obovoid- conical, c. 1 by 1/, mm. — 9 Flower ovoid, c. 2 by 1 mm; perianth lobes ovate-acute, c. 3/, by 1/, mm, sparsely pubescent outside; ovary c. 2 by 1 mm, stigmatic arms spreading, long-papillose, c. +/,- 1 mm. Drupe ovoid-globose, c. 3-4 mm @, turning orange to red when ripe. Endosperm copious. Distr. Malesia: New Guinea. Fig. 12. Ecol. Dominant pioneer tree in secondary vegetation on ridges, also in mossy forest and on limestone hills, 1000-2000 m. FI. fr. Jan.—Dec. Vern. Bésukan, Hattam lang., golan, Finsch- hafen dial., Morobe Distr. 2. Parasponia melastomatifolia J. J. SmitH, Nova Guinea 8, 2 (1914) 891, t. 158. — P. simulans MERR. & Perry, J. Arn. Arb. 22 (1941) 255. Shrub to small tree, up to 6 m, with spreading and brittle branches. Branchlets initially densely appressed-silvery-hairy, glabrescent and sparsely lenticellate. Stipules ovate-elliptic, 4-5 by 2-3 mm Leaves elliptic-lanceolate to ovate-elliptic, (5-)6-8 P. andersonii (-10) by (1'/,-)2"/,-3"/,(-41/.)cm (index 2-3), broadest at or slightly below the middle; charta- ceous to thin-coriaceous, above more or less glabrous, hardly scabrous and often with mineral deposits on the epidermis, beneath sparsely ap- pressed-hairy especially on midrib and nerves, or completely glabrous; base more or less rounded, symmetrical; margin finely serrulate to + entire; apex acute; midrib and nerves slightly raised beneath, flattish to impressed above; nerves 1-2 pairs, the basal ones running throughout the length of the leaf or nearly so, at an acute angle of less than 45°; reticulations subscalariform, inconspicuous on both surfaces; petiole (S—)8—12(-15) by 1 mm, sulcate, sparsely appressed-pubescent. Jnflores- cences 3 or 3g, paniculate, 10—20-flowered, at anthesis lax, 2-3 cm long and across, axes c. 1 mm @, densely appressed-pubescent; bracts ovate, c. 11/, by 1mm, sparsely appressed-pubescent outside. — 3 Flowers c. 1-2 mm @, perianth lobes ovate-acute, c. 1/,-1 by 1/, mm, sparsely appressed- hairy outside; filaments c. 1mm, anther sub- globose, c. 1 by !/, mm, pistillode obovoid, c. 4/,-1 by ?/,mm. — @ Flower ovoid, c. 1-2 by 1 mm; perianth lobes ovate-acute, c. +/,-1 by +/,—/, mm, sparsely appressed-hairy outside; ovary ovoid, c. 2 by 1 mm; stigmatic arms c. 1 mm, incurved. Drupe ovoid-globose, c. 3 by 2mm. Endosperm scanty. Distr. Malesia: New Guinea. Fig. 9. Ecol. Common and dominant in seral vegetation in gullies and river-banks, 200-1000 m. Fi. fr Jan.—Dec. Vern. Kwatoro, Onjob lang., Northern Distr., E. New Guinea. 3. Parasponia rugosa BL. Mus. Bot. 2 (1856) 66. — P. aspera BL. l.c. 66; Miq. Fl. Ind. Bat. 1, CA ge 218. — Trema vulcanica MERR. Philip. J. Sc. (1912) Bot. 260. — Trema philippinensis Fine, Leafi. Philip. Bot. 9 (1934) 3218. — P. parviflora (non Mia.) STEEN. Philip. J. Sc. 91 (1962) 507. — Fig. 10-11. 1977] ULMACEAE (Soepadmo) 45 Shrub or medium-sized tree, up to 20 m, 30 cm @, with spreading branches. Bark smooth, grey- brown; inner bark tough, brownish. Branchlets densely, silvery, appressed, long-hairy, subglab- rescent. Stipules ovate-lanceolate, (6—)8(—10) by 2-3 mm. Leaves ovate-lanceolate, (7—)8—10(—12) by (2-)2!/,-31/,(—5) cm (index 2'/,-3), broadest at or below the middle, thick-coriaceous; above rugose and scabrous, sparsely hairy especially on midrib and nerves, beneath densely silvery hairy by soft, erect hairs; base rounded to cordate, equal to slightly unequal; margin serrate, apex acute to acuminate, the acumen up to 2cm; midrib and nerves strongly raised and prominent beneath, flattish to impressed above; nerves 2-4 pairs, arcuate and ascending, the basal ones running up to 1/,7/, the length of the leaf; reticulations dense, areolate, prominent beneath; petiole 7-10 by 2-3 mm, subterete, densely appressed-pubescent. Inflorescences 3 or $2, 10-50-flowered, densely sil- very appressed-hairy, at anthesis condensed, axes c. */,-1"/, cm long, 1-2 mm @; bracts ovate-acute, c. 1*/,-1 mm. — ¢ Flower c. 1-2 mm @; perianth lobes ovate-acute, c. 1-2 by 1 mm, sparsely hairy outside; filaments 1-1!/, mm, anthers ovoid-reni- form, c. 1mm @; pistillode ovoid, compressed, c. 1-2 by 1 mm. — 2 Flower ovoid-conical, c. 1—14/, by 1 mm; perianth lobes narrow ovate-acute, c. 1- 1*/, by 1 mm, sparsely appressed-hairy outside; ovary ovoid, c. 1-2 by 1 mm, stigmatic arms c. 1mm, spreading. Drupe ovoid, 2-3 by 2mm, turning red when ripe. Endosperm copious. Distr. Malesia: East Java (Mts Kelud & Lamongan), Lesser Sunda Is. (Bali, Lombok), Philippines (Luzon, Leyte, Mindanao), Celebes (near Makassar; Tondano, Menado), Moluccas (Ternate, Banda), New Guinea (W. & E. High- lands and Morobe Distr., incl. New Britain). Fig. 12. Fig. 10. Pioneer vegetation on the volcanic ash of Mt Kelud, East Java, of Saccharum spontaneum and Parasponia rugosa BL. (Photogr. CLASON). Ecol. Rather common and often dominant or co-dominant pioneer plant in seral vegetation on various types of soils including volcanic ash, 50— 1900 m. Fi. fr. Jan.—Dec. Fig. 10, 11. Uses. Strips of the inner bark are used as ropes in house and fence building by local inhabitants. Vern. Java: anggring, anggris, J; Philippines: analdung, If.; Moluccas: kayu kuli, Banda; New Guinea: wanep, Enga lang., W. Highlands Distr., la karabi, W. Nakanai, New Britain. Fig. 11. Older pioneer forest on Mt Kelud of Parasponia rugosa BL., Trema, Cyathea contaminans, and Amomum (Photogr. CLASON). 46 FLORA MALESIANA 4. Parasponia parviflora Mig. Pl. Jungh. (1851) 69; Fl. Ind. Bat. 1, 2 (1859) 218, t. 16; BL. Mus. Bot. 2 (1856) 65, f. 35; PLANCH. in DC. Prod. 17 (1873) 194; J. J. SmirH in K. & V. Bijdr. 12 (1910) 663, p.p excl. syn. P. aspera BL.; BACK. & BAKH. f. Fl. Java 2 (1965) 12. — P. similis BL. Mus. Bot. 2 (1856) 66. Small to medium-sized tree, up to 15 m. Branch- lets initially densely silvery or grey appressed-hairy, glabrescent, smooth. Stipules ovate, 5-10 by 2-4 mm, sparsely hairy outside. Leaves lanceolate to narrow ovate-lanceolate, (3—)5-8(-10) by (1-)2-3(—3'/,) cm (index 3-4), broadest at or below the middle; chartaceous to thin-coriaceous, above + glabrous, not scabrous, beneath initially appressed-hairy, later glabrous except for the midrib and nerves; base rounded, more or less equal; margin finely serrate, apex acute; midrib and nerves slightly raised beneath, impressed and inconspicuous above; nerves 4-6 pairs, straight, ascending and parallel, at a narrow angle (30—40°), basal ones running up to + half the length of the leaf; reticulations fine, subscalariform, indistinct on both surfaces; petiole terete, densely appressed- hairy, 5-10 by 1 mm. Inflorescences 3, 2, or 38, 5-30-flowered, at anthesis condensed, shorter than or as long as the petiole, as the bracts densely short-hairy ; bracts ovate-acute, 1—11/, by '/,-1 mm. — 3 Flowers glabrous, 1-2 mm @; perianth lobes c. 1-11/, by ?/, mm; filaments 1/,-1 mm, anthers subglobular, c. 1 by +/,mm; pistillode ovoid- conical, compressed, 1-1'/, by ‘'/,mm. — 9 Flowers ovoid-conical, c. 2 by 11/, mm, + glabrous except for the inner base of the perianth lobes; perianth lobes ovate-acute, c. 1 by '/, mm; ovary ovoid, c. 11/, by 1 mm; stigmatic arms spreading, c. 1/,-1 mm. Drupe ovoid-conical, slightly com- pressed, 11/,-2 by 11/, mm. Endosperm scanty. Fig. 12. Localities of Parasponia rugosa BL. (dots), P. parviflora Mia. (triangles), and P. rigida MERR. & PERRY (squares). Distr. Malesia: S. Sumatra (Palembang, very rare), Java (common). Fig. 12. [ser. I, vol. 8? Ecol. In secondary or seral vegetation on ex- posed habitats, also in teak forest, often rather common and dominant locally especially on soils derived from volcanic ash, 500-2000 m. Fi. fr. Jan.—Dec. Vern. Kurai, k. lélaki, k. tiangkréng, S, anggring, anggris, anggrung, J 5. Parasponia andersonii (PLANCH.) PLANCH. in DC. Prod. 17 (1873) 193. — Sponia andersonii PLANCH. Ann. Sc. Nat. III, 10 (1848) 336; See- MANN, FI. Vit. (1867) 235; PARHAM, Pl. Fiji Isl. (1972) 133. — P. paucinervia MERR. & PERRY, J. Arn. Arb. 20 (1939) 324. Shrub to medium-sized tree, up to 15m and 30cm @. Branches spreading and drooping, initially densely set with erect but soft, silvery hairs, subglabrescent and sparsely warty lenticellate. Bark smooth to nodular, grey-brown; inner bark fibrous, tough, orange to brownish. Stipules ovate- acute, sparsely hairy outside, 6-10 by 3-4 mm. Leaves ovate to elliptic, thin-coriaceous, (5—)8—12 (-14) by (2-)3-4(-6)cm (index 2-3), broadest below or at the middle; above subglabrous, scab- rous, often covered with mineral deposits, beneath sparsely set (rarely rather densely) with short and soft hairs especially on midrib and nerves; base rounded to subcordate, equal, rarely unequal; margin serrate, apex acute to acuminate; midrib and nerves slightly raised beneath and impressed above; nerves 3-4 pairs, arcuating and ascending at an angle of 45-60°, basal extending up to c. 2/, the length of the leaf; reticulations fine, subscalari- form, rather distinct below; petiole (7—)10—15(—20) by 1-2 mm, densely set with silvery, soft, erect hairs, flat or sulcate. Inflorescences 3, 2, or rarely 32, 10-30-flowered, at anthesis condensed or lax, shorter than or as long as the petiole, including the bracts densely silvery, soft-hairy; bracts ovate- acute, c. 1 by !/, mm. — ¢ Flowers c. 11/,-2 mm @; perianth lobes ovate-elliptic, c. 1 by +/,mm; stamens glabrous; filaments c. 1mm, anthers subreniform to subglobular, c. 1 by +/,mm; pistillode subovoid-conical, c. 11/, by !/, mm. — Flowers ovoid-ellipsoid, c. 11/, by 1 mm; perianth lobes ovate-acute, c. 1 by !/,mm; ovary ovoid, slightly compressed, c. 1 by '/, mm; stigmatic arms c. ?/,mm, spreading and short-papillose. Drupe ovoid, slightly compressed, 2-4 by 2-3 mm. Endosperm copious. Distr. Polynesia (Tahiti), Melanesia (Fiji, New Hebrides, Solomons, very common), ? New Caledonia (no specimen seen but cf. GUILLAUMIN, Fl. Nouv.-Caléd. 1948, 94); in Malesia: New Guinea (several islands off Madang and Milne Bay) and New Britain. Fig. 9. Ecol. Primary as well as secondary forests, on various types of soils including limestone, 0-1500 m. Fi. fr. Jan.—Dec. Uses. In the Solomons the bark is reputed to have medicinal properties. Vern. New Britain: ip, ivu; Solomons: bulasisi, bulsisi, Kwara lang.; Fiji: ndroi, ndrou. 1977] ULMACEAE (Soepadmo) 47 3. TREMA Lour. Fl. Coch. 2 (1790) 562; BL. Mus. Bot. 2 (1856) 58; Bru. Fl. Austr. 6 (1873) 157; B. & H. Gen. Pl. 3 (1880) 355; ENGL. in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 65; BERNARD, Bull. Herb. Boiss. II, 6 (1906) 31, maps 19-21; J. J. SMirH in K. & V. Bijdr. 12 (1910) 649; RENDLE, FI. Trop. Afr. 6, 2 (1917) 10; De Wrrt, Bull. Bot. Gard. Btzg III, 18 (1949) 184; Hutcu. Gen. FI. Pl. 2 (1967) 148; Extras, J. Arn. Arb. 51 (1970) 37, f. 2; SOEPADMO in Whitmore, Tree Fl. Mal. 2 (1973) 420. — Sponia COMMERS. ex LAMK, Dict. 4 (1795) 138; ENDL. Gen. Pl. 4 (1837) 276; PLANCH. Ann. Sc. Nat. II, 10 (1848) 264; in DC. Prod. 17 (1873) 195. — Fig. 13, 16-17. Trees or shrubs, often buttressed and with spreading and drooping branches, monoecious. Jnnovations variously and densely set with simple bulbose-based hairs or/and with short multicellular capitate-glandular hairs. Terminal buds ovoid- conical, enclosed by overlapping but free extrapetiolar, caducous stipules. Leaves penninerved ; above + glabrous, variously scabrate, beneath glabrous, subglabrous, or variously densely set with bulbous-based hairs and/or with short multicellular glandular hairs; base triplinerved, cordate to acute, often unequal-sided; margin variously serrate or denticulate; apex acute to acuminate or caudate; petiole sul- cate. Inflorescence axillary, paniculate or thyrsoid, many-flowered, condensed or lax at anthesis, 3, 2, g2, densely and variously pubescent; bracts minute, ovate- acute, caducous. — ¢ Flower globular; perianth 4~S-lobed, lobes induplicate- valvate in bud, boat-shaped; stamens glabrous, introrse; filament subulate, glab- rous, incurved in bud; anthers subglobular to reniform, glabrous, dorsifixed near the base; pistillode present, hirsute at base. — 9 Flower ovoid; perianth 4~5-lobed; staminodes absent or very rarely present; ovary ovoid, (in Mal.) glabrous, slightly compressed, sessile; style short; ovule ana- to amphitropous. Drupe ovoid or sub- globose, (in Mal.) slightly compressed, glabrous; exocarp fleshy and fibrous, endocarp stony and very hard. Seed with a rather scanty or copious endosperm; embryo curved or nearly involute; hypocotyle ascending; cotyledons equal. Germination epigeal. Distr. About 10-15 spp., widely distributed throughout the tropics and subtropics. In Asia (with 6-7 spp.) from the warmer parts of the Himalayas, extending north-eastwards to China (incl. Hainan, Hong- kong, Formosa) and S. Japan and south and south-eastwards through India, Burma, Thailand, Indo- China, and Malesia to the tropical and subtropical parts of Australia and the Pacific islands as far east as Tahiti (31° N-37° S). In Africa (with 3-4 spp.) it occurs south of the Sahara to S. Africa and Madagascar (22° N-28° S). In America (with 4-5 spp.) the genus is known from Central & S. Florida and Mexico, extending south-eastwards through Central America, Bermuda, and the Bahamas, the Greater Antilles and southwards to South America as far south as the northern parts of Argentina (26° N-25° S). In Malesia: 4 spp., widely spread. Fig. 14, 15. Ecol. Throughout its range of distribution the genus seems to grow well and often gregariously in newly opened up habitats on various types of soils ranging from heavy laterite to limestone soils and soils derived from volcanic ash (fig. 17), from sea-level up to 2000 m. Pollination is probably affected by wind and small insects. The fruits which turn orange, red or black when ripe are dispersed by various species of bulbuls. In East Java CLASON (Bull. Jard. Bot. Btzg III, 13, 1935, 509, f. IIT) reported that following the frequent eruptions of Mt Kelud, Trema spp. together with Parasponia spp. formed a dominant association in the regrowths on volcanic ash. Taxon. The genus is homogeneous and closely related to Parasponia and Celtis. This is corroborated by the anatomy of the wood and leaves. Reports on the cytology are, however, suggesting that the number of chromosomes is not constant. svit Embryology. Very little is known about the sporogenesis and embryogenesis of the genus. A preliminary study carried out recently on Trema cannabina and T. tomentosa in the Malay Peninsula indicates that the development of the anther and microspores follow the so-called dicotyledon-type, and that of the embryo- sac conforms with the Polygonum-type. 48 FLORA MALESIANA [ser. I, vol. 8? en exer ‘ Fig. 13. Trema orientalis (L.) BL. a. Habit, with functionally 3 flowers, x 2/3, b-e. 3 flowers at various stages of development, all x 8, f. 2 inflorescence, x 2/3, g. 2 flower, h. older 2 flower, 2 tepals removed, i. ditto in LS, j-k. mature fruit, |. ditto, exocarp halfway removed, showing stone, m. ditto in LS,n.embryo, 0. detail of lower leaf surface, all x 8 (a-e BW 13889, fi BW 7019, j-n Brass 6496). 1977] ULMACEAE (Soepadmo) 49 meets q84q Wi! Nt Ss Bw = AN : 9 Fig. 15. Species density of Trema Lour. in Indo-Australia; above the hyphen the number of endemic spp., below it the number of non-endemic spp. Chromosomes. A few counts on the chromosome number which have been reported by various cytolo- gists suggest that cytogenetically the genus is rather variable. In Trema politoria from India n = 10+B (Mewra & GILL, Taxon 17, 1968, 574; J. Arn. Arb. 55, 1974, 663); in T. orientalis n = 18 (ArorA, Bull. Bot. Surv. India 2, 1960, 305), or n = 20 (GaAJAPaATHY, ibid. 3, 1961, 49; Hsu, Taiwania 13, 1967, 117), or n = 10 (MeHRA & Hans, Taxon 18, 1969, 310; HANs, Cytologia 36, 1971, 341); and in 7. tomentosa (cited as T. amboinensis) n = 10 or 80 (Hans, /.c.; MEHRA, Nucleus 15, 1972, 64). Specific delimitation has proved to be difficult and has led to more than 50 names in the genus. This was partly due to the various interpretations of the early described species. There is still no unanimity of opinion about the number of good species in the continents. In Africa, for example, ENGLER (Pfl. Welt Afr. 3 (1), 1915, 11) estimated the number for Africa at 5-7, following BLUME (1856), but ReNDLE (1917) and piemare th (Kew Bull. 19, 1964, 143) accept only one, either under the specific name 7. guineensis or T. orientalis. 50 FLORA MALESIANA [ser. I, vol. 8? In absence of a critical, reliable world monograph there is a similar uncertainty about the number of species in the neotropics and in Indo-Malesia. For Malesia out of 20-25 published names of species and varieties, only 4 spp. are recognized here. The proliferation of name giving in Malesia is mainly due to the fact that Trema spp. have a growth habit of continuously producing lateral and terminal new shoots on which flowers and fruits are borne. Many specimens collected were from these young shoots in which the indumentum and leaf-shape is often different from that of mature leaves. For accurate identification leaves, inflorescences, and fruits of mature specimens are essential. Besides, the indumentum was in earlier descriptions mostly derived from low magnification observations, but to differentiate sterile material of T. orientalis and T. tomentosa the difference in the indumentum becomes only clear under at least 40 x magnification. It is impossible to name young sterile specimens. KEY TO THE SPECIES (Based on mature leaves, inflorescences, and fruits) 1. Petiole (7—)10-15(-25) mm. Leaves broad ovate-elliptic, rarely narrow lanceolate (but then either glabrous or silvery brown tomentose beneath), (3—)5—10(—13) by (11/.-)2-4(-5'/,) cm (and then glab- rous or sparsely pubescent beneath) or (5—)10—15(-19) by (2—)3-7(—10) cm. Inflorescence 3, 2, or $9, at anthesis lax, 10—100-flowered, 11/,-5 cm long. Mature fruit 3-5 by 2-4 mm. 2. Leaves (3—)5—10(-13) by (11/,-)2-4(-5'/,) cm, glabrous or rarely sparsely pubescent beneath, char- taceous to thin-coriaceous; base rounded, rounded-attenuate, truncate, or very rarely subcordate, mostly symmetrical; nerves 2-4(-5) pairs. Inflorescence always with a slender axis. Mature fruit c. 3 by 2 mm, orange or red in colour 1. T. cannabina 2. Leaves (S—)10-15(-19) by (2-)3-7(-10) cm, densely and variously hairy beneath, thin- to thick- coriaceous; base cordate, subcordate or rounded, mostly asymmetrical; nerves 4-8 pairs. Inflores- cence with a stout or slender axis. Mature fruit black, 3-5 by 3-4 mm. 3. Leaves beneath (fig. 16) completely covered with matted glaucous to silvery straight appressed or curly, erect hairs and short, multicellular capitate-glandular hairs (epidermis invisible even under high magnification), very often distinctly discolorous with the upper surface darker in colour and weakly scabrate. 3 Inflorescence up to 2!/, cm. Fruit + globular. ....... 2. T. orientalis 3. Leaves beneath (fig. 16) densely or sparsely (but not completely) set with velvety greyish-brown erect hairs only (epidermis clearly visible between the hairs even under low magnification), mostly con- colorous, dark-brown to blackish in dried specimens; upper surface strongly scabrate. ¢ Inflores- ea oe te eS. eee Eee? de) ie neem, Sele Bem ld cence up to 5 cm. Fruits ovoid, compressed . CAG pay ese Me ely sy pf CS ae) e: . T. tomentosa 1, Petiole (2-)3-6 (-8) mm. Leaves narrow ovate-lanceolate, (4-)5—6(-11) by (11/,-)2-3(-4) cm, beneath densely tomentose by short, matted, rufous, 1-celled and multicellular glandular hairs (hoary). Inflorescence $ or $9, at anthesis condensed and shorter or as long as petiole, 5—15-flowered. Mature fruit subglobose, 2-3 by 2 mm. 1. Trema cannabina Lour. Fl. Coch. 2 (1790) 563; Me_rr. Trans. Am. Phil. Soc. 24, 2 (1935) 131; DE Wit, Bull. Bot. Gard. Btzg III, 18 (1949) 184; Back. & BAKH. f. Fl. Java 2 (1965) 12; SoepADMO in Whitmore, Tree Fl. Mal. 2 (1973) 421. — Celtis amboinensis WILLD. Sp. Pl. 4, 2 (1805) 997. — Celtis commersonii BRONGN. in Duperrey, Voy. Bot. Coq. Phan. (1829) 215. — Sponia commersonii (BRONGN.) Decne, Nouv. Ann. Mus. Hist. Nat. III, 3 (1834) 498; PLANcH. Ann. Sc. Nat. III, 10 (1848) 317; in DC. Prod. 17 (1873) 198. — Sponia timorensis Decne, Nouv. Ann. Mus. Hist. Nat. III, 3 (1834) 498; PLANcH. Ann. Sc. Nat. III, 10 (1848) 318; in DC. Prod. 17 (1873) 196; Mia. Fl. Ind. Bat. 1, 2 (1859) 216. — Sponia amboinensis (WILLD.) Decne, Nouv. Ann. Mus. Hist. Nat. III, 3 (1834) 498, quoad nomen. — Sponia virgata PLANCH. Ann. Sc. Nat. III, 10 (1848) 316; in DC. Prod. 17 (1873) 195, incl. var. major PLANCH. lI.c. 196; Miq. FI. Ind. Bat. 1, 2 (1859) 216; Kurz, For. Fl. Burma 2 (1877) 469. — Sponia glabrescens PLANCH. Ann. Sc. Nat. III, 10 (1848) 317; Mia. Fl. Ind. Bat. 1, 2 (1859) 217. — Sponia viridis PLANCH. Ann. Sc. Nat. IIT, 10 (1848) 319. — T. commersonii (BRONGN.) BL. Mus. Bot. 2 (1856) 60. — T. timorensis (DECNE) BL. /.c. 60; Hoox. f. Fl. Br. Ind. 5 (1888) 483; Hems-. J. Linn. Soc. Bot. 26 (1894) 452; Laur. Bot. Jahrb. 50 (1913) 317, incl. var. carinata (BL.) 4. T. angustifolia Laut. et var. pallida (BL.) LAuT.; HAND.-MAZz. Symb. Sin. 7 (1929) 107. — T. virgata (PLANCH.) BL. Mus. Bot. 2 (1856) 59; J. J. SMirH in K. & V. Bijdr. 12 (1910) 652; Laur. Bot. Jahrb. 50 (1913) 313, incl. var. pubigera (BL.) Laut. lc. 315; SCHNEIDER in Sargent, Pl. Wils. 3 (1917) 289; RIDL. Fl. Mal. Pen. 3 (1924) 319; GAGNepP. Fl. Gén. I.-C. 5 (1927) 686; HAND.-Mazz. Symb. Sin. 7 (1929) 106; CorNER, Ways. Trees (1940) 694; Li, Woody FI. Taiwan (1963) 109. — T. amboinensis (WILLD.) BL. Mus. Bot. 2 (1856) 61, guoad nomen, excl. syn. et sched.; MERR. Int. Rumph. (1917) 187. — T. viridis (PLANCH.) BL. Mus. Bot. 2 (1856) 58. — T. glabrescens (PLANCH.) BL. /.c. 58. — T. cari- nata BL. l.c. 59. — T. pallida BL. l.c. 60. — T. pubigera BL. l.c. 60. — T. morifolia BL. l.c. 59; Laut. Bot. Jahrb. 50 (1913) 318. — Sponia pallida (BL.) Mia. Fl. Ind. Bat. 1, 2 (1859) 215; PLANCH. in DC. Prod. 17 (1873) 196. — Sponia carinata (BL.) Mia. FI. Ind. Bat. 1, 2 (1859) 215; PLANCH. in DC. Prod. 17 (1873) 202. — Sponia pubigera (BL.) Mia. FI. Ind. Bat. 1, 2 (1859) 216; PLANcH. in DC. Prod. 17 (1873) 197. — Sponia morifolia (BL.) PLANCH. in DC. Prod. 17 (1873) 196. — Sponia vieillardii PLANCH. I.c. 201. — Sponia aspera var. viridis (PLANCH.) BTH. Fl. Austr. 6 (1873) 158. — T. orientalis var. amboinensis (WILLD.) KuRZ, For. Fl. Burma 2 (1877) 469, guoad nomen. — T. vieil- 1977] ULMACEAE (Soepadmo) 51 lardii (PLANCH.) SCHLTR, Bot. Jahrb. 36 (1905) 31. Shrub or small much-branched tree up to 6m, 15cm @. Bark smooth, grey-brown. Branchlets slender, spreading, often drooping, initially densely silvery-hairy, glabrescent and sparsely lenticellate. Stipules linear-lanceolate, 5-7 by 1-2 mm. Leaves chartaceous to thin-coriaceous, narrow ovate- caudate to broad ovate-acute, or elliptic-lanceolate, (3-)5-10(-13) by (11/,-)2-4(-5"/,) cm (index 2-3 (4)), broadest below or at the middle; base rounded to attenuate and acute, rarely subcordate, slightly contracted and more or less symmetrical; margin serrulate to denticulate for its entire length; apex with a sharp tip; above glabrous and variously scabrate, beneath glabrous or sparsely appressed- hairy; midrib and nerves raised beneath, impressed above; nerves (2—)3—4(—5) pairs, arcuate and sub- parallel, basal ones running up to + 2/, the length of the leaf; reticulations fine, subscalariform, obscure to visible beneath; petiole (S—)8—12(-15) by 1-2 mm, glabrescent. Inflorescence 3 or 39, with slender axes, 10—15-flowered, at anthesis lax, c. 1-2'/, cm long, densely greyish appressed-hairy ; bracts ovate-acute, c. 2-3 by 1 mm. — ¢ Flowers c. 1-2 mm @, outside sparsely hairy, glabrescent; perianth lobes 4-5, membranous, oblong-lanceo- late, c. 1-1'/, by */,-1 mm; filaments c. 1 mm, an- thers c. 1 by !/, mm; pistillode obovoid, compressed, c. 1 by '/, mm. — 9 Flowers c. 11/,-2 by 1-1'/, mm; perianth lobes mostly 5, membranous, glabrous, ovate-acute, c. 1—1"/, by !/, mm; staminode absent; ovary c. 1 by '/, mm; stigmatic arms spreading or incurved. Drupe 2-3 by 2 mm, turning deep-orange or red when ripe. Endosperm copious. Distr. Burma, China, Formosa, Hainan, Indo- China, Thailand, common throughout Malesia to Australia, Melanesia (Solomons, New Caledonia, New Hebrides), W. Polynesia (Fiji, Samoa), and Micronesia. Ecol. Common as a pioneer in newly opened up habitats along roadsides, edges of forests, re- growths, thickets, and in young secondary vegeta- tion, from sea-level up to 1200 m. FI. fr. Jan.—Dec. At least in Malaya pollination is affected by wind and by small insects (diptera). Ripe fruits are dis- persed by various species of bulbuls. Taxon. In Malesia there seem to be three rather but not completely distinct entities. These can be as follows: (i) specimens which have a completely glabrous, chartaceous and narrow- ovate leaf of (3—)5—8(—10) by (1!/,-)2-3(-4) cm with an index of 2'/,-3, more or less non-scabrate upper surface, and 2-3 lateral nerves (7. cannabina and T. virgata); (ii) specimens with a thin-coriaceous, narrow ovate-lanceolate to elliptic-lanceolate leaf of (6-)7-10(-12) by (1'/,-)2-3(-3'/,) cm, with an index of 4-5, slightly scabrate upper surface and sparsely hairy lower surface, and 4-5 lateral nerves which are straight and ascending and forming a narrow angle (less than 40°) with the midrib (7. timorensis, T. virgata var. scabra, and T. cannabina var. scabra) ; (iii) those with a broad ovate and coria- ceous leaf of (8—)9-11(-13) by (3-)4-4'/,(—5'/,) cm, with an index of 2'/,-3, rugose and slightly scab- rate upper surface and sparsely pubescent beneath, and 3-4 lateral nerves forming a broad angle (more than 45°) with the midrib (7. glabrescens, T. viridis, and T. vieillardii). Various intermediates are how- ever present, making it difficult to recognize them as distinct infra-specific taxa. Vern. Malaya: ménérong, méngkirai, M; S. Sumatra: délung, M, Palembang; Java: anggrung, J; N. Borneo: bintanong, Murud; Lesser Sunda Is. : rédong kué, Flores, pépaka, Alor: Moluccas: loli-sawu, Halmaheira; Solomons: bulasisi, Kwara. 2. Trema orientalis (L.) BL. Mus. Bot. 2 (1856) 62; Bru. FI. Austr. 6 (1873) 158; Hook. f. Fl. Br. Ind. 5 (1888) 484; J. J. SmMirH in K. & V. Bijdr. 12 (1910) 655, p.p., excl. syn. T. commersonii et T. griffithii; Laut. Bot. Jahrb. 50 (1913) 320, p.p., incl. var. rigida (BL.) Laut. l.c. 322, excl. var. viridis et var. amboinensis; CORNER, Ways. Trees (1940) 694, pl. 211, p.p. excl. syn.; DE Wit, Bull. Bot. Gard. Btzg III, 18 (1949) 189, p.p., incl. var. bicornis DE Wir, /.c. 190, excl. var. bicolor et syn. T. angusti- folia, T. imbricata et T. velutina; BACK. & BAKH. f. Fl. Java 2 (1965) 12, p.p., excl. syn. T. amboinensis auct. non (WILLD.) 'BL.; SOEPADMO in Whitmore, Tree Fl. Mal. 2 (1973) 421. — Celtis orientalis LINNE, Sp. Pl. 2 (1753) 1044; Roxs. Fl. Ind. ed. Carey 2 (1832) 65. — Celtis rigida BL. Bijdr. (1825) 486. — Celtis discolor BRONGN. in Duperrey, Bot. Voy. Coq. Phan. (1829) 215, pl. 47B. — Sponia discolor (BRONGN.) DECNE, Nouv. Ann. Mus. Hist. Nat. III, 3 (1834) 498; PLANcH. Ann. Sc. Nat. III, 10 (1848) 324; in DC. Prod. 17 (1873) 201. — Sponia orientalis (L.) DECNE, Nouv. Ann. Mus. Hist. Nat. III, 3 (1834) 498; PLANCH. Ann. Sc. Nat. III, 10 (1848) 323; in DC. Prod. 17 (1873) 200. — Sponia rigida (BL.) DECNE, Nouv. Ann. Mus. Hist. Nat. III, 3 (1834) 498; PLANcH. Ann. Sc. Nat. III, 10 (1848) 336; Mra. FI. Ind. Bat. 1, 2 (1859) 217. — Sponia argentea PLANCH. Ann. Sc. Nat. III, 10 (1848) 323; in DC. Prod. 17 (1873) 201. — Sponia wightii PLANCH. Ann. Sc. Nat. III, 10 (1848) 322; WicuT, Ic. 6 (1853) t. 1971. — T. argentea (PLANCH.) BL. Mus. Bot. 2 (1865) 58. — T. bur- mannii BL. I.c. 62. — T. rigida (BL.) BL. /.c. 61. — T. scaberrima BL. l.c. 63. — T. wightii (PLANCH.) BL. /.c. 58. — T. discolor (BRONGN.) BL. /.c. 58; Laut. Bot. Jahrb. 50 (1913) 319. — Sponia scaber- rima (BL.) Mig. Fl. Ind. Bat. 1, 2 (1859) 217; PLANCH. in DC. Prod. 17 (1873) 202. — Sponia burmannii (BL.) PLANCH. in DC, 17 (1873) 200. — Fig. 13, 16. Shrub to large tree, 3-36 m, 10-90 cm @. But- tresses, if present, up to 1'/,m. Bark smooth to finely fissured, lenticellate, grey-brown or whitish- grey. Branchlets, stipules, petioles, and inflores- cences densely set with appressed and matted or erect silvery to glaucous 1-celled hairs and short multicellular glandular hairs. Stipules linear- lanceolate to ovate-acute, 3-4 by 1-2 mm. Leaves thin- to thick-coriaceous, often rigid and brittle, ovate, ovate-lanceolate to narrow elliptic, lan- ceolate, (6-)10—-15(-18) by (1'/,-)2'/,-6(-10) cm, index (2-)3—4(—5'/,), broadest at or mostly below the middle, mostly discolorous, above dull grey- brown or grey-green in dried specimens, scabrate and sparsely set with bulbous-based hairs, beneath densely oamsds by a combination of silvery, glaucous or grey-brown, appressed 1-celled hairs and shorter multicellular glandular hairs (fig. 16); base cordate, rounded, or sometimes truncate, often contracted, asymmetrical or symmetrical ; margin serrate to denticulate for its entire length; 52 FLORA MALESIANA [ser. I, vol. 8? apex acute to acuminate-caudate; midrib and nerves raised beneath and impressed above; nerves 4-6(-8) pairs, the lowest pair arcuating and run- ning up to !/,—*/, the length of the leaf; reticulations subscalariform to subareolate, sometimes strongly raised and distinct beneath; petiole (7—)10—15(-18) by 1-2 mm, densely short pubescent. Inflorescences either ¢ or 2 borne on separate vegetative branches, a much-branched panicle or thyrse, at anthesis lax or condensed, axes 1-2 mm thick; bracts ovate- acute, 2-3 by 1mm. Flowers 5-merous. — ¢ Inflorescences up to 3—5 cm long, 20-100-flowered ; 3 flower c. 11/,-2 mm @; perianth lobes ciliate, 11/,-2 by 1mm; filaments 1-1'/,mm, anthers c. 1 by 1/, mm, pistillode obovoid-conical, com- pressed, 1-1!/, by '/,-1 mm. — 2 Inflorescences 5-15-flowered, 11/,-2!/,cm long, axes 1-2 mm thick; 2 flower c. 2-3 by 1-2 mm; perianth lobes ovate acute, c. 1-1!/, by 1/, mm, ciliate and densely short pubescent, glabrescent; staminode absent; ovary ovoid-conical, c. 2 by 1 mm; stigmatic arms slender, c. 1-11/,mm., spreading. Drupe 3-5 by. 2-4mm, turning black when ripe. Endosperm scanty to copious. Distr. ?Tropical Africa, SE. Asia (Ceylon, India: from W. Himalayas to Bombay and Mala- bar; Burma, Thailand, Indo-China, China, also Hainan, Formosa, to S. Japan), through Malesia to Queensland, Melanesia (Solomons), Micronesia (Marianas), and Polynesia (Fiji, Tonga, Tahiti). In Malesia: Malay Peninsula and Sumatra (rather rare), Java (rather common in the hills and sub- montane regions), Lesser Sunda Islands (rare), Borneo (common), Philippines (rather common), Celebes (rather rare), Moluccas (rare), and New Guinea (incl. New Britain, rare). Ecol. In W. Malesia and continental Asia the species is more common in the hills and montane regions between 600-2000m, whereas in E. Malesia, Australia, and Pacific Islands it is more common in the lowlands. The ripe fruits which turn to deep purple or black are dispersed by, ~\.s Le . i — —" various species of birds, particularly bulbuls. Fl. fr. Jan.—Dec. Taxon. Three rather but not completely distinct entities may be recognized. These are: (i) specimens from continental Asia and W. Malesia which have been variously identified as T. orientalis, rigida, argentea, and wightii by previous authors. They are characterized by: thick-coriaceous, broadly ovate to ovate-elliptic leaves with grey-brown to glaucous indumentum, slightly asymmetrical to symmetrical cordate, subcordate or rounded base, rugose upper surface, and acute to acuminate apex; and by a relatively larger fruit of c. 4-5 by 3-4 mm and stouter inflorescence axes. (ii) Specimens from S. Japan, Formosa, Hainan, the Philippines, New Guinea, Micronesia, Mela- nesia, and Polynesia, and Australia, which have been included in the so-called T. discolor, charac- terized by: thin-coriaceous, narrow-ovate leaves with strongly asymmetrical cordate base, hardly scabrate upper surface, short and matted silvery to grey-brown indumentum, lax inflorescence with slender axes, and fruits c. 3-4 by 2-3 mm. (iii) A few specimens from scattered localities in S. China, Thailand, Sumatra, and Borneo, which have been described by DE Wir (1949) as T. orien- talis ssp. bicornis, characterized by: very narrow, ovate-lanceolate thin-coriaceous leaves with silvery appressed and matted dense indumentum on the lower surface and non-scabrate upper surface, 6-8 pairs of nerves, and the shorter and few- flowered inflorescence. Several intermediates are present however, making formal infraspecific distinction not ad- visable. Vern. ndérung, Karo, Sumatra: endrung, indarung, Pajakumbu, bandorung, Tapanuli, lando- jung, Simelungun, éndélung, Palembang, Bencoo- len, magélong, nélung, Bencoolen, nériung, Lam- pong. Java: gorai, kuraj, S, anggrung, njampu, J. Lesser Sunda Is.: /énggung, Bali, rédong, Flores, tabélah, W. Sumbawa. Borneo: randagong, x */ a - ; ‘i oS * Dey La Fig. 16. Indument of lower leaf-surface, strongly enlarged. Left: only unicellular hairs of Trema tomentosa (Roxs.) HARA, between which the epidermis is visible. Right: 7. orientalis (L.) BL., with long unicellular hairs and crowded multicellular crisped hairs covering the epidermis. 1977] tandago, Dusun; béngkirai, E. Kutai. Philippines: anadgong, Bis. Celebes: ngawoi, Malili, mawa, Bonthain, kantu, Toradja, tajapu. Moluccas: rufu, Ternate, /aei, Tidore, soka soka, E. Ceram. West New Guinea: bésuwai, Hattam lang., karara, Ambai, kKaniem, mier, Kebar lang. 3. Tremia tomentosa (RoxB.) HARA, Fl. E. Himal. 2 (1971) 19; So—PADMo in Whitmore, Tree Fl. Mal. 2 (1973) 423. — Celtis orientalis (non L.) BL. Bijdr. (1825) 485. — Celtis amboinensis (non WILLD.) BRONGN. in Duperrey, Bot. Voy. Coq. Phan. (1829) 212, pl. 47A, p.p., excl. specim. ex Ventenat, Amboina. — Celtis tomentosa Roxs. Fl. Ind. ed. Carey 2 (1832) 66. — Sponia amboinensis (WILLD.) Decne, Nouv. Ann. Mus. Hist. Nat. III, 3 (1834) 498, quoad specim.; PLANCH. Ann. Sc. Nat. III, 10 (1848) 321; Mia. Fl. Ind. Bat. 1, 2 (1859) 216; PLANCH. in DC. Prod. 17 (1873) 198. — Celtis lima (non Sw.) BLANCO, FI. Filip. 2 (1837) 139.— Sponia griffithii PLANCH. Ann. Sc. Nat. III, 10 (1848) 324. — Sponia tomentosa (ROxB.) PLANCH. /.c. 336. — Sponia velutina PLANCH. I.c. 327, p.p., excl. specim. Cuming 1232 ex Luzon. — Sponia blancoi PLANCH. l.c. 327; Mia. Fl. Ind. Bat. 1, 2 (1859) 218. — 7. griffithii (PLANCH.) BL. Mus. Bot. 2 (1856) 58. — T. blancoi (PLANCH.) BL. /.c. 58. — T. imbricata BL. /.c. 63. — T. velutina (PLANCH.) BL. l.c. 58; GaGnepP. Fl. Gén. I.-C. 5 (1927) 689; L1, Woody FI. Taiwan (1963) 109. — T. amboinensis (WILLD.) BL. Mus. Bot. 2 (1856) 61, quoad specim.; BTH. FI. Austr. 6 (1873) 159; Hook. f. Fl. Br. Ind. 5 (1888) 484; K. Sco. & Laut. Fl. Schutzgeb. (1900) 264; J. J. SmirH in K. & V. Bijdr. 12 (1910) 659, p.p., excl. syn. Celtis amboinensis WILLD. et Trema burmannii Bi.; Merr. En. Born. (1921) 217; Ript. Fl. Mal. Pen. 3 (1924) 319. — Sponia imbricata (BL). PLANCH. in DC. Prod. 17 (1873) 199. — T. orientalis var. amboinensis (WILLD.) Kurz, For. Fl. Burma 2 (1877) 469, quoad specim.; Laut. Bot. Jahrb. 50 (1913) 321. — T. orientalis (non (L.) BL.) MERR. Sp. Blanc. (1918) 121. — T. dielsiana HAND.-MAzz. Symb. Sin. 7 (1929) 106; P’e1, Bot. Bull. Ac. Sin. 1 (1947) 289. — Fig. 16. Shrub to medium-sized tree of 5—15(—24) m, 5—30(-50) cm @. Bark grey-brown, smooth to finely fissured, lenticellate. Branchlets, inflores- cences, petioles, stipules, and underside of leaves densely and thickly set with greyish, erect, velvety hairs. Stipules linear-lanceolate, c. 5 by 1 mm. Leaves thin- to thick-coriaceous, broadly ovate to ovate-elliptic, (5—)8-15(-19) by (2-)4~7(-9) cm, index 2'/,-3, broadest mostly below the middle; more or less concolorous, drying dark-chocolate brown to blackish brown; above strongly scabrate; base cordate, rarely subcordate or rounded, mostly strongly asymmetrical, rarely symmetrical; margin serrate throughout, apex acute to acuminate- caudate, acumen sharp, 1-3 cm; midrib and nerves raised beneath (often very strongly), impressed and hairy above; nerves pairs, ascending and solicavatiel, at an angle of + 45°, the lowest pair running to + */,/, the length of the leaf ; reticu- lations subscalariform to subareolate, often rather distinct beneath; petiole 1-1'/,cm by 1-2 mm, densely pubescent. Inflorescences 3,9, or 39, either on the same or on different vegetative branches; bracts ovate-acute, c. 1 by '/, mm. — At anthesis dé and 3° axes of the inflorescences lax, 2'/,-4'/, cm ULMACEAE (Soepadmo) 53 long, 20—100-flowered; 3 flower c. 1'/,2mm @; perianth lobes mostly 5, elliptic, c. 11/, by 1 mm; filaments c. 1 mm, flat, glabrous, anthers c. 1 by ‘/,mm; pistillode obovoid-ellipsoid, compressed, 1'/, by '/, mm. — 2 Inflorescence 1-2 cm long, axes 1-2 mm thick, 5—15-flowered; 2 flower c. 2 by 1 mm; perianth lobes 4-5, ovate-acute, c. 1 by ‘/,mm; staminode mostly absent, if present strongly reduced in size and non-functional; ovary c. 1"/, by '!/,-1 mm, stigmatic arms slender, c. 1 mm, spreading. Drupe c. 3 by 2mm, maturing black. Endosperm copious. Distr. East tropical Africa, Madagascar, SE. Asia: Pakistan, India, Bangladesh, Burma, Thai- land, Indo-China, China (incl. Hainan), Hong- kong, Formosa, Ryu Kyu Is. (Okinawa), through- out Malesia to Queensland, Melanesia (New Caledonia), Micronesia, and Polynesia (Fiji, Tonga, and Hawaii). Ecol. Common in the lowlands and hills, at sea- level up to 1000 m, as a pioneer plant invading and occupying newly opened up habitats on all kind of soils, including limestones. F/. fr. Jan.Dec. At least in Malaya pollination is affected by wind and small insects (diptera). The ripe black fruit is dis- persed by various species of birds. Taxon. Evidently, 7. tomentosa is closely allied to T. orientalis, and it is possible that, when more field data become available in the future, the former may prove to be only representing a juvenile ontogenetical form of the latter. Except for a few specimens from the Philippines (e.g. WHITFORD 681, BS 37313, 48355, ELMER 8417) and from New Guinea (e.g. ANU 2075, 2752, 6240, HARTLEY 10937, MANNER & STREET 270, NGF 29353, SCHODDE 1419, and BW 16510) in which the leaves are thick-coriaceous and with a more or less sym- metrical base and pale grey-brown in colour, specimens of 7. tomentosa can be easily dis- tinguished from those of T. orientalis by the charac- ters mentioned in the key. Fig. 16. It is also interest- ing to note that according to HANs (Cytologia 36, 1971, 341) and MEHRA (Nucleus 15, 1972, 64) the chromosome number in T. tomentosa is n = 10 or 80, whereas that of T. orientalis isn = 10, 18, or 20. Vern. Malay Peninsula: ménarong, méndarong, méngkirai, M. Sumatra: béngkirai, Gajo, éndélung, Palembang, eémaha, Enggano, hana(w)e, Batak, kamésén silai, Simalur, mangkirai, Pajakumbu, manghirei, ménkirei, Lingga, mudén sabu, Djambi, (n)dér(r)ung, Karo-Batak, randurung, Toba, sang- kiraja, Batak, tindjau, Riouw. Java: anggrung, J, kuraj, k. awéwena, S. Lesser Sunda Is.: rédong, damot, Flores, ruka parak, Sumba. Borneo: Sara- wak: murieng, Bidajuh, kérénénog, Iban, tuku baroh, Land Dayak; N. Borneo: anjalakat, Kedayan; Brunei: balék balék angin jantan, balik angin, rundagong, Brunei, bintanong, Murut, damai, Suluk, entimon, Iban, lindagong, Kedayan, landa- gong, ‘Dusun Tambato & Kayan, /undagong, sali- muak, Dusun, randagong, Tenggara, réndagong, Dusun Labuk; E. Borneo: bangérai, bangkirai, tjalundung, E. Kutai; W. Borneo: éngkirai, butu. Philippines: anaginong, Mang., anugdon, Tag-Bis., anabiong, hanagdong, Tag., ‘arangyan, karayang- yang, Tagb. Moluccas: mandalirung’ a, Talaud, pohon rupong, Banda, rufut, Buru. West New Guinea: fidukwa, Manokwari, hormas, Sorong; East New Guinea: komukai, Maring name, natua, 54 FLORA MALESIANA | Fig. 17. Young blukar (regrowth or secondary forest) on an abandoned tea estate near Tapos, West Java, c. 1000 m, consisting of three layers: 2 m high stand of Eupatorium inulifolium, 5-6 m high tree ferns of Cyathea contaminans, above which is an open canopy of Trema orientalis (L.) BL. (Photogr. VAN STEENIS). Kainantu, seraun, Daga-Bonenau, wanip, Enga lang., wantip, Medlpa, Wahgi, wan’um, Mendi. 4. Trema angustifolia (PLANCH.) BL. Mus. Bot. 2 (1856) 58; Hook. f. Fl. Br. Ind. 5 (1888) 484; GAGNEP. FI. Gén. I.-C. 5 (1927) 686; HAND.-MAzz. Symb. Sin. 7 (1929) 108. — Sponia angustifolia PLANCH. Ann. Sc. Nat. III, 10 (1848) 326; Mia. FI. Ind. Bat. 1, 2 (1859) 215; PLANcH. in DC. Prod. 17 (1873) 202. — Sponia acuminatissima MiQ. Sumatra (1861) 410; PLANcH. in DC. Prod. 17 (1873) 202. — Sponia sampsonii HANCE, Ann. Sc. Nat. V, 5 (1866) 242. — T. acuminatissima (MIQ.) BOERL. Handl. 3 (1900) 358. — T. lanceolata MerR. Lingn. Sc. J. 7 (1931) 302. — T. sampsonii (HANCE) MERR. & CHUN, Sunyatsenia 5 (1940) 40. — T. orientalis var. ut DE WIT, Bull. Bot. Gard. Btzg III, 18 (1949) Shrub or small tree with spreading and drooping branches, 3-7 m, 5-15 cm @. Branchlets densely set with rufous multicellular glandular hairs and glaucous short and matted 1-celled hairs, sub- glabrescent. Stipules linear-lanceolate, 3-4 by 1mm. Leaves chartaceous to thin-coriaceous, narrow ovate-lanceolate to lanceolate, (3—)5—8(-10) by (1-)2-3(-4) cm, index 3-3/,, broadest below or at the middle; discolorous, upper surface strongly scabrate, dark chocolate-brown to blackish-brown, [ser. I, vol. 82 lower surface densely set with short and matted rufous to glaucous 1-celled and multicellular glandular hairs; base rounded to attenuate, sym- metrical; margin finely serrate throughout; apex acute to acuminate; midrib and nerves slightly raised beneath, impressed above; nerves 4-5 pairs, straight, ascending at 30-40°; reticulations fine, subscalariform to subareolate, obscure above and faintly visible beneath; petiole (2—)3-6(-8) by 1 mm, densely short hairy. Inflorescences 3 or 39, densely set with a short rufous indumentum, much- branched, (5—)10-15(—30)-flowered, at anthesis condensed, shorter than or as long as the petiole; bracts narrow ovate-acute, c. 4/,-1 by 1/,—'/, mm. — 6 Flowers c. 1-11/,mm @; perianth lobes 5, elliptic, c. 1-11/, by 1 mm; filaments c. 1 mm long, c.1/,-1 mm @; pistillode obovoid-ellipsoid, strong- ly compressed, c. 4/,-1 by 1/,mm. — 9 Flowers ovoid-conical, c. 2 by 1 mm; perianth lobes 5, narrow-lanceolate, acute, 1/,-1 by 1/,—'/,mm; staminode absent; ovary c. 1-2 by 1 mm; stig- matic arms c. 1/,-1 mm, spreading or incurved. Drupe c. 1/,-2 mm @, turning orange to red when ripe. Endosperm copious. Distr. China (Yunnan, Hainan), Thailand, and Indo-China; in Malesia: Malay Peninsula (com- ae Sumatra (rare), Borneo (rare), Celebes (very rare). 1977] ULMACEAE (Soepadmo) 55 Ecol. Scattered in newly available habitats in Excluded the lowlands to submontane regions, from sea-level : er rate eae . Trema (Parasponia) lancifolia Ripv. J. Mal. Br. to 1200 m. Fi. fr. Jan.—Dec. Ripe fruits are dis- R. As. Soc. 1 (1923) 91 = Debregeasia longifolia persed by various species of bulbuls. ‘ Vern. Malaya: ménarong, méngkirai, M; (Burm. f.) WEDD. (Urticaceae). Sumatra: kayu anggurung, M, Eastcoast, mang- kirai kétjil, M, Palembang. 4.CELTIS LINNE, Gen. PI. ed. 5 (1754) 467; Sp. Pl. 2 (1753) 1043; PLANCH. Ann. Sc. Nat. III, 10 (1848) 262; BL. Mus. Bot. 2 (1856) 70; Mia. Fl. Ind. Bat. 1, 2 (1859) 220; PLANCH. in DC. Prod. 17 (1873) 168; B. & H. Gen. Pl. 3 (1880) 354; ENGL. in E. & P. Nat. Pfl. Fam. 3, | (1888) 63; BERNARD, Bull. Herb. Boiss. II, 5 (1905) 1112, maps 9-15; J. J. SMitH in K. & V. Bijdr. 12 (1910) 639; Leroy, Fl. Madag. et Com. Fam. 54 (1952) 3; POLHILL, Kew Bull. 19 (1964) 139; Hutcu. Gen. FI. Pl. 2 (1967) 147; Extas, J. Arn. Arb. 51 (1970) 32; SOEPADMO in Whitmore, Tree Fl. Mal. 2 (1973) 414. — Solenostigma ENDL. Prod. Fl. Norf. (1833) 41; BL. Mus. Bot. 2 (1856) 66; Mia. Fl. Ind. Bat. 1, 2 (1859) 219. — Fig. 18, 20, 22-23. Small to large monoecious or polygamo-monoecious trees, often buttressed. Bark smooth or finely fissured, often conspicuously warty lenticellate. Branches (in Mal.) unarmed, initially densely yellow-brown or rufous-hairy, glabrescent; hairs l-celled. Buds enclosed by the overlapping stipules or naked. Stipules thick and tough, peltately attached or free and scarious, caducous. Leaves entire dr not, 3-nerved at base, semideciduous or persistent. /nflorescenes 3, %, or 3%, branched racemes or panicles, few- to many-flowered, axillary or subterminal on the new shoot; staminate inflorescences borne on the lower and leafless part or in the axil of leaves of the new shoot; in the ¢% inflorescence the &% flowers are borne on the distal ends of the axes; bracts minute, caducous. — Staminate (3) flowers globular, pedicelled or sessile; perianth lobes 4-5, imbricate in bud, membraneous, boat- shaped, outside sparsely pubescent, at anthesis recurved, caducous; stamens glabrous, inserted on the densely pilose receptacle; filaments subulate, incurved in bud and spreading elastically, exserted at anthesis; anthers ovoid to subreniform, dorsifixed just above the emarginate base, extrorse; pistillode present or absent. — % Flowers ovoid, pedicelled; perianth lobes 4-5, imbricate in bud, connate at base, membranous, outside sparsely pubescent, boat-shaped, at anthesis recurved, cadu- cous; stamens well-developed and functional or rudimentary, other characters as in 3 flowers; ovary ovoid-ellipsoid, sessile, style short or + absent; stigmatic arms elongate, divergent, the tips entire to deeply bifid; ovule anatropous. Drupe fleshy, ovoid, ellipsoid or globose; exocarp thick and firm, mesocarp thin and fleshy, con- taining slimy substances; endocarp hard and persistent, smooth or variously ridged or pitted. Seed: coat membranous, chalazal area broad, dark-coloured and close to the minute hilum; endosperm scanty or wanting, oily or gelatinous, nearly enclosed between the folds of the cotyledons. Embryo curved, cotyledons broad, foliaceous, equal or unequal in thickness, flat or conduplicate, variously folded, incumbent on or embracing the short superior and ascending radicle. Germination epigeal. Distr. About 50-60 PP: widely distributed in tropical and temperate regions of the world, the majority of species (30-40) in the Old and New World tropics, throughout Malesia (9 spp.). Fig. 19, 21. 56 FLORA MALESIANA [ser. I, vol. 8? a A eee Fig. 18. Celtis philippensis BLANCO. a. New shoot bearing flowers, x 10, f. fruits, x 2/5, g. fruit, exocarp in LS, x 11/3, and chalaza, nat. size, j. ditto in LS showing folded in LS, x 1*/;. — C. tetrandra Roxs. 1. New shoot bearing flowers, x 2/3, flowers, x 10 (a—c MERRILL 52, d-e BS 1920, f-k BARTLETT 15071, -o Scumutz 1666). x 2/3, b-c. 3 flowers, x 8, d-e. 3 flowers, h. stone, endocarp in LS, x 11/3, i. seed showing testa cotyledons, shaded oily endosperm, x 1'/3, k. embryo m-n. § flowers, x 8, 0. ¢. 1977] ULMACEAE (Soepadmo) 57 Fig. 19. Approximate range of Celtis L. Fossil records are indicated by dots. Fossils. Numerous fossilized wood-fragments, leaf-impressions, drupes and pollen grains have been discovered in various localities in Europe, the U.S.A., and in Asia. According to ELtas /.c. the first (oldest) records are apparently from the early Eocene in Wyoming and the late Eocene in Georgia, U.S.A. Continuing through the Oligocene, Celtis spp. are best known from the Miocene, fossilized material of younger age is relatively sparse at least in the U.S.A. — References: A. GRAHAM (ed.), Floristics and Paleofloristics of Asia & Eastern North America (1972) 147; GreGuss, Tert. Angios. Hung., Ak. Kiado Budapest (1969) 83; La Motte, Mem. Geol. Soc. Am. 51 (1952) 112. Fig. 19. Ecol. Malesian species of Celtis may be classified into two rather distinct ecological groups, i.e. the C. philippensis var. philippensis and C. tetrandra groups. The first group, which includes also C. hilde- brandii, C. latifolia, C. luzonica, C. paniculata, and C. rigescens, is found mainly in the lowland forests, both primary and secondary, and is an important constituent of the understorey tree community in moist areas. The C. tetrandra group, which includes C. rubrovenia, C. timorensis, and C. philippensis var. wightii, is confined to areas which are subject to a rather pronounced seasonal climate, or if they occur in wetter regions, they grow on strongly drained substrates, e.g. rocky shores, limestone, efc. In tune with this environmental preference, the second group shows a more prominent flush-wise growth habit and its species are completely or partly deciduous. In Malaya, species of Celtis are producing flowers around July-September, while fruit ripens January—March. How pollination is affected is not certain, but judging from the structure and position of the inflores- cence, some species (e.g. C. philippensis var. philippensis, C. hildebrandii, and C. latifolia) may be pollinated Hd = while others (C. tetrandra, C. timorensis, C. rubrovenia, and C. rigescens) may be pollinated wind. The ripe fleshy drupes which turn to orange, red or bluish-black may be dispersed by birds, or alter- lose they may be dispersed by water as the embryo is protected by the hard, persistent and durable endocarp. Morph. Except for C. paniculata the stamens of the pistillate flowers are well-developed and functional. In the male or staminate flowers the pistillode is rudimentary or completely absent in C. tetrandra, C. timorensis, and C. rubrovenia. In the other species the pistillode is present and relatively rather well- developed though non-functional. Chromosomes. The chromosome numbers reported are: n = 10 (2n = 20) (C. australis var. eriocarpa, C. inguana, C. laevigata, C. occidentalis, C. sinensis, and C. timorensis (under C. cinnamomea)); 2n = 22 (C. spinosa); 2n = 28(C. occidentalis);2n = 40(C. australis and C. tupalangi). — References: Sax, J. Arn. Arb. 14 (1933) 82; Bowpen, Am. J. Bot. 32 (1945) 195; DARLINGTON & WyLie, Chromos. Atlas (1955) 182; MeHra & GILL, Taxon 17 (1968) 574; GADELLA et al. Acta Bot. Neer]. 18 (1969) 74; MEHRA & HANs, Taxon 18 (1969) 310; Feporov (ed.), Chromos. Numbers Flow. Pl. (1969) 710; MeHra & GILL, J. Arn. Arb. 55 (1974) 663. : According to Sax I.c. there seems to be at least in C. occidentalis a high degree of pollen sterility and a high incidence of meiotic irregularity. This may be one of the causes why in Celtis there is a very high percentage of barren seeds production, even among tropical species. f Embryology. No detailed study on the microsporogenesis, megasporogenesis and embryogenesis of Aye aig has ever been carried out. In Malesia the solitary ovule is bitegmic, anatropous and inserted just below the apex of the locule. After fertilization both integuments develop into thin membranous seed coats with a broad, dark-coloured, more or less circular chalaza. The endocarp becomes woody and very hard and impregnated by mineral deposits. It is persistent and becomes variously sculptured (ridged, 58 FLORA MALESIANA [ser. I, vol. 8? pitted, or nearly smooth). The embryo is strongly curved with the hypocotyle superior and ascending, situated in between or nearly enclosed by the broad, thick, foliaceous cotyledons. The cotyledonar lobes are somewhat unequal in thickness, and they are either induplicate or variously folded. Endosperm is very scanty to absent and either gelatinous or oily. Especially in C. paniculata and C. tetrandra, at least 70-80% of the fruits produced are barren. Though the fruits are developed normally, the embryo fails to grow and becomes shrivelled. As a result the fruits are empty. KEY TO THE SPECIES (Measurements of leaf and fruit based on fully mature material) 1. Leaves entire or nearly so. ¢ Inflorescence a much-branched many-flowered panicle with up to 150 flowers. Pistillode rather well-developed, c. 1—-11/, by 1/,-1 mm. Stigmatic arms bilobed or bifid at the tip. 2. Leaves rugose, brittle, sparsely pubescent beneath; midrib and lateral nerves strongly raised beneath. Stipules not peltately attached, free from one another. Fruit densely appressed-hairy. 1. C. rigescens 2. Leaves not rugose, not brittle, glabrous; midrib and nerves only slightly raised beneath. Stipules peltately attached, overlapping. Fruit glabrous. 3. Leaves with (2—)3—5 pairs of nerves. 4. Leaves elliptic-orbicular or elliptic-oblong, index (11/,—)1*/,(—2); midrib and nerves slightly raised beneath; lowest pair of nerves running to 7/,—*/, the length of the leaf, upper pairs of nerves ascend- ing and arcuating. Stigmatic arms shallowly bilobed at the tip. Fruit globose, 16-20 by 14-18 mm. 2. C. luzonica 4. Leaves elliptic or ovate-eliptic, index (11/,—)2—2'/,(—3); midrib and nerves flattish beneath; lowest pairs of nerves running up to !/;—'/,(—?/,) the length of the leaf; upper pairs of nerves weak, sub- horizontal. Stigmatic arms deeply bifid at the tip. Fruit ovoid or ellipsoid, 7-12 by 5-8 mm. 5. Nerves 3-5 pairs; lowest pair running to '/;-'/, the length of the leaf. Inflorescence mostly d or ¥. Stamens of § flower rudimentary, non-functional. Fruit ovoid, feebly 4-5-angular in CS; endo- carp with reticulate ridges... .... . PES ee ee 3. C, paniculata 5. Nerves 1—2(—3) pairs; lowest pair running up to 2/, the length of the leaf. Inflorescence g or ¢ 4. Stamens of % flower well-developed and functional. Fruit ellipsoid, + terete; endocarp smooth. 3. Leaves with 1 pair of nerves. 4b;).C. philippensis var. wightii 6. Leaf symmetrical. Inflorescence ¢ or 38. 3 Inflorescence 15—-40-flowered. Cotyledons curved but not folded. 7. Leaves (4-)8-14(-18) by (2—)3—6(-8) cm; reticulations fine, dense. Ovary glabrous. Fruit 8-15 by 7-12 mm; endocarp smooth 4a. C. philippensis var. philippensis 7. Leaves (8-)15—18(—25) by (6—)8—-12(-18) cm; reticulations coarse, wide-spaced. Ovary densely appressed-pubescent. Fruit 15-25 by 10-18 mm; endocarp pitted. ..... 5. C. latifolia 6. Leaf oblique. Inflorescence f and %. 3 Inflorescence 60-150-flowered. Cotyledons cure and many 6. times folded. hildebrandii 1. Leaves serrulate to denticulate at least in the upper half. g¢ Inflorescence racemose, 5—20-flowered. Pistillode strongly reduced in size or absent. Stigmatic arms entire. 8. Leaf not strongly oblique in outline; nerves 1—2(—3) pairs. Inflorescence 3 and 3%; 3 flower 5-merous. 9. Leaves (6-)10—12(-17) by (21/.-)4-S(-8) cm, index 11/,-2'/. g Inflorescence 10—20-flowered. Ovary glabrous. Infructescence 4-5 cm long. Fruit ovoid, strongly beaked, 5-10 by 3-6 mm. 7. C. timorensis 9. Leaves (3—-)4-5(-6!/,) by (11/,-)1!/,-2'/,(-3'/,) cm, index 2-3.3. 3 Inflorescence 5—7-flowered. Ovary densely hairy. Infructescence c. 1'/, cm. long. Fruit globose, not beaked, c. 3 by 3 mm 8. Leaf strongly oblique; nerves 3-4 pairs. Inflorescence gj and %; ¢ flower 4-merous.. 1. Celtis rigescens (M1Q.) PLANCH. in DC. Prod. 17 (1873) 182; So—EPADMO in Whitmore, Tree Fl. Mal. 2 (1973) 416. — Solenostigma rigescens Mia. Sumatra (1861) 411. — Solenostigma sumatrana Mia. /.c. 411. — C. sumatrana (MiQ.) PLANCH. in DC. Prod. 17 (1873) 181. — C. nymanii K.Scu. in K.Sch. & Laut. Fl. Schutzgeb. Nachtr. (1905) 240; Laur. Bot. Jahrb. 50 (1913) 311. — C. asperi- folia MeRR. Philip. J. Sc. 17 (1920) 246; En. Philip. 2 (1923) 32. — Fig. 20e, 23a-b. Large tree up to 45 m, 1 m ©. Buttresses up to 6m tall, 3m out, 5cm thick. Bark grey-brown, smooth, finely fissured to pustulate and lenticellate. Innovations densely yellowish brown to rufous simple hairy. Older twigs glabrous, finely striate 8. C. rubrovenia 9. C. tetrandra and sparsely minute lenticellate. Terminal buds ovoid-conical, c. 3 by 2 mm, scales densely yellow- ish-brown tomentose. Stipules linear-lanceolate, 4-5 by 1-1'/, mm. Mature /eaves thick-coriacecus, strongly rugose, stiff and brittle when dry, ovate- elliptic to elliptic-oblong, (5S—)8-12(-15) by (2%/,-) 3—5(-6!/,) cm (index 11/,-2'/,), broadest at or slightly below the middle; above glabrous, shining, beneath sparsely yellowish-brown pubescent es- pecially on midrib and nerves; base rounded to subcordate, symmetrical, rarely attenuate-rounded and slightly asymmetrical; margin undulate, en- tire or distantly serrate in the upper half, very often incurved; apex rounded-acute to acuminate; midrib and nerves strongly raised beneath, flattish 1977] ULMACEAE (Soepadmo) 59 Fig. 20. Celtis paniculata (ENDL.) SPACH. a. 2 Inflorescence, x 8, b. flower, b. var. korthalsiana 1. Inflorescence subglabrous to puberulous. 4. Fruit with prominent persistent disk, over 21/, by 11/, cm. Leaves 10-24 by 5-12 cm. Inflores- cence robust, up to 15cm, branched 5 (to 6) times, not terminating in a dichasium. d. var. rhynchocarpa 4. Fruit without prominent persistent disk, up to 2 by 1 cm. Leaves usually 5-12 by 2-4'/, cm. Inflorescence slender, up to 10 cm, branched 3(-4) times, often terminating in a dichasium e. var. cClarkeana a. var. trichotoma — M. laxa BL. — M. trichotoma BL. var. laxa Mig. — M. laxa BL. var. angustifolia BL. — M. acuminatissima BL. — M. trichotoma BL. var. acuminatissima DANSER — M. caesia BL. —M. kimanilla Bt. incl. var. caesia Miq. Branchlets rather slender, yellowish, puberulous to velutinous. Leaves elliptic to oblong, 5-15 by 3-8 cm, chartaceous to subcoriaceous, sub- glabrous to puberulous; base cuneate to attenuate; apex acute to acuminate; nerves 5-8 pairs, promi- nulous to prominent below, seldom arcuate; petiole 11/,—2(—2!/,) cm, rather slender. Inflores- cence up to 8 cm, compact, branched 3(-4) times, villous to woolly; basal bracts under 5mm; terminal bracts often deeply boat-shaped. Sepals 4. Petals 4. Stamens 4. Fruit elongate-ovoid, 11/.-3 by 1977] 1-11/, cm; persistent disk inconspicuous; sepals prominent. Distr. Malesia: N. Sumatra, W.-E. Java, Borneo, Lesser Sunda Is. (Bali). Fig. 5. Ecol. In primary forest from low altitude up to 1800 m. Fi. May—Jan., fr. July—March. Galls occur on stem and fruit. Vern. Java: djérét, huru hiris, h. minjak, kéndu, kibéntéli, kibunting, kidédak, kilumlum, kilun glum, kiménjan, kiténjo, (huru) méhmal, mémah, palaglar minjak, ténggau, ténju. b. var. korthalsiana (WANGERIN) DANSER, Blumea | (1934) 63; MATTHEW, Blumea 23 (1976) 70. — M. korthalsiana WANGERIN. — Fig. 3g (galls). Branchlets rather slender, yellowish, subglab- rous to velutinous. Leaves subovate-elliptic, 10-12 by 3-5 cm, subcoriaceous; base long attenuate; apex acuminate; nerves 5-6 pairs, seldom arcuate; petiole 1/,—-13/,cm, rather slender. Inflorescence up to 10cm, very lax, branched 3(-4) times, few- flowered, velutinous to woolly; bracts under 3 mm. Sepals 5. Petals 5. Stamens 5. Fruit elongate-ovoid, 2'/,-3 by 1-1'/, cm, persistent disk inconspicuous, sepals prominent. Distr. Malesia: Sumatra, Borneo. Fig. 5. Ecol. In primary forest from low altitude up to 640 m, often scattered. Rather prone to galls. Vern. Sumatra: médang kladi. E. Borneo: sérgam pipit, Sangkulirang I. Note. The lax and few flowered inflorescence, the 5-merous flowers, and elongate fruit distinguish this variety from var. trichotoma to which it is closely allied. ce. var. maingayi (CLARKE) DANSER, Blumea 1 (1934) 63; MATTHEW, Blumea 23 (1976) 70. — M. maingayi CLARKE, incl. var. subtomentosa KING — M. junghuhniana (non MiQ.) CLARKE — M. rostrata (non BL.) Rip_. — M. propinqua RIDL. Branchlets very stout, yellowish, woolly. Leaves ovate to elliptic, 9-20 by 4-11 cm, thick coriaceous, tough and stiff; base obtuse to truncate; apex acute to acuminate; nerves 5-6 pairs, deeply impressed above, very prominent below, often arcuate, numerous parallel intermediary veins conspicuous; petiole 1'/,-2'/,cm, stout, villous to woolly. Inflorescence up to 15cm, compact, branched 4(-5) times, velutinous to golden woolly; basal bracts up to 1 cm, persistent. Sepals 4. Petals 4. Stamens 4. Fruit ovoid, 2'/,-3'/, by 11/,-2.cm; persistent disk inconspicuous, sepals prominent. Distr.Malesia: Sumatra, Malay Peninsula (also Penang), Banka, Borneo. Fig. 5. Ecol. In primary forests with Dipterocarps; also in secondary or marsh forests, from the lowland to 1400 m. Fl. Febr.-Au . (Nov. ), Sr. April—Dec. Galls occur on leaves and fruit. Vern. Malaya: karu nuri, kayu béngkal bukit, kayu maura, médang. Banka: médang puntung, m. pusér. Borneo: médang kanigara. Note. Easily recognized by the woolly indumen- tum, large and stiff leaves with prominent veins, and the numerous massive fruits. d. var. rhynchocarpa DANseR, Blumea | (1934) 64; MATTHEW, Blumea 23 (1976) 71. — M. trichotoma . var. benculuana et var. simalurana DANSER. — Fig. 3h (galls. CORNACEAE (Matthew) 93 Branchlets stout, dark, subglabrous. Leaves elliptic to oblong, 10-24(-28) by 5-12cm, coriaceous, tough and stiff; base cuneate to obtuse; apex acute, acuminate to caudate; nerves 8-15 pairs, very prominent below, seldom arcuate; petiole 11/,-2'/,(—3'/,) cm, stout. Inflorescence up to 15cm, compact, profusely branched up to 5(-8) times, not terminating in a dichasium, sub- glabrous to puberulous; basal bracts up to 5 mm. Sepals 4(—5). Petals 4. Stamens 4. Fruit ovoid to elongate-ovoid, 2!/,-3'/, by 11/,-2 cm; persistent disk prominent, bulging; sepals prominent. Distr. Malesia: Sumatra (incl. Simalur I.), W Java, Borneo, NE. Celebes, Moluccas (Ambon, Ceram). Fig. 5. Ecol. Common in primary lowland and mossy forest, up to 1800 m. Fi. April—Aug., fr. June- March. Galls. This variety is very prone to fruit galls and the largest ones in the genus (over 1!/, cm @) occur here. Vern. Sumatra: ahélat, awa ahélat uding, awa énti, awa simangurach, tutun simangurah, Simalur I.; bung, médang tima, tanah, Bencoolen. Borneo: médang aima. Moluccas: soya. Note. This variety is noted for the generally large dimensions of leaves and inflorescence, though there is a reduction in size from Borneo to Moluccas. e. var. Clarkeana (KING) DANSER, Blumea 1 (1934) 62; MATTHEW, Blumea 23 (1976) 72. — M. clarkeana KING, incl. var. macrophylla Kinc — M. korthalsiana WANGERIN var. macrophylla WANGERIN — Vitex eg lab od ELMER — M. prem- noides HALL. f. — M. trichotoma BL. var. tenuis DANSER. Branchlets slender, grey, subglabrous. Leaves oblong to elliptic-oblong, 5—12(-18) by 2—4(-8'/,) cm, coriaceous; base cuneate to obtuse; apex acute to acuminate; nerves 5-7 pairs, seldom arcuate; petiole 1-1'/,cm, slender. Inflorescence up to 10cm, compact, branched 3(—4) times, often ter- minating in a dichasium. Sepals 4. Petals 4. Stamens 4. Fruit ovoid, 1*/,-2 by */.-1 cm; per- sistent disk inconspicuous; sepals prominent. Distr. Peninsular Thailand (Pattani) and Malesia: Sumatra, Banka, Malay Peninsula, Borneo, Philippines (Mindanao). Fig. 5. Ecol. Primary forest, from low altitude to 1100m. Fl. Jan.—Aug., fr. July—Febr. Galls occur on stem and fruit. Vern. Philippines: /amog. 4. Mastixia eugenioides MATTHEW, Blumea 23 (1976) 73. Tree up to 30 m; d.b.h. up to 30 cm; branchlets stout, opposite, glabrous. Leaves opposite, elliptic to oblong-elliptic, 4-12 by 2-5'/,cm, thick- coriaceous, glabrous; base cuneate; apex acumi- nate to caudate; nerves 5-7 pairs, prominent beneath, with intermediary ones and distinct veins; petiole 1'/,—2'/, cm, stout. /nflorescence up to 8 cm, rather stout and compact, glabrous, up to 4 times branched, at times terminating in a dichasium; branches ‘of the first order opposite; higher order bracts triangular, under 3mm; lower bracts lanceolate, up to 5mm, all glabrous. Submature flower bud 2mm @. Sepals 4, broader than long, 94 FLORA MALESIANA [ser. I, vol. 8? thick, glabrous. Petals 4, thin, glabrous outside. Stamens 4. Ovary glabrous. Fruit (unripe) ovoid, 2'/, by 1'/,cm; persistent disk and sepals incon- spicuous. Distr. Fig. 4. Ecol. Primary (often Dipterocarp) lowland forests, up to 400 m. Fl. July—Aug., fr. Sept. Notes. Leaf scars conspicuous; inflorescence notably erecto-patent when young, spreading later. The inflorescence and flowers somewhat resemble those of M. rostrata ssp. rostrata, but the stout branchlets with strictly opposite leaves and stout petiole, prominent intermediary veins, and fruits of different shape with thick pericarp, make this species quite distinct. All the 9 collections are from a restricted area. Malesia: Borneo (Sarawak, Brunei). 5. Mastixia rostrata BL. Mus. Bot. 1 (1850) 258; Mia. Fl. Ind. Bat. 1, 1 (1856) 773, (1858) 1095; K. & V. Bijdr. 5 (1900) 92; WANGERIN, Pfi. Reich Heft 414 (1910) 22; Koorp. Atlas 1 (1913) t. 191; DANsER, Blumea 1 (1934) 52; Burk. Dict. (1935) 1428; Doct.v.LEEUWEN, Ned. Kruidk. Arch. 51 (1941) 207; Back. & BAKH. f. Fl. Java 2 (1965) 159; MevER, Bot. News Bull. Sandakan 8 (1976) 65; MATTHEW, Blumea 23 (1976) 73. — M. jung- huhniana Mia. FI. Ind. Bat. 1, 1 (1856) 772. — M. margarethae WANGERIN in Fedde, Rep. 4 (1907) 335; Pfil. Reich Heft 41* (1910) 21. — M. cuspidata BL. var. margarethae HALL. f. Beih. Bot. Centralbl. 34, 2 (1916) 41. — M. caudatifolia MerR. Pl. Elm. Born. (1929) 233. Tree up to 30 m; d.b.h. up to 50 cm; branchlets slender, (sub)opposite or scattered, glabrous. Leaves (sub)opposite or scattered, elliptic to oblong- elliptic, 4-10 by 2-5cm, chartaceous to sub- coriaceous, glabrous; base cuneate; apex caudate over 1 cm; nerves 4-6 pairs, prominulous beneath; veins obscure; petiole 1—2'/, cm, slender. Inflores- cence up to 6cm, slender, compact or lax, sub- glabrous, up to 4 times branched, branches of the first order (sub)opposite or scattered; bracts triangular, under 3mm, glabrous. Submature flower bud 1-2!/,mm @. Sepals 4, broader than long, thin, glabrous. Petals 4, glabrous outside. Stamens 4. Ovary glabrous. Fruit ovoid to oblong, 1+/,-3 by 3/,-1 cm, persistent disk prominent or inconspicuous, sepals inconspicuous. Distr. Malesia: Sumatra, Banka, Java, Borneo, Lesser Sunda Is. (Sumbawa, Flores). Fig. 4. KEY TO THE SUBSPECIES 1. Submature flower bud 2!/, mm @. Inflorescence compact, branches of the first crder (sub)- opposite. Leaves (sub)opposite. Galls absent a. ssp. rostrata 1. Submature flower bud 1 mm @. Inflorescence lax, branches of the first order scattered. Leaves scattered. Galls frequent. . b. ssp. caudatifolia a. ssp. rostrata — M. junghuhniana Mia. Branchlets (sub)opposite. Leaves (sub)opposite, less often scattered ; petiole 11/,-2 cm. Inflorescence compact, generally under 4 cm, glabrous; primary branches (sub)opposite. Submature flower bud 2//,mm @. Distr. Malesia: West & Central Java, Lesser Sunda Is. (Sumbawa, Flores). Fig. 4. Ecol. Moist forest, from low altitude up to 1400 m. Fl. March, very fragrant, fr. May-July. Galls absent. Ripe fruit dark blue. Vern. Java: daun kaju téndjo, daun kitadjas, huru gading, kilburoy, kiléjas, kiténdjo, kitindjo, lalakina, tjangkar. Flores: bumis, rau, tapadeke. Notes. Specimens from the Lesser Sunda Is. have larger leaves than those from Java. The reference in KANJILAL & Das (Fl. Assam 2, 1938, 371) to this species seems erroneous, as it does not occur on the Asian continent. If the state- ment “‘stamens 3”’ is correct, the plant can even not belong to Mastixia. b. ssp. caudatifolia (MERR.) MATTHEW, Blumea 23 (1976) 74. — M. margarethae WANGERIN — M. caudatifolia MERR. — Fig. 3f (galls). Branchlets scattered. Leaves scattered; petiole up to 1'/, cm. Inflorescence very lax, up to 6 cm, puberulous at the nodes; primary branches scattered. Submature flower bud 1 mm @. Distr. Malesia: northern half of Sumatra, Banka, Borneo. Fig. 4. Ecol. Primary forest, from the lowland up to 1600 m. F/. June—Oct., fr. Aug.—March. Globose to elongate galls are common, specially those on fruits. Sometimes they resemble a legume and can be up to 31/, cm long. Vern. Borneo: patoli entelit, Iban lang. Note. It is almost impossible to separate sterile materials of ssp. caudatifolia from those of M. cus- pidata, though their flowers are entirely different. 2. Series Alternae MATTHEW, Blumea 23 (1976) 75. Inflorescence branches of the first order scat- tered; branchlets and leaves scattered; nodes terete; fruit generally ellipsoid or oblong. Distr. Ceylon and continental Asia; through Seti but absent in New Guinea and Solomon S. 6. Mastixia macrocarpa MATTHEW, Blumea 23 (1976) 75. Tree up to 21 m; d.b.h. up to 20 cm; branchlets stout, scattered, woolly. Leaves scattered, elliptic- oblong to oblong, 13-30 by 51/,-15 cm, subcoria- ceous, villous, especially below; base cuneate, at times slightly oblique; apex acuminate; nerves 7-10 pairs, prominent below, villous; veins promi- nulous, puberulous to villous; petiole 4-7 cm, stout, woolly. Inflorescence up to 9cm, stout, woolly, branched up to 4 times; branches of the first order scattered ; bracts triangular to lanceolate, up to 8 mm, densely woolly. Submature flower bud 4mm @. Sepals 5, broader than long, villous. Petals 5, velvety outside. Stamens 5. Ovary densely villous. Fruit oblong-ovoid, 4-4'/, by 2 cm; per- sistent disk inconspicuous, sepals prominent, up to 5 mm. Distr. Malesia: Borneo (Sarawak), Philippines (Luzon); 2 collections. Fig. 6. Ecol. Lowland forest. Fi. Oct., fr. June. Sticky resin on the branches; fruits very pale green. Note. Leaves and fruits are the largest known in the genus; lenticels up to 3 by 1 mm; leaf scars up 1977] CORNACEAE (Matthew) 2 to 4 by 4mm; peduncles of terminal cymes up to 6mm; pedicels 2mm; sepals 2 mm broad at the base; filaments 3 mm; anthers 1'/, mm; receptacle 3-31/, by 2mm; style 4mm; stigma 5-lobed, appearing bifid. ®@ M. mecrocarpa Matthew O m. glauca matthew @ M. cuspidate Slume Fig. 6. Localities of three species of Mastixia. 7. Mastixia glauca MATTHEW, Blumea 23 (1976) 76 Tree up to 15m; branchlets stout, scattered, glabrous. Leaves scattered, obovate, 7-16 by 4'/,-8'/, cm, thick-coriaceous, glaucous and waxy below, glabrous; base obtuse; apex apiculate; nerves 4-5 pairs, with intermediary ones, all obscure; veins obscure; petiole 2-3'!/, cm, stout, glabrous. Inflorescence up to 5 cm, rather stout and compact, subglabrous to sparsely puberulous, branched up to 5 times; branches of the first order scattered; bracts triangular, under 3mm. Sub- mature flower bud 3 mm @. Sepals 4(-5), as long as broad, sparsely puberulous. Petals 4(—5), thick, appressed-hairy outside. Stamens 4(-5). Ovary sparsely puberulous. Distr. Malesia: Borneo (Sarawak: Mt Santu- bong), 3 collections Fig. 6. Ecol. Lowland forest. F/. April-May. Note. Tender bark of branchlets yellowish; inflorescence clearly broader than long; 5-merous flowers only occasional. 8. Mastixia tetrapetala Merr. Philip. J. Sc. 13 (1918) Bot. 42; En. Philip. 3 (1923) 242; MATTHEW, Blumea 23 (1976) 76, f. 5 (map). — M. pachyphylla Mere. Philip. J. Sc. 13 (1918) Bot. 325; En. Philip. 3 (1923) 241. — M. crassifolia MeRR. Philip. J. Sc. 26 (1925) 486. Tree up to 8(-15)m; branchlets very stout, scattered to subopposite, subglabrous. Leaves scattered to subopposite, crowded at apices of branchlets, obovate to oblanceolate, 5-15 by 2-7 cm, thick coriaceous, glabrous; base cuneate; apex acute to acuminate; nerves 6—8(-12) pairs, usually prominent below; veins prominulous below; petiole 1-2'/,cm, stout, glabrous. IJn- escence up to 3-(6) cm, stout, very compact, berulous to villous, 2(—3) times branched; lee of the first order scattered; higher order bracts triangular, under 3 mm; lower ones lanceo- late up to 5 mm, puberulous. Submature flower bud 5 mm @. Sepals 4, broader than long, glabrous to appressed-hairy. Petals 4, thick, glabrous to appressed-hairy. Stamens 4. Ovary glabrous to appressed-hairy. Fruit ellipsoid, 2-3 by 1%/,— 11/, cm; persistent disk and sepals inconspicuous. Distr. Malesia: Philippines (Luzon, Catan- duanes), 7 collections. Ecol. Primary forest, from low altitude up to 2300 m. Fi. Febr.—March, fr. Sept.—Febr. Notes. Branchlets stout with conspicuous leaf scars and fibrous bark. Phyllotaxis tends to be obscured owing to congestion of parts: leaves are generally crowded towards the apices of branch- lets. Inflorescence branches do not always elongate as in other species. The species is quite distinct and stands rather isolated from others in the stoutness of parts, the large, 4-merous flowers, and the large, ellipsoid fruits. Two other species of MERRILL, M. pachyphylla and M. crassifolia are considered conspecific with M. tetrapetala. There are indeed certain differences: leaves of M. crassifolia generally have 8-12 nerves per side prominent below, and massive fruits. M. pachyphylla has (sub)opposite leaves and pri- mary inflorescence branches. However, when examined together, it is seen that both M. pachy- phylla and M. crassifolia are extreme variations of M. tetrapetala. 9. Mastixia cuspidata BL. Mus. Bot. 1 (1850) 256; Mia. FI. Ind. Bat. 1, 1 (1856) 772; HALL. f. Beih. Bot. Centralbl. 34, 2 (1916) 41; DANseR, Blumea 1 (1934) 55, excl. var. margarethae; MATTHEW, Blumea 23 (1976) 79. — M. pentandra BL. var. cuspidata Miq. Fl. Ind. Bat. 1, 1 (1858) 1095; WANGERIN, Pfi. Reich Heft 414 (1910) 26. — M. bracteata CLARKE, FI. Br. Ind. 2 (1879) 746; KING, J. As. Soc. Beng. 71, ii (1902) 73; WANGERIN, Pfl. Reich Heft 41* (1910) 26, f. 1 G-K, N-O; DANSER, Blumea 1 (1934) 54. Tree up to 24 m; d.b.h. up to 40 cm; branchlets very slender, scattered, subglabrous. Leaves scattered, obovate, elliptic or oblong, 4-12(—16) by 2-4(—6) cm, subcoriaceous, glabrous; base cuneate; apex abruptly cuspidate (over 1 cm), oblique; nerves 4 (or 5) pairs, arcuate, imp above; veins obscure; petiole 1—1'/, cm, slender. Inflores- cence up to 4cm, rather slender, subglabrous to puberulous, 2(—3) times branched; branches of the first order scattered; higher order bracts subulate, lower ones foliaceous, over 10 mm, passing into foliage leaves. Submature flower bud 3 mm @. Sepals 5, broader than long, subglabrous. Petals 5, thick, densely appressed-hairy outside. Stamens 5. Ovary densely appressed silky-hairy. Fruit oblong, 1'/,-2 by °/,-l1.cm; persistent disk and sepals inconspicuous. Distr. Malesia: Sumatra, Banka, Malay Pensinula, Borneo. Fig. 6 Ecol. Primary and secondary forests, from low altitude up to 900 m. Vern. Sumatra: bébung, kundur. Banka: mén- kapas. Malay Peninsula: dadaru. Borneo: biansu- gunong, Sarawak. 10. Mastixia BL. Bijdr. (1826) 654; DC. Prod. 4 (1830) 275; BL. Mus. Bot. 1 (1850) 256; Mig. Fl. Ind. Bat. 1, 1 (1856) 771, (1858) 1095; 96 FLORA MALESIANA K. & V. Bijdr. 5 (1900) 88; Merr. Philip. J. Sc. 1 (1906) Suppl. 111; DANserR, Blumea 1 (1934) 49; BACK. & BAKH. f. Fl. Java 2 (1965) 159; MATTHEW, Blumea 23 (1976) 80, f. 5 (map), 6. — M. arborea [non (WIGHT) BEDD.] CLARKE, FI. Br. Ind. 2 (1879) 745, p.p.; KANJILAL & Das, Fl. Assam 2 (1938) 370; HUNDLEY & CuitT, Trees Shr. Burma ed. 3 (1961) 119. — M. cambodiana Pierre, Fl. Coch. (1892) t. 260 B; WANGERIN, Pfl. Reich Heft 41 (1910) 29; EvrarD, FI. Gén. I.-C. 2 (1923) 1195. — M. scortechinii Kinc, J. As. Soc. Beng. 71, ii (1902) 73; WANGERIN, Pfi. Reich Heft 41* (1910) 27, f. 1 L-M; DANseER, Blumea 1 (1934) 56. — M. Philippinensis WANGERIN in Fedde, Rep. 10 (1912) 273; Merr. En. Philip. 3 (1923) 241; CHAo, Taiwania 5 (1954) 94, 99, f. 37; Lt & CHAO, Quart. J. Taiwan Mus. 7 (1954) 124, f. 19. — M. parvi- folia HALL. f. Beih. Bot. Centralbl. 34, 2 (1916) 41; DANSER, Blumea 1 (1934) 51. — M. subcaudata Me_rr. Philip. J. Sc. 13 (1918) 43; En. Philip. 3 (1923) 242. — M. megacarpa RIDL. FI. Mal. Pen. 1 (1922) 891. — M. chinensis MERR. Sunyatsenia 3 (1937) 256; Li, Taiwania 1 (1938) 94. — M. alterni- folia Merr. & CHUN, Sunyatsenia 5 (1940) 153. — M. poilanei TarpigEvu, Fl. Camb. Laos & Vietnam 8 (1968) 16. — Fig. 3c—e (galls). Tree up to 37 m; d.b.h. up to 75 cm; branchlets slender or stout, scattered, subglabrous to puberu- lous. Leaves scattered, obovate, elliptic to oblong- elliptic, 4-16 by 2-8cm, chartaceous to thick- coriaceous, subglabrous: base cuneate to attenuate; apex acute or acuminate to caudate; nerves 4~7(-9) pairs, distinct below; veins distinct below; petiole stout or slender, 1-4 cm. Inflorescence up to 8 cm, slender or stout, subglabrous to densely appressed- hairy, up to 3(-4) times branched; branches of the first order scattered; bracts either all triangular, under 3 mm, or lower ones lanceolate, up to 15 mm; basal inflorescence axes of the first order subtended by ordinary bracts or by leaves. Submature flower bud up to 3!/, mm @. Sepals 4 or 5, broader than long or as long as broad, thick, puberulous, to appressed-hairy. Petals 4 or 5, thick, glabrous to appressed-hairy. Stamens 4-5. Ovary puberulous to appressed-hairy. Fruit ovoid to oblong, 2!/,—31/, by 1-11/, cm; persistent disk con- spicuous or not; sepals inconspicuous. Distr. Continental SE. Asia (NE. India, Bhutan, Burma, Thailand, Tonkin, S. Yunnan) and throughout Malesia; not yet recorded from the Lesser Sunda Is. and New Guinea. KEY TO THE SUBSPECIES 1. Flowers basically 4-merous. Inflorescence stout, rusty-puberulous. Corolla glabrous outside. Leaves obovate, coriaceous; apex acute; base b. ssp. moluccana 1. Flowers basically 5-merous. pairs; veins distinct beneath. Fruit ovoid. 3. Basal bracts lanceolate, up to 15 mm. Fruit larger than 3 by 1'/,cm . a. ssp. pentandra 3. All bracts triangular, under 3 mm. Fruit up to Dae Oy ein os co -< CSSD. CHINeNSIS 2. Leaves up to 4-12 by 11/,-5 cm; nerves up to 6 pairs; veins obscure beneath. Fruit oblong. {[ser. I, vol. 8? 4. Length of sepals up to half as long as wide. Leaves chartaceous to subcoriaceous. Fruit 2'/,-3 cm long : . d. ssp. philippinensis 4. Length of sepals almost as long as wide. Leaves coriaceous to thick-coriaceous. Bracts uniformly triangular. Fruit 1°/,-2cm long e. ssp. scortechinii a. ssp. pentandra. Tree up to 34 m; branchlets stout. Leaves elliptic to oblong-elliptic, 8-16 by 4-8 cm, coriaceous; base cuneate; apex acuminate; nerves 6~-7(-9) pairs; veins distinct below; petiole stout, 2-4 cm. Inflorescence up to 8 cm, stout, densely appressed- hairy; basal bracts lanceolate, up to 15 mm. Sepals 5, broader than long. Petals 5, appressed-hairy outside. Stamens 5. Fruit ovoid, 3—-31/, by 11/,-11/, cm. Distr. Malesia: West & East Java. Ecol. In humid, mixed forest, 400-500 m. Fi. July—Dec. Vern. Java: Auru lilin, tenjau. b. ssp. moluccana MATTHEW, Blumea 23 (1976) 81. Tree up to 15 m; d.b.h. 20 cm; branchlets stout. Leaves obovate, 8-15 by 3-8 cm, coriaceous; base attenuate; apex acute; nerves 5-6 pairs; veins distinct below; petiole stout, 2!/,-3 cm. Inflores- cence up to 5cm, stout, rusty puberulous; basal bracts up to 4 mm. Sepals 4(—5), broader than long. Petals 4(—5), glabrous outside. Stamens 4(—5). Fruit (immature) ovoid, 11/, by 3/4 cm. Distr. Malesia: Moluccas (Morotai). Ecol. Mixed rain-forest, up to 1000 m. Fl. May. Once a leaf-gall was noted. Notes. The basal pair of lateral inflorescence branches often occur in the axils of normal leaves, a tendency noted in ssp. philippinensis. Flowers relatively large, yellowish to greenish; corolla dome-shaped (in bud). Calyx margin wavy; petals 4 by 3 mm; filaments 3 mm; anther 1 mm; style 11/, mm. The only fruit seen is detached and im- mature. The arrangement of the primary inflorescence branches is at times obscure. The soft, coriaceous texture of the leaves, dark above, and pale below, the stout inflorescence with rusty indumentum and the few, large, 4-merous flowers with glabrous dome-shaped corolla (in bud) distinguish this subspecies from the others. It is yet only known from Morotai I. c. ssp. chinensis (MERR.) MATTHEW, Blumea 23 (1976) 83. — M. chinensis MERR. — Fig. 3¢ (galls). Tree up to 20m; branchlets stout. Leaves elliptic to elliptic-oblanceolate, 8-20 by 4-8 cm, coriaceous; base attenuate; apex acute; nerves 6-8 pairs; veins distinct below; petiole stout, 1%/,- 2'/, cm. Inflorescence up to 8 cm, subglabrous to appressed-hairy; all bracts uniform, under 3 mm. Sepals 5, broader than long. Petals 5, appressed- hairy outside. Stamens 5. Fruit oblong, 2-2'/, by 1 cm. Distr. NE. India, Bhutan, N. Burma, Thailand, S. China (Yunnan), Tonkin; in Malesia: Malay Peninsula (Kedah, once). Ecol. Mixed forests up to 1900 m. Fl. May- June, fr. Aug.—May. In India galls and domatia occur. 1977] d. ssp. philippinensis (WANGERIN) MATTHEW, Blumea 23 (1976) 85.— M. philippinensis WANGERIN —WM. subcaudata MERR. — Fig. 3d-e (galls). Tree up to 22'/,m; brachlets slender. Leaves obovate to elliptic, 4-12 by 1'/,-5 cm, chartaceous to subcoriaceous; base attenuate; apex acuminate; nerves 4-6 pairs; veins obscure beneath; petiole slender, 1-2 cm. Inflorescence up to 7 cm, slender, subglabrous to puberulous; all bracts triangular, under 3 mm. Sepals (4-)5, broader than long. Petals (4-)5, glabrous to appressed-hairy outside. Stamens (4-)5. Fruit oblong, 2!/,-3 by 1*/,—1"/, cm. Distr. Malesia: throughout the Philippines. Ecol. In forests, from low altitude up to 1350 m. Fl. May-Sept., fr. Jan.Dec. Galls are rather frequent on stem, leaf and fruit. Note. Ssp. philippinensis is distinguished from ssp. scortechinii in the generally smaller height and smaller and thinner leaves, more slender inflores- cence axis, the lower 1 or 2 inflorescence axis (axes) a the axil(s) of normal leaves, and the oblong Tuits. e. ssp. scortechinii (KING) MATTHEW, Blumea 23 (1976) 86. — M. scortechinii KING — M. mega- carpa RipL. — M. parvifolia HALL. f. Tree up to 37m; branchlets stout. Leaves obovate to oblong, 4-12(-15) by 3—5(—6) cm, thick- coriaceous; base cuneate to attenuate; apex acute to acuminate; nerves 4-6 pairs; veins obscure beneath; petiole stout, 11/,-2'/, cm. Inflorescence up to 8 cm, stout, puberulous to villous; all bracts triangular, under 3 mm. Sepals (4-)5, as long as broad. Petals (4~-)5. Stamens (4-)5. Fruit oblong, 2!/,—-2!/, by 13/,-2 cm. Distr. Thailand (once); in Malesia: S. Sumatra, Banka, Malay Peninsula, Borneo, Celebes. Ecol. Primary forests, from low altitude up to CORNACEAE (Matthew) 97 2400 m. Fi. fr. Jan.—Dec. Inflorences and fruit galls occur. Vern. Malaya: médang pisang. Banka: mén- kapas. Borneo: kaju wulu, médang surugan. Note. Ssp. scortechinii is distinguished from ssp. pentandra by the generally obovate and smaller leaves, less stout inflorescence, uniformly short bracts and oblong fruits with thick pericarp. Excluded Mastixia cuneata BL. Mus. Bot. Lugd. Bat. 1 (1850) 257 = Notaphoebe umbelliflora (Bu.) BL. Cf. HALL. f. Beih. Bot. Centralbl. 34, 2 (1916) 42; DANSER, Blumea | (1934) 68. Mastixia gracilis KiNG, J. As. Soc. Beng. 74, ii (1902) 73; WANGERIN, Pfi. Reich Heft 41+ (1910) 28; DANsER, Blumea 1 (1934) 68; MATTHEW, Blumea 23 (1976) 90 = Vaccinium bancanum Mia. var. tenuinervium J. J. S. (Ericaceae), according to the type number Wray 1528 mentioned by SLEUMER, Blumea 11 (1961) 76. — Ed. Mastixia heterophylla BL. Mus. Bot. Lugd. Bat. 1 (1850) 257; WANGERIN, Pfl. Reich Heft 414 (1910) 28; DaANsER, Blumea 1 (1934) 69; MATTHEW, Blumea 23 (1976) 90. — HALLItER f. suggested this to be Gomphandra capitulata BEcc., but this was questioned by SLEUMER, Blumea 17 (1969) 193. According to us this sterile sheet (L 901, 169-350) collected by PRAETORIUS in Palembang, is not a Mastixia but we cannot give a proper identification. Mastixia tetrandra (THW.) CLARKE. — DANSER, Blumea 1 (1934) 56, referred two Sumatran speci- mens to this species, which is hitherto only found in Ceylon and the Andaman Is. One of these is sterile and the other is in bud; they can equally well be referred to M. rostrata ssp. rostrata, and their identification remains doubtful. Cf MATTHEW, Blumea 23 (1976) 77. Excluded Cornus caudata Hassk. & ZOLL., nom. illeg., C. ilicifolia (BL.) HAssK. & ZOLL., C. serrulata (BL.) Hassk. & ZOLv., and C. stricta ZoLL. & Mor. are all combinations or names made by HAssKARL and ZOLLINGER based on Polyosma BL., because of their opinion that this genus of the Saxifragaceae would be synonymous with Cornus L. Cf. HAssKARL, Cat. Hort. Bog. (1844) 168 and ZoLLINGER, Natuur- & Geneesk. Arch. Neérl. Ind. 2 (1845) 10. ONAGRACEAE (P. H. Raven, St. Louis)! Annual or perennial herbs (in Mal.), occasionally somewhat woody near the base, sometimes aquatic. Leaves spiral or opposite. Stipules absent or reduced, deltoid. Flowers mostly 4-merous, rarely 5-merous (in Mal.), solitary or arranged in a terminal racemose inflorescence, subtended by (often reduced) leaves or bracts. Bracteoles absent or 2 at the base of the ovary. Floral tube short or absent. Sepals erect, persistent. Petals caducous, contorted in aestivation, white, pink or yellow, sometimes emarginate. Stamens 4, 5, 8, or 10, in 2 whorls, rarely with an inter- mediate number, epipetalous ones sometimes shorter. Anthers usually versatile, sometimes seemingly basifixed by reduction: pollen single or in tetrads. Ovary inferior, (in Mal.) 4- or 5-celled and with c~ ovules; summit of the ovary (disk) flat to conical (in Mal.), sometimes with depressed nectaries surrounding the bases of the epipetalous stamens. Style simple; stigma capitate, clavate or globose, often 4-lobed. Ovules with axial placentation, |-pluriseriate. Fruit (in Mal.) a mostly long and slender loculicidal or irregularly rupturing capsule. Seeds rounded or elongate, in Ludwigia sometimes embedded in powdery or surrounded by cork-like endocarp tissue, in Epilobium with a chalazal plume of trichomes (coma); endosperm absent; embryo straight. Distribution. About 17 genera and more than 600 spp. in tropical and temperate regions, with a dis- tinct centre of diversity on the northern hemisphere in the New World, in Malesia two native genera which are both almost ubiquist. Ecology. Ludwigia is largely confined to the hot lowland and hills usually in wet or damp localities, Epilobium is confined to the higher mountain regions. Dispersal. Epilobium spp. are manifestly wind dispersed by virtue of their coma. Ludwigia spp. depend on dispersal by water and possibly incidental exozoic dispersal by water birds; in Ludwigia hyssopifolia there are two kinds of seed, one of which is enveloped by a corky tissue derived from the endocarp, enhancing their buoyancy. Pollination. Almost all of the Malesian species are self-pollinated, shedding pollen directly on the stigma at or before anthesis and rarely visited by insects. In Ludwigia peruviana, introduced in the Old World, the anthers are extrorse and shed pollen away from the stigma; thus outcrossing is predominant. Some outcrossing probably also occurs in the relatively large-flowered L. adscendens and L. octovalvis, which are known to be visited by insects, and in Epilobium detznerianum, in some populations of which the stigma is even held above the anthers. In our area, Here (Dansk Bot. Ark. 5, 1927, 18) reported Melipona sp., a bee, visiting the flowers of Ludwigia peruviana (as Jussieua peruviana), and Bombus rufipes at the flowers of the locally naturalized Oenothera stricta (as O. lamarckiana) and Fuchsia magellanica (as F. coccinea). The Melipona bees were not observed to contact the anthers or stigma of the large-flowered Ludwigia, but would certainly do so in visiting smaller-flowered species. All of the Malesian species are genetically self-compatible. Morphology & Anatomy. The Onagraceae are distinctive in their monosporic, 4-nucleate, ‘Oenothera type’ embryo sac development; in the nearly universal presence of viscin threads among the pollen; and in the loose construction of their pollen exine. Most species of Epilobieae and about half of Ludwigia shed their mature pollen in tetrads; these include all Malesian species except L. adscendens and L. hyssopifolia, in which the pollen is shed singly. Intraxylary phloem occurs throughout the family adjacent to primary xylem, and interxylary phloem (included phloem) is found in many genera but not in Ludwigia (CarLquist, Ann. Mo. Bot. Gard. 62, 1975, 386); in these features Onagraceae resemble many other Myrtales. The stomata are surrounded by three or more subsidiary cells, sometimes resembling those of Cruciferae. All Onagraceae have an inferior ovary and a floral tube, which is prolonged beyond the ovary in all except Ludwigia and Epilobium sect. Chamaenerion. In several species of Ludwigia half-submerged parts of the stem are covered by a whitish aerenchyma; in L. adscendens short roots at the nodes are transformed into inflated, elongate aerophores enhancing floating on water. Chromosomes. All species of Epilobium sect. Epilobium, a taxon that includes all Malesian species, which have been examined have had a gametic chromosome number of n = 18. Species of Ludwigia have a gametic chromosome number of m = 8 and multiples. These genera differ from most others in Onagra- (1) With co-operation of the General Editor in framing the manuscript. ’ The author gratefully acknowledges the support of the U.S. National Science Foundation to the studies of Onagraceae. (98) 1977] ONAGRACEAE (Raven) 99 ceae in having small chromosomes that are heteropycnotic and dark-staining throughout the mitotic cycle. Naturally occurring interchange heterozygotes, abundant in the tribe Onagreae, are not known to occur in either group. The original basic chromosome number of the family is x = 11, as in Fuchsia, Circaea, and others. Hybridization. Hybrids are rare between the recognized species of Ludwigia. In Epilobium sect. Epilobium they are occasional where two or more entities come together, but their occurrence is limited by the predominant autogamy or cleistogamy of most species and to some extent by ecological differentiation also. A wide range of fertilities is characteristic of these hybrids, as explained in detail in our monograph of the Australasian species (RAVEN, D.S.I.R. New Zeal. Bull. 216, 1976), and cytoplasmic differences sometimes occur. Two of the Malesian species occur together in N. Luzon, and a few individuals suggest hybridization; four occur together in the mountains of New Guinea, with hybrids probably occasional but poorly studied so far. All species of Epilobium found in Malesia have the same chromosome arrange- ment that is predominant in Eurasia, from where they doubtless came. Chemotaxonomy. Raphides, needle-like crystals of calcium oxalate, are ubiquitous in the vegetative parts of Onagraceae. The few reports of alkaloids are doubtful and seem to indicate rather the presence of secondary amines. Ellagic acid occurs. Among the flavonoids reported from the family, flavonols based quercetin are ubiquitous, whereas kaempferol and more highly oxygenated types based on myricetin are frequent. The anthocyanins include predominantly malvidin and cyanidin derivatives, with the latter predominant in the rose-purple petals of Epilobium. The yellow petals of most species of Ludwigia are colored by carotenoids, with the chalcone isosalipurposide forming a non-ultraviolet-reflective centre in many species, including L. peruviana. Uses. Only some species of Ludwigia are mentioned to be in use for minor medicinal purposes; see under Ludwigia spp. KEY TO THE GENERA 1. Capsule loculicidally dehiscent with 4 valves; axis persistent. Seeds comose. Floral tube present. Petals 4, white, pinkish or red. Stamens 8. Stem-base without aerenchyma. . . . . 2. Epilobium 1. Capsule irregularly dehiscent; axis not persistent. Seed not comose. Floral tube absent. Petals yellow, if white or creamy then flowers 5-merous and floating aquatic plant adorned with short spongy aeren- chyma-roots at the nodes. Stamens 4-10. Stem-base not rarely covered by aerenchyma. 1. Ludwigia 1. LUDWIGIA Linn£, Gen. Pl. (1754) 55; Sp. Pl. 1 (1753) 118; Munz, Bull. Torr. Bot. Cl. 71 (1944) 152; HaRA, J. Jap. Bot. 28 (1953) 289; A. & R. FERNANDES, Garcia de Orta 5 (1957) 109; RAVEN, Reinwardtia 6 (1963) 327. — Jussiaea LINNE, Gen. PI. (1754) 183; Sp. Pl. 1 (1753) 388; BAcK. Trop. Natuur 3 (1914) 59; Fawcett, J. Bot. 64 (1926) 10; MuNz, Darwiniana 4 (1952) 179. — Nematopyxis Mia. FI. Ind. Bat. 1, 1 (1855) 600. — Fig. 1, 4, 5. Slender herbs, erect or creeping and rooting at the nodes, to large shrubs. Underwater parts often swollen and spongy or bearing inflated white spongy aero- phores. Leaves alternate or opposite, mostly entire. Stipules absent or reduced, deltoid. Flowers borne singly, clustered, or arranged in an inflorescence. Brac- teoles lacking or conspicuous, usually two, at or near the base of the ovary. Floral tube absent. Sepals 3-7, persistent after anthesis. Petals as many as the sepals or absent, caducous, yellow or white, with contorted aestivation. Stamens as many as or twice as many as the sepals, or flowers very rarely with an intermediate number of stamens; anthers usually versatile but sometimes apparently basifixed by reduc- tion. Pollen shed in tetrads or singly. Disk (summit of the ovary) flat to conical, often with depressed nectaries surrounding the bases of the epipetalous stamens. Stigma hemispherical or capitate, the upper 1'/,-*/, receptive, often lobed, the number of lobes corresponding to the number of locules. Ovary with a number of cells equal to the number of sepals, very rarely more; placentation axial; ovules pluriseriate or uniseriate in each cell, in one species uniseriate below, pluriseriate above; if uniseriate, the seeds sometimes embedded in powdery or woody endocarp 100 FLORA MALESIANA [ser. I, yokes from which they detach easily or with difficulty. Capsule irregularly dehiscent, or by a terminal pore, or by flaps separating from the valve-like top. Seeds rounded or elongate, the raphe usually easily visible and in some sections equal or nearly equal in size to the body of the seed. Distr. According to my synopsis (Reinwardtia 6, 1963, 329) 75 spp., all over the world; in Malesia 6 spp., one of which is certainly introduced. Ecol. One aquatic and the other species mostly in swampy or damp places, often in rice-fields, from the lowland up to c. 2100 m, mostly below 1000 m. Flowers last only one day. Taxon. I have divided the genus into 17 sections, the largest of which (sect. Myrtocarpus) is neotropical. They are often shrubby with large, 4- or 5-merous flowers, dimerous stamens, prominently 4- or 5-ribbed capsules, free seeds and pollen grains shed in tetrads. They appear phylogenetically central in the genus. In Malesia this section is represented by an introduced weed, L. peruviana. Close to this section are one African (sect. Africana) and one American section (sect. Macrocarpon) with terete capsules. Following these is a series of small Old World sections which have the stamens reduced to one whorl; in one African section flowers are 3-merous. L. hyssopifolia forms a monotypic section unique in having two kinds of seeds, those in the lower part of the capsule uniseriate and embedded in the endocarp, those in the upper part pluriseriate and free, while pollen grains are single, Other sections, not represented in Malesia, have all of the seeds loosely embedded in powdery endocarp. The structure of the seed is important in the dis- crimination of sections. The second major line of the genus consists of species in which the seeds are embedded in coherent chucks of woody endocarp which render the capsule a tough unit from which it is difficult to separate the seeds. The two sections belonging to this line have basically 5-merous flowers and pollen shed singly. Through the disentangling of these relationships it appears that the number of stamens is not decisive for dividing the genus into two genera as this would go across relationships and lead to heterogeneous assem- blages of species. Each Malesian species belongs to a different section and being so small in number it seems not useful to give descriptions of these sections; I refer to my revision (1963). The cradle of the genus is probably South America with an important secondary centre of evolution in Africa. It is one of the most primitive genera in the family. Note. It has appeared that seeds retain viability in the herbarium in unpoisoned, not too old speci- mens; flowering plants can thus be raised from fruiting herbarium specimens. KEY TO THE SPECIES 1. Stamens twice as many as sepals. 2. Seeds pluriseriate, free (not embedded in endocarp). 3. Plant subglabrous to appressed-pubescent. Capsule terete. Petals 5-17 by 4-17 mm. Style 1.5- 3.5 mm. Raphe equal in diameter to the body of theseed. . ......... 2. L. octovalvis 3. Coarse, strongly pubescent or villous plant. Capsule strongly 4-angled, villous, with flat sides. Petals 15-24 by 16-26 mm. Style 1 mm long. Raphe not more than '/, the diameter of the body of the Sr MERE eee | Oc eee. dea ame soy = whe: « SIS yay! acngssl zyciai uSGuis: 41 Me, ie Ase Ae 1. L. peruviana 2. Seeds at least below uniseriate and embedded in endocarp. 4. Seeds in the c. 1/, upper part of the capsule pluriseriate and free. Sepals 4. Petals 2-3 mm long _ 5. L. hyssopifolia 4. Seeds all uniseriate in each cell of the capsule and embedded in endocarp. Sepals 5—7 (rarely 4). Petals 4.5-23 mm long. 5. Aquatic, with floating branches forming erect clusters of spongy, spindle-shaped aerophores (aerenchyma). Petals white or creamy, with yellow atthe base. . ...... 6. L. adscendens 5. Plant not forming such aerophores on the decumbent branches. Petals bright golden-yellow with a darker spot at the base. New Zealand, Australia, Pacific Is., Formosa, China, and Japan, also in the New World, might possibly be found in East Malesia. Cf. Aston, Aquat. Pl. Austr. (1973) 144, f. 55 L. peploides (H.B.K.) RAVEN 1. Stamens as many as sepals, very rarely more in some flowers. 6. Seeds pluriseriate in each cell of the capsule. Petals elliptical, 1-3 by 0.7-2 mm. Capsule terete. Seeds ira egy Oa NI) oe 8. | hab 5a so beet te, Patera late tetas dace 3. L. perennis 6. Seeds uniseriate in each cell of the capsule. Petals narrow spathulate, 1.3-2.2 by 0.4-0.9 mm. Capsule Slightly 4-anpied. Seeds'0:5—0)6 by: O:3Simm- 20) ee ee eo BO 4. L. prostrata 1. Ludwigia peruviana (L.) Hara, J. Jap. Bot. 28 Steen. Fl. Sch. Indon. (1949) 305. — Oenothera (1953) 293; RAvEN, Reinwardtia 6 (1963) 345, hirta LInNé, Sp. Pl. ed. 2, 1 (1762) 491. — Jussieua map 14. — Jussiaea peruviana LINNE, Sp. Pl. 1 hirta (L.) Sw. Obs. Bot. (1791) 142, non LAmMK, (1753) 388 ; Back. Trop. Natuur 3 (1914) 61; Onkr. 1789; Back. Ann. Jard. Bot. Btzg Suppl. 3 (1909) Suiker. (1930) 470, Atlas t.445; ALstoNin Trimen, 406. — Jussiaea speciosa RiDL. J. Bot. 59 (1921) FI. Ceyl. 6 (1931); Munz, Darwiniana 4 (1942) 131; 259; Fl. Mal. Pen. 1 (1922) 828. 1977] ONAGRACEAE (Raven) 101 Shrub 0.5-3 m, entirely villous, the hairs often multicellular, especially in the inflorescence; long inflated aerophores arising from submerged, buried roots. Leaves lanceolate to broadly lanceo- late, 4-12 by 0.3—-1.5 cm, narrowly cuneate at base, apex acute to acuminate; nerves 12-22 pairs; sub- marginal vein not prominent; petiole 3-12 mm. Flowers in upper leaf axils. Bracteoles lacking or up to 7 mm long, subulate. Sepals 4 or 5, lanceolate, irregularly serrulate, 10-18 by 4-8 mn, villous. Petals bright yellow, veiny, suborbicular, 15-24 by 16-26 mm, shallowly emarginate, with a claw 1-3 mm. Stamens 8 or 10, subequal; filaments 2-3.5 mm; anthers 3-4.5 mm long, extrorse and not shedding pollen directly on the stigma at anthesis. Pollen shed in tetrads. Disk elevated 1-2 mm, with a depressed densely white-hairy nectary around the base of each epipetalous stamen. Style c. 1 mm; stigma broadly elongate-hemispherical, 2-3 mm high. Capsule villous, 1.2-3 by 0.6—-1 cm, light yellowish brown with 4 prominent dark brown ribs, 4-angled, thin-walled, readily and irregularly loculicidal; pedicel 2-4.5 cm. Seeds pluriseriate in each cell, free, light brown, finely striate and cellu- lar pitted, obovoid, 0.6—-0.8 mm long; raphe */, to 1/, the width of the body. Gametic chromosome number (Old World populations), n = 40. Distr. Native of the New World, from the SE. United States throughout South America, intro- duced and naturalized in Malesia since the 2nd half of the last century, collected in Malaya, Sumatra, Java (common in West), but obviously still absent from many areas. Ecol. Along ditches and in moist places, mostly in the lowland but ascending to c. 1400m. Fi. Jan.—Dec. Vern. Banka: pitjanket, M; Java: tjatjabéan, S, lombokan, J. Note. BAckKer (Onkr. Suiker. 1930, 470) observed that in inundated situation the plant produces aerophores which are emitted by shallow, horizontal roots; they are erect but their tips usually float on the water. 2. Ludwigia octovalvis (JAcQ.) RAVEN, Kew Bull. 15 (1962) 476; Reinwardtia 6 (1963) 356, maps 19-20, incl. ssp. brevisepala (BRENAN) RAVEN ef ssp. sessiliflora (MicH.) RAVEN; HENRY & PRIT- CHARD, Bot. Div. Lae, Bot. Bull. 7 (1973) 132, fig.; Everist, Pois. Pl. Austr. (1974) 393. — Jussiaea suffruticosa LINNE, Sp. Pl. 1 (1753) 388; Bru. Fl. Austr. 3 (1867) 307; F.v.M. Descr. Not. Pap. Pl. 4 (1876) 60; CLARKE, FI. Br. Ind. 2 (1879) 587; Koorp. Exk. Fl. Java 2 (1912) 703; Rip. Trans. Linn. Soc. Bot. II, 9 (1916) 57, incl. var. hirta Ripv.; GaGn. Fl. Gén. L.-C. 2 (1921) 986; C. T. Wuite, Proc. R. Soc. Queensl. 34 (1922) 48; Back. & BAKH. f. Fl. Java 1 (1963) 261. — Oeno- thera octovalvis JACQ. En. Syst. Pl. (1760) 19. — Jussiaea pubescens LINNE, Sp. Pl. ed. 2, 1 (1762) 555. — L. perennis (non L.) Burm. /. Fl. Ind. (1768) 37. — Jussiaea angustifolia LAMK, Encycl. 3 (1789) 331. — Jussiaea villosa LAMK, I.c.; RipL. Fl. Mal. Pen. 1 (1922) 828. — Jussieua octovalvis (JACQ.) Sw. Obs. Bot. (1791) 142. — Jussiaea angustifolia Bi. Bijdr. (1826) 1132, non Lamxk, 1789; BAcK. Trop. Natuur 3 (1914) 62. — Jussiaea blumeana DC. Prod. 3 (1828) 55. — Jussiaea burmannii DC. l.c. 57. — Jussiaea exaltata Roxs. (Hort. Beng. 1814, 33, nomen) Fl. Ind. ed. Carey 2 (1832) 401. — Jussiaea costata Pr. Epim. Bot. (1849) 217. — Jussiaea junghuhniana Mia. FI. Ind. Bat. 1, 1 (1855) 627; VAL. Bull. Dép. Agric. Ind. Néerl. 10 (1907) 41. — Jussiaea erecta (non L.) RIDL. J. Bot. 59 (1921) 258; Fl. Mal. Pen. 1 (1922) 827, incl. var. exaltata (Roxs.) Ripi.; Back. Onkr. Suiker. (1930) 470, Atlas t. 446; STEEN. Arch. Hydrobiol. Suppl. 10 (1932) 314. — L. pubescens (L.) HARA, J. Jap. Bot. 28 (1953) 293. — Fig. 1. Usually robust, well-branched herb, sometimes woody at the base, up to 4 m, subglabrous or with sparse or dense appressed or spreading pubescence. Leaves lanceolate or narrowly lanceolate, to narrowly ovate, or subovate, 2-14 by 0.5-4 cm, narrowly to broadly cuneate at the base and attenuate at apex; nerves 11-20 pairs, sub- marginal vein well developed; petiole up to 1 cm. Bracteoles reduced or to 1 mm long. Sepals 4, ovate or lanceolate, 6-15 by 1-7.5 mm. Petals yellow, broadly obovate or cuneate, emarginate, 17 by 2-17 mm. Stamens 8, epipetalous ones shortest; filaments 1-4 mm long; anthers 0.5-4 mm long, extrorse but soon crumbling and shedding pollen directly on the stigma. Pollen shed in tetrads. Disk slightly raised, with a white-hairy sunken nectary surrounding the base of each epipetalous stamen. Style 1.5-3.5 mm; stigma subglobose, shallowly 4-lobed, 1.2-3 mm @. Capsule thin-walled, 1.7- 4.5cm by 2-8mm, terete, pale brown with 8 darker ribs, readily and irregularly loculicidal; pedicel up to 10mm. Seeds pluriseriate in each cell, free, brown, rounded, 0.6-0.75 mm long, 0.5-0.7 mm wide including the inflated raphe which is equal in size to the body of the seed and evenly transversely ridged. Gametic chromosome numbers, n = 16, 24. Distr. Throughout the tropics of the world, between c. 32° N and 30° S. Ecol. Mostly in humid places, damp grassland, rice-fields, along ditches and water-courses, in swamps, lakes and pools, drains, sandy or silty floodbanks, gravelly riverbeds, on floating islands in lakes, on floating logs in lagoons, sago swamps, mountain peat swamps with sedges, also in old native gardens and coconut plantings, from the lowland up to c. 1000 m, in Java and Celebes up to 1400 m, in New Guinea up to 2100 m. Fi. Jan.— Dec. Several collectors mention that it propagates by runners and that old leaves turn reddish. The lower part of the stem is at times coated by aerenchyma. In inundated condition aerophores are produced; see under |. L. peruviana. DOocTERS VAN LEEUWEN (Zoocecidia, 1926, 427 f. 808; Ned. Kruidk. Arch. 51, 1941, 204) recorded fruit galled by beetles and aphid galls on the ter- minal leaves of the branches. Uses. In Java minor medicinal qualities are ascribed to this species, amongst others against sprew. Rumpuius, who described it under the name herba vitiiginum (Herb. Amb. 6, p. 49) did not mention uses (Heyne, Nutt. Pl. 1927, 1206). BurkiLL (Dict. 1935, 1274) reported that the mucilaginous leaves, after which the plant is called ‘lakomayer’ = ‘water Vitis’, are used for poulticing in a variety of complaints; it has also been recorded as used for headaches, orchitis, glands in the neck, 102 FLORA MALESIANA [ser. I, vol. 8? i Me: oom ay \} Fig. 1. Ludwigia octovalvis (JACQ.) RAVEN. a. Habit in flower, b. withered fruits, both x #/, (a after BACKER, 1940, b JUNGHUHN CCXxII). 1977] diarrhoea, dysentery, nervous diseases, and as a vermifuge. In WILKINSON’s Dictionary it is said that a kind of tea is made from the leaves. Also in India medicinal properties are ascribed to it. Vern. Malaya: buyang samalam, lakom ayér, pujang malam, yénlidah, M; Sumatra: urang aring itém, Simalur I.; Java: gagabusan, tjatjabéan, S, kalaménja, Md, salah njowo, J; Lesser Sunda Is.: pangambo, E. Sumbawa; Philippines: tayilakton, Tag., balansuit, Mag., malapako, tubong-talapan, Bik., pachar-pachar, Sul., palangdisin, Ig., talang- kau, Ilk., halangot, naudyawa tubig, If.; Celebes: kélétélé téngén, Tonsaw. dial., Minahassa; Moluccas: daun panu, Ambon; New Guinea: ewo, onarenare, Kapauko lang., kampur, Sakaj bivouac, Merauke, pfauhanu, Kutubu, mayenke, Orne lang., Kaiye, rowijetwi, Enga lang., Yogos, togorarah, Wapi lang., Marok, rama-rama, Matapaili lang. Note. In my revision (1963) I distinguished three subspecies, more or less geographically defined, which I wish to withdraw here. 3. Ludwigia perennis LINNg, Sp. Pl. 1 (1753) 119; A. & R. FERNANDES, Garcia de Orta 5 (1957) 114, 475; RAVEN, Reinwardtia 6 (1963) 367, map 21. — L. parviflora Roxs. (Hort. Beng. 1814, 11, nomen) Fl. Ind. 1 (1820) 440; Brn. Fl. Austr. 3 (1867) 307; Kurz, J. As. Soc. Beng, 46, ii (1877) 91; CLARKE, Fl. Br. Ind. 2 (1879) 588; TRIMEN, FI. Ceyl. 2 (1894) 234; Rip-. Fl. Mal. Pen. 1 (1922) 828; BAck. Onkr. Suiker. (1930) 471, Atlas t. 447; BAck. & Baku. f. Fl. Java 1 (1963) 261. — L. lythroides BL. Bijdr. (1826) 1134. — L. gracilis Mig. Fl. Ind. Bat. 1, 1 (1855) 629. — Jussiaea perennis BRENAN, Kew Bull. 8 (1953) 163. Annual herb up to | m, subglabrous or minutely puberulent on younger parts. Leaves narrowly elliptic to lanceolate, 1-11 by 0.3—-2.7 cm, narrowly cuneate at base, apex subacute; nerves 6-12 pairs; submarginal vein weakly developed; petiole 2-15mm, winged. Sepals 4, rarely 5, deltoid, (1.3-)2-3.5 by (0.5-)0.7-1.8 mm, glabrous or minutely puberulent. Petals yellow, elliptic, 1-3 by 0.7-2 mm. Stamens usually 4 or 5, rarely more; filaments 0.3-0.7 mm; anthers 0.5-0.7 by 0.5- 0.7 mm, shedding pollen directly on the stigma at anthesis. Pollen shed in tetrads. Disk slightly elevated, glabrous. Style 0.7—1.5 mm; stigma glo- bose, 0.4-0.5 mm @. Capsule thin-walled, glabrous or puberulent, 3-16 (-19) mm long, terete, pale brown, readily and irregularly loculicidal; capsule sessile or on a pedicel up to 6 mm, often + nodding. Seeds pluriseriate in each cell, free, brown with fine brown lines, ellipsoid-rounded, 0.3-0.5 by 0.2-0.25 mm; raphe very narrow and inconspi- cuous. Gametic chromosome number, 1 = 8. Distr. Tropical Africa (from Senegal, Lake Chad, and the Sudan south to E. Congo and N. Natal), Madagascar, through continental SE. Asia (Ceylon to S. China, Hongkong, and Hainan), and throughout Malesia (except Borneoand Moluccas) to tropical Australia and New Caledonia. Fig. 2. Ecol. Sunny, humid or marshy situations fallow rice-fields, sawah dikes, along ditches, rivers and water-courses, dry riverbeds, sugarcane fields on heavy clay, damp places in Eucalypt savannah —" aero from the lowland to c. 500 m. Fi. an.-—Dec. ONAGRACEAE (Raven) 103 In Java and Madura I. decidedly preferring regions subject to a dry monsoon. Fig. 2. Range of Ludwigia perennis L. (after RAVEN, 196 4. Ludwigia prostrata Roxs. (Hort. Beng. 1814, 11, nomen) Fl. Ind. 1 (1820) 441; Bru. FI. Austr. 3 (1866) 308; CLARKE, Fl. Br. Ind. 2 (1879) 588; TRIMEN, FI. Ceyl. 2 (1894) 234; Merr. FI. Manila (1912) 355; Rpt. Fl. Mal. Pen. 1 (1922) 829; Me_rr. En. Philip. 3 (1923) 220; Back. Onkr. Suiker. (1930) 472, Atlas t. 448; Back. & BAKH. f. Fl. Java 1 (1963) 261; RAvEN, Reinwardtia 6 (1963) 374, map 23. — L. fruticulosa BL. Bijdr. (1826) 1133. — L. leucorhiza BL. I.c. — Nema- topyxis pusilla Mia. FI. Ind. Bat. 1, 1 (1855) 630. — Nematopyxis prostrata Mig. l.c. — Nematopyxis fruticulosa Mia. l.c. Annual herb 0.1-0.6m, subglabrous, often reddish-tinged. Leaves elliptic or narrowly elliptic, 1-13 by 0.3-2.7 cm, glabrous or with a few minute hairs along the veins, narrowly cuneate at the base, apex acute; submarginal vein weakly developed; petioles 4-25 mm, distinct. Sepals 4, deltoid, 1.3- 2.5 by 0.7-1.1 mm, glabrous. Petals yellow, narrow- ly spatulate, 1.3-2.2 by 0.4-0.9mm. Stamens 4; filaments 0.8-1.2 mm; anthers 0.4-0.5 mm wide, broader than long, closely appressed to the stigma and shedding pollen directly on it at anthesis. Pollen shed in tetrads. Disk slightly elevated, glabrous. Style c. 1 mm; stigma globose, c. 0.5 mm, the upper half receptive. Capsule thin-walled, glabrous, 12-22 by 0.8-I1 mm, +4-angled, pale brown, readily and irregularly loculicidal, the seeds showing plainly as indentations in the walls at maturity. Seeds uniseriate in each cell, free, pale brown, speckled or striped transversely with narrow darker brown stripes, plump, ovoid, apiculate at one end, 0.5-0.6 by 0.3 mm; raphe narrow, linear. Gametic chromosome number, 7 = 8. Distr. Tropical SE. Asia (Ceylon and S. Deccan to N. India, Assam, S. China and Andamans); in Malesia: Malay Peninsula, Java, Lesser Sunda Is. (Timor), Borneo, and Philippines (Palawan, Luzon, Negros, Mindanao). The single record from Australia (N. Queensland: Mossman R.) recorded by C. T. WuiTe (Proc. R. Soc. Queensl. 50, 1939, 78) is a misidentification of L. hyssopifolia. On the anon L. prostrata is in Malesia a rare species. ig. 3. Ecol. Paddies, fallow and planted, by ditches, along riverbanks, in swampy places, rather rare, from lowland up to c. 800 m. Fl. Jan.-Oct. VERN. Philippines: alubihud, P.Bis. Fig. 3. Range of Ludwigia prostrata Roxs. (after RAVEN, 1963). 5. Ludwigia hyssopifolia (G.DON) ExELL, Garcia de Orta 5 (1957) 471; A. & R. FERNANDES, /.c. 471, 474; RAVEN, Reinwardtia 6 (1963) 385, map 30; HENTY & PRITCHARD, Bot. Div. Lae, Bot. Bull. 7 (1973) 131, fig. — Jussiaea linifolia VAHL, Eclog. Am. 2 (1798) 32, non L. linifolia Por. 1813; BACK. Trop. Natuur 2 (1913) 20, fig.; ibid. 3 (1914) 61; Onkr. Suiker. (1930) 469, Atlas t. 444; STEEN. Arch. Hydrobiol. Suppl. 10 (1932) 314; Back. & BAKH. f. Fl. Java 1 (1963) 260. — Jussiaea hyssopi- folia G.Don, Gen. Syst. 2 (1832) 693. — Jussiaea suffruticosa (non L.) RIDL. J. Bot. 59 (1921) 257; Fl. Mal. Pen. 1 (1922) 827. — L. prostrata (non Roxs.) C. T. Wuite, Proc. R. Soc. Queensl. 50 (1939) 78. — Fig. 4. Annual herb 5 cm to 3 m, often becoming woody at the base; young growth and inflorescence minutely puberulent; elongate aerophores arising from buried submerged roots. Leaves lanceolate, 1-9 by 0.2-3 cm, narrowly cuneate at the base, apex acuminate; nerves 11-17 pairs; submarginal vein not prominent; petiole 2.5-18 mm. Sepals 4, lanceolate, 2-4 by 0.7-1.2 mm, finely puberulent, 3-nerved. Petals yellow, fading orange-yellow, elliptic, 2-3 by 1-2 mm. Stamens 8, pale greenish yellow, epipetalous ones shorter; filaments of episepalous stamens 1-2 mm, those of epipetalous ones 0.5-1 mm; anthers 0.4-0.6 mm wide, 0.2- 0.3mm high, shedding pollen directly onto the stigma at anthesis. Pollen grains shed singly. Disk + elevated, with a depressed ciliate nectary sur- rounding the base of each epipetalous stamen. Style pale greenish yellow, 1-1.5 mm; stigma de- pressed-globose, c. 0.6-1.2mm @, 0.5-0.8 mm high, shallowly 4-lobed, the upper portion recep- tive. Capsule relatively thin-walled, finely puberu- lent, 1.5-3cm by 1-1.2 mm, subterete, enlarged in the upper ?/, to 4/3, subsessile. Lower seeds uni- seriate in each cell of the capsule, nearly vertical, brown, oblong, 0.7-0.85mm long, each firmly embedded in a cube of relatively hard endocarp; raphe '/, the diameter of the body. Seeds in upper inflated portion of the capsule multiseriate, free, ovoid, 0.35-0.5 mm long, paler brown than the lower seeds and with a narrower raphe. Lower part of capsule at first marked by distinct bumps corresponding to the position of the uniseriate seeds, but as the endocarp hardens and swells, the capsule becomes smooth. FLORA MALESIANA [ser. I, vol. 82 Gametic chromosome number, n = 8. Distr. Tropical Africa (Dakar to Lake Chad, S. Sudan and S. Congo), continental SE. Asia (Ceylon to Hainan), throughout Malesia to Micronesia and N. Australia (Cape York Penin- sula and Arnhem Land). It is difficult to fix the native country of this now widely spread palaeotropical weed which has no close relatives. It might have been introduced in tropical Africa where it is relatively local and con- fined to the west, but it was collected in Sao Tomé as early as 1822. Ecol. A very common weed of pools, along drains, shallow ditches, water and river edges, in paddies and humid, fallow garden land, on waste land, fallow sawahs, and in clearings, both on clay and humid white sand (Borneo), in pools in Euca- lypt savannah (Wetar I.), on Mt Kelud (Central Java) as a pioneer on old volcanic mudstreams (lahars), from the plains up to c. 1000 m. Fi. Jan.- Dec; BACKER (Trop. Natuur 2, 1913, 133) described in detail the biology of this species, of which the flowers open at 7 a.m. Seeds are gradually released by decay of the pericarp of which the vessels remain persistent. To his surprise both kinds of seed were buoyant for 16 days, after which they sank. On the 17th day he observed one buoyant seedling, but unfortunately no mention was made from which kind of seed; his experiment was through inter- ference discontinued. He made also notes on the aerophores emitted by the shallow roots as occur also in L. peruviana and other species. DOCTERS VAN LEEUWEN (Zoocecidia, 1926, 428; Ned. Kruidk. Arch. 51, 1941, 204) reported galled fruits caused by beetles. Leaves are often attacked by a blue beetle: Graptodera cyanea (BACKER, Trop. Natuur 2, 1913, 132). Uses. HEYNE (Nutt. Pl. 1927, 1206) noted that in N. Celebes it is used for poulticing pimples. According to BuRKILL (Dict. 1935, 1273) it is in Malaya generally stocked by Chinese herbalists, but its use is not clear; it was once recorded that an infusion of the root is swallowed by Malays for syphilis. In the Philippines the whole plant is used for black dye (QUISUMBING, Medic. Pl. Philip. 1951, 676). Vern. Sumatra: méligai, M, Banka; Java: (djukut) anggéréman, mainang, tjatjabéan, S; Philippines: pasau-na-hapai, sila sila, Tag., ilam- num-wiliyan, Mag., kakaggin diloba, If., barigaud, Bik., manakatud, Ik., talang-duron, Pamp., tohod-téhod, Bik.; Celebes: kayu ragi, Manado. Note. The dimorphous seeds are very unusual and it would be most interesting to have informa- tion on the properties of the two seed types with respect to germination. 6. Ludwigia adscendens (L.) Hara, J. Jap. Bot. 28 (1953) 290; A. & R. FERNANDES, Garcia de Orta 5 (1957) 475; RAVEN, Reinwardtia 6 (1963) 387, maps 31, 33; Aston, Aquat. Pl. Austr. (1973) 142; HENTy & PRITCHARD, Bot. Div. Lae, Bot. Bull. 7 (1973) 130, fig. — Jussiaea repens LINNE, Sp. Pl. 1 (1753) 388, non L. repens Forst. 1771; F.v.M. Descr. Not. Pap. Pl. 4 (1876) 60; CLARKE, FI. Br. Ind. 2 (1879) 587; O. K. Rev. Gen. Pl. 1 (1891) 251, incl. var. pilosa O. K. et var. glaberrima O. K.; TRIMEN, FI. ONAGRACEAE (Raven) 105 1977] ( u N a i TN SV LZ a Y] AS 7 f any "TS J re iN x "/, (a after Fig. 4. Ludwigia hyssopifolia (G. Don) Exe . a. Habit, in flower, b. withered fruits, both Backer, 1940, b VerBoom 9). 106 Ceyl. 2 (1894) 233; Koorp. Exk. Fl. Java 2 (1912) 703; BACK. Trop. Natuur 3 (1914) 56, 60, f. 1-5; RIDL. Fl. Mal. Pen. 1 (1922) 827; GAGNn. Fl. Gén. I.-C. 2 (1925) 987; Hocur. Candollea 3 (1925) 479, incl. ssp. glabrata Hassv. f. albiflora Hocnr.; HARTSEMA, Flora (Allg. Bot. Z). n.s. 22 (1927) 242, t. 3; Back. Onkr. Suiker. (1930) 469, Atlas, t. 443; STEEN. Arch. Hydrobiol. Suppl. 10 (1932) 314, f. 62; Fl. Sch. Indon. (1949) 305; Back. & BAKH. f. Fl. Java 1 (1963) 260. — Jussiaea adscendens LinnE, Mantissa 1 (1767) 69. — Jussiaea fluviatilis BL. Bijdr. (1826) 1132. — Fig. 5. Herb with prostrate or ascending stems, rooting at the nodes, with conspicuous white, erect, spindle-shaped, mucronate aerophores arising in clusters at the nodes of the floating stems and from the roots, the more or less erect stems to 60 cm; floating stems to 4m; plants normally glabrous, but the branches growing on dry ground densely FLORA MALESIANA [ser. I, vol. 8? villous and rarely flowering. Leaves broadly oblong-elliptical, 0.4-7 by 0.7-4cm, narrowly cuneate at base, apex acute or obtuse; nerves 6-13 pairs; submarginal vein not prominent; petioles long. Flowers in upper leaf axils. Bracteoles present near base of capsule, deltoid, c. 1.2 by 1.3- 1.5mm. Sepals 5, deltoid-acuminate, 5-11 by 2-3.2 mm, glabrous or villous. Petals creamy white, yellow at base, obovate, rounded at apex, 9-18 by 6-10 mm. Stamens 10, epipetalous ones slightly shorter; filaments white, 2.5-4mm; anthers 1.2-1.8mm long, apparently shedding pollen directly on the stigma at anthesis. Pollen grains shed singly. Disk + elevated, with a depressed white-hairy nectary surrounding the base of each epipetalous stamen. Style white, 4-8 mm, densely long-hairy in lower half; stigma globose, green, 1.5-2 mm @, 1-1.2 mm high, the upper 2/; recep- tive. Capsule glabrous or villous, 1.2-2.7 cm by Che: “a y i SANY FD > Zan Fig. 5. Ludwigia adscendens (L.) HARA, X 2/; (after BACKER, 1940). 1977] ONAGRACEAE (Raven) 107 344 mn, light brown, with 10 conspicuous darker brown ribs, terete, the seeds evident between the ribs as bumps c. 1.5mm apart; capsule thick- walled, very tardily and irregularly dehiscent; pedicel 2.5—5.5 cm. Seeds uniseriate in each cell of the capsule, pale brown, 1.1-1.3mm long, + vertical, firmly embedded in coherent cubes of woody endocarp 1.2—1.5 mm high, 1—-1.2 mm thick, the endocarp firmly fused to the capsule wall. Gametic chromosome numbers, 7 = 8, 16. Distr. Continental Asia (from Ceylon to S. China), throughout Malesia, in Australia one locality in W. Arnhem Land. Fig. 6. Ecol. Freshwater pools, ditches, swamps, fallow and wet paddies, very common, from the lowland up to c. 1600 m. Fi. Jan.—Dec. Fig. 6. Range of Ludwigia adscendens (L.) HARA (after RAVEN, 1963). BACKER (Trop. Natuur 3, 1914, 56) depicted and described in detail the biology. The root system consists of three kinds, long + unbranched anchor roots, shorter much-branched feeding roots, and the erect spongy aerophores. BACKER cut the latter, but the plant remained (only very slightly less) buoyant. After pollination the pedicel bends down and the fruit ripens in the water (as in several other aquatics); the fruit decays and releases the cork- winged seeds which are buoyant. On desiccated muddy soils a never-flowering terrestrial form often occurs, marked by very small crowded leaves and a stronger pubescence. Uses. Malays in Perak use it for poulticing inskin complaints (BURKILL, Dict. 1935, 1273). Batak people use this (and other aquatics, like also do the Chinese) to feed pigs, and it is recorded to be eaten as salad in Indo-China. QUISUMBING (Medic. Pl. Philip. 1951, 677) reported that it is used in a decoction as an astringent for dysentery. Vern. Water primrose, E; Sumatra: buang buang, Toba-Batak; Java: pangeor, M, ganggéng landeuj, kambang peutjit, ruba silah, (rumput) kologa, S, krangkong, kréma, patjar banju, tapak doro, J; Philippines: sigang-dagat, Tag., gabi-gabi, Mag., tabagan, If.; New Guinea: agidahano, Kutubu lang. Note. L. adscendens appears to be allied more closely to the American L. helminthorrhiza (MART.) Hara than to any Old World species. Together with the mostly yellow-flowered African L. Stolonifera (GUILL. & PERR.) RAVEN, these three are the only species that regularly produce clusters of erect inflated aerophores at the floating nodes, although other species have descending root-like aerophores at these nodes and may have long, spongy aerophores from the submerged under- ground parts. Excluded or Doubtful ae i tenella BurM. f. Fl. Ind. (1768) 103, t. 34, 2. There is no unanimity of opinion about the identity of BURMAN’s plant which he said to have come from Java. MERRILL (Philip. J. Sc. 19, 1921, 369) suggested it to be a form of L. octovalvis which I doubt in view of the specific epithet and the species with which it is being contrasted. ALSTON (in Trimen, Fl. Ceyl. 6, 1931, 130) took it up for L. hyssopifolia, and this suggestion seems the most plausible. From BuRMAN’s description, which contains almost certainly errors, it cannot be identified. Unfortunately, or perhaps fortunately, the type at G could not be found. Ludwigia erigata LINNE, Mantissa 1 (1767) 40. — L. triflora Desr. in Lamk, Encycl. 3 (1792) 615, nom. illeg. subs., belongs to the Rubiaceae. Ludwigia trifolia Buro. f. Fl. Ind. (1768) 36; Houtt. Nat. Hist. 2, 7 (1777) 344 is according to MERRILL (J. Arn. Arb. 19, 1939, 368) Oldenlandia biflora L. (Rubiaceae). 2. EPILOBIUM Linné£, Gen. Pl. ed. 5 (1754) 164; Sp. Pl. 1 (1753) 347; HAUSSKNECHT, Monogr. Epilob. (1884); RAVEN, Bull. Br. Mus. Nat. Hist. Bot. 2 (1962) 325, 13 maps, pl. 33- 39; Brockig, New Zeal. J. Bot. 4 (1966) 366, 2 fig.; Bothalia 9 (1967) 309, 7 fig.; RAVEN, Blumea 15 (1967) 269, 7 fig.; BRockir, New Zeal. J. Bot. 8 (1970) 94; Raven, D.S.I.R. New Zeal. Bull. 216 (1976) 321 pp., 158 fig. — Fig. 7. Perennial herbs, often flowering in the first year, occasionally somewhat woody near the base. Leaves (in Mal.) opposite below, spirally arranged above. Flowers in the axils of the often greatly reduced upper leaves. Floral tube short (in Mal.) or essentially absent. Sepals 4, erect. Petals 4, white, pink, or purple, emarginate 108 FLORA MALESIANA [ser. I, vol. 8? en VII\ VI BASS PY) A Ha VO aw. . “s 7. SVE (Pu ere we Oka LL SSeS AUVN \- } Via PX Fig. 7. Epilobium hooglandii RAVEN. a. Habit, nat. size, b. node, x 10, c. seed with base of coma, x 30. — E. detznerianum SCHLTR ex Diets. d. Node, x 10. — E. prostratum Wars. e. Seed, x 30 (a—c HOOGLAND & PULLEN 5540, d HOOGLAND & PULLEN 5687, e WOMERSLEY c.s. 6103). After Raven, 1967. 1977] ONAGRACEAE (Raven) 109 (in Mal.). Stamens 8, in 2 whorls, the epipetalous ones shorter. Stigma (in Mal.) clavate or rarely globose, usually surrounded by the shedding anthers at maturity. Ovary 4-locular, the ovules very numerous. Fruit a long, slender, loculicidal cap- sule. Seeds very numerous, small, with a chalazal plume of trichomes (coma). Distr. About 200 spp., well-represented in temperate regions, mostly on the northern hemisphere, with the greatest centre of morphological diversity in the western U.S.A., at relatively high latitudes and alti- tudes; in the tropics confined to the mountains; in Malesia: rare and local, in W. Central Sumatra, E. Java (Mt Tengger), Lesser Sunda Is. (Lombok, Sumbawa, Timor), Philippines (N. Luzon, Panay), SW. Central Celebes (Latimodjong Range), Moluccas (Central Ceram), and New Guinea. Ecol. Characteristic of open, disturbed habitats or grassland or the alpine zone, not normally found in primary forest. Dispersal takes easily place by wind as the light seeds are provided with a silky coma. It is still rather surprising that, whereas open habitats are very common in the Malesian mountains, with their numerous volcanoes and alpine habitats, the number of localities is so very restricted. It must probably be considered that though dispersal may be easy, the coma only acts efficiently in dry air and this is in the tropics with their frequent rain and cloud formation and nightly fogs during most of the year a rather rare climatic situation. In this respect the west-monsoon, blowing from Asia southeastwards (Nov.— March) is most unsuitable, as these winds are laden with moisture. The east-monsoon, blowing from Australia northeastwards (May-Sept.) is in this respect definitely more favourable as this is a dry wind. This wind regime is a consequence of the situation of the Asian and Australian continents. Local dispersal by water might also occur, as the lowest localities in New Guinea are all along stream- sides, but this may also be due to the downward air-stream over rushing water. For hybridization see under the family. Genesis. The species occurring in Sumatra (1 sp.) and in Luzon (2 spp.) also occur in continental SE. Asia, the one in E. Java and the Lesser Sunda Is. (E. hirtigerum) also occurs in Australia, Tasmania, and New Zealand. The four remaining species are all endemic in East Malesia, 3 being confined to New Guinea and 1 also occurring in Central Celebes and Ceram. They are closely related to species found in Australia and New Zealand, from which they, and E. hirtigerum, were probably derived via the east-monsoon dis- cussed above. I have argued that the Australasian species ultimately show affinity to those of continental Asia and their ancestors must have reached Australia across the tropical mountains of Malesia. My assumption is that this southeastward penetration of the genus occurred in the Pliocene. KEY TO THE SPECIES 1. Buds and flowers erect, the inflorescence erect or somewhat drooping. 2. Stems pubescent only along conspicuously elevated lines running down from the margins of the DEES. Meee GLY, fe Ow, E. wallichianum 2. Stems pubescent all around. 3. Upper leaves strongly reduced. Inflorescence slightly nodding (Luzon). ween waves ovate, 0:4-1;5. cnt wide. i) 22. arte ie Lee ea oe 1. E. brevifolium 4, Leaves very narrowly elliptic to linear, 0.1-O.5Scm wide... .... . 2. E. platystigmatosum 3. Upper leaves not notably reduced. Inflorescence erect (Java, Lesser Sunda Is., New Guinea). 5. Plants finely glandular-pubescent; leaves usually ericoid in appearance; petals steerer ae ; . E. keysser og Maenwe? se | ot bhp We Sa Se el Kae Oe Bee ee Ga 5. Plants densely strigulose; leaves broader, not ericoid; petals white or very pale pink 4. E. hirtigerum 1. Buds and flowers drooping, becoming erect in fruit. 6. Stems with elevated glabrous lines running from the center of each petiole and elevated pubescent lines from their angles; petals 7-14 mm long 6. E. detznerianum 6. Stems without elevated lines, uniformly pubescent all around; petals 2.5-8.5 mm long. 7. Petals 6-8.5 mm long; seeds 1-1.4mmlong. .......... 7. E. hooglandii 7. Petals 2.5—5(-6) mm long; seeds 0.7-0.9mm long. . . . . 2-1 ee eee 8. E. prostratum efi tol .t? g 1. Epilobium brevifolium D. Don, Prod. Fl. Nepal. (1825) 222; RAveN, Bull. Br. Mus, Nat. Hist. Bot. 2 (1962) 361. ssp. trichoneurum (HAUSSKNECHT) RAveN, Bull. Br. Mus. Nat. Hist. Bot. 2 (1962) 362. — E. tricho- neurum HAUSSKNECHT, Oest. Bot. Z. 29 (1879) 54; Monogr. Epilob. (1884) 208; H. LéveiLvé, Ic. Gen. Epilob. (1910) t. 84. — E. philippinense C.B. Ros. Philip. J. Sc. 3 (1908) Bot. 209; Merr. En. Philip. 3 (1923) 220, excl. syn. Robust, perennial herb, 15-40 cm, the under- ground parts not scaly; plants strigulose, stems pubescent all around. Leaves mostly opposite, alternate in and near the inflorescence, the upper ones reduced, strigulose along the veins and margins especially below, the nerves evident, ovate, 1-3 by 0.4-1.5cm, serrate; petiole 1-2 mm. In- florescence slightly nodding. Flowers erect, borne in the axils of reduced upper leaves. Floral tube 1.82.4 mm across, |-1.3 mm deep. Sepals 2.5-5.5 by 1-1.3 mm. Petals obovate, 4.5-8 by 3-5.5 mm, 110 rose purple, the notch 1-1.5 mm deep. Anthers 1-1.2 mm long; filaments of the longer stamens 1.2-2.5 mm, those of the shorter 1-1.5 mm. Style 2.5-4 mm. Stigma clavate, 1.8-2.3 mm high, c. 1 mm thick, surrounded by the anthers at anthesis. Capsule 3.5-7cm long, on a pedicel 0-1 cm. Seeds 0.9-1.2 by 0.4-0.5 mm, papillose, blackish brown, not beaked, obovoid, blunt at both ends, the coma 5—7 mm long, white. Gametic chromosome number, n = 18. Distr. Continental SE. Asia (SE. Tibet: Chumbi Valley, to Assam, Burma, and W. China); in Malesia: Philippines (Mountain Province of N. Luzon). Ecol. Wet open slopes in the pine region, along streams and by springs, 1400-2100 m. Fi. July, Oct. Note. E. brevifolium ssp. trichoneurum is one of three subspecies of a species which ranges from Himachal Pradesh in the Western Himalaya east- wards throughout the Himalaya and southern China to Formosa, northern Luzon, northern Vietnam, and northern Burma. In northern Luzon, it occurs together with the other species found in the Philippines, E. platystigmatosum, and one plant of the collection CLEMENS 16385, suggests hybridi- zation between these two entities, which are usually widely distinct morphologically, although doubt- less more closely related to one another than to other Malesian species. 2. Epilobium platystigmatosum C. B. ROBINSON, Philip. J. Sc. 3 (1908) Bot. 210; MerrR. En. Philip. 3 (1923) 221; RAVEN, Blumea 15 (1967) 272. — E. cephalostigma var. linearifolium HisauT1, J. Jap. Bot. 14 (1938) 143, f. 3. — E. formosanum MaAsum. Trans. Nat. Hist. Soc. Formosa 29 (1939) 62; OnwIl, Fl. Japan (1965) 657. — E. sohayakiense Korpz. Act. Phytotax. Geobot. 8 (1939) 61. Slender, perennial herb, 15—40 cm tall, the under- ground parts not scaly; plants strigulose, densely so in the inflorescence, the stems pubescent all around. Leaves mostly opposite, alternate in and near the inflorescence, the upper ones reduced, strigulose along the veins and margins especially below, the nerves evident, very narrowly elliptic or linear, 1-3(-4) by 0.1-0.5 cm, weakly and sparsely ser- rulate; petiole 1-4mm. Inflorescence slightly nodding. Flowers erect, borne in the axils of reduced upper leaves. Floral tube c. 1 mm across, c. 0.8 mm deep. Sepals 3-4 by 1.2-1.6 mm. Petals narrowly obovate, 3.3-4.5 by c. 2mm, white or pale pink, the notch c. 1 mm deep. Anthers 0.2-0.3 mm long; filaments of the longer stamens c. 1.8 mm, those of the shorter c. 1.2mm. Style c. 2mm. Stigma broadly clavate, c. 0.8 mm high, c. 0.4 mm thick, surrounded by the anthers at anthesis. Capsule glabrescent, 3-5 cm long, on a pedicel 0-1.8 cm. Seeds 0.8-1 by 0.3-0.4 mm, papillose, not beaked, obovoid, blunt at both ends, the coma 4-6 mm long, white. Gametic chromosome number, n = 18. Distr. Japan, China (Hupeh, Kiangsu), For- mosa, and Malesia: Philippines (N. Luzon: Benguet Prov.; Panay: BS 31439). Ecol. Chiefly along small streams and about cliffs, 1200-2400 m. F/. April-June. Note. I can find no difference between the species generally known as E. formosanum and the sup- posed Philippine endemic populations of E. philip- FLORA MALESIANA {[ser. I, vol. 8? pinense. This species is apparently not common on the Asian mainland. 3. Epilobium wallichianum HAUSSKNECHT, Oéest. Bot. Z. 29 (1879) 54; RAVEN, Bull. Br. Mus. Nat. Hist. Bot. 2 (1962) 365; Blumea 15 (1967) 272. — E. nepalense HAUSSKNECHT, Oest. Bot. Z. 29 (1879) 53, p.p.; Monogr. Epilob. (1884) 218, p.p.; H. LEVEILLE, Ic. Gen. Epilob. (1910) t. 120. — E. duclouxii H. LEvEILLE in Fedde, Rep. 6 (1908) 110; Ic. Gen. Epilob. (1910) t. 144. — E. sarmen- taceum (non HAUSSKNECHT) BUNNEMEJER, Trop. Natuur 10 (1921) 57, f. 9. Erect perennial herb 15-SOcm, from a long rhizomatous base from which leafy shoots arise; plants strigulose, more densely so above, with elevated pubescent lines running down from the sides of the petioles, stems thick and hollow. Leaves opposite in lower half of the plant, alternate above, the margin and nerves densely strigulose, narrowly ovate or lanceolate, subacute at the apex, obtuse at the base, sharply and densely serrulate, 1.5—4 by 0.5—1.5 cm; petiole short but distinct, up to 2 mm. Inflorescence densely strigulose with an admixture of glandular trichomes, somewhat nodding in bud. Floral tube c. 2 mm across, c. 0.8mm deep. Sepals c. 5 by c. 2 mm, apiculate. Petals obovate, c. 8 by 3.5-4 mm, pale violet, the notch shallow, c. 1 mm deep. Anthers 1.3-1.5 mm long; filaments of the longer stamens c. 2 mm, those of the shorter c. 1 mm. Style 3-3.5 mm. Stigma globose, c. 1.5 mm thick, surrounded by the anthers at anthesis. Capsule densely strigulose with an admixture of erect, glandular trichomes, erect, 5-9.5 cm long, on a pedicel 1-2 cm. Seeds 1.2 by 0.6 mm, coarsely papillose, dark brown, the ei 6-7 mm long, white, tinged with brown at the ase. Distr. Continental SE. Asia (W. Nepal to Yunnan, south to the Khasya & Naga Hills), in Malesia: Central W. Sumatra (Mt Kerintji), one collection. Ecol. Along river-bank, c. 2000 m. 4. Epilobium hirtigerum A. CUNN. Ann. Mag. Nat. Hist. 3 (1839) 33; HAUSSKNECHT, Monogr. Epilob. (1884) 291; H. LEveILLE, Ic. Gen. Epilob. (1910) t. 18; ALLAN, Fl. New Zeal. 1. (1961) 279; Bur- BIDGE & Gray, Fl. A.C.T. (1970) 276; WiL.is Handb. Pl. Vict. 2 (1972) 464; RaveN, D.S.I.R. New Zeal. Bull. 216 (1976) 141, f. 63-64, 65-66 (maps). — E. brasiliense HAUSSKNECHT, Oéest. Bot. Z. 29 (1879) 119. — E. sarmentaceum (non HAUSSKNECHT) Koorp. Exk. FI. Java 2 (1912) 704; Back. & BAKH. f. FI. Java 1 (1963) 262. — E. cinereum (non A. RicH.) RAVEN, Blumea 15 (1967) 273, pro specim. mal. Robust, erect, clumped perennial herb, 15-100 cm, the underground parts not scaly; plants densely strigulose, the stems pubescent all around. Lower- most /eaves opposite, the rest alternate, densely strigulose, especially below and along the veins, nerves evident, narrowly lanceolate, 0.6-3 by 0.2-0.5 cm, coarsely serrate, subsessile. Inflores- cence erect. Flowers erect, borne in the axils of upper leaves. Floral tube c. 1 mm across, c. 1 mm deep. Sepals 3—5.5 by 0.8-1.6 mm. Petals obovate, 3-5 by 2-3.2 mm, white or very pale pink, the notch c. 1mm deep. Anthers 0.7-1 mm long; 1977] ONAGRACEAE (Raven) 111 filaments of the longer stamens 2-2.8 mm, those of the shorter 1-1.8mm. Style 1-3mm. Stigma clavate, 1.5-2.7 mm _ high, 1-1.5 mm thick, sur- rounded by the anthers of anthesis. Capsule 3-5(-6)cm long, on a pedicel 0-2cm. Seeds 0.9-1.2mm long, 0.35-0.45mm thick, finely papillose, brown, not beaked, obovoid, blunt at both ends, the coma 5-8 mm long, white. Gametic chromosome number, 7 = 18. Distr. Very widely distributed, South America (Argentina, Uruguay, Brazil: Santa Catarina), New Zealand, Australia, and Malesia: Lesser Sunda Is. (Timor, Sumbawa, Lombok) and East Java (Mt Tengger). Ecol. Moist places, grasslands; in E. Java the only locality is near the single small well on the otherwise dry and barren extinct volcanic cone of Mt Widodaren on Mt Tengger caldera, at 2100 m. In Lombok in Casuarina forest; in Timor in Eucalypt savannah; 1800-2200 m. F/. Oct.—Jan. Note. Variable species. The Malesian specimens belong to a highly autogamous Australian form with small white flowers which occurs in swampy places throughout the lowlands of Australia (from Queensland to temperate W. Australia and Tas- mania) and is predominant in New Zealand. I have assumed that it came there from Australia and has spread again from New Zealand to South America, either by natural dispersal or by man. 5. Epilobium keysseri DreLs, Bot. Jahrb. 62 (1929) 486; HooGL. Blumea Suppl. 4 (1958) 223; Borc- MANN, Z. Bot. 52 (1964) 124, 143; RAVEN, Blumea 15 (1967) 274, f. 1 (map). Clumped erect perennial herb 12-60 cm, often ericoid in aspect, + woody at the base, the under- ground parts not scaly; plants finely glandular- pubescent. Lowermost /eaves opposite, the rest alternate, coriaceous, subglabrous, the nerves obscure, margin revolute, narrowly elliptic to elliptic, 0.5-1.3 by 0.1-0.2(-0.4) cm, with a few coarse teeth on each side, subsessile. Inflorescence erect. Flowers erect, borne in axils of upper leaves. Floral tube 1-2 mm across, 0.7—1 mm deep. Sepals 3-5.6 by 1-2.5 mm. Petals obovate, 4.5—-8 by 2.8-4 mm, purplish rose, the notch 1.5-2mm deep. Anthers 0.7-1 mm long; filaments of the longer stamens 1.8—2.5 mm, those of the shorter 0.5-1.5 mm. Style 2.5-4mm. Stigma clavate, 1.5-2 mm high, 0.5—0.7 mm thick, surrounded by the anthers at anthesis. Capsule 3-6 cm long, glabrescent, on a pedicel 1-2.5cm. Seeds 0.9-1.2 by 0.4-0.6 mm, papillose, brown, with a short pellucid beak, the coma 5-7 mm long, white. Gametic chromosome number, 1 = 18. Distr. Malesia: New Guinea (Mt Wilhelmina in West, common on many summits in East), many collections. Fig. 8. Ecol. Subalpine and alpine meadows and grass- lands, open shrubberies, swampy treefern grass- land, occasionally epiphytic on treefern trunks, forest glades, often in succession after ground-fires, (1600-)1950—3800 m. F/. (Jan.—)June—Aug.(—Dec.). Vern. Aingum, Tomba, gonema, Chimbu, yandepai, Enga lang., Wabag, papal, Enga lang., Poio, tamitan, Mendi lang., Giluwe. Notes. A distinctive species but clearly belong- ing to the Australasian assemblage. Most plants, with their narrow leaves, are dis- Fig. 8. Range of Epilobium keysseri Diets (after RAVEN, 1967). tinctly ericoid in appearance, and thus parallel representatives of many other typically non-ericoid groups that occur in alpine and subalpine regions of New Guinea. 6. Epilobium detznerianum SCHLTR ex DIELs, Bot. Jahrb. 62 (1929) 485; RAVEN, Blumea 15 (1967) 277, f. 3, 5 (map). — E. papuanum RIDL. var. alpestre RiDL. Trans. Linn. Soc. Bot. II, 9 (1916) 58. — E. papuanum (non RID.) HooGL. Blumea Suppl. 4 (1958) 228. — Fig. 7d. Clumped perennial herb 3-15 cm, the under- ground stems not scaly; plants mostly glabrous, with elevated, strigulose lines running down from the margins of the petioles and glabrous ridges running down from back of petioles. Leaves mostly opposite, alternate in the inflorescence, coriaceous, nerves obscure, broadly elliptic or ovate, obtuse at apex and base, entire, 0.3-1 by 0.2-0.7 cm; petiole 1-1.5mm, short but distinct. Flowers nodding, the ovaries erect, borne in the axils of upper leaves. Floral tube 1.5-3 mm across and about as deep. Sepals 4.5-7 by 1-2.5 mm. Petals obovate, 7-14 by 3-6 mm, bright purplish rose, the notch c. 2 mm deep. Anthers 0.8-1.2 mm long; filaments of the longer stamens 4—5.5 mm, those of the shorter 3.3-4.5 mm. Style 5.5-8 mm. Stigma broadly clavate, 1.3-1.7 mm high, c. 1 mm thick, surrounded by or held just above the anthers at anthesis. Capsule erect, subglabrous, 4-5 cm long, on a pedicel 2.5-8.5cm. Seeds (0.9-)1-1.5 by 0.5-0.7 mm, not beaked, finely papillose, pale brown, the coma c. 8 mm long. Gametic chromosome number, 7 = 18. Distr. Malesia: New Guinea (Mts Carstensz & Wilhelmina in West, Telefomin, Mts Sarawaket & Wilhelm in East). Fig. 9. Fig. 9. Range of Epilobium detznerianum SCHLTR ex Dies (after RAVEN, 1967). 112 FLORA MALESIANA [ser. I, vol. 8? Ecol. Subalpine and alpine grasslands and ridges, open slopes and bogs, earth screes, sub- alpine forest and its grassy edges, rock clefts, cliff crevices in alpine thickets, on dripping slate land- slides, more rarely on sandy or gravelly gully beds; 3000-4500 m. Fi. (Jan.—Febr.—)May-—Sept. (—Dec.). Note. An attractive alpine species reaching the highest elevations in the genus in New Guinea. It is usually more condensed in habit than E. hooglandii and differs at once from that species in the glabrous ridge decurrent from the center of each petiole and the elevated pubescent lines decurrent from the edges of the petiole. It grows sympatrically with E. hooglandii and E. keysseri, but no intermediates have been observed; intermediates with the former species would be very difficult to detect. 7. Epilobium hooglandii RAVEN, Blumea 15 (1976) 278, f. 2,6 (map). — E. pedunculare (non A. CUNN.) F.v.M. Trans. R. Soc. Vict. I, 2 (1889) 7. — E. detznerianum (non SCHLTR ex DiELS) HOoc_. Blumea Suppl. 4 (1958) 228. — Fig. 7a—c. Caespitose perennial herb with decumbent branches, the erect portions 10—25(—45) cm long; plants glandular pubescent along elevated lines running down from the margins of the petioles below, more densely and uniformly so above, and with an admixture of strigulose pubescence in the inflorescence. Leaves mostly opposite, alternate in the inflorescence, subcoriaceous, nerves + visible in dried material, broadly elliptic to ovate, acute or obtuse at apex and base, entire or with a few teeth on the margins, 0.5-1.3 by 0.3-0.9 cm; petiole 1-3 mm, short but distinct. Flowers nodding, the Ovaries erect, borne in the axils of upper leaves. Floral tube 1.4-2mm across, 1-1.2mm deep. Sepals 3-4 by 1-1.6 mm. Petals 6-8.5 by 2.5—4.5 mm, pink to purplish rose, the notch c. 1.5 mm deep. Anthers 0.7-1 mm long; filaments of the longer stamens 3—6 mm, those of the shorter 2-4 mm. Style 2.5-6 mm. Stigma clavate, 2—2.5 mm long, 1-1.2 mm thick, surrounded by the anthers at anthesis. Capsule erect, glabrescent, 5-8 cm long, on a pedicel 4-12 cm. Seeds 1-1.4 by 0.3-0.45 mm, finely papillose, brown; coma 5-8 mm long, white. Gametic chromosome number, n = 18. Distr. Malesia: New Guinea (Mt Wilhelmina in West, many localities in East). Ecol. Subalpine and alpine meadows, fire- induced alpine grassland, near boulders, in alpine shrubbery, on peaty grassland, near waterfalls, in secondary forest on limestone cliffs, on stream- bank gravel, stony creek beds, along forest tracks, between grass tussocks in old lake basin, in pen- dent masses on rocks, occasionally in moist, eG areas; 2000-4150m. FI. May—Nov. —Jan.). Vern. Dirimpia, Chimbu, Masul, nonami, Mairi, Mondo. Note. Differs from the closely related E. detznerianum by its evenly pubescent stems, and from E. prostratum by large flowers and seeds. It grows sympatrically with the other three New Guinean species. Occasional collections are inter- mediate with E. prostratum and obviously result from incidental hybridization. However, the two species keep in general amply distinct and are as well differentiated as most recognized species of the genus. 8. Epilobium prostratum Wars. Bot. Jahrb. 16 (1893) 15, 23; RAVEN, Blumea 15 (1967) 280, f. 4, 7 (map). — E. papuanum Rip-. Trans. Linn. Soc. Bot. II, 9 (1916) 57. — Fig. 7e. Similar to E. hooglandii, but differing as follows: habit lax, spreading. Leaves 0.4-0.8 by 0.2-0.4 cm. Floral tube 0.8-1.1 mm across, 0.6-0.9 mm deep. Sepals 1.5-3 by 0.6-1.4 mm. Petals 2.5—-5(-6) by 1.3-2.5 mm, very pale pink to purplish rose, the notch c. 1mm deep. Anthers 0.5-0.7 mm long; filaments 1.8-3.5 mm. Stigma clavate, 0.8—1.5 mm long, 0.50.9 mm thick, surrounded by the anthers at anthesis. Capsule 3.5—-5 cm long, on a pedicel 5-10 cm. Seeds 0.7-0.9 by c. 0.3-0.4 mm, finely papillose, brown, the coma 5-8 mm long. Gametic chromosome number, n = 18. Distr. Malesia: SW. Central Celebes (Latimod- jong Mts), Moluccas (Ceram), and throughout New Guinea. Fig. 10. Fig. 10. Range of Epilobium prostratum WARB. Ecol. Moist open places, especially on rocky alluvium bordering streams, stream-banks across treefern grassland, along river-bank in limestone scree, on fallen trees on open ridge, rarely in montane forest, sometimes colonizing landslips; (1200-)1900-3400 m, in Celebes and Ceram at c. 2750-3400 m. Fl. Jan.—Dec. WARBURG’s type specimen was found by HELL- WIG at an exceptional low altitude of 1200 mina streambed. Vern. New Guinea: kimbil, Enga lang., Poio, dirimpia, Chimbu, Masul, kokorabadi, Mandi lang. Notes. Closely similar to E. hooglandii, but readily distinguished for the most part by smaller flowers and seeds. Occasionally intermediate specimens (obviously hybrids) are found where the two occur together. In New Guinea it descends to lower altitudes and has correspondingly a much wider range. Introduced Fuchsia boliviana CARR. Rev. Hort. (1876) 150; as F. corymbiflora (non R. & P.) BACK. & BAKH. f. FI. Java 1 (1963) 264. Locally cultivated and perhaps established in the mountains of West and East Java (Mts Malabar and Tengger) between 1500 and 2000 m. 1977] ONAGRACEAE (Raven) 113 Easily distinguished from the following species by its drooping inflorescences of bright red flowers with a floral tube 5-6 (instead of less than 1) cm long. Fuchsia magellanica LAMK var. gracilis (NICH.) BAILEY: BACK. & BAKH. f. Fl. Java 1 (1963) 264; as F. coccinea (non SOL. ex AITON) CuRT. — BUNNEMEWER, Trop. Natuur 10 (1921) 56, fig.; Wisse, ibid. 11 (1922) 480, fig.; Hocur. Candollea 2 (1925) 480; BAck. & BAKH. f. Fl. Java 1 (1963) 264. This species, native of temperate South America, has repeatedly been reported to occur cultivated but also naturalized in anthropogenous places and on volcanic ash on the mountains of West, Central, and East Java (Mts Patuha, Malabar, Diéng, Sindoro, and Tengger) between 1000 and 2100 m, and in the mountains of Sumatra (Karo Lands). Oenothera stricta LEDEB. ex Link, En. Hort. Berol. 1 (1822) 377, ssp. stricta; as O. erythrosepala (non BorsAs) BACK. & BAKH. f. Fl. Java 1 (1963) 262. — Oenothera sp. Doct.v.LEEUWEN, Verh. Kon. Ak. Wet. A’dam II, 31 (1933) 191. Docrers VAN LEEUWEN /.c. introduced this species from seed he collected in Hawaii and sowed on the summit of Mt Pangrango in West Java, at 3000 m altitude, in 1921. It is maintaining itself there and, although self-pollinating, is visited by Bombus rufipes (HEIDE, Dansk Bot. Ark. 5, 1927, 18) and doubtless by nocturnal insects as well. It is a native of temperate South America, widely cultivated and naturalized. It may be distinguished from all other species of the family in Malesia by its combination of a long floral tube and yellow petals; the flowers open at sunset. Cultivated Representatives of several genera — Clarkia and Gaura among them — are cultivated, mainly in the mountains. BACKER & BAKHUIZEN VAN DEN BRINK f., Fl. Java 1 (1963) gave an account of these. BIGNONIACEAE (C. G. G. J. van Steenis, Leyden) Trees, shrubs, lianas, very rarely herbaceous (extra-Mal.); twigs often lenticellate and nodes with gland fields; spines very rare (extra-Mal.). Stipules absent. Leaves simple or mostly compound (digitate or impari-1—4-pinnate), (in Mal.) decussate, rarely in whorls of 3-4, often provided with glands underneath, in the New World often provided with terminal tendrils, rarely scattered or in pseudo-whorls (extra- Mal.); domatia sometimes present (fig. 8b, 23h). Inflorescences bracteate, cymose, but not rarely thyrses contracted to racemiform or racemose inflorescences, or even reduced to solitary flowers (extra-Mal.), terminal, axillary or from the old wood. Pedicels mostly with 1-2 bracteoles. Flowers usually very showy, rather large, bisexual, articulate with the pedicel or not. Calyx connate, closed in bud and later (not rarely irregularly) splitting into lobes, or cupular, or spathaceous, or lobed from the beginning and with equal or unequal, valvate lobes, developing earlier than the corolla, often glandular outside and inside with water and slime producing glands and hydathodes, persistent or circumscissile caducous along an abscission line. Corolla sympetalous, campanulate, tubular, funnel- or salver-shaped, mostly zygomorphic, lobes equal or unequal, valvate or imbricate in bud, tube often with a narrow cylindrical (constricted) lower part (basal tube) and a widened upper part (upper tube). Stamens 5 almost equal, or mostly 4 didynamous, the 5th sterile, rudimentary, adnate to the corolla tube, mostly inserted at the rim of the basal tube and not rarely (glandular) hairy at the insertion, more rarely inserted higher up. Anthers basifixed, 2-celled, rarely one cell barren or 1-celled, introrse, dehiscing lengthwise, usually the anthers connivent in pairs; anther cells often free and diver- gent, connective not rarely produced. Disk intrastaminal, mostly annular, rarely absent. Ovary superior, 2-celled, rarely 1- or 4-celled (extra-Mal.); style filiform, stigma usually 2-lipped, sensitive. Ovules (in Mal.) in each cell on the septum in two or more rows of 3-04, mostly on 2 placentas. Capsule 2-valved, either loculicid with the septum perpendicular to the valves — sometimes provided with an addi- tional transverse false septum — or septicid with the septum parallel with the valves, or (extra-Mal.) an indehiscent, 1-celled, soft or hard-shelled, pulpy berry. Seeds in each cell attached to the dissepiment in one or more rows, inserted transverse to axis of fruit, anatropous, mostly on both sides with hyaline wings; embryo exal- buminous, the cotyledons mostly notched, sometimes on both sides. Germination always epigeal. . Distribution. About 120 genera and some 650 spp., mainly in the tropics and subtropics, roughly between 40° N and 30-35° S, very few in the warm-temperate zone; in Malesia: 14 native genera of which 2 are endemic, viz Hieris in Penang and Lamiodendron in Papuasia. Among the remaining 12 one occurs through the Old World (Dolichandrone), 7 are shared with continental SE. Asia (two of which extend also to Africa and Madagascar: Fernandoa, Stereo- spermum) and 4 with Australia and Melanesia; the latter occur in Malesia only in the east except Deplanchea which ranges westward to Sumatra. In the family tropical Asia and Africa share a few genera (Markhamia, Fernandoa, Stereosper- mum, and Dolichandrone), but Africa and America share only one, viz Tecoma. This latter affinity goes further, though very disjunct via Campsidium (Chile) and Campsis (N. America and E. Asia) to Tecomanthe-Pandorea-Neosepicaea (Moluccas to Three Kings Is. and E. Australia). Otherwise there appear to be only two other transoceanic ties, viz tribe Crescentieae which is shared by Africa and the Americas, and the genus Catalpa which occurs in E. Asia and the Caribbean area. As GENTRY (Brittonia 25, 1973, 227-230) has shown, the average number of species per genus (114) 1977] BIGNONIACEAE (van Steenis) 115 is only 5, which is very small in comparison with many other families, but can only partly be explained by a possibly small generic concept. There are quite a number of monotypic genera (in Malesia 5), but they are well defined in many characters and stand very apart. This, and the worldwide distribution of the family, and the disjunctions in ranges, definitely point to relict survival and ancient origin, onwards of which period the three tribes have under- gone a separate, independent development on the continents, mainly leading to differentiation in Indo-Australia and in the New World, with the greatest abundance in the latter. Unfortunately, the fossil evidence (only Tertiary) is meagre and untrustworthy (SCHUMANN in E. & P. Nat. Pfl. Fam. 4, 3b, 1894, 208), both to macrofossils and to pollen. Ecology. Within the family there is a fairly wide coverage of habitats and there are quite a few which are confined to arid conditions (Rhigozum and Catophractes in S. Africa, Tecomella in Arabia, Phyllarthron bernierianum in Madagascar, Dolichandrone filiformis, D. heterophylla and the linear-leaved drought form of Pandorea pandorana in Australia). A few are rheophytic (Astianthus and Chilopsis in the Americas). A few are warm-temperate (Catalpa) or subtropical- alpine (U/ncarvillea incl. Amphicome in the Sino-Himalayas and Argylia and Campsidium in the Andes). One species is bound to the mangrove (Dolichandrone spathacea in Indo-Melanesia). The majority, however, belongs to the tropical forest, mostly the everwet type, but a fair num- ber in the seasonal type, below c. 2000 m; only Tecomanthe ascending to c. 3100m in New Guinea. In Malesia most are evergreen, but Oroxylum indicum (fig. 5) and Dolichandrone spathacea can stand leafless in the dry season for many months. Fernandoa macroloba is also deciduous, as are the species of Stereospermum. Habit. The majority of the Malesian genera are small or large trees, the only climbing genera being Nyctocalos, Hieris, Tecomanthe, Pandorea, and Neosepicaea; in subalpine heathland Tecomanithe may be forced to creep on other vegetation. Most trees are of medium size, but species of Stereospermum, Fernandoa, Pajanelia, and also Radermachera gigantea may attain quite good dimensions. Oroxylum is a short-lived nomad tree. Dominance. Almost all species occur scattered in the forest and several are very rare indeed. An exception is Dolichandrone spathacea, bound to the swampy, brackish inner mangrove, which according to CORNER (Wayside Trees, 1940, 164) is in the North of the Malay Peninsula, in Perlis, a feature of the country, flanking roads and standing as an upright poplar in the rice- fields. Fig. 16. In Great Natuna I. (NW. off Sarawak) I found Pajanelia longifolia locally very common in coastal forest, but this was probably encouraged by devastation. In secondary forests, on earth slides, abandoned fields, and on fresh volcanic ash Radermachera glandulosa and R. gigantea may be frequent in the pioneer upgrowth in Java (also on Krakatao), but this high frequency is local; KOoRDERS mentioned it for G. Telemojo and Pringombo in Central Java. Even Oroxylum indicum which is a nomad plant bound to secondary growths is always found in only a few specimens. Lamiodendron magnificum was once mentioned by Brass as forming a com- munity on a gravel bank behind the beach, in Normanby I., but this tree is extremely rare other- wise. Flower biology. In many species (like in several Gesneriaceae, Verbenaceae, Solanaceae) the calyx develops much earlier than the corolla and is closed in bud. Inside of the calyx with water and slime producing glands and hydathodes in which the corolla develops. These so-called waterbuds are very characteristic, especially in such large-flowered species as Spathodea campanu- lata, the tulip tree, which derives a Dutch and Malay name from this feature (spuitjesboom, panchot) which is enjoyed by children to play with. It is one of the few biological phenomena which are entirely confined to the tropics, as far as I am aware. Treus (Ann. Jard. Bot. Btzg 8, 1889, 38-46, t. 13-15) made an anatomical study of the glands inside the calyx of Spathodea campanulata and on his instigation GREsHOFF examined the exudate, dissolved organic and inorganic substances, which appeared similar to those of leaf-hydathodes. Koorpers (ibid. 14, 1897, 354-469, t. 21-27) extended this subject with research on some other genera of Bignoniaceae (Parmentiera, Kigelia, Crescentia, Fernandoa adenophylla, Radermachera gigantea) and some other plants, confirming TrReus’s results. Pollination. Flower-shape, -colour, -position, and -scent are very different in the mostly showy flowers of Bignoniaceae, and the syndromes attract different visitors. 116 FLORA MALESIANA (ser. I, vol. 8? Bats are frequently visiting species of certain genera, another phenomenon restricted to tropi- cal plants. According to FAEGRI & VAN DER PuL (Principles Pollination Ecology, ed. 2, 1971, 154) the attraction syndrome is: nocturnal anthesis, whitish, creamish or drab greenish or dark purple colour, stale or sour, unpleasant smell reminiscent of fermentation at night, large quantity of nectar and pollen in large anthers, large-mouthed and coarse flowers on strong stalks sticking out of the foliage or cauliflorous to flagelliflorous flowers, thus coming into easy reach for landing. This is found in Malesia in several cultivated genera (Kigelia, Crescentia, Parmentiera, Mark- hamia, etc.) but occurs also in the native Fernandoa adenophylla, Pajanelia, and Oroxylum. Fig. 7. Notwithstanding the many papers and records of observation — corollas show claw marks after these visits — it is not proved to my satisfaction that visits of bats are compulsory for polli- nation cq. fertilisation, experimenting being in this field deplorably meagre. My doubt is streng- thened by observations by HARRIS & BAKER in Ghana where Kigelia is native (J. West Afr. Sc. Assoc. 4, 1958, 28) and can set fruit in absence of bats; they observed also frequent visits by sphingids but they doubt effective pollination by these. Birds, humming birds and sun-birds, frequently visit certain species, the attraction syndrome being: tubed, vividly coloured (orange, scarlet), diurnal, mostly odorless, nectar-producing tubular flowers. Here also many observations are made, e.g. in Tecoma (Tecomaria). To this class belong in Malesia some species of Radermachera (R. ramiflora), Neosepicaea, Tecomanthe, and it can be expected for Deplanchea. Also the cultivated Spathodea campanulata is frequented by birds (cf. BEUMEE, Trop. Natuur 14, 1925, 28-30), notably kutilans and ?djalaks, at Bogor; they severely damage the corolla. Here again the question whether bird-visits are compulsory for pollination cg. fertilisation is inadequately supported by experiments. Caution is necessary to conclude to the necessity of cross-pollination, as e.g. HUNTER (Rec. Auckl. Inst. Mus. 6, 1967, 169-170, t. 24) recorded that in Tecomanthe speciosa, of which cuttings of a single plant led to its cultivation, fertilisation — that is selfing — could be effected by hand-pollination, but later also naturally by bees, although far from its native habitat. Moths. A few species have the moth-attraction syndrome for flower visitors, which implies: nocturnal, very fragrant flowers with abundant honey, in mostly pale or white, long-tubed or salver-shaped corollas. Fig. 1, 8, 15. This is found in species of Nyctocalos, Dolichandrone, Hieris, Millingtonia, and some species of Radermachera (e.g. the Chinese R. sinica, R. pentandra, and R. peninsularis). Probably long-tongued sphingids (hawk-moths) visit these flowers. Bees and butterflies. Possibly bees visit flowers of species not belonging to the three categories mentioned above. The ecological role of the many sorts of extra-floral nectaries in unexplained. It is a fact that in general fruit setting is scarce in Bignoniaceae; in several the fruit was only occasionally found long after the plant had been described in flower. In Malesia the fruit is still unknown of Hieris and Lamiodendron. Even after abundant flowering fruit production is often very low with 1 or 2 fruits in each inflorescence, except in Radermachera glandulosa and Tecoma stans. With all these flower visit devices one would expect otherwise. Dispersal. Bignoniaceae occur throughout the tropics and several are still found in the sub- tropics of the whole world. One might ascribe this to their having winged seed (except Crescentieae and a few other exceptions), but against expectations they are almost absent from oceanic islands. Bignoniaceae occur all along the coasts of the West Pacific, notably in New Guinea and in Australia species of Tecomanthe and Pandorea are not rare, but the only occurrence in the West Pacific islands is a common Australian Pandorea in New Caledonia and Lord Howe lI., anda peculiar Tecomanthe in a single locality of the Three Kings Is., the northernmost territory of New Zealand. Obviously wind dispersal has not been as effective as one would expect. Dispersal by seawater is common in Dolichandrone spathacea, a back-mangrove species, ranging from the western Deccan Peninsula to North Luzon, south to Timor and southeastwards to New Caledonia; the range is almost continuous without gaps. Fig. 14. It is most peculiar, however, that so far it has never been found in the mangroves of northern Australia. Its seeds have thickish corky wings instead of flimsy wings as usual in most members of the family (except the fleshy indehiscent fruits of the Crescentieae and a few other exceptions as e.g. Pauldopia) and are most excellently adapted to be dispersed by seawater. 1977] BIGNONIACEAE (van Steenis) 117 Seedlings. These are very uniform in all tribes of the family; in the embryo the foliaceous cotyledons are flat in one plane, mostly emarginate at both ends, hypocotyle and rootlet are small. Germination is epigeal by stretching of the hypocotyle. The first leaves are mostly simple, as in most compound-leaved families; they are often dentate. In a very few genera with thick seeds there may develop — possibly by intrusion of the testa — a false septum in the seed, in Malesia notably in Stereospermum and here also the cotyledons are folded. The only exception is men- tioned by LusBock (Contr. Knowl. Seedlings 2, 1892, 334, fig. 569) for ‘“Bignonia insignis’ with fleshy connate cotyledons and hypogeal germination; the name is evasive and at Kew the identity could not be traced; presumably the record rests on an error. Literature: E. BureEAu, Monogr. Bign. (1864); J. LusBock, Contr. Knowl. Seedlings 2 (1892) 332-345, fig. 569-575; R. S. Troup, Silvic. Indian Trees 2 (1921) 684-693; J. A. Duke, Ann. Mo. Bot. Gard. 52 (1965) 349, pl. 20; G. DE LA MENSBRUGE, La germination et les plantules essences arbres Céte d'Ivoire; Techn. For. Trop. Nogent-sur-Marne, Paris (1966) 332-333; D. BURGER, Seedlings (1972) 52-54; C. S. SCHOPMEIJER, Seeds Woody PI. U.S., Agric. Handb. 450 (1974) 260, 281, 321. Juvenile plants of Pandorea pandorana show leaves very different from the mature foliage, in having many jugae and being coarsely dentate. Tecoma filicifolia NICHOLS. was based on such material. This led also to a serious misinterpretation of Tecoma leptophylla Bu., from New Guinea, of which the juvenile leaves (fig. 37b) are Pandorea pandorana but the flowers belong to Neosepicaea. Taxonomy. Since the basic work on the systematy by E. BuREAU (Monographie, 1864), the treatment of the family in Flora Brasiliensis by BUREAU & SCHUMANN (1896-97), and the treat- ment of SCHUMANN in the Pflanzenfamilien (1895) the traditional subdivision of the family in 5 tribes has proved satisfactory. Crescentieae with 1-celled berries occur in Africa and the Americas, two other monogeneric tribes are South American, while the bulk of the family belongs to Big- nonieae and Tecomeae, of which the latter are about balanced as to number of genera in the Old and New World, but Bignonieae are predominantly American. These two tribes are largely dis- tinguished on the dehiscence of the fruit, loculicid in Bignonieae and septicid in Tecomeae. In passing it may be remarked that Gentry (PI. Syst. Evol. p. 126, 255) recently advocated that Crescentieae of the neo-tropics and of Africa-Madagascar are of separate descent and would represent two parallel evolutionary lineages; this suggestion is more based on geographic argu- ment and evolutionary hypotheses than on morphological arguments. The delimitation against other families of Sympetalae is well-defined, but there are a few genera, notably Wightia and Paulownia, which are sometimes referred to Bignoniaceae, though FENZL (Denkschr. K. Bay. Bot. Ges. Regensburg 3, 1841, 227-230), BUREAU, SCHUMANN, VON WETT- STEIN, and other specialists referred them to Scrophulariaceae. A survey of opinions I gave in my paper on Wightia (Bull. Bot. Gard. Btzg III, 18, 1949, 214-216), in which I excluded it from Bignoniaceae. Even recently Paulownia is sometimes casually treated as Bignoniaceous (e.g. SCHOPMEVER, Agric. Handb. U.S.A. 450, 1974, 527), although the embryo is embedded in endosperm; furthermore the stigma is different from that in Bignoniaceae, the anthers have no prolonged connective, there is no rudimentary stamen and the seeds are provided with several wings and seem to be laterally attached, not transverse as in Bignoniaceae. For Wightia I tabulated (1.c.) the relation to both families. Its seeds have no endosperm, but the absence of a staminode, the structure of the stigma, the central placenta and the absence of a produced connective on the anthers point distinctly to Scrophulariaceae. The seed is quite differently attached as compared with Bignoniaceae, viz laterally and the wing surrounds the entire seed. Its wood has two kinds of medullary rays, narrow and broad ones, a character which, at least in Malesian Bignoniaceae, is absent. Though the capsule in Wightia is septicid and in Paulownia loculicid, both genera have the same kind of axile placentation, in which the thickened placenta becomes detached from the valves as a subquadrangular seed-cake, showing their close affinity, completely differing from the situation in Bignoniaceae. According to SURYAKANTA (J. Palyn. 9, 1973, 73) the pollen of both genera differs from that in Bignoniaceae and resembles that of Scrophulariaceae. NAKAI (J. Jap. Bot. 24, 1949, 13) accommodated Paulownia in Paulowniaceae, probably in- 118 FLORA MALESIANA [ser. I, vol. 8? duced mostly by its arboreous habit and fruit; they certainly merit to be placed in a separate tribe or subtribe of Scrophulariaceae. We regard nowadays the arboreous habit as primitive in herbaceous families and we might conclude that they are ancient relicts from a period when Bignoniaceae and Scrophulariaceae had a common matrix. Also in South America there are two woody genera of the Scrophulariaceae which were at times referred to Bignoniaceae, viz Schlegelia (syn. Dermatocalyx) and Gibsoniothamnus, accord- ing to GENTRY (Fieldiana, Bot. 34, 1971, 55; Ann. Mo. Bot. Gard. 61, 1974, 533-537); see also LEINFELLNER (Oest. Bot. Z. 121, 1973, 13-22). They are (hemi-?) epiphytic shrubs or lianas, a similar habit as in Wightia. Genetics. Chromosomes. DARLINGTON & WyLIiE (Chrom. Atlas, 1955) and Moore (ed.) (Regn. Veg. 90, 1973) gave for 26 genera x = 20 (2n = 40) and they belong to Tecomeae, Bignonieae and Crescentieae, both from the palaeo- or neotropics. There is one higher number x = 22 (Amphilophium, South America, Niedzwedzkia = Incarvillea) and several lower ones: Pandorea, and some doubtful countings in Tecoma x = 19, Tecomanthe dendrophila 2n = 36 (Christine BRIGHTON in litt.), Jacaranda x = 18, Tecoma capensis x = 17, Oroxylum, Milling- tonia, Argylia (from South America) x = 15, Spathodea x = 13, and Incarvillea x = 11. In supplement indices Campsis is also given as 16 and Oroxylum as 14. I have scanned the numbers of Scrophulariaceae, Gesneriaceae and Verbenaceae, but can find no reliable ties, Bignoniaceae being obviously more homogeneous than those. The number given for Paulownia, 2n = 40, x = 10, might as well fit Bignoniaceae as Scrophu- lariaceae. Hybridisation. Not many species hybrids are known to me, but those known are interesting, as there are at least two between species of East Asia and SE. North America which are now very disjunct after the Pleistocene Ice Age; it is not impossible that they formed part of more con- tinuous populations in the warmer Pliocene via Beringia. This idea is supported by the fact that in both cases the hybrids are fertile. E. C. SmitH (J. Arn. Arb. 22, 1941, 219) reported on Catalpa ovata DON x C. bignonioides WALT. (= x C. syringifolia Sims). Haploid all have 20 chromosomes (SAx, J. Arn. Arb. 14, 1933, 274). Then there is x Campsis tagliabuana (VIVIAN1) REHDER, a hybrid between the Chinese C. grandiflora (THUNB.) K. Scu. (C. chinensis (LAMK) Voss.) and C. radicans (L.) SEEM. which pro- duces fertile progeny (cf. STEARN, Bot. Mag. 169, 1953, t. 198). The third one is also bi-continental, Tecoma smithii W. WATSON (Gard. Chron. 14, 1893, 649, fig. 104; E. Smitu, ibid. 16, 1894, 64; cf. also Gartenflora 44, 1895, 51, fig. 14). This is a reputed hybrid, which E. SMITH made at Adelaide, in 1882, between 7. velutina (a hairy variety of T. stans) and T. capensis. It was propagated by cuttings, but it produced seed and its offspring of seedlings diverged in size and flower colour. Curiously SPRAGUE, in a succinct note (Fl. Cap. 4, 2, 1904, 448) reduced it to T. alata DC., without referring to its hybrid nature. Anatomy. Wood. Of the Malesian Bignoniaceae only a small proportion of tree species is known wood-anatomically; the climbing and scandent species are fully unexplored in this respect. Anomalous structure has, however, been described for several genera outside Malesia. As far as known, the Malesian tree genera are wood-anatomically rather homogeneous: with simple, rarely also reticulate, perforations to the vessels, homogeneous rays, mainly paratracheal, aliform or confluent parenchyma, and fibres with simple to minutely bordered pits. Except for its unusually narrow rays, Dolichandrone spathacea from the mangrove swamps does not differ appreciably from the inland genera in its wood structure. Leaves. Very poorly known for the Malesian representatives. Diversity of stomatal type and indumentum (non-glandular and glandular hairs in a variety of forms) certainly deserves detailed comprehensive studies, which will probably yield important additional taxonomic characters. References: General surveys: SOLEREDER, Syst. Anat. Dicot. Stuttgart (1899, 1908); METCALFE & CHALK, Anat. Dicot. Oxford (1950). — Wood: JaNssonius, Mikrographie des Holzes 4 (1926) 721-753 (Dolichandrone, Oroxylum, Stereospermum); PANSHIN, Philip. J. Sc. 48 (1932) 143-205 (Dolichandrone); DrescuH, Mal. For. Rec. 15 (1941) 50 (Deplanchea, Dolichandrone, Pajanelia, Stereospermum); JANSSONIUS, Blumea 6 (1950) 450-452 (affinities); SEBASTINE, J. Ind. Bot. Soc. 34 (1955) 299-306 (Pajanelia); JuTTE, Nova Guinea 10 (1959) 242 (Deplanchea). — 1977] BIGNONIACEAE (van Steenis) 119 Leaves: SIEBERT, Ann. Mo. Bot. Gard. 35 (1948) 123-136 (glands); PALIWAL, Flora 159 (1970) 124-132 (stomata). — P. BAAs. Pollen morphology. Bignoniaceae have a long history of pollenmorphological study, starting with the pioneer studies of H. MOHL (1835). The first author to present a detailed pollen- morphological survey of the family, drawing attention to the taxonomical significance of the pollen characters, was URBAN (1916). He concluded that (1) any attempt to base the main sub- division of the family on pollen characters would group together taxonomically unrelated genera, (2) for generic delimitation pollen had limited value. Later studies by several authors have confirmed this (cf. BUURMAN, in the press). Inaperturate, tricolpate, stephanocolpate and pericolpate apertural types occur in the family. In a few genera tetrads are found. Size ranges between 25 um in Astianthus antisyphilitica and 100 um in Nyctocalos cuspidata, shape varies between suboblate and subprolate. Remarkable is the amount of variation which may occur within genera or even intraspecifically. In Stereospermum inaperturate, tricolpate and perisyncolpate pollen is found, while in Anemo- paegma longepetiolatum inaperturate, stephanocolpate, pericolpate grains as well as tetrads occur. In such cases sculpture affords more constant characters. The tricolpate type is dominant in the family and is found in all four tribes. It rarely shows well developed equatorial endoapertures. Instead, characteristically ruptured aperture membranes are present, especially in Tecomeae. Operculate colpi occur in Argylia. A subdivision of the tri- colpate type is possible on sculpture, which mostly varies between perforate and reticulate. In some genera very complex pollen grains are present, such as those of Nyctocalos (fig. 2). The tricolpate pollen grains in Bignoniaceae resemble those in Scrophulariaceae and Myopora- ceae. The similarities with Pedaliaceae, stressed by ERDTMAN (1952) refer to a rather specialized pollen type and may not reflect close affinity. References: MOHL, Ann. Sc. Nat. II, 3 (1835) 304-346; URBAN, Ber. Deut. Bot. Ges. 3 + (1916) 728-758; ERDTMAN, Pollen morphology and plant taxonomy. Angiosperms. Almqvist & Wiksell, Stockholm (1952) 73-74; BUURMAN, Contribution to the pollenmorphology of the Bignoniaceae, with special reference to the tricolpate type (in the press). — J. MULLER. Chemotaxonomy. Bignoniaceae share a number of biochemical tendencies with Verbenaceae, Labiatae, Scrophulariaceae and with several other families of WETTSTEIN’s Tubi- florae. Most of their outstanding chemical characters were already mentioned and discussed in my ‘Chemotaxonomie der Pflanzen’ vol. 3 (1964) 268-281, 645-646, to which the reader is referred. Much phytochemical information, however, became available only in more recent time. Recent results confirm the trends already apparent in 1963; they are summarized in the following pages. Chemical characters of Bignoniaceae may ultimately prove to be very useful in tracing inter- and intrafamiliar relationships. (1) Most members seem to produce and accumulate iridoid glucosides (formerly often called pseudoindicans). Since a long time Bignoniaceae are known to contain labile glycosidic bitter principles. Such a compound was isolated in 1888 from the bark and fruits of Catalpa bignonioides WALTER and called catalpin (name changed later to catalposide). The structure of catalposide was definitely established in 1962; it is an aucubin-type (C,-aglucone) ester glucoside and one of the first pseudoindicans for which clearcut structural and biogenetic relationships with iridodial and nepentalactone were demonstrated (hence the name iridoid glucosides for a presently very large group of constituents of dicotyledonous plants). Catalposide (tastes bitter) is an ester of p-hydroxybenzoic acid with catalpol (= 7,8-epoxy-aucubin). Catalpol and catalposide occur in all species of Catalpa (leaves, stems, fruits) and catalpol (= catalpinoside) was also isolated from barks of Paulownia tomentosa Steup. and P. fargesii FRANCH. where it occurs together with syringin (V. PLouvier, C. R. Ac. Sc. Paris 272D, 1971, 1443). Probably catalpol and catalposide occur in many more members of the family. In most recent times some related glucosides were isolated from Bignoniaceae. Vanilloyl-catalpol (= amphicoside) is a constituent of Amphicome emodi LinvL, and veratroyl-catalpol occurs in Tecomella undulata Seem. 5-Hydroxycatalpol (= macfadyenoside) was isolated from Macfadyena cynanchoides MORONG. All iridoid glucosides mentioned hitherto have structures based on the aucubin-derivative catalpol. The first non- aucubin-type glucoside described from Bignoniaceae is tecomoside with a C,o-aglucone; it was 120 FLORA MALESIANA [ser. I, vol. 8? isolated from Tecoma capensis LINDL. (A. BIANCO et al. Gazz. Chim. Ital. 105, 1975, 195). It is to be expected that much more iridoid glucosides will be detected in the family in future. (2) Some Bignoniaceae produce alkaloids. So far only pyridine-type and piperidine-type alkaloids with an iridoid Cy 9- or rarely Cy-skeleton were identified definitely in species belonging to this family. This fact strengthens the belief that the tendency to produce iridoid compounds is a very important character of Bignoniaceae. Thusfar simple iridoid alkaloids were described for species of Campsis (boschniakine), /ncarvillea (plantagonine, indicain), Tecoma (tecomanine, tecostidine, tecostanine, boschniakine, 4-noractinidine and several derivatives of skytanthine). The basic constituents of Amphicome (now reduced to Incarvillea), Newbouldia and other genera may belong to the same group of alkaloids. A recent review of the chemistry, distribution and systematic meaning of all presently known main groups of iridoid plant constitutents was published by S. ROSENDAL JENSEN et al. (Bot. Notis. 128, 1975, 148-180). (3) Many Bignoniaceae synthesize naphthaquinones and corresponding anthraquinones by prenylation of o-succinylbenzoic acid. This pathway to quinonoid naphthalene- and anthracene- type secondary metabolites is presently known from taxa belonging to Rubiaceae, Verbenaceae, Scrophulariaceae, Bignoniaceae and possibly Acanthaceae and Gesneriaceae. In roots, woods and barks of Bignoniaceae lapachol, lapachonone, «- and B-lapachone and dehydro-«-lapachone occur frequently. These monomeric naphthaquinonoid compounds are often accompanied and sometimes replaced by more complex dimeric constituents like tectol, guayacanine and guayine and by corresponding anthraquinones such as tectoquinone and 2-methyl-3-hydroxyanthra- quinone. Woods which contain appreciable amounts of these quinonoid compounds are more or less resistant to marine borers, white ants and Fungi. At the same time such woods may be the causes of skin irritations and of allergic skin diseases in man. Lapachol- and tectoquine-type substances are presently known from species of the genera Catalpa, Heterophragma, Kigelia, Paratecoma, Phyllarthron, Stereospermum, Tabebuia, Tecoma, Tecomella, and Zeyhera. R. H. THOMSON has reviewed the chemistry and distribution of quinones and related compounds in his book ‘Naturally occurring quinones’ (2nd ed. 1971). The phthalide catalpalactone from the wood of Catalpa bignonioides WALTER and C. ovata G. Don arises from the same pathway as lapachol and its congeners (H. INouyE et a/. Chem. Pharm. Bull. Tokyo 23, 1975, 384, 392, 2523). On the other hand it should be stressed that the red-coloured naphthaquinones of Boraginaceae (e.g. alkannin) which are structurally very similar to lapachol are produced along a totally different biosynthetic pathway (cf. E. LEISTNER, Chinoide Farbstoffe, Ber. Deut. Bot. Ges. 88, 1975, 163-178). (4) The “‘tannins’”’ mentioned for many Bignoniaceae in the older phytochemical literature (e.g. DEKKER, 1913) seem to be glycosides and esters of o-diphenolic compounds. Orobanchin (= verbascoside)-type ester glycosides were definitely demonstrated to occur in species of Campsis, Catalpa, Eccremocarpus and Pandorea. A review of this type of polyphenolic plant constituents which simulate true tannins in some respects is to be found in my ‘Chemotaxonomie der Pflanzen’ vol. 5 (1969) 250-252. Orobanchin yields a molecule of caffeic acid, 3,4-dihydroxy- phenylethanol, glucose and rhamnose each. Just as in most other families of Sympetalae true tannins are replaced in Bignoniaceae by more or less complex esters and glycosides of o-diphenolic cinnamic acid derivatives. Moreover, simple esters of caffeic acid and biosynthetically related derivatives of cinnamic and benzoic acids are present in large amounts in many Bignoniaceae. The recent investigations of V. B. PANDEY and B. DasGupTA with the bark of Tecomella undulata SEEM. (veratroylglucose = tecomin: Experientia 26, 1970, 1187) and of M. SUGUMARAN et al. with leaves of Tecoma stans H.B.K. (16 aromatic acids: Ind. J. Exper. Biol. 13, 1975, 93) exemplify this trend. p-Hydroxybenzoic acid is present as an ester in all species producing catalposide; probably this phenolic acid is rather ubiquitous in the family. The presence of appreciable amounts of hydroquinone (in living cells as the glucoside arbutin?) in leaves of Jacaranda mimosaefolia D. Don (S. SANKARA SUBRAMANIAN ef al. Phytochemistry 12, 1973, 220) might be connected with a strong tendency to produce and accumulate p-hydroxybenzoic acid; if this is actually the case hydroquinone (and arbutin?) may be detected in much more Bignoniaceae in future. Jacaranone, a quinonoid compound which exhibits antitumor and cytotoxic activity was recently isolated from leaves and twigs of Jacaranda caucana PITTIER (M. OGurRa et al. Lloydia 1977] BIGNONIACEAE (van Steenis) 121 39, 1976, 255); it seems to be derived from tyrosine and is chemically very similar to the Cornus quinol glucoside (= cornoside) which is also present in leaves of Digitalis purpurea (Bot. Notis. 128, 1975, 174). (5) According to J. B. HARBORNE (Phytochemistry 6, 1967, 1643) leaf flavonoid patterns of Bignoniaceae are close to those of Acanthaceae, Gesneriaceae, Labiatae and Scrophulariaceae. Features which support such a statement are the replacement of flavonols by flavones in many species, the relatively frequent occurrence of flavones with an unsubstituted B-ring (e.g. chryson, baicalein), of 6-hydroxylation of chrysin (baicalein), apigenin (scutellarein) and luteolin (6- hydroxyluteolin) and of 0-methylation of flavones. The latter trend is illustrated by Zeyhera tuberculosa BUR. ex VERLOT which contains 5,6,7-trimethoxyflavone and 5,6,7,8-tetramethoxy- flavone in leaves (J. P. KuTNEY & H. W. HANSSEN, Phytochemistry 10, 1971, 3298). The bitter principle of the fruits of Sparattosperma vernicosum Bur. & K. SCH. was shown by J. P. KUTNEY et al. (Phytochemistry 9, 1970, 1877) to be the 7-neohesperidoside of pinocembrin (= 2,3- dihydrochryson). (6) Free triterpenic acids occur in appreciable amounts in leaf waxes of many families of Tubiflorae (especially Verbenaceae, Labiatae and Plantaginaceae) and reJated orders. It is of interest in this respect that ursolic acid was isolated in recent time from leaves of Bignonia diversifolia H.B.K., Campsis radicans SEEM., Catalpa bignonioides WALTER, Heterophragma quadriloculare K. Scu., Jacaranda mimosaefolia D. Don (not definitely identified) and Paulownia tomentosa STEUD. The bark of Jacaranda mimosaefolia yielded lupenon. (7) Many members of Verbenaceae, Labiatae, Scrophulariaceae and Plantaginaceae replaced starch by stachyose-type oligosaccharides as storage carbohydrates. The same trend seems to exist in Bignoniaceae. Large amounts of stachyose occur in species of Catalpa (roots, wood, bark), Newbouldia laevis SEEM. (roots) and Paulownia tomentosa (stem). (8) Most representatives of Tubiflorae produce starch-free seeds which are rich in proteins and oils. The seed oils are often characterized by a high degree of unsaturation. In this respect Bignoniaceae conform to the rule. Their seeds generally contain 20-35% of oil. In some taxa oleic and (or) linolic and (or) linolenic acid are the only major fatty acids of the seed oils (e.g. species of Crescentia, Niedzwedzkia = Incarvillea, Paulownia and Stereospermum). In other taxa the ‘normal’ fatty acids are accompanied or replaced by large amounts of unusual fatty acids such as conjugated trienoic acids (species of Catalpa, Chilopsis, Jacaranda), Cy -keto-acids (Cuspidaria pterocarpa DC.), octadeca-trans-3,cis-9,cis-12,cis-15-tetraenoic acid (Tecoma stans H.B.K.) or hexadec-9-enoic and octadec-11l-enoic acid (Doxantha unguis-cati Miers). M. J. CHISHOLM and C. Y. Hopkins discussed the chemistry of seed oils of 11 species representing 4 tribes (Canad. J. Chem. 43, 1965, 2566). The preceding phytochemical picture places Bignoniaceae phytochemically very close to a number of families of Tubiflorae, especially Verbenaceae, Labiatae and Scrophulariaceae. Still other constituents are known from Bignoniaceae. Lack of acquaintance with their structures and (or) with their distribution, however, does not yet allow a systematic evaluation. Saponins, which seem to be rather widespread in the family but were never investigated in detail, belong to these chemical characters. The same holds for a number of phenolic compounds isolated in recent time, such as the lignans sesamin and paulownin from Paulownia tomentosa StEuD. and Phyl- larthron comorense DC., the dilignol (a lignan-type compound) zeyherol from Zeyhera digitalis HoeHNeE and the dihydroisocoumarins 6-methoxymellein, kigelin and 6-demethylkigelin from Kigelia pinnata DC. Concluding it may be stated that the intimate relationships between Bignoniaceae and Scrophu- lariaceae which are indicated by genera like Catalpa and Paulownia (often placed in Scrophu- lariaceae) are confirmed by phytochemistry. At the same time phytochemistry stresses a very close coherence of a core of families of Tubiflorae; this core comprises Scrophulariales sensu Cronouist (1968) and Lamiales sensu TAKHTAJAN (1969). — R. HEGNAUER. Uses. There are no outstanding qualities marking Bignoniaceae as useful plants, otherwise than ornamentals and these concern mostly the introduced species for which I refer to the special key and account at the end. There are magnificent native species notably of Tecomanthe but they have as yet not become in general use. Good roadside trees are Millingtonia hortensis and Spathodea campanulata. A highly esteemed 122 FLORA MALESIANA [ser. I, vol. 8? vegetable (/alab) with the Sundanese is Oroxylum indicum (flowers, buds, and very young pods). For re-afforestation and holding terraces on slopes the pioneer qualities of species of Rader- machera and Deplanchea might be useful. The timber is in general not valuable and in nature not available in sufficient quantity. The soft wood of Millingtonia hortensis was advertized as useful for tea-boxes. The only species yielding sizeable timber of good quality are: Fernandoa macroloba, Pajanelia longifolia, Radermachera gigantea, and the three species of Stereospermum, which all may be valuable for silviculture. Terminology. The shape of the corolla has been defined as tubular (fig. 4b, 10e) in which case there can be a distinction in a basal tube (on apex of which inside the stamens are inserted) as in fig. 4b; funnel-shaped (fig. 23d, 26c), hypocrateriform or salver-shaped (fig. 1a, 8c, 15) or infundibuliform (fig. 32a). Notes. Since my thesis (Rec. Trav. Bot. Néerl. 24, 1927, 787-1049), here always cited as ‘Thesis (1927)’, and subsequent revision in Bull. Jard. Bot. Btzg III, 10 (1928), I have remained always much interested in this family and have published some revisions and many notes precursory to the present treatment. I have to thank the late Mr. N. Y. SANDWITH (Kew) for namings of cul- tivated species, and Dr. A. L. GENTRY (St. Louis) for recent information on them, Dr. H. HEINE (Paris) for assistance in various matters, Mr. Michael GALorE (Lae) and Prof. E. J. H. CORNER (Cambridge) for photographs, Miss Christine BRIGHTON (Jodrell Lab., Kew) for the first chromo- some count in Tecomanthe, while I gratefully acknowledge precursory work performed by Mr. J. C. DEN HARTOG on Tecomanthe and Pandorea in 1969/70 at the Rijksherbarium where he worked as a graduate student. KEY TO THE GENERA! 1. Erect trees or shrubs. ee Peaved aitc. THOSUY I WHOLIS, 7 © oe See ss se es ea ee eee 6. Deplanchea 2. Leaves compound, almost always decussate. 3. Leaves 1-pinnate. 4. Leaf rachis with a sharp, keel-like ridge above. Leaflets 8-12 pairs. Capsules winged 12. Pajanelia 4. Leaf rachis not keeled above. Leaflets less than 6 pairs. Capsules not winged. 5. Corolla salver-shaped, white, nocturnal, the tube 12-18cm long, narrow-cylindric. Calyx spathaceous to the base, 3-6 cm. Leaves nigrescent, with domatia. A flat pseudoseptum developed parallel to the valves. Seeds rectangular, with corky wings as thick as the seed . 7. Dolichandrone 5. Corolla much shorter. No pseudoseptum. Seeds with hyaline wings. Leaves not nigrescent. 6. Mature leaflets serrate; crenate, or pmnatiid ~~ 5 ©." 2 <2 2". a>. ee 5. Tecoma 6. Mature leaflets entire. 7. Calyx regularly 5-lobed, rather thin, reticulately veined, short-hairy, 2-3 cm 11. Lamiodendron' 7. Calyx lobes unequal, often less than 5; calyx not thin and reticulate-veined. 8. Capsule rather flattened, with a flat septum, if terete with 10 ribs .. . . 10. Fernandoa 8. Capsule terete, with a terete septum, never with many ribs, sometimes one ridge on each valve. No domatia in Mal. spp. 9. Seeds thick, in one row in each cell, each fitting in deep notch in the septum. Valves rather ards Ovplesiince2 TOWS IN iGACH COM) ac ah as, aie eae ee ees 8. Stereospermum 9. Seeds thick, in many rows, the septum without notches. Valves thin. Ovules in many rows Be cach cede 5 Lah I are ei Gs SRE ae 9. Radermachera 3. Leaves 2—3(—-4)-pinnate. 10. Flowers fleshy, very coarse: calyx 2-4 cm, corolla wide, dirty violet or reddish purple to liver brown, 7-10 cm. Capsule flat, sword-shaped, 60-120cm. .......... 3. Oroxylum 10. Flowers not fleshy, much smaller or thinner, white or pink. Capsule smaller and of different shape. 11. Leaflets with domatia. Corolla white, salver-shaped, the tube 6-8 cm long, cylindric, 2 mm wide. Capsule compressed parallel to the septum, the latter parallel to the valves; dchicoanead septicidal. 11. Leaflets without domatia. Corolla tube otherwise, wider. Capsule cylindric, with a terete corky ane” septum attached perpendicular to the valves; dehiscence loculicidal. . . . . 9. Radermachera . Lianas. 12. Corolla salver-shaped, with a narrow cylindric tube, 5-19 cm long. Capsule flat, large, with a median lengthwise ridge. Septum parallel to the valves, dehiscence loculicid. . . . 1. Nyctocalos 12. Corolla not salver-shaped, tubular, infundibuliform or funnel-shaped. (1) Of Lamiodendron and Hieris the fruit is still unknown. The key given here is only to native and thoroughly naturalized species (only Tecoma stans). A tentative key to the cultivated species is added in an appendix on page 180. 1977] BIGNONIACEAE (van Steenis) 123 13. Corolla tube + geniculate above the basal tube, upper tube slightly curved and flattened with a prominent fold. Calyx c. '/, cm, below the very short lobes with a short spur-like tooth. Ovules 6-8 per cell, in + two rows. Leaflets 5, those of the lower pairsessile ...... 2. Hieris’ 13. Corolla tube not geniculate, without a fold. Ovules ~ per cell in several rows. Capsule loculicid, with boat-shaped valves. 14. Leaves digitate, with 3(—-5) leaflets. Corolla lobes valvate ......... 13. Neosepicaea 14. Leaves pinnate, with 1 or more pairs of leaflets. Corolla lobes imbricate. 15. Flowers in racemes, axillary, mostly on the old wood, rarely terminal (in a high-mountain sp.), the rachis at the base usually with several crowded pairs of minute sterile bracts. Calyx large, 15-40 mm, distinctly lobed. Corolla large, mostly red, 5-12 cm long incl. the lobes, not bearded in the mouth and upper part of the tube, but stuppose hairy at the insertion of the stamens (rim of the basal tube). Corolla lobes usually deltoid, very narrowly overlapping. Anther-cells eset bones. Vee . Se oo a ae 14. Tecomanthe 15. Flowers in thyrses, only occasionally depauperate into racemes, usually terminal, sometimes axillary or on old wood. No sterile bracts at base of peduncle. Calyx small, mostly stunted or very shortly lobed, 2-7 mm. Corolla incl. lobes up to c. 3'/, cm (in a single Australian species with large lobes c. 5 cm), usually white or yellowish with red dots or streaks inside lobes and mouth, often bearded in the mouth and upper part of the tube, sometimes with a hair-ring at the insertion of the anthers. Corolla lobes widely overlapping, often roundish. Anther-cells 1*/,- RE ey ae re Sof FT ae 15. Pandorea Tribe 1. Bignonieae B. & H. Gen. Pl. 2 (1876) 1027; K.Scu. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 209; BUREAU, Fl. Bras. 8, 2 (1896) 16. — Eubignonieae ENDL. Gen. Pl. (1839) 712. — Subtribe Eubignonieae DC. Rév. Bign., Bibl. Univ. Genéve (1838) 122; FENZL, Denkschr. K. Bay. Bot. Ges. Regensburg 3 (1841) 262; DC. Prod. 9 (1845) 143. Capsule septicid, the septum parallel to the valves. Frequently lianas, with tendrils, mostly in the neotropics. Note. Boyer (Hort. Maur. 218) and DC. (/.c.) included the two present tribes in tribe Bignonieae. 1. NYCTOCALOS T. & B. in Mig. J. Bot. Néerl. 1 (1862) 366; BUREAU, Mon. (1864) 52; Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 201; ibid. 3 (1867) 249; SEEM. J. Bot. 8 (1870) 147; K.Scu. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 219 (‘Nycticalos’); STEEN. Thesis (1927) 805; Bull. Jard. Bot. Btzg III, 10 (1928) 178; Acta Bot. Neerl. 2 (1953) 306; SANTISUK, Kew Bull. 28 (1973) 182. — Fig. 1-2. Lianas, without tendrils. Leaves pinnately 3-foliolate (in one extra-Mal. sp. I-pinnate with 5 leaflets). Leaflets herbaceous to chartaceous, entire, + elliptic, acuminate, with a few scattered crateriform glands along and spaced from the midrib underneath, above very fine punctate-pitted glandular. /nflorescence a short lateral or terminal 8—12-flowered raceme. Pedicels bracteolate. Flowers nocturnal, erect, fragrant; lobes widely imbricate. Calyx cup-shaped, truncate, with 5 horn- like teeth, the latter with glands on both sides. Corolla almost actinomorphic, salver-shaped, the long narrow basal tube dilated in the upper part, with 5 rounded unequal or subequal lobes. Stamens inserted in the throat 4 with or without a rudi- ment or 5, equal or 2 anterior ones sometimes longer, not exserted; anther-cells divergent, versatile; filaments glabrous and no hairs near their insertion. Disk annular, fleshy. Ovary with c~ rows of c ovules along the margins of the dissepi- ment; style long, filiform. Capsule large, flat, stalked and acuminate but with 124 FLORA MALESIANA [ser. I, vol. 8? v 4 —_ Fig. 1. Nyctocalos cuspidata (BL.) Miq. a. Habit, in flower, b. fruit, opened, showing seeds and dissepiment, x 1/, (after MIQUEL, 1867) parallel edges, valves with a central prominent rib; dissepiment thinnish, flat. Seeds flimsy winged, roundish. Distr. Three spp. in SE. Asia (Assam, Burma, Thailand, Yunnan) and West Malesia (Java, Borneo, Celebes, and the Philippines). Fig. 3. Ecol. Rare rain-forest lianas at low altitude. Taxon. The genus stands isolated in the Old World flora. In many aspects the small New World genus Tanaecium Sw. is very similar, but in this genus at least part of the leaflets carries a terminal tendril; besides its fruit is not flat, but said to be cylindric or quadrangular with convex woody valves. Still I believe it to be an ally in the New World. There is also a similar resemblance with the monotypic South American genus Macranthisiphon BUREAU but that has 2-ranked ovules and a more elongate, funnel-shaped corolla tube. For a moment I thought that Nyctocalos pinnata STEEN. (from Yunnan, only known in fruit, /.c. 1953, 306) might belong to Hieris, but the very numerous seeds defeat this, as far as H. curtisii is concerned. Hieris is, of course, the most intimate related genus, with the same punctate glands on the leaves and a deceptively similar calyx; its pollen is quite different. Too much importance has been ascribed to the structure of the androecium: 5 equal stamens to 4 didynamous; this varies as is explained under N. cuspidata. It led SEEMANN (/.c.) even to the inclusion of the Australian Hausmannia jucunda into the genus, which belongs in fact to the Tecomeae with quite different fruit and valvate corolla lobes. Nomencl. The generic name is female, being derived from the Greek nux. KEY TO THE SPECIES 1. Corolla tube c. 15-16 cm long, white. Calyx teeth horn-like, with a linear apex. Lateral petiolules oe Se a ne ae een ee er ree ae Fy ea ee Se 1. N. cuspidata 1. Corolla tube c. 5-6 cm long, tinged palish yellow suffused with pinkish shade. Calyx teeth triangular, acute.Lateral petiolules:2=3 mimi... «/i)4.. go ease See ee 2. N. brunfelsiiflora 1977] 1. Nyctocalos cuspidata (BL.) Mia. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 249, t. 8B (‘cuspidatum’); Merr. Philip. J. Sc. 1 (1906) Suppl. 237; C. B. Ros. ibid. 6 (1911) Bot. 211; MeERR. En. Philip. 3 (1923) 443; STEEN. Thesis (1927) 813, incl. var. oblongum STEEN.; Bull. Jard. Bot. Btzg III, 10 (1928) 180; SANTISUK, Kew Bull. 28 (1973) 183. — Tecoma cuspidata BL. Rumphia 4 (1849) 35. — N. macro- siphon T. & B. Cat. Hort. Bog. (1856) 155, nomen. —N. brunfelsiaeflorus (non T. & B.) Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 201, pro specim. celeb. — N. thomsonii Hook. f. Bot. Mag. 93 (1867) t. 5678; CLARKE, FI. Br. Ind. 4 (1884) 377; STEEN. Thesis (1927) 809; Bull. Jard. Bot. Btzg III, 10 (1928) 180; SANTISUK, Kew Bull. 28 (1973) 183. — Gelseminum cuspidatum O. K. Rev. Gen. Pl. 2 (1891) 479. — N. assamica Hook. f. ex. K.ScH. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 221, nomen, lapsus. — Fig. 1-2. Leaflets elliptic, ovate, obovate, or narrow oblong, rounded at base, acuminate to cuspidate at apex, 6-11(—18) by 31/,-7(-10) cm; petiole 3-5 cm; rachis 2!/,—31/,cm; petiolules !/,-1 cm. Pedicels c.+/,-1 cm. Calyx c. 6mm. Corolla in bud pale green, later creamy, the tube c. 15-19 cm; dilated part c. 3-6cm; lobes rounded, c. 1'/,-2 cm. Stamens 4, didynamous, with or without a filiform rudimentary Sth one; anther-cells 6-10 mm; con- nective with a filiform appendage 2-3 mm. Capsule 16-24 by 33/,-47/,cm. Seeds (including - Ped wings) rounded to obovate, 3-4 by 2!-3 cm. Fig. 2. Nyctocalos cuspidata (BL.) MiQ. Pollen grain, SEM x 500 (BS 10396). Distr. SE. Asia (Assam: Mikir & Gowhatty Hills) and Central Malesia: Philippines (Luzon, Polillo, Palawan, Biliran, Mindanao, Basilan) and Celebes (Manado, Kema, Bantaeng). The type was said to have been collected in the Moluccas by ZipPeL but this must be doubted. Fig. 3. Ecol. Lowland rain-forests. Notes. Hitherto importance was laid in keys on the difference between WN. cuspidata and N. thomsonii in that the former was described with 5 fertile stamens and the latter with 4 didynamous BIGNONIACEAE (van Steenis) 125 stamens and a filiform rudiment. This was suspi- cious as there were hardly any other differences. In material on spirit of Celebes specimens cultivated in Hort. Bog. I have now found flowers with didynamous stamens whether or not accompanied by a staminode. Furthermore, in RIEDEL s.n. from Manado and in BS 10396 from Polillo there are 5 perfect stamens; but even here in one flower the two anterior stamens were somewhat longer than the others. There is thus variation in the degree of tendency to zygomorphism. This is also visible in the difference in size of the corolla lobes of which one is mostly larger than the others. The taxono- mical implication is the reduction of N. thomsonii. In the Malesian specimens the calyx teeth appear to be somewhat longer and more horn-like than in the Assam specimens depicted by Hooker f. 2. Nyctocalos brunfelsiiflora T. & B. in Mig. J. Bot. Néerl. 1 (1862) 367 (brunfelsiaeflorus’); Mia. Choix (1863) t. vu; Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 201, excl. syn. et specim. celeb.; ibid. 3 (1867) 248, t. 8A; STEEN. Thesis (1927) 811; Bull. Jard. Bot. Btzg III, 10 (1928) 179 (‘brunfelsiiflorus’) ; Acta Bot. Neerl. 2 (1953) 306; BAck. & BAKH. f. Fl. Java 2 (1965) 536; SANTISUK, Kew Bull. 28 (1973) 183. — N. shanica MACGREGOR & W. W. SmiTH, Rec. Bot. Surv. India 4 (1911) 280; STEEN. Thesis (1927) 811; SANTISUK, Kew Bull. 28 (1973) 182, 183; Thai For. Bull. Bot. 8 (1974) 88. Leaflets elliptic, oblong, or obovate, acuminate to cuspidate, 7-13 by 3-6cm; petiole 4-7 cm; rachis 11/,-3cm; petiolules 1-2mm. Racemes 5-10-flowered. Pedicels 1—-11/, cm. Calyx 5-6 mm, suffused with reddish tinge in anthesis (ex coll.). Corolla whitish afterwards yellowish, later apically suffused with pink; tube 5—7 cm, the dilated part some 2-2'/,cm; lobes rounded to obovate or truncate, c. 2cm. Stamens (so far as known) 5. fertile, equal or subequal. Capsule 10-13 by 3-4 cm, Fig. 3. Range of the genus Nyctocalos T. & B.: N. brunfelsiiflora T. & B. (triangles), N. cuspidata (BL.) Mig. (dots), and N. pinnata Steen, (circle). 126 FLORA MALESIANA [ser. I, vol. 8? NIf ZS "J tA) 4 ps Y ea LE = 2 a8 Fig. 4. Hieris curtisii (RIDL.) STEEN. a. Habit, x 4/2, b. corolla, nat. size, c. LS of ovary, disk and recep- tacle, x 2!/,, d. calyx, x 2!/,,e. CS of ovary, x 7'/2, f. glands on ovary (HENDERSON S.n.). 1977] BIGNONIACEAE (van Steenis) 127 Distr. SE. Asia (Upper Burma: S. Shan States; Thailand: N. & SE.), in Malesia: Borneo (Sabah, near Kudat; Sarawak, near Niah), SW. Java (Wijnkoops Bay). Fig. 3. Ecol. Lowland rain-forests, even in Burma below 300 m. Uses. TEYSMANN found it a beautiful ornamen- tal; as far as nocturnally flowering plants can be. He could easily propagate it by marcottes. VAN HASSELT (in sched.) noted that in SW. Java crushed leaves are rubbed against head and stomach aches. Vern. Kakatjangan, S, SW. Java. Notes. N. shanica was distinguished by having glabrous anthers; these had in N. brunfelsiiflora been described and depicted as hairy to the base. This is, however, a lapsus: the base of the filaments and tube in the vicinity of the insertion is only dotted with small sessile granular glands. SANTISUK (l.c.) said that the pollen would be different from that of N. shanica, but I cannot accept this for specific distinction. The colour of N. shanica was described as white, but field data enumerated by SANTISUK mention also creamy white, buds pur- plish, and pale yellow flowers. Of N. brunfelsiiflora MIQUEL mentioned them to be pale pinkish ‘tirant légérement vers le jaune’, more purplish to anthesis. BACKER said: corolla at first white, afterwards yellowish; tube apically suffused with red. I do not ascribe much importance to these faint colour variances, especially nocturnal flowers often dis- colour with age. In comparing the scanty flowering material of both species it seems that in N. shanica the widen- ing of the corolla tube starts lower than in N. brun- felsiiflora, but I can not accept this for specific distinction. 2. HIERIS STEEN. Bull. Jard. Bot. Btzg III, 10 (1928) 279, f. 13. — Fig. 4. Slender woody twiner. Leaves 1-pinnate, with (1—)2(-3) pairs of entire leaflets. Racemes (axillary or) terminal. Pedicels bracteolate. Flowers scattered. Calyx cupular, articulated with the pedicel, short- or indistinctly 5-lobed, just below the margin with 5 spur-like, upcurved teeth at the base of each lobe, each tooth with a few glands on each side of its base. Corolla zygomorphic, the basal narrowed tube c. 1/, of its length, + geniculate with the upper */, which is widened, flattened, and gently curved; lobes subequal, suborbicular, finely capitate-glandular, papillose- hairy outside at base. Stamens 4, didynamous, inserted at the end of the basal tube, included, 5th rudimentary; anthers connivent in pairs, divaricate, connective apiculate. Disk entire, pulvinate-annular, fleshy. Ovary ovate, subterete, with 2 grooves, microscopically glandular; ovules in + two rows of 3-4 in each cell. Distr. Monotypic. Malesia: Malaysia: Penang I. (near village on north coast). Taxon. Outstanding by the few ovules and the geniculate tube of the corolla. The structure of the inflorescence seems to be racemose though flowers are articulated. Notes. Unfortunately the fruit and seed of this most interesting plant is unknown; from the cross- section of the ovary one might assume it to belong to tribe Bignonieae as the grooves of the ovary corres- pond with the edges of the septum, while furthermore the calyx structure is a replica of that of Nyctocalos, with which genus it seems closest related. In Nyctocalos, however, there is a very large number of ovules in each cell, which are very few in the Penang plant, like in the South American genus Memora. By the pinnate leaves Hieris shows resemblance with Nyctocalos pinnata STEEN. from Yunnan, which is unfor- tunately only known in fruit, but has abundant seeds in each cell. Mr J. MULLER told me (Febr. 1975) that the pollen of Hieris is not in the least resembling the showy pattern of Nyctocalos. 1. Hieris curtisii (RipL.) Steen. Bull. Jard. Bot. Btzg ILI, 10 (1928) 280, f. 13. — Tecoma curtisii Ruiw-. J. As. Soc. Str. Br. 49 (1908) 26. — Pandorea curtisii Ripc. Fl. Mal. Pen. 2 (1923) 553, f. 125; Steen. Nova Guinea 14 (1927) 301; Thesis (1927) 846, f. 4(2). — Fig. 4. Glabrous. Twigs terete, with very many small lenticels ; nodes with glands and a dark transversal line. Leaves (10—)15-20 cm pe petiole 2'/,-8 cm; rachis c. 4-5 cm; petiolules of lower pair of leaflets 4-12 mm, of upper pair (0-)1-2 mm. Leaflets herbaceous, slightly unequal-sided, ovate-oblong, long-acuminate, 5-8 by 2~3 cm; nerves 4-5 pairs; undersurface with scattered, small, rimmed-crateri- form glands, upper surface with microscopical pitted-punctate glands. Rachis rather densely flowered, microscopically puberulous, 2-7 cm; peduncle 2 cm, with barren bracts. Bracts linear, 2mm. Pedicels thin, 5-8 mm, with 1-2 minute bracteoles in the lower half. Ca/yx 6 mm, purplish, bluntly 5-ribbed, inside with dark red microscopical capitate-glandular hairs. Corolla 4'/,-5 cm, tube yellow, lobes whitish turning pale lilac, c. 1 cm @, 2 upper recurved, 3 lower erect; outside on transi- tion of tube and lobes scattered, rather large, brown red glands; basal tube 1'/,cm by 2mm, 128 FLORA MALESIANA [ser. I, vol. 8? Distr. Malesia: Penang I., see above. Ecol. Not well noted, 3 collections, all from 1898-1902; ff. June, July, Nov., and a cultivated specimen in Hort. Sing., Lawn 0, in Febr. near the insertion of the stamens scattered micro- scopical red-tipped, capitate-glandular hairs as in the calyx tube. Filaments glabrous, 12 and 14 mm; rudiment 5mm, with reflexed apex; anther-cells 2!/,mm, connective appendage linear, 1 mm. Ovary elliptic 1'/, by 1 mm; style 2 cm; stigmatic lobes elliptic, 1'/, mm. 3. OROXYLUM VENT. Dec. Gen. Nov. (1808) 8; K.ScuH. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 225 (‘Oroxylon’), I.c. 212 in clavi; STEEN. Thesis (1927) 816; Bull. Jard. Bot. Btzg III, 10 (1928) 181. — Calosanthes BL. Bijdr. (1826) 760; DC. Prod. 9 (1845) 177; BUREAU, Mon. (1864) 45, t. 9. — Hippoxylon RaFin. Sylv. Tellur. (1838) 78, nom. illeg. — Fig. 5, 7. Glabrous tree, robust in all its parts. Leaves 2—3(—4)-pinnate, all nodes with in sicco shrinking articulations; leaflets entire. Flowers very large, fetid, nocturnal, in large terminal racemes (by exception in a thyrse). Calyx persistent, not articulated, coriaceous, closed in bud, with a fine apical pore, later opening campanulate, + entire. Corolla funnel-shaped, lobes 5, subequal, imbricate in bud. Stamens 5, subequal, all fertile; anthers 2-celled, cells free, + parallel. Ovary with ~ rows of ovules in both cells. Capsule flat, very large, sword-shaped, linear; dissepiment flat, coriaceous. Seeds large in ~ rows; insertion linear, 1 cm wide. Distr. Probably monotypic. From Ceylon, the Deccan and Himalayas through SE. Asia (also in S. China: Yunnan, Kwangsi, Setchuan, Kweichou) and Malesia eastwards to the Philippines, Celebes, and Timor. Fig. 6. Ecol. Acharacteristic, short-lived nomad tree, nowhere gregarious, not in mature rain-forest but always in openings, secondary growths and thickets, rather indifferent to climate (also in teak forest under seasonal conditions) and soils, mostly below 1000 m, but in S. China up to 1375 m (HANDEL-MAZZETTI). Taxon. A second species has been described, raised from seed, collected by A. HENRY in Yunnan, in 1889, in the Arnold Arboretum, and named O. flavum REHDER (in Sargent, Trees & Shrubs 1, 1904, 193, t. 92). REHDER discriminated this from O. indicum chiefly by the sulphur yellow colour of the nearly symmetrical flowers, the plain not toothed or crisped corolla lobes, the splitting calyx, and the oblong leaflets. Several of these characters are not valid, especially if we take into consideration that REHDER’s plant was an unbranched sapling of 3 m high. In such saplings the leaves are always somewhat longer and thinner. The sulphur-yellow corolla is also rarely found in O. indicum from where I described it (1928) as var. citrinum STEEN. on a cultivated specimen at Bogor so annotated by J. J.SmirH (C.H.B. XV.K.B.IX- 11). The calyx is indeed different from that in O. indicum, in being thinner and having 5 faint ribs, but it is lobed by tearing, and this is also sometimes found in fruiting specimens of O. indicum. The corolla in O. flavum is also regular and somewhat smaller than usual but an examination of the type showed an exactly similar occurrence of hairs at the anther bases, the patelliform glands outside and the granular- glandular hairs inside. Remains the plain, entire corolla lobes, and an other character figured by REHDER but not mentioned by him, viz that the inflorescence is not a raceme, but a thyrse, with the lower stalks 5-flowered in double triads and the upper ones in simple triads, a situation never recorded or seen by me in O. indicum. 1 cannot well account for these two differences, but they could be due to cultivation; in on seperience tropical plants in hothouses often deviate from those in the wild, certainly in first-flowering saplings. 1. Oroxylum indicum (L.) Kurz, Fl. Burma 2 (1877) 237; CLARKE, Fl. Br. Ind. 4 (1884) 378; K. & V. Bijdr. Booms. 1 (1894) 66, Atlas 2 (1914) t. 358; RIDL. Fl. Mal. Pen. 2 (1923) 548; Merr. En. Philip. 3 (1923) 444; STEEN. Thesis (1927) 816; Bull. Jard. Bot. Btzg III, 10 (1928) 181, incl. var. citrinum STEEN. /.c. 184; OCHSE & BAKH. Ind. Groent. (1931) 77, f. 46; HAND.-Mazz. Symb. Sin. 7 (1936) 888; CORNER, Ways. Trees (1940) 166, Atlas t. 29. — Palega-pajaneli RHEEDE, Hort. Mal. 1 (1686) 77, t. 43. — Bignonia indica var. « LINNE, Sp. Pl. (1753) 625; Roxs. Fl. Ind. ed. Carey 3 (1832) 110. — Bignonia pentandra Lour. Fl. Coch. 2 (1790) 379. —Bignonia tripinnata NORONHA, Verh. Bat. Gen. 5 (1790) art. 4, p. 8, nomen. — Spathodea indica Pers. Syn. 2 (1807) 273. — Calosanthes indica BL. Bijdr. (1826) 760; WiGut, Ic. Pl. 4 (1850) t. 1337- 1338; Mig. Fl. Ind. Bat. 2 (1858) 752; BUREAU, Mon. (1864) 45, t. 9. — Bignonia quadripinnata Bianco, FI. Filip. (1837) 499, ed. 3, t. 219. — 1977] BIGNONIACEAE (van Steenis) 129 RRNA. te +o * 44 cy Sain fe nd “ne 4 Fig. 5. Oroxylum indicum (L.) Kurz, the ‘midnight horror’. Pole on left is a sapling that has flowered and fruited and is temporarily leafless. Different branches of the same tree may be in leaf, flower or fruit at the same time (Tg. Bukit, Sg. Sedili Ketchil, photogr. CorNER, June 1934), Hippoxylon indicum RarFin. Sylv. Tellur. (1838) 78, nom. illeg. — Arthrophyllum ceylanicum Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 27. — Arthro- phyllum reticulatum BL. ex Mia. |.c., et corr. 318. — Fig. 5, 7. Smallish, glabrous, sparingly branched, semi- deciduous tree, 6—-20(—27) m; trunk 10-40 cm @, with grey bark and large leaf-scars; twigs thick, (as the trunk at least at apex) pithy, later hollow, lenticellate, as the leaf-rachis. Leaves tufted at twig-ends, with a long petiole, '/,-2 m; leaflets long petioled, ovate to oblong, acuminate, 4-11(—15) by 3-9 cm, cuneate, rounded or reniform at the trip- linerved mostly oblique base, underneath distinctly reticulate-veined, with some scattered gland fields near the axils of the nerves and scattered, micros- copical scales. Innovations of leaves and racemes viscid. Racemes terminal, erect,'/,—1'/, m long, pith of twig-apex, peduncle and rachis partitioned. Pedicels long, with a few bracteoles in lower part, 2-4 cm. Calyx coriaceous, becoming almost woody in fruit, containing water in bud, truncate or irregularly shallow lobed by tearing, campanulate, brown or dirty-violet, 2-4 by 1'/,-2 cm. Corolla reddish purple to liver-brown to dirty violet out- side, dirty yellowish to pinkish inside, with a foxy stench, 7-10 cm long, the lobes subequal, in young buds strongly folded into a massive apex, + 130 crisped or undulate-crenate, in flower patent to + reflexed, outside with scattered patelliform glands, the lobes inside with dense, almost sessile capitate gland-hairs; basal tube wide, widened to base, c. 1'/,cm. Stamens inserted in throat, their base long hairy. Style 4-6 cm, dark violet as the sub- entire, large disk. Capsule pendent, 45-120 by 6-10 cm, valves flat, almost woody, finally black. Seeds incl. wings 5—9 by 2'/,—-4 cm. Distr. As the genus. Fig. 6. Fig. 6. Range of the genus Oroxylum VENT. Ecol. As the genus. As a consequence of its short-lived, short-sized nomad habit relatively rare in tracts with largely high primary forest, e.g. Borneo. In Malaya chiefly by villages and by rice- fields (CORNER). Also not particularly common in open but seasonally very dry tracts, such as the Lesser Sunda Is. and in teak forest largely confined to mixed forest stands. F/. Jan.—Dec., according to Koorpbers in Java at the start of the dry season; fr. July—May, the fruit remaining during the dry season on often leafless stems. CORNER (l.c.) gave a lively account of his obser- vations on this grotesque treelet. He remarked: “that each leaf develops as a unit and when it withers it breaks up gradually in regular order from the tip to the base: the leaflets fall off singly and the main stalk and its side-stalks break up at the joints: the bits accumulate round the base of the trunk like a collection of limb-bones, so that we may call it the ‘Broken Bones Plant’. The leaves are crowded near the end of the stem or its branches, and saplings, which remain unbranched until after their first flowering at a height of some 15 ft., look like gigantic umbrellas. When the saplings flower, the inflorescence develops from the apical bud and therefore further upward growth of the main stem is prevented. When the inflores- FLORA MALESIANA [ser. I, vol. 8? cence has finished flowering, the leaves below it fall off and the leafless stem is left as a pole with a few sabre-like pods dangling from its extremity: wherefore, we may call it the ‘Tree of Damocles’. Then, after 3-4 weeks ina leafless state, one or more lateral buds on the stem break out and grow into side-branches which, in due course, flower, fruit, shed their leaves and branch in their turn: and, thus, the big trees are constructed sympodially with open irregular crown and a few lanky ascend- ing limbs. Each branch seems to flower independ- ently of the others so that flowers, fruits and grow- ing twigs may be found on the same tree.” Flower biology. The flowers are nocturnal; on each raceme 1-2 flowers open on one night. According to CorRNER (i/.c.): “‘The corolla begins to open about 10 p.m., when the tumid, wrinkled lips part and the harsh odour escapes from them. By midnight, the lurid mouth gapes widely and is filled with stink. Before sunrise the corolla is detached and slips off over the long style. The flowers are pollinated by bats which are attracted by the smell and, holding to the fleshy corolla with the claws on their wings, thrust their noses into its throat: scratches, as of bats, can be seen on the fallen flowers of the ‘Midnight Horror’ next morning.” Fig. 7. Dispersal. The gauzy seeds slip out of the opened pods and flit away on the breeze with the jerky motion of a butterfly: so in noon-tide, we may call the tree the ‘Midday Marvel’ (Corner, /.c.). Uses. Popular with the Sundanese as a veget- able (/alab), fresh young leaves and flowers; even unripe capsule valves are eaten after being cooked (HASSKARL). In Bawean I. flowers are used against inflam- mation of the eyes. The bitter bark is chewed in Java for depurative purpose, especially after delivery. In Sarawak used for dyeing rattan of black shiny baskets. In West Java (Priangan) local people are con- vinced that the tree is a protection of the house against thieves, a superstition probably derived from the sword-like shape of the capsules. Vern. Midnight horror, E; Malaya: (beka) kampong, blalai, blonglai (kaya), bulai kaju, kain, merlai, poko bulai, Malacca; Sumatra: bolai, Minangkabau, habreng, Atjeh, '(ka)kapung, M, :. Sum., méngleo, Simalur I., abang-abang, Asahan; Borneo: gimurai, Sarawak, Bidayan name; Java: ki tongtorang, pongpor(r)ang, S, (kayu) lanang, mungli, wongli, wungli, J, pedangan, Japara, délég, kadjén djalér, kéok, padangan, raon, J (all once noted), bunglo, punglo, Md; h. lema kaba, kowa, Flores; Celebes: buli, Bantaeng, pohon padang, Manado; karu kadang, kayu, Kutai; Philippines: balilang-uak, pingka-pingka, p.-pinkahan, taghilau, Tag., abang-abang, P.Bis., abong-abong, Bis., sakayan-bakus, Tagb., balay-uak, bunglui, Sul., baliuag, bungoi, C.Bis., banloi, Sub., barangau, kamkampilan, Ilk., kampilan, Neg., maidbaid, Bik. BIGNONIACEAE (van Steenis) ee eeeUUUE a EEE ESS SEES UENSSE SEINE EERE ERS Fig. 7. Oroxylum indicum (L.) Kurz. a. Top of raceme with two open flowers, * '/, b. bat arriving on a flower, c. landed bat on a flower sucking honey (Old Bot. Garden, Univ. Malaya, Kuala Lumpur, photogr. SoEPADMO, Sept. 1973, 9.30 p.m.). 132 FLORA MALESIANA [ser. I, vol. 8? Fig. 8. Millingtonia hortensis L. f.a. Habit, x */2, b. detail of underside of leaflet, showing domatia, x 5, c. flower, nat. size, d. anthers, one in CS, e. capsule, x 1/,, f. seed, nat. size (a after WALLICH, c-d after BUREAU, b, e—~f SPANOGHE s.n. TIMOR). 1977] BIGNONIACEAE (van Steenis) 133 4. MILLINGTONIA LINNE f. Suppl. (1781) 45, 291, non DONN, 1807, nec Roxs. 1820; K.Scu. in E. & P. Nat. Pfi. Fam. 4, 3b (1894) 226, f. 89 j-k; STEEN. Thesis (1927) 825; Bull. Jard. Bot. Btzg III, 10 (1928) 186. — Nevrilis RAFIN. Sylv. Tellur. (1838) 138, nom. illeg. — Fig. 8. Medium-sized evergreen or deciduous tree with corky bark. Leaves 2-3-pinnate, with domatia. Thyrses lax, co-flowered, terminal. Flowers white, fragrant, noctur- nal. Calyx small, truncate-campanulate, + 5-lobed, persistent. Corolla salver- shaped, glabrous, with a very long, slender, basal tube at apex widening towards the limb, limb at base short funnel-shaped, zygomorphic (+ 2-lipped), 5-lobed, imbricate in bud. Stamens 4, didynamous, glabrous, inserted at the base of the widened part of the tube (throat), shortly exserted, no staminode; anthers with 1 fertile cell, the other spur-like, barren; connective dorsal, swollen. Disk cup- shaped, crenate. Capsule linear, compressed parallel to the septum, septicid- dehiscent, valves flat. Seeds ~, thinly discoid, winged. Distr. Monotypic. SE. Asia (India, Burma, Thailand, Indo-China, Yunnan) and Malesia: probably native, in E. Java, Madura and Kangean Is., Lesser Sunda Is. (Bali, Sumbawa, Sumba, Flores, Timor), and S. Celebes (SW. Peninsula; Muna I.), in many places also cultivated (Penang, Sumatra, Java, etc.) and in dry areas running wild, may be wild also in N. Malaya (Perlis and Kedah, Corner, /.c.). Fig. 9. Ecol. Lowland monsoon forest. 1. Millingtonia hortensis LiINNé f. Suppl. (1781) 291; Decne, Herb. Timor. (1835) 32; Span. Linnaea 15 (1841) 326; Mia. Fl. Ind. Bat. 2 (1858) 753; BUREAU, Mon. (1864) 45, t. 8; F.-VILL. Nov. App. (1880) 150, cult. Manila; CLARKE, FI. Br. Ind. 4 (1884) 377; K. & V. Booms. Java 1 (1894) 65; STEEN. Thesis (1927) 826; Bull. Jard. Bot. Btzg III, 10 (1928) 187; CorNER, Ways. Trees (1940) 165; Merr. J. Arn. Arb. 25 (1944) 316; BAcK. & BAKH. f. Fi. Java 2 (1965) 234. — Bignonia suberosa Roxs. Cor. Pl. 3 (1811) 11, t. 214, nom. illeg. — Big- nonia cicutaria MART. Denkschr. K. Ak. Wiss. Minchen 6, KI. Math. Phys. (1820) 153, t. D. — M. dubiosa SPAN. in Hook. Comp. Bot. Mag. 1 (1835) 348, nomen. — Nevrilis suberosa RAFIN. Sylv. Tellur. (1838) 138, nom. illeg. — Fig. 8. Evergreen (or deciduous?) tree, 5-25m, to 30 cm @; bark corky, very rough, cracking; twigs lenticellate. Mature leaves herbaceous, nearly glabrous, 3—S-jugate, lower pairs pinnate, up to 1 m; leaflets ovate-lanceolate, acuminate, sinuate or crenate, or entire, 2'/,-6 by 1'/,-3 cm; domatia haired. Thyrse erect, 10-40cm, puberulous, flowers fragrant, only few open at a time. Calyx 2-4 mm, teeth short, broad, obtuse, margin revo- Fig. 9. Range of Millingtonia L. f.; delimitation in Asia is slightly arbitrary. lute. Corolla tube 6-8 cm by 2mm, widened to mouth, limb 4-5cm @, lobes ovate, acute, out- side with crateriform glands, c. 1'/, cm. Filaments c. 10 and 14mm long; anthers 2mm, with a small appendage at the base. Style to 8 cm long. Stigmatic lobes ovate-acute, 1'/, mm. Capsule 30- 35 by 1'/,-1°/, cm. Seeds thin-discoid, 1'/,-3'/, by 1-1"/, cm including the wings. Distr. As the genus. Fig. 9. Ecol. A characteristic tree of regions subject to annual drought (‘monsoon flora’), companion of teak, fire-resistant by its thick corky bark and pro- fuse capacity of suckering from roots, below 750 m alt. In Timor common in Ziziphus stands (MEVER Drees). F/. Jan.—Sept., mostly June. Father ScumMurTz reported that it is in Flores not deciduous; flowers appear at the end of the dry season, before the first rains set in. Uses. The soft, even-grained timber was some- times advertized for tea-boxes but is not of high quality. Tree sometimes used for parks or road- sides, leaves as a poor substitute of opium in cigarettes, sometimes received from the opium factory in Java under the vernacular name gendje. 134 Vern. Indian cork tree, E, kurkboom, D, kahombu, M (Sum.), amfiunan, sékar pétak, sékar putih, J, karpoti, Kangean, kanongoh, Bali, kétangar, Sumba, takah, Dawang lang., Timor, ai katong inggar, takah, toka hau, Timor, ai kakassa, Tetun lang., Port. Timor, katangka, Bug., Makas- FLORA MALESIANA [ser. I, vol. 8? Notes. From Sumba the flowers have once been noted to be red (IBoET 264), never confirmed. MEER DreEES (Comm. For. Res. Inst. Bogor 33, 1951, 39) recorded that Millingtonia is deciduous in the driest regions of Timor, but the scant field notes do not confirm this. sar, Kaulolo, Muna I. Tribe 2. Tecomeae ENDL. Gen. Pl. (1839) 711; FENZL, Denkschr. K. Bay. Bot. Ges. Regensburg 3 (1841) 261; B. & H. Gen. Pl. 2 (1876) 1029; K.Scu. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 209; BuREAU, FI. Bras. 8, 2 (1897) 300. — Subtribe Catalpeae DC. Rév. Bign., Bibl. Univ. Genéve (1838) 123; Prod. 9 (1845) 203. Capsule loculicid, the septum attached transverse to the valves. Mostly trees or shrubs, rarely lianas, by exception with tendrils. 5. TECOMA Juss. Gen. (1789) 139; REHDER, Mitt. Deut. Dendr. Ges. 22 (1913) 262; Britton, Bull. Torr. Bot. Club 42 (1915) 372; URBAN in Fedde, Rep. 14 (1916) 313; MELcuior, Ber. Deut. Bot. Ges. 59 (1941) 18-31. — Stenolobium D.Don, Edinb. Phil. J. 9 (1823) 264; Seem. J. Bot. 1 (1863) 87; STEEN. Thesis (1927) 964; Bull. Jard. Bot. Btzg III, 10 (1928) 217. — Tecomaria Spacu, Hist. Nat. Vég. 9 (1840) 137; FENZL, Denkschr. K. Bay. Bot. Ges. Regensburg 3 (1841) 266; SEEM. J. Bot. 1 (1863) 19-23; SPRAGUE, FI. Cap. 4, 2 (1904) 448; STEEN. Thesis (1927) 831; Bull. Jard. Bot. Btzg III, 10 (1928) 193; BRUMMITT, Bull. Jard. Bot. Nat. Belg. 44 (1974) 421. Erect or scrambling shrubs or small trees. No gland fields at the nodes. Leaves l-pinnate, sometimes 1-jugate, or more rarely simple; leaf or leaflets incised or serrate, densely microscopically glandular-punctate and with hairy domatia under- neath. Pedicel with minute bracteoles. Flowers in terminal racemes or more often raceme-like thyrses, yellow, orangish or scarlet. Calyx cupular or campanulate, with 5, often apiculate deltoid lobes, glands scattered. Corolla tubular, with a short basal tube, funnel-shaped, + straight or + curved, widened to the mouth, lobes almost equal to unequal, imbricate in bud, minutely ciliate. Stamens 4, didyna- mous, exserted or included; anther-cells divergent, often finally standing out trans- versally, free or partly connate, sometimes hairy; Sth rudimentary. Disk cupular- pulvinate to shallowly cup-shaped. Ovary narrow cylindric or oblong, compressed, lepidote; ovules 2—-4-seriate in each cell. Capsule linear, -- compressed parallel to septum; valves smooth. Seeds hyaline-winged all round, insertion punctiform. Distr. Some dozen species in the New World, from extreme S. Arizona and S. Florida to northern Argentina, especially in the Andes, and one species in southern Africa. Some species widely cultivated in the tropics and subtropics and one of these locally naturalized in Malesia. Taxon. I can see not sufficient reason to keep Tecomaria generically apart from Tecoma. It often is said to differ by the exserted stamens and orange-red to scarlet flowers, but it has appeared that among the many taxa of South American Tecoma (Stenolobium) there are taxa with exserted stamens and in some the flowers are orangish. Inadvertently SEEMANN (J. Bot. 1, 1863, 19-23) also united them, but curiously later distinguished Stenolobium (l.c. 87). 1977] BIGNONIACEAE (van Steenis) 135 According to SPRAGUE (FI. Cap. 4, 2, 1904, 448) there are only two valid characters, viz the number of the rows of ovules in each cell (2 in Tecoma, 4 in Tecomaria) and the anthers. As to the first character, in a dozen American genera this number varies, from 2-4, 2-6 and in Tabebuia even from 2-many; its value seems therefore to be rather low. The second character holds: in American Tecoma the anther-cells are completely free causing them in full anthesis to stand often perpendicular to the filament; in Tecomaria they are connate in the upper 3rd or 4th part, so that they can not diverge so widely. The intimate relationship between Tecoma and Tecomaria is emphasized by a reputed fertile hybrid, x Tecoma smithii W. WATSON (see p. 118), between Tecoma velutina and Tecomaria capensis. As the differences between the genera coincide with the geographical disjunction I am prepared to 9 at sectional level and refer Tecomaria to Tecoma sect. Tecomaria (SPACH) ENDL. Gen. mt if From southern Africa 3 spp. were described but F. Wuirte (For. Fl. N. Rhod. 1962, 380) and BRUMMITT (Bull. Jard. Bot. Nat. Belg. 44, 1974, 419) distinguish only one. In South America a thorough revision probably will also show reduction to fewer variable and raciated species. 1. Tecoma stans (L.) H.B.K. Nov. Gen. Sp. 3 (1819) 144; DC. Prod. 9 (1845) 224; F.-VILL. Nov. App. (1880) 151; MerR. Fl. Manila (1912) 428; JOHNSTON, Proc. Cal. Ac. Sc. IV, 12, 2 (1924) 1166; SANDWITH in Pulle, Fl. Surinam 4, 2 (1938) 79; CORNER, Ways. Trees (1940) 170, f. 44, pl. 159; Back. & Baku. f. Fl. Java 2 (1965) 539; GENTRY, Ann. Mo. Bot. Gard. 60 (1973) 958, f. 38, with full synonymy. — Bignonia stans Linne, Sp. Pl. ed. 2 (1763) 871; Juss. Gen. (1789) 139; RECHINGER, Denkschr. K. Ak. Wiss. Wien 85 (1911) 356. — Stenolobium stans SEEM. Ann. Mag. Nat. Hist. 10 (1862) 30; J. Bot. 1 (1863) 88, incl. var. pinnata SEEM. type var.; BUREAU, Mon. (1864) t. 13; MerRrR. En. Philip. 3 (1923) 444; STEEN. Thesis (1927) 905; Bull. Jard. Bot. Btzg III, 10 (1928) 218. Shrub, up to c. 1-4 m. Leaflets 1-3 pairs (cult. sometimes 1-foliolate) lanceolate, acuminate, serrate, glabrous, but often along midrib laxly hairy, 3-10 by 1-4cm, cuneate at the base, no proper petiolules; petiole 2-5cm. Racemes glabrous, c. 5-15cm. Pedicels 5-10 mm. Calyx campanulate, 5-7 mm, usually with some im- pressed plate-shaped glands in middle part or upper half, lobes short-ciliate. Corolla yellow, 3'/,-5 cm, limb up to 3'/,cm @. Stamens included, anther-cells + pilose. Capsule acute, often lenti- cellate, 10-22 by '/,-*/, cm. Seeds (incl. wings) 2 by 1/, cm, inserted in two rows on the margins of the septum. Distr. From Florida through Central and South America to N. Argentina, widely cultivated in the tropics, also in Malesia, and sometimes run wild, naturalized e.g. in Tahiti, the Society Is. (Raiatea), and the Marquesas (Nunuhiva), often together with tree ferns and Gleichenia. Notes. Vegetatively a rather variable species. The normal form is with pinnate leaves, but some- times there are 3-foliolate and even simple leaves intermixed in one sheet. In Tahiti the specimens have 5—6 pairs of leaflets. (This may be var. multi- jugum R. E. Fries, Ark. Bot. 1, 1903, 401). In exico a sheet had almost entire leaflets (Sumi- CHRAST 1885). There is in America a form with underneath woolly-hairy leaflets: 7. stans var. velutina DC. (T. mollis H.B.K.), but the density of the tomentum varies considerably in degree and I am not very much in favour to recognize this; this is also the opinion of STANDLEY (Trees, Shrubs Mexico, 1926, 1319). JOHNSTON (J. Arn. Arb. 21, 1940, 264) said that in Mexico the normal-leaved form occurs in coastal regions, the incised-leaved form ininland places and the tomentose form south of these two, all three replacing, suggesting subspecific segregation. It is rather peculiar that, though the normal- leaved form is widely cultivated throughout Malesia, the only naturalized one is a fairly con- stant form with deeply incised leaflets, which seems to be rather rare in the Americas. var. incisa G. Don, Gen. Syst. 4 (1838) 224; J. K. MAHESWARI, Bull. Bot. Surv. India 3 (1961) 357. — T. incisa Sweet, Hort. Brit. ed. 1 (1827) 284, nomen. — T. stans var. apiifolium DC. Prod. 9 (1845) 224; Back. & BAKH. f. Fl. Java 2 (1965) 539. — Stenolobium stans var. apiifolium SEEM. J. Bot. 1 (1863) 89; STEEN. Thesis (1927) 906; Bull. Jard. Bot. Btzg III, 10 (1928) 218. — ? Stenolobium incisum RosE & STANDLEY, Contr. U.S. Nat. Herb. 16 (1913) 174. — ? T. stans var. angustatum REHDER, Mitt. Deut. Dendr. Ges. 24 (1915) 227. — T. incisa (ROsE & STANDLEY) JOHNSTON, J. Arn. Arb. 21 (1940) 264. Leaflets (2—3—)4(—5) pairs, very coarsely toothed to deeply pinnately incised, not rarely to the mid- rib, making acute-triangular lobing, 5-10 by 1-—2'/, cm (incl. teeth). Distr. Central America, widely cultivated in the tropics, also in Malesia: naturalized in Timor, Ternate, and SW. New Guinea (near Uta); also naturalized in the Concan and N. Kanara (TALBOT). Ecol. In Timor (Kupang and Baucau Plateau) characteristic for red calcareous soils and lime- stone, in the latter place gregarious in shrubberies, flowering already at an early age; 5-500 m. FI. mostly Aug.—Sept. (Oct.), fr. Oct.-Dec. Vern. Yellow bells, E, Malaya; ai funan, Tétu lang., E. Timor; dufa dufa, Ternate. 6. DEPLANCHEA VIEILLARD, Bull. Soc. Bot. Normandie 7 (1862) 96; BUREAU, Bull. Soc. Bot. Fr. 9 (1862) 164; Beauvis. Gen. Montrouz. (1901) 90; STEEN. Thesis (1927) 906, f. 8-9; 136 FLORA MALESIANA [ser. I, vol. 8? Bull. Jard. Bot. Btzg III, 10 (1928) 218, f. 2-3; Heine, Fl. Nouv.-Caléd. 7 (1976) 71, f. 16-17. — Diplanthera BANKS & SOL. ex R.BR. Prod. (1810) 448, non THouaRs, 1806, nec SCHRANK, 1819; SCHEFFER, Nat. Tijd. N. I. 31 (1870) 332; B. & H. Gen. Pl. 2 (1876) 1048; STEEN. Nova Guinea 14 (1927) 293. — Bulweria F.v.M. Fragm. 4 (1864) 147. — Fig. 10, 12-13. Fig. 10. Deplanchea bancana (SCHEFFER) STEEN. a. Flower, b. fruit on thickened rachis, c. opened capsule, seeds covering the septum, d. seed, all nat. size. — D. glabra (STEEN.) STEEN. e. Flower, nat. size, f. LS of ovary, enlarged. — D. tetraphylla (R.BrR.) F.v.M. g. FLOwEr, nat. size (a—-d C.H.B. XIII-J—65, e-f GIELLERUP 583, g after VAN STEENIS, 1927). Trees, with thick, pithy branches, robust in all parts. Leaves simple, in whorls of 3-4, + tufted to end of branches, entire, at base above with a few large crateriform or saucer-shaped glands, underneath often fine-punctiform dotted and sometimes with scattered larger crateriform glands; glabrous or with a yellow indument of simple hairs. Thyrses terminal, erect, a thick short rachis with crowded, horizontal, long-stalked triads or twice-forked triads. Flowers erect, showy, yellow, stalked. Calyx on a 2-3 mm high, obconical, solid hypanthium, articulate with the pedicel, closed in bud, with short lobes hairy at tip, in anthesis with 5 rather regular, acute lobes or tearing into 2-5 unequal, irregular lobes, inside fine-glandular, outside not rarely with few large crateriform glands, tip penicellate. Corolla imbricate in bud, lobes ciliate, zygomorphous to degree, hardly with a distinct basal tube, tube just or far exceeding the calyx. Stamens 4, didynamous, exserted, rarely a 5th rudiment, + erect or recurved to one side, together with a style inserted shortly 1977] BIGNONIACEAE (van Steenis) 137 above the base of tube, base capitate-glandular hairy; filaments ribbon-shaped; anther-cells free, wide-divergent. Disk annular, crenate. Ovary subsessile, glabrous, 2-celled, each cell with 2 closely placed placentas; style very long; stigma with 2 narrow lobes. Ovules o¥, in many rows. Capsule short-stalked, ellipsoid, with hard, boat-shaped valves, erect; septum flattened, lens-shaped, thick. Seeds very many, roundish, very thin hyaline-winged all around, punctate-inserted. Distr. Probably 5 spp., 1 in West Malesia, 2 in New Guinea (of which | sp. also in N. Australia and the other also in E. Borneo and Central Celebes), and 2 in New Caledonia. Fig. 11. Ecol. Rain-forests with preference for light and secondary forest, kerangas forest, others in woodland savannahs and invading grasslands, from sea-level to 1000 m. Notes. The much increased collections gave a better understanding in specific delimitation and varia- bility of characters, leading to reduction in the number of species. Especially the hairiness occurs to degree and is occasionally deviating; in occasional specimens of D. bancana the calyx may possess dense long hairs inside the calyx. For this reason I have reduced D. tubulosa STEEN. and D. coriacea STEEN. The Australian D. hirsuta BAILEY I have reduced tentatively to D. tetraphylla; I believe it to be a juvenile form which accounts for its sinuate leaf margin and occurrence of deviating phyllotaxis, decussate or whorls of 3. For brevity’s sake the characters mentioned in the key are not repeated in the descriptions. Specimens in fruit or in bud, or without corolla are difficult to identify. Affinity. Deplanchea has no affinity to other Old World genera. BUREAU (Mon. 1864, 51) compared it with the genus Delostoma from Andine South America with which it shares several characteristic features: thick twigs, terminal inflorescences, simple leaves, and boat-shaped fruit valves. Delostoma differs in having the valves said to be unequal, one flat, one boat-shaped, and further by triplinerved leaves, a regular, dentate (sometimes ‘double’) stunted calyx, and pink or violet flowers. KEY TO THE SPECIES 1. Corolla tubular, the tube + twice as long as the calyx, straight or slightly curved. Stamens and style erect, + straight in anthesis. Leaves in whorls of 3, underneath almost always very laxly hairy on midrib and nerves as is the petiole. Calyx lobes + equal, corolla lobes ditto : . 3. D. glabra 1. Corolla tube only for 2-5 mm exceeding the calyx, the limb distinctly zygomorphous, 2 lobes higher connate, patent and longer than the others. Stamens and style patent-curved over this lobe or recurved. Leaves in whorls of 3-4. 2. Branches of the thyrse triads, or flowers solitary. Pedicels 1'/,-2!/, cm long. Bud + cylindric in shape, rarely pear-shaped, often with 5 faint ribs below the lobes. Calyx with a few to several large crateri- form glands, the lobes + equal, in anthesis c. 1/,;-"/, as long as the tube. Corolla tube almost cylindric, c. 10 by 5 mm. Stamens c. 3-3!/, cm. Leaves in whorls of 3 (by exception 4), hairy underneath or glabrous bancana 2. Branches of the inflorescence often 2(—3) times forked. Pedicels !/,-2 cm. Buds pear- or spindle-shaped, or obovoid, smooth. Calyx without crateriform glands, distinctly widened + campanulate in anthesis, the lobes mostly unequally tearing, sometimes only 2 or 3, '/,-*/; as long as the tube in anthesis. Corolla tube widened almost from the base, c. 1-1!/, cm high, 1'/, cm wide at the mouth in anthesis. Stamens 4-4'/, cm. Leaves in whorls of (3—)4, always hairy underneath 2. D. tetraphylla Se wae hoe Soe Se, See? eee VY ee es ee A ee ey a ee emer Nees 2 es ge Noe Oe We ee ee pe ee . D. 1 bancana (SCHEFFER) STEEN. Thesis . Deplanchea orange, darker than corolla. Fruit 10-14 by 3'/, cm. (1927) 921, incl. var. glabra STEEN. I.c. 923; Bull. Seeds c. 3/,4cm @, incl. wings 3 by 2 cm. Distr. Jard. Bot. Btzg III, 10 (1928) 221, f. 2b, 3. — Diplanthera bancana Scuerrer, Nat. Tijd. N. I. 31 (1870) 334; Hassk. Flora 53 (1870) 219; CLARKE, Fl. Br. Ind. 4 (1884) 385; RipL. Fl. Mal. Pen. 2 (1923) 552. — D. coriacea Steen. Bull. Jard. Bot. Btzg III, 10 (1928) 224, f. 2c-e. — Fig. 10a-d. Small to large tree, 4-36 m; bole to 20 m; d.b.h. 15-150 cm, with small or larger buttresses; bark finely fissured, flaky, wood soft, white. Leaves chartaceous to coriaceous, obovate to elliptic, 9-34 by 5'/,-20 cm, apex rounded, rarely short-wide- acuminate; base cuneate to cordate; glabrous to yellow hairy in various degree, as is the thyrse; petiole 3-6 cm. Peduncle 5-20 cm; rachis 2-5 cm; primary lateral stalks of triads 2-4 cm. Calyx 12- 18 mm. Corolla tube inside at base densely capitate- glandular hairy on insertion of stamens. Anthers West Malesia: Sumatra (Palembang, Asahan, Bencoolen, Indragiri, Tapanuli), Riouw Is. (Karimon I., P. Temiang, P. Kedondong), Malaya (also Penang I.), Banka (common), Billiton, and Borneo. Fig. 11. Ecol. In primary and secondary forests, in Borneo not rare in heath forest, mostly on sandy soils, podsols and wet kerangas, slopes of podsol terraces, from sea-level up to 1000 m. F/. Jan.—Oct. Vern. Méndjanbing, méngkubéng, méngkubung, M (Banka), kayu chéndéru, Malacca, labu, Palem- bang, méngkubong, mértapa, P. Temiang, kayu si martim, baha, Batak, tui, M (P. Karimon, in error with Dolichandrone?), éndjabiengien, Billiton. Notes. A fairly variable plant. In addition to the yellow-tomentose or velutinous haired typical form as described by Scuerrer there occur glab- FLORA MALESIANA [ser. I, vol. 8? = Mh -. ; *! Ae, | | e Saeed : < Xe a aS e 24 fi Fig. 11. Range of the genus Deplanchea VIEILLARD: D. tetraphylla (R.BR.) F.v.M. (dots), D. glabra (STEEN.) STEEN. (triangles), D. bancana (SCHEFFER) STEEN. (line). In New Caledonia 2 endemic species. rous forms, but in degree, with few hairs, or the hairs still more reduced and confined to the lower part of the petiolar groove, or only to the axillary bud. There are also specimens of which the calyx is inside long pubescent with 1-seriate hairs, notably S$ 11989, 16427, 17591, 25411, SAN 32195, ANDERSON 8398, HALLIER B 2507, VAN NIEL 4019, : but they are hairy as the type or glabrous, or with small glabrous leaves, rounded leaves or with short acuminate tip. This is also the reason that I cannot maintain D. coriacea, as the leaf-base varies from cordate to rounded to cuneate, without correlation with other sets of characters. Also the number of crateriform glands varies and these are also found in other specimens. I will not exclude the possibility that in the field certain biotypes may be bound to certain soil types, but I see no possibility to defini- tions and formal recognition from the herbarium. In ANDERSON 8398 from Sarawak the leaves are in a whorl of 4. Fruits are extremely scarce in the herbarium. 2. Deplanchea tetraphylla (R.BR.) F.v.M. Second Syst. Cens. Austr. Pl. 1 (1889) 167; STEEN. Thesis (1927) 916, incl. var. novoguineensis STEEN. l.c. 917; Bull. Jard. Bot. Btzg III, 10 (1928) 220; Proc. R. Soc. Queensl. 41 (1929) 55; Webbia 8 (1952) 435. — Diplanthera tetraphylla R.Br. Prod. (1810) 449; Bru. Fl. Austr. 4 (1869) 540; SCHEFFER, Nat. Tijd. N. I. 31 (1870) 335; BANKS & SOL. Ill. Cook’s Voy. 2 (1901) 72, t. 229; BatLey, Queensl. Fl. (1901) 1137; Compr. Cat. Q. Pl. (1909) 368; WuiTE, Proc. R. Soc. Queens}. 34 (1922) 52; ibid. 38 (1927) 259; LANE-PooLe, For. Res. (1925) 137; STEEN. Nova Guinea 14 (1927) 293. — Bulweria nobilis- sima F.v.M. Fragm. 4 (1864) 147. — D. bulwerii F.v.M. ibid. 5 (1865) 72, (1866) 214. — ? Diplan- thera hirsuta F. M. BAtLey, Bot. Bull. Dep. Agric. Queensl. 14 (1896) 11; Queensl. Fl. (1901) 1137; Compr. Cat. Q. Pl. (1909) 368. — Faradaya chrysoclada K.Scu. & Laut. Nachtr. Fl. Schutzgeb. (1905) 370; Beer & H. J. Lam, Blumea 2 (1936) 225, cf. LAM & MEEUSE, Blumea 3 (1938) 201. — D. hirsuta (F. M. BAILEY) STEEN. Proc. R. Soc. Queensl. 41 (1929) 56. — Fig. 10g, 12-13. Tree, without buttresses, 4-25 m; d.b.h. 10 to over 100cm; bole 1-17m; bark grey or grey- brown, corky, furrowed and rectangular-fiaking; wood pale straw-coloured. Leaves chartaceous to coriaceous, usually obovate or oblong-obovate, underneath yellow-velutinous, base somewhat cuneate to stunted, exceptionally cordate, on the base above with 1-7 cup-shaped large glands, 11-23(-60) by 7-14(-30) cm; petiole 2!/,-5 cm. Peduncle 4-12 cm; rachis 3-9 cm; branches 2-71/, cm; pedicels 1-2 cm. Calyx 12-14 mm. Fruit 5-11 by c. 2!/, cm. Seeds incl. the wings 2 by 1*/, cm. Distr. NE. Queensland (incl. Thursday L., Fitzroy I.) and East Malesia: New Guinea and the Aru Is. (Trangan and Wokam Is.). Fig. 11. Ecol. Predominantly in the periodically dry belts of New Guinea, also in gallery forest, very rarely in rain-forest, almost confined to grassland and wooded savannahs and associated with Eucalyptus tereticornis (Central Distr.) or Mela- leuca, but also in mixed savannahs (Antidesma, Schefflera, palms, etc., at Merauke), not rarely common, also a pioneer in fired areas, from sea- level to c. 600 m, rarely at 1200 m (Mafulu). Fi. May-Oct., fr. July—Oct., often flowers and fruits together, but fruiting specimens very rare in the herbarium. Dwarf specimens may in places flower and fruit. C. J. Srerers (Verkenningsrapport Berari Komebwaller. Mimeo, Fak Fak, 1956, p. 6, 7, 10, phot. 2) reported D. tetraphylla from sandy soils, often inundated through an impervious subsoil in a heathy forest of Melaleuca. Field notes: style greenish yellow, filaments yellow, anthers brown. According to VAN ROYEN the flowers have a sourish-sweet scent and are eaten by ‘luries’, lorikeets (at Merauke). Uses. At Fak Fak the timber is used for prahus by the Papuans. Vern. Laargola, Trangan, Aru Is., kapul, M, Merauke, bas, Sorong, Mooi lang., tembako d’ora, Fak Fak, Ersania lang., pwan, Mumuni, Orokaiva lang., pakawa, Maipa, Mekeo lang. Notes. The phyllotaxis is not constant, several specimens have whorls of 3, reminding of D. bancana with which this species is closest related. In 3m high saplings (DOCTERS VAN LEEUWEN n. 38, cultivated in Hort. Bog. sub XVI.I.F.8) all leaves were opposite. Similarly saplings in Queensland may have opposite leaves and, moreover, narrow oblong to lanceolate leaves with wavy, even toothed margin (L. S. SmitH 12382). Such plants were described as D. hirsuta and may precociously flower; also very small normal-leaved specimens may flower, obviously at an early age, in New Guinea, possibly stimulated by open, pyrogenous habitat. Such specimens may also sucker. 3. Deplanchea glabra (STEEN.) STEEN. Thesis (1927) 919, f. 8f, 1; Bull. Jard. Bot. Btzg Ili, 10 (1928) 225. — Diplanthera glabra STEEN. Nova Guinea 14 (1927) 293. — D. tubulosa STEEN. Thesis (1927) 926, f. 8g, k, m; Bull. Jard. Bot. Btzg III, 10 (1928) 226. — Fig. 10e-f. Tree, 11/,-22 m; d.b.h. 12-60 cm; bole 3-12m; bark grey, scaly; mostly + glabrous in all its parts. Leaves obovate-oblong to elliptic-oblong, coria- ceous, usually very laxly haired on the midrib (and nerves) below, very rarely yellow short-velutinous on inflorescence, midrib, nerves and petiole, fine dark-dotted beneath, 9-40 by 4'/,-25 cm, rarely with some scattered larger glands; base rounded to BIGNONIACEAE (van Steenis) Fig. 12. Deplanchea tetraphylla (R.Br.) F.v.M. Inflorescence from above, capsules, partly opened, showing dissepiment (fl. photogr. HOOGLAND 4249; fr. photogr. WomERSLEY, 1956, Sogeri). 140 FLORA MALESIANA [ser. I, vol. 8? Fig. 13. Rather young tree of Deplanchea tetraphylla (R.BR.) F.v.M. in Bot. Garden Lae (photogr. M. GALORE). 1977] BIGNONIACEAE (van Steenis) 141 cuneate or subcordate, apex rounded; petiole 21/.-7 cm. Peduncle 3-7 cm; rachis 1-11 cm; triads 1/,-2 cm stalked; pedicels 4-15 mm. Bud spindle- shaped to obovoid. Calyx glabrous, very rarely haired on the mid-sepaline ribs, pustular towards apex but the pustules hardly ever opening as crateriform glands, 15-17 mm long, lobes 5 sub- equal, 3-6 by 2'/.-5 mm, c. 21/,-3'/, times as short as the tube. Corolla tube inside near the stamens capitate-glandular hairy. Placentas in each cell 2, nearly confluent. Capsule 6-9 by c.2-2'/, cm. Seeds incl. wings 11/, by 1 cm. Distr. Malesia rather common in North New Guinea in the vicinity of Hollandia and Mt Cyclops, also found twice in Central Celebes (Malili, Tobela, Palopo) and once in E. Borneo (Mt Nijapa, Kelai R., Berau). Fig. 11. It is remarkable that the species is obviously never found in other parts of New Guinea. Ecol. Both in high forest and in savannah, but flowering and fruiting already when small, 10-700 (—1000) m. Field notes: pedicels red, filaments and style yellowish green, anthers orange, glands on leaf- base orange. No buttresses. Fi. fr. March—Oct. Vern. Celebes: momo, Malili, kKalambutoh, Toradja. Notes. Although this species is usually almost glabrous, except for some lax hairs on the midrib beneath, KOSTERMANS & SOEGENG 444, from Hol- landia, has very hairy inflorescences, midrib and petioles, so that obviously the indumentum may vary as it does in D. bancana. Also the single Bornean specimen known so far (KOSTERMANS 21491) is similarly hairy; it was found on a mountain ridge at 1000 m alt. The two specimens from Central Celebes I refer to this species, although one is in fruit and the other in bud, because: the calyx is + regularly lobed, with a few pustules but without crateriform more commonly in pyrogenous _ grassland (Gleichenia-Ischaemum), aS a _ pioneer, often glands and the small pod and seed do not match D. bancana; both spp. have leaves in whorls of 3. 7. DOLICHANDRONE (FENZL) SEEM. [Ann. Mag. Nat. Hist. III, 10 (1862) 31, nomen; J. Bot. 1 (1863) 226, nomen] J. Bot. 8 (1870) 379, nom. cons.; K.SCH. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 240, f. 92B-D; SpraGuE, Kew Bull. (1919) 303; STEEN. Thesis (1927) 928; Bull. Jard. Bot. Btzg III, 10 (1928) 227. — Pongelia RaFIN. Sylv. Tellur. (1838) 78, nom. rejic. — Dolichandra sect. Dolichandrone FENZL, Denkschr. Bay. Bot. Ges. Regensb. 3 (1841) 265. — Fig. 15-16. Trees with 1l-pinnate leaves (or scattered simple leaves, extra-Mal.), leaflets entire (or serrulate, extra-Mal.). Flowers in few-flowered terminal racemes, salver- shaped, white, fragrant, nocturnal. Calyx not articulate, closed in bud, later spathaceous, caducous. Basal tube of corolla long, narrow-cylindric, upper part inflated, lobes mostly crisped. Stamens 4, didynamous, 5th rudimentary, inserted at the throat; anther-cells divergent. Disk annular. Ovules ~ in 4-6 rows, inserted on 2 placentas in each cell, on the septum. Capsule elongate subcylindric to + compressed, septum very narrow, false septum very broad and parallel with the valves. Seeds hyaline-winged, in the Mal. sp. rectangular with thick corky wings. Distr. Spp. 9, one in tropical E. Africa, 4 in tropical SE. Asia, 3 in tropical N. Australia, and one ranging widely from Malabar through Indo-Malesia to New Caledonia. Fig. 14. Ecol. Allinland species, except the wide-ranging D. spathacea which is a back-mangrove tree. Typif. In the Code and in Ind. Gen. D. spathacea (L.f.) K.Scu. has been accepted as the type, following SEEMANN (J. Bot. 1, 1863, 226). However, the lectotype must be chosen from the original materials incorporated by Fenzt in Dolichandra sect. Dolichandrone, elevated by SEEMANN to generic rank. He referred to Spathodea b of ENDLICHER and Spathodea R.Br. Under the first reference D. spathacea is not represented, at most a Dolichandrone represented by Bignonia spathacea (non L.) sens. Roxsp. Corom. t. 144, a wrong identification for D. falcata (WALL. ex DC.) Seem. (cf. SPRAGUE, Kew Bull. 1919, 308). This then must be the type of the genus. Taxon. The Australian species are in habit very different from the African and Indo-Malesian ones: characteristic shrubs or small trees of xerophytic habit and xeromorphous structure, the leaves sometimes obviously not decussate, coriaceous, sometimes entire, with fine parallel ascending veins, or leaflets even needle-like. In addition I find the pods terete and the pseudoseptum not flat, but irregularly, corky swollen with impressions of the seeds. According to URBAN (Ber. Deut. Bot. Ges. 34, 1916, 755) these species would also be different in pollen from the other species. I have arranged them in subg. Coriaceae STEEN. es 1927, 931, f. 10). They should probably be better arranged in a separate section, rather than in a subgenus. 142 FLORA MALESIANA [ser. I, vol. 8? Fig. 14. Range of the genus Dolichandrone (FENZL) SEEM. and its species, the largest range being that of D. spathacea (L. f.) K.ScH., a mangrove tree with buoyant seeds. Three spp. are distinguished; D. filiformis (DC.) F.v.M. is a fairly constant one, with 3-5 filiform leaflets, but D. heterophylla (R.BR.) F.v.M. is very variable, with simple to pinnate leaves (3-7 leaflets) also varying in width, whereas D. alternifolia (R.BR.) SEEM. with ovate, simple leaflets shows a tendency to split the leaf. Already SEEMANN (J. Bot. 8, 1870, 382) remarked that the latter two probably belong to one variable species, which opinion I now tend to share; he accepted the epithet heterophylla. 1. Dolichandrone spathacea (L. f.) K.Scu. FI. Kais. Wilh. Land (1889) 123; Merr. Fl. Manila (1912) 429; Int. Herb. Amb. (1917) 469; Sp. Blanc. (1918) 349; SPRAGUE, Kew Bull. (1919) 304; STEEN. Thesis (1927) 937; Bull. Jard. Bot. Btzg III, 10 (1928) 227; C. T. Wuite, J. Arn. Arb. 10 (1929) 265; MerRR. Comm. Lour. (1935) 355; CORNER, Ways. Trees (1940) 163, Atlas pl. 26-27; HEINE, Fl. Nouv.-Caléd. 7 (1976) 81, pl. 18. — WNiir Pongelion RHEEDE, Hort. Mal. 6 (1686) 53, t. 29. — Lignum equinum RumpH. Herb. Amb. 3 (1750) 73, t. 46. — Bignonia spathacea L. f. Suppl. (1781) 283; REtTz. Obs. Bot. 5 (1788) 5; BLANco, FI. Filip. (1837) 499. — Bignonia longissima Lour. FI. Coch. (1790) 380, nom. illeg., non JAcQ. 1760. — Bignonia javanica THUNB. Mus. Nat. Ac. Upps. 17 (1794) 150, nomen; Fl. Ceil. (1825) 7, nomen, cf. STEEN. Blumea 6 (1950) 359. — Spathodea longi- flora VENT. Choix (1803) 40; SPAN. Linnaea 15 (1841) 326. — Spathodea rheedii SPRENG. Syst. 2 (1825) 835, quoad syn.; WALL. Cat. (1832) n. 6516; DC. Prod. 9 (1845) 206; Mia. FI. Ind. Bat. 2 (1858) 754. — Pongelia longiflora RaFin. Sylv. Tellur. (1838) 79. — Bignonia longiflora WILLD. ex DC. Prod. 9 (1845) 206. — Spathodea loureiriana DC. l.c. 209. — Spathodea luzonica BLANCO, FI. Filip. ed. 2 (1845) 350; ed. 3, 2 (1878) 284, t. 242. — Spathodea diepenhorstii Miaq. Fl. Ind. Bat. 2 (1858) 754. — D. rheedii SEEM. J. Bot. 8 (1870) 380; Kurz, Fl. Burma 2 (1877) 234; CLARKE, FI. Br. Ind. 4 (1884) 379; K. & V. Bijdr. (1894) 69; Gams -e, J. As. Soc. Beng. 74, ii (1905) 377; Rip. Fl. Mal. Pen. 2 (1923) 549. — D. longissima K.Scu. in E. & P. Nat. Pfi. Fam. 4, 3b (1894) 240. — Fig. 15-16. For fuller references see STEEN. (1928). Evergreen, glabrous tree, 5-20 m; 10-40cm @; wood soft, white. Leaves usually 3-4-jugate, 15-35 cm, stalked, in the herbarium nigrescent as all other parts; young parts + viscid, young leaves slightly pinkish in the field (CORNER); leaflets thin, ovate-oblong to lanceolate, unequal-sided, entire, long-tipped (in seedlings sometimes toothed), 6-16 by 3-7cm, underneath with hairy domatia. Racemes 2-8-flowered. Rachis 2-3cm. Bracts caducous. Bracteoles 0. Pedicels 2-4 cm. Flowers not articulated. Calyx conical, coriaceous, usually arcuate, beaked, circumscissile caducous, with many microscopical glands and a field with large crateriform glands at apex, 3—6(—8'/,) cm. Corolla tube 12-18 cm long, the mouth 7-12 cm @; basal tube 9-12 cm, gradually funnel-shaped expanded above the throat for 4cm; lobes 5, broad, sub- equal, frilled round the edge, with large glands, 2'/,-3 cm. Stamens not exserted. Style exserted. Capsule flattened-cylindrical, + ribbed, straight or + arcuate, or twisted, tipped, 25-70 by 2-3 cm; valves hard leathery, pseudoseptum flattish, hard corky, c. 11/,-13/,cm wide. Seeds dark grey, rectangular, in many rows, 12-18 by 68mm including the thick corky wings; attachment a fine line, 8-10 mm long. Distr. From the coast of Malabar throughout tropical SE. Asia and the whole of Malesia to New Guinea, Micronesia (W. Carolines: Korror; Yap: Tomil I.), the Solomons, the New Hebrides and New Caledonia, not found in Australia and Poly- nesia. Fig. 14. Ecol. Confined to the back-mangrove and banks of tidal rivers and estuaries. RIDLEY (Kew Bull. 1910, 203; J. As. Soc. Str. Br. 59, 1911, 40, 146) BIGNONIACEAE (van Steenis) 143 Fig. 15. Dolichandrone spathacea (L. f.) K.Scu. Flowers and twigs in bud and fruit, magnification */;; upper capsules opened (photogr. CORNER). recorded it common in low-lying rice-fields near Kanga village, Lower Siam, as the predominant tree, which he ascribed as “‘relics of the time when this whole country was a tidal swamp, gradually filled up after the disappearance of the sea”. Other seashore plants were also found in these paddy fields, such as Euphorbia atoto. This inland occur- rence is also stressed by Corner (/.c. 164) who found it “frequently in coastal rice-fields; in Perlis it is indeed a feature of the country; also in North Kedah, as soon as one reaches Kodiang it attracts attention, standing in the paddies as an upright poplar and flanks the roads which lead to Kangar and Singgora. Old tree trunks are massive and fluted at the base, the crown tapering upward. The old, opened twisted pods remain for a long time on the tree.” Brass found it very abundant in Daru I. (S. New Guinea), while K. J. Wire found almost pure stands in Umboi I. (Morobe Distr.) in swamps behind the mangrove. He recorded it also from freshwater swamps in the Markham Valley. At the Bogor Botanic Gardens it is successfully cultivated in freshwater. The calyx is filled with water in bud. The very young inflorescence and developed ovary is often slightly glossy varnished in the herbarium, similarly as is found in Radermachera, certainly by the exu- date of glands which are found at the apex of the calyx. Fl. fr. Jan.—Dec., flowers and fruits not rarely found together. Koorpers (1894 /.c.) and HEYNE (Nutt. Pl. 1371) say that in the dry season it may be at times nearly leafless fruiting in Central and East Java. 144 FLORA MALESIANA [ser. I. vol. 8? Fig. 16. Trees of Dolichandrone spathacea (L. f.) K.Scu. in the coastal rice-fields of Perlis where it is a feature of the country (photogr. CORNER). Pollination. Flowers open at dusk and drop before sunrise; they must be pollinated by hawk- moths with very long tongues to attain the honey. In each inflorescence one flower is open at a time (CORNER). Dispersal. The corky seeds float readily and must be dispersed by seawater. In this respect it is strange that the species is not found in northern Australia and Polynesia. Uses. Of little use other than fire-wood; in N. Borneo a collector deemed the wood useful for making clogs and matches; in the Carolines (Koror I.) leaves and fruit are said to be used as a sub- stitute for betel leaves in chewing. HEYNE (Nutt. Pl. 1371) said that the wood is not durable, but light and easy to work for small things in the house; pieces of branches are sometimes used for floats of fishing nets in East Java and the Karimon Djawa Is.; in the Minahassa it is used for scabbards, in Madura I. for masks for the topéng. In Madura a cold concoction of the leaves is also used against mouth sprew. RUMPHIUs said that in Ambon twigs of lignum equinum (translation of kaju kuda) were used for making hedges. Vern. Malaya: poko kulo, tuj, M; Sumatra: tuwé-éj, Atjeh, kudo-kudo uwi, Simalur, kuda kuda, Pariaman, ki arak, Palembang; Java: kaju or ki djaran, M, djarang, S, djaram, djaran pélok, djaranan, kadjéng kapal, kaju pélok, kapal, J, kadju djharan, kaju djaran binék, Md; Borneo: kélaju, tuwi, Kutai, toi, tui, Brunei, Bajau lang., towi, Kedayan; Celebes: fojet, kaju pélumping, sangi, tomana, Minahassa; Talaud Is.: sansarangi; Alor: bombila; Ternate: djodjamé; Tidore: djamé; Ambon: kaju kuda, kati kati; Philip.: tua, tui, Tag., pata, Ilk., tafgas, Tagb., tanhas, C. Bis., tanghas, P.Bis., tewi, Mbo., tiwi, Tag., Bik., C.Bis. ; New Guinea: rie, Holtekang, Wembie lang., pide, pier, S. New Guinea, Asmat lang., aisumbu, Mani- kiang lang., asember tiy, Oransbari, Hatam lang., daud, Cape Vogel, Wanigela; New Britain: /atiu, W. Nakanai, favituviti, Gazelle Pen., Boava lang.; Solomons: kwae kwaele, Guadalcanal, kwe kweale, Malaita, kwe’ekwe’eali, Kolombangara, ririge, Small Nggela, Kwara’ae lang. Note. Especially leaflets of saplings may, at times, show some serrulations on the margin and may also be slightly hairy. 1977] BIGNONIACEAE (van Steenis) 145 hh EEE 8. STEREOSPERMUM Cuam. Linnaea 7 (1832) 720; A.DC. Prod. 9 (1845) 210; B. & H. Gen. Pl. 2 (1875) 104, pro sect. Eustereospermum ; STEEN. Thesis (1927) 946; Bull. Jard. Bot. Btzg III, 10 (1928) 233; CHATTERJEE, Bull. Bot. Soc. Beng. 2 (1948) 68. — Hieranthes RAFIN. Sylv. Tellur. (1838) 79. — Dipterosperma HASSsK. Flora 25, 2 (1842) Beibl. 1, p. 28; Cat. Hort. Bog. (1844) 152; Pl. Jav. Rar. (1848) 507. — Fig. 17, 19. Fig. 17. Stereospermum personatum (HASSK.) CHATTERJEE. a. Habit, b. capsule, both x */,, c. seed, nat. size (a after WicHT, Ic. 4, t. 1341, b-c BeuseKom & GEESINK 3662). 146 FLORA MALESIANA [ser. I, vol. 87 Rather tall, deciduous trees. Leaves 1-pinnate; leaflets a few pairs, diminishing in size downwards, underneath usually with flat, dish- or cup-shaped glands or glandular spots; no domatia (in Mal.). Thyrses well-branched, paniculiform, mostly terminal, or on old wood. Flowers fragrant (in Mal.). Calyx usually short-lobed. Corolla infundibuliform; basal tube mostly concealed in the calyx, upper part usually funnel-shaped; mouth bilabiate, upper lip 2-, lower 3-cleft, lobes subequal, rounded, crisped, toothed or laciniate. Stamens 4, didynamous, included, 5th rudimentary; anthers glabrous, cells divergent. Disk cupular to annular. Ovary cells each with 2 rows of many ovules. Capsule long linear, terete, mostly twisted, usually 4-angular in section; septum thick, corky, terete, with alternating notches to fit the thick seeds which appear in two rows; valves coriaceous. Seeds o, thick, trigonous, wedge-shaped, with a cross-groove, on both sides thinly winged; coty- ledons folded, 2-lobed, radicle straight. Distr. Over a dozen spp., in tropical Africa and Madagascar, in SE. Asia as far east as Yunnan, in Malesia: 2 spp. in Malaya, possibly also in Sumatra, and a doubtfully indigenous record of a third in East Java. Fig. 18. Ecol. Largely confined to regions subject to a seasonal climate, all in the lowlands, in everwet rain- forest obviously deciduous and flowering after a dry spell. Note. Besides the 3 Malesian spp. distinguished here, P. Dop mentioned S. cylindricum PIERRE from Malaya (Fl. Gén. I.-C. 4, 1930, 582), but this must rest on an error as that species is only known from Thailand and Indo-China (cf. SANTISUK, Thai For. Bull. Bot. 8, 1974, 22). KEY TO THE SPECIES 1. Inflorescence (incl. flowers) viscid-hairy by patent capitate-glandular hairs. Leaves not glabrous. eta tube gradually funnel-shaped widened, straight, Stamens glabrous at base. Capsule not 4-ridged. 2. Leaflets 0-5 mm stalked, base cuneate-attenuate. Capsule faintly 3-ridged on each valve, c. 15-18 mm @, septum 8-13 mm @. Calyx campanulate, 5-7 mm. Corolla dull purple, yellow-streaked within, c. 3cm (stretched); lobes crenulate; tube c. 13/,cm. Filaments towards the insertion with small Seanlar glands. cot o Matas eee cee eee as ea eae Sea eee 2. S. chelonoides 2. Leafiets 5-11 mm stalked, rounded to subcordate at the base. Capsule only with 1 median ridge on each valve, 8-12 mm @, septum 6-9 mm @. Calyx + cylindric, 8-12 mm. Corolla pale lilac, 6-7 cm; lobes deeply fringed- -laciniate; tube c.4cm. Filaments glabrous ....... 1. S. fimbriatum 1. Inflorescence and leaves glabrous (or very rarely minutely puberulous). Corolla c. 3 cm, suddenly widened and curved above the basal tube, yellowish, the limb with reddish veins and markings, lobes crenulate. Throat and base of filaments densely hairy. Leafiets 5-15 mm stalked, long-acuminate. Calyx campanulate, 4-5 mm. Capsule 4-ridged, 8-10 mm @; septum 4-5mm @ . -3. S. personatum 1. Stereospermum fimbriatum (WALL. ex G. Don) A.DC. Prod. 9 (1845) 211; Kurz, Fl. Burma 2 (1877) 231; CLARKE, Fl. Br. Ind. 4 (1884) 383; GaMBLE, Man. (1902) 516; J. As. Soc. Beng. 74, ii (1905) 378; Rip. Fl. Mal. Pen. 2 (1923) 550; STEEN. Bull. Jard. Bot. Btzg III, 10 (1928) 234; Dop, Fl. Gén. I.-C. 4 (1930) 578; CoRNER, Ways. Trees (1940) 172, pl. 33; CHATTERJEE, Bull. Bot. Soc. Beng. 2 (1948) 69; SANTISUK, Thai For. Bull. Bot. 8 (1974) 23. — Bignonia fimbriata WALL. (Cat. (1832) n. 6500, nomen] ex G. Don, Gen. Syst. 4 (1838) 221. — Fig. 19. Very upright, deciduous tree, to 27-30m by 30-160 cm @; crown narrow, cylindrical, rather : open; bark light grey, rough and flaky; young Fig. 18. Range of the genus Stereospermum CHAM. _ leaves purple or violaceous. Leaves 30-75 cm, with In Asia and Africa generalized, and numbers of rather sticky hairs, stalks yellowish; leaflets species an approximation. Occurrence in Sumatra (2-)3(-4) pairs, rounded at the asymmetric base, and Java doubtful. ovate-oblong, long-tipped, 8-16 by 3-7cm; BIGNONIACEAE (van Steenis) 147 Fig. 19. Stereospermum fimbriatum (G.Don) A.DC. Its slender habit; some trees deciduous; on the churchyard in Malacca (photogr. CORNER). petiolules 5-10 mm. Flowers in large spreading viscid-pilose clusters, 8-30cm 2, on the bare twigs or with the new leaves. Calyx tubular, with 5 very short pointed lobes, c. 8-12 mm. Corolla dull white to pale pink or pale pinkish lilac, narrow funnel-shaped, without a distinct basal tube, the tube 4-5 cm long, the lobes beautifully long-fringed (as a dainty night cap), c. 2 cm. Filaments glabrous, inserted at c. 11-13 mm from the base. Capsule more or less quadrangular, snake-like twisted, 35-60 cm by 8-12 mm; septum c. 5mm @. Seeds c. 21/, cm long, 7mm wide, with rather thick wings. Distr. Continental SE. Asia (Burma; Thailand: Chiengmai to Peninsular Thailand); in Malesia: Malay Peninsula (incl. Langkawi, Penang, and Tioman Is.), possibly also in Sumatra. The record from Sumatra rests on an un- published, beautiful plate in a collection of RAFFLES in the India Office Library & Records (NHD 49/20), which might have been made in Bencoolen, but according to Mr R. DEsMoND the provenance is uncertain. It might be rare in Sumatra, similarly as S. personatum. Ecol. In the lowland and hill forests, in Burma up to 1000 m, in Malaya in high forest and open country: frequent in villages and belukar from Malacca to Perlis and Kelantan, often on rocky coasts and headlands. Fi. Febr.—June, fr. March- Nov. “After the first spell of dry weather the leaves are shed and the flowers appear on the bare boughs in delicate clusters until the new foliage is mature; in the early morning the corollas spin down like snow-flakes and carpet the ground with pale lilac blossom. There are many trees in the Christian Cemetery at Malacca, and a fairer one for a grave- yard would be hard to come by” (CORNER, /.c.). Uses. A hard and durable fairly large timber rather dark coloured, used for beams and posts and said to be durable in the soil. Roots and leaves are used medicinally for some minor ailments: juice of leaves is dropped into the ear for ear-ache; leaves pounded with lime are applied to the skin for itch; a decoction of roots is given as a protective medicine after childbirth (BURKILL, Dict. 1935, 2082). Vern. Malaya: chac(h)a(h), chachar, chechar, chicha(r), M, lempoyan, beka(k) (BURKILL), snake tree, E (CORNER). 2. Stereospermum chelonoides (L. f.) A.P.DC. Bibl. Univ. Genéve II, 17 (1838) 125, pro comb., excl. ref. RHEEDE t. 26; A.DC. Prod. 9 (1845) 210, pro basion.; HAINES, Kew Bull. (1922) 121, in text; SANTISUK, Kew Bull. 28 (1973) 176; non auct. al. — Bignonia chelonoides L. f. Suppl. (1781) 282, pro typ., excl. ref. RHEEDE; non auct. al. — Bignonia suaveolens Roxs. FI. Ind. ed. Carey 3 (1832) 104. — Tecoma suaveolens G. Don, Gen. Syst. 4 (1838) 224. — Hieranthes fragrans RaFIn. Sylv. Tellur. (1838) 79, nom. illeg. — S. suaveolens A.DC. Prod. 9 (1845) 211; WiGurt, Ic. 4 (1848) 9, t. 1342; Kurz, Fl. Burma 2 (1877) 231; CLARKE, FI. Br. Ind. 4 (1884) 382; TrIMEN, FI. Ceyl. 3 (1895) 284; GAMBLE, Man. (1902) 515; BRANDIS, Ind. Trees (1906) 495; Haines, Fl. Bihar Orissa (1922) 656; Beumée, Fl. Anal. Onderz. (1922) 33; GAMBLE, Fl. Madras 2 (1924) 998; STEEN. Thesis (1927) 948, incl. f, verticillatum STEN. l.c. 950; Bull. Jard. Bot. Btzg III, 10 (1928) 236; Dorp, Fl. Gén. I.-C. 4 (1930) 588; CHATTERJEE, Bull. Bot. Soc. Beng. 2 (1948) 70; BACK. & BAKH. f. FI. Java 2 (1965) 540. Deciduous tree, up to 30m, 80cm @; timber dark, hard. Innovations viscid hairy. Leaves opposite (rarely in whorls of 3), 30-50 by 15-25 cm; leaflets 3—4 pairs, viscid-hirsute, glabrescent, rough above and brittle when mature, ovate to obovate to broadly oblong, acute to short-acuminate, entire or fine-dentate, 523 by 3-10 cm; glandless or with a few scattered spots; midrib finally puberulous 148 FLORA MALESIANA [ser. I, vol. 8? above, venation sparsely hirsute beneath; petiolules thick, 2-3(-5) cm. Thyrse up to 25cm @, viscid- pubescent with capitate-glandular hairs. Flowers dull crimson to dull purple, yellow streaked within, very fragrant. Calyx campanulate, viscid pubescent, 6-8 mm, shortly acutely 5-lobed. Corolla viscid- hairy, 2!/,-3 cm long, the tube rather gradually funnel-shaped, mouth long pubescent; lobes subentire, + as long as the tube. Filaments in- serted at 4-5 mm from the base, towards the inser- tion with small sessile granular glands. Ovary 4-ribbed, sometimes sparsely glandular. Capsule smooth or valves obscurely 3-ribbed, to 45 by 14/,-13/, cm; valves woody; septum 8-13 mm @. Seeds 3/, by 3/4 cm. Distr. Widely distributed in continental tropi- cal SE. Asia, from Ceylon and the Deccan to Assam and Burma, not yet reported for Thailand and erroneously so for Indo-China (SANTISUK, l.c.); in Malesia: very locally found in East Java, but somewhat doubtful whether native. BeuMEE /.c. recorded this tree for the first time for East Java, where it was found locally in some places in the (teak) forest districts S. Surabaya and E. Tuban. He suggested that this occurrence would fit the theory of a number of forest officers of early import by Hindus of teak and some associated trees (Butea monosperma, Schleichera oleosa, etc.) and several other plants. I certainly agree that in the Hindu period (roughly 800-1400 A.D.) plants have come from India, especially those favoured for sacred purposes; for example Cochlospermum religiosum (L.) ALSTON, and others went to India vice versa, as for example Santalum album L. (see C. E. C. FiscHer, J. Bomb. Nat. Hist. Soc. 40, 1938, 458-467). The first is still only found near Hindu temples in Bali and the latter is still spread- ing in India. The disjunction between the localities in East Java and India-Burma is in these cases certainly caused by intentional dispersal by man in historic time. There are, however, a large number of other plants showing this same disjunction, and all bound to a seasonal climate, that is, subject to a distinct annual period of drought. In a succinct analysis I found these to belong to 4 classes (Hand. 8th Ned. Ind. Natuurwet. Congr. Surabaya 1938, 1939, 408-409). Later I have further elaborated this problem and tried to solve it (Reinwardtia 5, 1961, 420-429, maps 1-6). From this it appeared that the ecological disjunction of the seasonal climate between the colossal area it covers in SE. Asia (south as far as Tenasserim) and a similar ecology in Central & East Java and the Lesser Sunda Is. is shared by a homologous plant-geographical dis- junction of many hundreds of plants which do not occur in everwet West Malesia, or only in very local seasonal spots in Celebes and the Philippine Islands. A fair number extend their range south- eastwards to Australia. This proves that such pat- terns are quite natural; I have assumed they originated during the Pleistocene Glacial Period, which created a temporary pathway for drought plants between SE. Asia and Australia, to vanish in the Late Holocene. It could thus well be that also S. chelonoides does occur in the native state in the East Javanese teak forest. As a matter of fact no fruit has yet been collected, although flowering was abundant. I cannot subscribe to the opinion of BEUMEE that its dispersal is here by vegetative means, because I cannot well see by what vegetative means and furthermore because it is difficult to see how it would have maintained itself vegetatively in this way for many centuries. On the other hand the existence of a Javanese vernacular name is no argument that it is native; experience tells us that such names are often invented quickly. If it is native, it remains curious that, though it is obvious- ly of rare occurrence, it was only recently dis- covered. It cannot be disproved, however, that its seed was inadvertently introduced by the Forest Service with teak seed from India or Burma. Ecol. Seasonal forest and savannahs. FI. Sept.- Oct. (India: April-June), fr. (Asia) Nov.—Dec. Uses. GAMBLE, /.c. 516, said it is in SE. Asia rather an important large tree by its durable timber which is easy to work and good for building, though the amount of heartwood is small. It also is an excellent fire-wood and makes good charcoal. The root and bark are used as a favourite tonic native medicine. It also is important in sylviculture for its very free seed reproduction; the fruit remains long unopened on the tree and seed gets dispersed at the very end of the hot season after the danger of fire is nearly over, and can germinate with the first rains. Even on exposed slopes and among grass its good natural reproduction is noticeable. BurRKILL (Dict. 1935, 2082) mentions that it yields a gum of the tragacanth class. He also men- tioned that it is referred to as a plant of magic in Sanskritic India, patala, being the Sanskrit name, of which modern vernaculars have been derived. In this respect it is noteworthy that the Javanese name ‘bedali’ is a name for Radermachera spp. TRIMEN reported it in Ceylon planted near Buddhist temples. Vern. Djati tékén, kaju tékén, J. Notes. In East Java one specimen had leaves in whorls of three. Leaves of saplings and suckers may show a serrate-dentate leaf margin. Such leaves are sometimes also rather narrow and acuminate; those of mature trees are broader and more wide at apex. Under S. personatum I referred to the lamentable name change caused by erroneous interpretation of the type of Bignonia chelonoides L. f. The first to observe this was G. Don, /.c., who put ‘B. chelo- noides Kon. under the synonyms of Tecoma suaveolens. Then HAINES remarked that the type of Bignonia chelonoides L. f. was currently named S. chelonoides (Kew Bull. 1922, 121). I myself (Thesis, 1927, 951) was of the same opinion. But these observations were not evaluated nomen- claturally until recently by CHATTERJEE and SANTISUK. 3. Stereospermum personatum (HAssK.) CHATTERJEE, Bull. Bot. Soc. Beng. 2 (1948) 70; SANTISUK, Kew Bull. 28 (1973) 178; Thai For. Bull. Bot. 8 (1974) 26. — Padri RHEEDE, Hort. Mal. 6 (1736) 47, t. 26. — Bignonia chelonoides (non L. f.) Rox. Fl. Ind. ed. Carey 3 (1832) 106, p.p. — S. chelonoides [non (L.f.) A.P.DC.] A.P.DC. Bibl. Univ. Genéve II, 17 (1838) 125, quoad ref. RHEEDE, t. 26; A.DC. Prod. 9 (1845) 210, p.p.; Kurz, Fl. Burma 2 (1877) 230; CLARKE, FI. Br. Ind. 4 (1884) 382; TriMEN, FI. Ceyl. 3 (1895) 283; Ripi. Fl. Mal. Pen. 2 (1923) 1977] BIGNONIACEAE (van Steenis) 149 550; STEEN. Thesis (1927) 951; Bull. Jard. Bot. Btzg III, 10 (1928) 237; Dorp. Fl. Gén. I.-C. 4 (1930) 579; CorNER, Ways. Trees (1940) 172, f. 43. — Dipterosperma personatum Hassk. Flora 25, 2 (1842) Beibl. 1, p. 28; Cat. Hort. Bog. (1844) 152; Pl. Jav. Rar. (1848) 507. — S. tetragonum A.DC. Prod. 9 (1845) 210; Harnes, Fl. Bihar Orissa (1922) 655; Kew Bull. (1922) 121; GAMBLE, Fl. Madras 2 (1924) 998; HaNpD.-MaAzz. Symb. Sin. 7 (1936) 889. — Bignonia caudata A.DC. Prod. 9 (1845) 166. — S. hasskarlii Z. & M. ex ZOLL. Syst. Verz. 3 (1855) 54, nom. illeg., based on Dipterosperma personatum Hassk.; Miq. Fl. Ind. Bat. 2 (1858) 756; Ann. Mus. Lugd. Bat. 1 (1864) 200. — S. caudatum Mia. /.c. 200. — Fig. 17. Deciduous, glabrous tree, up to 30 m, 75cm @; bark pale pinkish grey becoming rather coarsely fissured and flaky but not ridged. Young leaves purplish or pinkish (CORNER). Leaves glabrous, 20—50 cm; leaflets 3-6 pairs, elliptic-oblong, gra- dually tapering to the base, tip acuminate to cau- date; 5-15 by 2'/,-6cm; underneath minutely glandular-punctate, often with a few large flat spot-glands (black in dry state); petiolules slender, 5-15 mm. Thyrses widely branched, paniculiform, to 40 cm long. Flowers dingy yellow, cream within with dark red stripes, in slender, erect, lengthening thyrses 15—40 cm long, on the bare twigs with the new leaves, c. 3 cm long, 1°/, cm wide at the limb. Calyx campanulate, 6-8 mm, with 3-4 short acute lobes, purple. Corolla with a narrow basal tube 4-5mm long, then campanulately widened and curved, the bell-shaped part compressed with the mouth closed and the underside grooved; tube c. 13/,cm long; lobes crisped, the upper two re- curved, the lower bearded at the mouth, ochre- buff with brownish or purplish lines, pinkish purple on the outside (CORNER). Filaments with a dense hair tuft at the base. Capsule linear, + terete, 4-ribbed, curved or twisted, 8-45 cm by 8-10 mm; septum 4-5mm @. Seeds 2 by */, cm, incl. wings ,cm. Distr. Widely distributed from Ceylon through entire continental tropical SE. Asia to Yunnan and Indo-China; in Malesia: Malaya (very rare, only in Penang, e.g. on Glugor Road, and Singapore), possibly also in Sumatra. Unfortunately there is no certainty about the records in Indonesia: HASSKARL described his type from trees cultivated in the Botanic Gardens at Bogor; ZOLLINGER described S. hasskarlii also from a cultivated tree in these gardens (ZOLLINGER 3069) but noted that it would have originated from Bantam, West Java, adding the Sundanese verna- cular ‘ki langir’; a duplicate of this number in Paris is said by SANTISUK to have been annotated to come from the Lampong Distr. in S. Sumatra, which then probably is an error. Then there is a collection said to have been col- lected by KORTHALS with ‘Borneo’ printed labels. This provenance is very unlikely, as the use of these old labels has been proved to be often erratic. These specimens may have come from West Central Sumatra but may also have been collected by KoRTHALS in the Bogor Botanic Gardens. Plant-geographically the species might occur (or have occurred) in Sumatra and West Java, but probably as rare as in Malaya because of its preference for seasonal forest conditions. Ecol. Preferring lowland forests, up to 1000 m. Fl. March-July (at Bogor Aug., Nov.—Febr.), fr. Febr. (Asia), June (Malaya). Uses. According to BurRKILL in India an im- portant timber tree, especially in the northeast where it is common; the hard grey wood is moder- ately durable and easy to work, good for furniture, used but less good for building; in Assam and E. Bengal padri-wood is used for canoes and tea- boxes. In S. India a cooling drink, from roots and flowers, is given in fevers. The fragrant flowers are offered in temples. Nomencl. There has been a most unfortunate confusion about the identity of Bignonia chelo- noides L. f. (1781). This emanated from LINNE f. who described it as a hairy plant (type herb. Konig, in LINN), but added the reference to Padri of RHEEDE, an other glabrous species with long petiolules. This probably misled RoxBURGH who applied LINNE’s epithet to the latter. This interpretation was followed by almost all subse- quent authors. In 1922 Harnes concluded that two species were involved and he adopted for the present one the name S. tetragonum DC. In 1948 CHATTERJEE replaced this by an older epithet of HASSKARL. Notes. As in several other members of this family leaves from suckers and saplings may be toothed or serrate at the margin; leaves of mature trees are entire. In a few continental Asiatic specimens a very minute puberulous indument occurs on nerves underneath in the inflorescence (KERR 1167, KOSTERMANS 1056, LESCHENAULT 157). 9. RADERMACHERA ZoLL. & Mor. in Zoll. Syst. Verz. 3 (1855) 53; BUREAU, Adansonia 2 (1861) 192, t. 2; Mon. (1864) 50, t. 28; Mig. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 250; SEEM. J. Bot. 8 (1870) 145; Jackson, Ind. Kew. 2 (1895) 679 (‘Radermachia’); STEEN. Thesis (1927) 953; Bull. Jard. Bot. Btzg III, 10 (1928) 238; Dop, Fl. Gén. I.-C. 4 (1930) 583; CHATTERJEE, Bull. Bot. Soc. Beng. 2 (1948) 71; SANTISUK, Thai For. Bull. Bot. 8 (1974) 27; STEEN. Blumea 23 (1976) 121. — Lagaropyxis Mig. Ann. Mus. Bot. Lugd.-Bat. | (1864) 198. — Mayodendron Kurz, Prel. Rep. For. Veget. Pegu, App. D (1875) pl. | & 2; Fl. Burma 2 (1877) 232. — Stereospermum sect. 150 FLORA MALESIANA [ser. I, vol. 82 Fig. 20. Radermachera peninsularis STEEN. a. Leaflet, b. apex of thyrse, c. calyx and style, d. corolla, all nat. size. — R. glandulosa (Bu.) Mig. e. Leaf base, underneath with dense glandfield, x 2, f. empty capsule, with 2 valves and septum, x 1/3, g. section of septum, x 5, h. seed, x 2 (a—d LARSEN c.s. 31239, e-h KOSTERMANS 6360A). 1977] BIGNONIACEAE (van Steenis) 151 Radermachera et Xylocarpaea B. & H. Gen. Pl. 2 (1876) 1047; CLarkg, FI. Br. Ind. 4 (1884) 383; K. & V. Bijdr. Booms. | (1894) 71. — Radermachera sect. Alatae STEEN. Acta Bot. Neer]. 2 (1953) 307. — Fig. 20, 22. Trees, rarely shrubs, evergreen (except 2 Indian and 2 Chinese spp.). Twigs and floral rachis mostly distinctly lenticellate. Innovations sticky-lacquered. Leaves 2-3-pinnate; stalks and leaflets articulated; rachis sulcate; leaflets underneath minutely but densely dotted, furthermore with gland-fields or scattered glands underneath, mostly acuminate, very rarely laxly puberulous. 7hyrses terminal, very rarely ramiflorous, in one sp. a raceme; bracts and bracteoles inconspicuous, very rarely leafy; thyrses (and calyx) very rarely short-hairy. Flowers white, pink with yellow streaks in mouth, more rarely greenish yellow, or golden yellow to orange-red. Calyx closed in bud, rarely lobed from the beginning, splitting spatha- ceously (not to the base) or mostly irregularly lobed, rarely stunted, often with microscopical scale-like glands, moreover mostly with larger crateriform glands; after anthesis almost always circumscissile-dehiscing at the base, along an abscis- sion line, in one sp. persistent. Corolla either salver- or narrow funnel-shaped or more commonly with a mostly short basal tube and often rather suddenly widened into an upper tube; lobes mostly rounded unequal, not rarely ciliate. Stamens didynamous with a Sth rudiment, but in one Chinese sp. 5 equal stamens, not exserted; another-cells V-shaped divergent; filaments inserted at the apex of the basal tube, except in two spp. capitate-glandular hairy at the insertion and in the basal part, for the rest glabrous; connective produced. Ovary elongate often minutely lepidote, glabrous, or tuberculate, never hairy; in both cells with several rows of ovules; style filiform, mostly exceeding the anthers; stigma 2-lipped. Capsule linear, terete, up to 75 cm; valves smooth, pergamentaceous, rarely + woody, in one sp. tuberculate; septum terete, brittle, corky, but with shallow im- pressions of the flat seeds, a narrow line on both sides testimony of attachment to the middle of the valves. Seeds very «, flat, small, narrow, on both ends with a hyaline wing. Distr. About 15 spp. in Indo-Malesia, from the Deccan to S. China, Hainan, Formosa, and the southern Ryu Kyus, most abundant in tropical SE. Asia; throughout Malesia, but not in the Moluccas proper and New Guinea. Fig. 21. Ecol. Lowland primary and secondary rain-forests, up to c. 1500 m, not rarely pioneering in disturbed forest and on slopes. Flower colour and corolla shape vary from pure white to orange-red and from hypocrateriform (R. sinica (HANCE) HeEmsL. and R. frondosa CHUN & How) to tubular or campanulate. For the narrow- tubular orange-flowered species pollination by birds can be expected (R. ramiflora) and for the pure white, possibly nocturnal R. peninsularis hawk-moths may be the pollinating insects. Ramiflory is found in Malesia in R. ramiflora; it occurs also occasionally in poor forms of R. pinnata after leader-shoots have been damaged. In continental Asia it is also found in R. ignea (KURZ) STEEN. and in R. hainanensis MERR. Uses. Minor uses, see under the species. Syst. Related to Stereospermum, which differs radically in having only two rows of ovules in each cell, and thick trigonous seeds with folded cotyledons fitting in cavities of the septum; moreover, its leaves are always 1-pinnate and most species are deciduous and prefer a seasonal climate. Two small genera have recently been split off, both from SE. Asia, viz the monotypic Pauldopia STEEN. (Acta Bot. Neerl. 18, 1969, 425) which has winged rachises, a curved corolla tube, wingless thickish seed, a very thin septum, and a truncate calyx open in bud; and Barnettia SANTISUK (Kew Bull. 28, 1973, 172) with 2 spp. in Thailand, with 1-pinnate leaves in pseudowhorls, a short-ellipsoid, -+- compressed capsule, and tuberculate or immersed-glandular calyx and capsule and a cruciform septum; the latter genus shows relationship with Heterophragma. The closest ally of Radermachera is obviously the Afro-Asian genus Fernandoa (cf. Steen. Blumea 23, 1976, 133) from which it differs by the terete septum, the usually not ribbed or striate and thinner, 152 FLORA MALESIANA [ser. I, vol. 8? narrower fruit valves, mostly 2-4-pinnate leaves, absence of domatia underneath the leaflets, and mostly crateriform glands. Notes. Specific delimitation, especially in Malesian material, proved difficult, as several species appear to be variable and many were only known from the type, notably in the Philippine Islands. Degree of pinnation is variable, often in one collection. Also the number and place of the larger glands on under- surface of leaflets and on the calyx is liable to variation, as well as the flower colour and the way in which the calyx splits. A critical scrutiny showed only few tangible characters and resulted into a severe reduc- tion of species in Malesia. KEY TO THE SPECIES 1. Calyx strongly lengthwise 5-6-ridged, 13/, cm, narrow, one side cleft down + halfway, 3-toothed at apex. Leaves 1-pinnate, with 5 leaflets, coriaceous, very glossy above, with recurved edge, 7-14 by 3-4 cm. Corolla 4 cm, with narrow tube, slightly enlarged above; lobes c. 1 cm, obtuse. 6. R. coriacea 1. Calyx not lengthwise 5-6-ridged. 2. Corolla narrowly funnel-shaped, without distinction of basal and upper tube. Flowers in cauli- florous racemes, with yellow tube and red limb. Leaves 2-pinnate. ..... . 1. R. ramiflora 2. Corolla with a cylindric basal tube, widened rather abruptly into an upper tube. Flowers in terminal thyrses which are rarely raceme-like depauperated. 3. Calyx very short (3-5 mm), with a stunted rim, persistent. Leaves always 1-pinnate 2. R. glandulosa 3. Calyx longer, irregularly lobed, after anthesis circumscissile-caducous. 4. Corolla 2-3'/,(-4+/,) cm long (incl. lobes). Calyx 5-10(-13) mm long 3. R. pinnata 4. Corolla 5-6 cm long. Calyx 10-25 mm. 5. Filaments and inside of basal tube glabrous. Corolla tube outside towards apex and on lobes with minute sessile glands. Leaves above puberulous on midrib and nerves, beneath with some scattered glands. Corolla white or cream-coloured 4. R. insularis 5. Filaments and inside of basal tube near insertions capitate-glandular hairy. Corolla tube outside towards apex mostly laxly and very short-hairy. Leaves glabrous above, beneath mostly with at least a gland-field at base. Corolla mostly pinkish, sometimes white, with yellow streaks in the mouth 1. Radermachera ramiflora STEEN. J. Bot. 72 (1934) 5; Blumea 23 (1976) 129. Large tree, 24-30 m, stem 30-60 cm @; fluted at base; bark grey, fissured. Innovations and racemes lacquered, resinous sticky. Leaves (2—)3-pinnate, up to 1 m, more or less crowded at the twig-ends; pin- nae 4-6 pairs; leaflets elliptic-lanceolate, acuminate at both ends, stalked, 31/,-8'/, by 11/,-31/,cm, under- neath fine glandular-punctate and with scattered small, shallow glands especially near the base, and a few scattered flat glandular spots. Flowers thickly set in closely placed ramiflorous pendent racemes to 20 cm long, erect on curved pedicels 1-2 cm long and with 3 bracteoles + halfway on an articulation. Calyx in bud pear-shaped, closed, reddish green, tubular, 2—3-lobed, eglandular outside, microsco- pically glandular-papillose inside, c. 21/,—2'/, cm long, 1 cm @ at mouth, lobes 4-6 mm, 1-2 split + halfway. Corolla with yellow tube and red limb; tube slightly curved, narrow salver-shaped, without narrowed basal tube, 5-7 cm, the basal 11/,cm densely pubescent with thick hairs, mouth 11/,- 2cm @; lobes rounded, subequal, + Icom @, + papillose inside. Stamens (and style) reaching the mouth, inserted halfway the tube, glabrous but their adnate base lax glandular-papillose, thecae + divergent, 4-5 mm; no produced connective. Disk thick, annular-cup-shaped, faintly crenate. Ovary ribbed, pistil c. 4cm, stigmatic lobes 2 mm, very narrow. Capsule straight or twisted, 35-70 cm, c. 5mm @; septum 2!/,-3mm @. Seeds 4-5 by 2'/, mm, the wings 6-7 mm. Distr. Malesia: Sabah, Mt Kinabalu (Peni- bukan, Dallas, Mesilau, Tenompok, Kota Belud, Kp. Kiau I resthouse), not rare. 5. R. gigantea Ecol. Rain-forest, also in disturbed forest on hill side, 950-1500 m. Fl. Aug., Jan.—March, fr. April, Dec. Notes. A characteristic, isolated species: rami- florous, flowers in racemes, shape of corolla. Leaflets of suckers and saplings dentate. NOOTE- BOOM & ABAN 1603 has only 2-pinnate leaves, CLEMENS 30364 has them 2-3-pinnate. Fig. 21. Range of the genus Radermachera Z. & M. 2. Radermachera glandulosa (BL.) Mia. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 250; K. & G. J. As. Soc. Beng. 74, ii (1905) 380; Koorp. Atlas Baum. 2 (1914) t. 356; STEEN. Thesis (1927) 965; Bull. Jard. Bot. Btzg III, 10 (1928) 241; CorNeER, Ways. Trees (1940) 168, f. 43; CHATTERJEE, Bull. Bot. Soc. Beng. 1977] BIGNONIACEAE (van Steenis) 153 2 (1948) 74; STEEN. Acta Bot. Neerl. 2 (1953) 307; Back. & BAKH. f. Fl. Java 2 (1965) 541; SANTISUK, Thai For. Bull. Bot. 8 (1974) 27, f. 15; STEEN. Blumea 23 (1976) 126. — Spathodea glandulosa Bi. Bijdr. (1826) 762. — Bignonia porteriana WALL. ex DC. Prod. 9 (1845) 165. — R. stricta Z. & M. ex ZOLL. Syst. Verz. 3 (1855) 53; Rw . FI. Mal. Pen. 2 (1923) 550. — Stereospermum glandu- losum Mig. Suppl. (1860) 240, 565; CLarKE, FI. Br. Ind. 4 (1884) 383; K. & V. Bijdr. Booms. 1 (1894) 74; KANyJILAL & Das, Fl. Assam 3 (1939) 404. — Lagaropyxis glandulosa Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 199. — Fig. 20e-h. Small, glabrous, evergreen, crooked tree, up to 12m, 40 cm @; bark slightly pimply and peeling, but not flaky or fissured, bitter; young leaves deep purple, acrid. Leaves 1-pinnate; leaflets 2-5 pairs, large, chartaceous, elliptic (mostly broad-), rarely oblong-lanceolate, short-tipped, with a conspicuous dark, dense gland-field at the oblique base underneath (often bulging on upper surface), 10-30 by 5-17 cm. Thyrse narrow, 6-50 cm, gradu- ally elongating, sometimes the upper part still flowering while lower are in fruit. Calyx 3-5 mm, cup-shaped, without abscission line, persistent, truncate, purple, spotted with 5—7 purple glands in a crescent. Corolla narrow, slender, tube narrow, slightly curved, halfway rather gradually widening, 3(-4) cm, pinkish purple outside, white inside, with a gland-field at outside of the ciliate lobes, basal part of lower tube short capitate-glandular hairy inside. Stamens hairy at insertion. Pods hanging in bunches, straight, 15-30cm; valves 5-7 mm wide; septum 3-4 mm @. Seeds 10-16 by 11/,-2 mm. Distr. Continental SE. Asia (Assam, Burma, Thailand, Laos, China: Kwangsi, Kwantung) and West Malesia: Malay Peninsula (also Penang), Sumatra, Krakatao, West to East Java. The two records from Borneo are erroneous: the KoORTHALS specimens are mislocalized and BECCARI 811 (mentioned by MERRILL, En. Born. 1921, 525) is a PS number. Both are from Sumatra. Ecol. Primary and secondary forests and thick- ets, frequently by streamsides, even rocky Saraca-streams (CORNER), both under everwet and under seasonal conditions (in Central and East Java), from sea-level to 900 m, once reported from 5000 ft (Cameroon Highlands). Fi. April, July—Nov., fr. Jan.—Dec. The fine seed would indicate easy dispersal, but though the species is found in Penang and Krakatao Is., it has never been collected in the Riau Is., not in the islands west of Sumatra and also not in those close to East Java (Madura I., Bali, and Kangean) though it is found in Java as far east as Bali Straits and R. gigantea extends east as far as Alor. The glands at the leaflet basis are often black from sooty moulds indicating actual glandular excretion. Uses. The timber is small and of negligeable value. Vern. Tuwi gadang, M, Minangkabau; kia bako, ki hapit, ki langhit, kipahit, ki sakat, ki sikap, 8; ambal, bangking, bangkongan, djelibru, godong ambol, hambal, paras = gabret, klaju, lambal, padali, pedali, pudang, J; sekar pote, Md. 3. Radermachera pinnata (BLANCO) Seem. J. Bot. 8 (1870) 147; Merr. Philip. J. Sc. 3 (1908) Bot. 336, incl. var. glabra MERR. l.c.; Sp. Blanc. (1918) 350; En. Philip. 3 (1923) 446; STEEN. Thesis (1927) 973; Bull. Jard. Bot. Btzg IIT, 10 (1928) 248; Blumea 23 (1976) 129. — Millingtonia pinnata BLANCO, FI. Filip. (1837) 501. — Méillingtonia quadripinnata BEANGO; Ics: ed! 3,;, 2 (1878): 286, t. 252. — R: banaibana BUREAU, Adansonia 2 (1861) 194. — R. quadripinna Seem. J. Bot. 8 (1870) 147. — Stereo- spermum pinnatum F.-VILL. Nov. App. (1880) 151; Ro re, J. Linn. Soc. 23 (1884) 314; VipAL, Phan. Cuming. Philip. (1885) 132; Rev. Pl. Vasc. Filip. (1886) 203. — Stereospermum quadripinnatum F.-VILL. Nov. App. (1880) 151; VmDAL, Syn. Atlas (1883) 35, t. 73. — Stereospermum banaibanai Ro.rFe, J. Linn. Soc. 23 (1884) 314. — Stereo- spermum seemannii ROLFE, /.c. — R. acuminata Merk. Philip. J. Sc. 3 (1908) Bot. 335; En. Philip. 3 (1923) 445; STEEN. Thesis (1927) 980. — R. mindo- rensis MERR. Philip. J. Sc. 3 (1908) Bot. 338; En. Philip. 3 (1923) 446; STEEN. Thesis (1927) 971. — R. fenicis MERR. Philip. J. Sc. 3 (1908) Bot. 335, 434; En. Philip. 3 (1923) 446; STEEN. Thesis (1927) 979, incl. var. acuminata STEEN.; Jard. Bot. Btzg III, 10 (1928) 261. — R. whitfordii MerR. Philip. J. Sc. 7 (1912) 352; En. Philip. 3 (1923) 447; STEEN. Thesis (1927) 963. — R. brachybotrys MERR. Philip. J. Sc. 26 (1923) 489; E_mer, Leafl. Philip. Bot. 10 (1939) 3809. — R. sorsogonensis ELMER ex STEEN. Thesis (1927) 973; ELMer, Leafl. Philip. Bot. 10 (1939) 3809. — R. elegans STEEN. Bull. Jard. Bot. Btzg III, 10 (1928) 252, f. 8. — R. fenicis (non MERR.) STEEN. /.c. 261, f. 11. See for other synonyms under ssp. acuminata. Distr. Malesia: Sumatra (also Simalur, Banka, and Siberut Is.), Malaya, Borneo, Philippines, Celebes (also Muna I.), and W. Moluccas (Sula Is.: Taliabu). Notes. It has appeared impossible to separate the material into smaller species, as there is gradual transition of the many populations, especially in the Philippine islands, notably in vegatative characters. R. brachybotrys is merely a depauperate ridge facies. In one specimen 1- and 2-pinnate or biternate leaves are not seldom found together. For brevity I have omitted from the synonymy the many pro parte citations under R. amoena, hypostictum and gigantea. Some speci- mens from Borneo and Celebes show a tendency in leaf-shape towards ssp. acuminata. KEY TO THE SUBSPECIES 1. Leaflets usually chartaceous, elliptic, acuminate to caudate; basal gland field mostly distinct, at apex eglandular or with a few scattered glands ssp. pinnata 1. Leaflets firmly chartaceous to coriaceous with consequently less marked prominent venation beneath, usually obovate to elliptic-obovate, at apex mostly short- and blunt-tipped or blunt or rounded; basal gland field dense, often one also BEDDGKinn bhai’ Winn ae Ras ssp. acuminata ssp. pinnata. Tree to 20m, 15-40cm @. Leaves (1-)2-3- pinnate, 25-50(—70) cm; leaflets elliptic-oblong, acuminate, mostly at both ends, to caudate, (3-)5—16 by (1'/,-)2-5(-8) cm, usually chartaceous, 154 FLORA MALESIANA {ser. I, vol. 8? io Fig. 22. Radermachera pinnata (BLANCO) SEEM. ssp. acuminata (STEEN.) STEEN. Habit, in flower, Aug. 1970 (photogr. B. C. STONE). with a basal gland field, apical gland field absent or of scattered glands. Thyrses glabrous, sometimes puberulous, (3—6—)15-60cm, in odd specimens sometimes on the old wood. Calyx #/,-1(-11/,?) cm, lobes unequal, (1—)2—4, glabrous, usually glandless. Corolla pink to pale purplish, with yellow mark- ings in the throat, 2—3'/,(—4"/,) cm, rarely slightly lax short-hairy. Stamens glandular-hairy at their insertion. Ovary and style glabrous, 12-17(-22) mm. Capsule (6-)30-50cm; valves 4-6(—7) mm wide; septum 2-2!/,mm @. Seeds (7-)13-15 by 2-3 mm. Distr. Malesia: Philippines (the most common species), Celebes (also in Muna I.), Moluccas (Sula Is.: Taliabu). Ecol. Lowland and montane primary and secondary forests, on streamsides, up to 600 m. Fl. Aug.—May, fr. Febr.—Nov. Uses. Minor uses only, for carving and fuel. Vern. Philippines: banai-bdnai, Tag., the com- mon name, kalapuing, salai, tuing-huld, ulimbabon, yabang-ydabang, Tag., banoi-banoi, Bag., ansohan, badlan, Bis., labayanan, C.Bis., paling-udk, Bik., pagalayan, Bon., bunglai, Buk., atiatip, Ig., lanunisi, lasilak, Ybn., barangauan, I\k., bunlai, Mbo,. banaibayan, paitan, pata, Pang., bani-bani, Sbl., kutokong, Sub., hali-hdali, Sulu; Celebes: ririh, Muna. Note. In a few specimens some inflorescences are ramiflorous, obviously due to damage of the leader shoot. ssp. acuminata (STEEN.) STEEN. Blumea 23 (1976) 129. — Spathodea lobbii T. & B. Nat. Tijd. N. I. 25 (1863) 413. — R. lobbii Mig. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 250; SEEM. J. Bot. 8 (1870) 147; STEEN. Bull. Jard. Bot. Btzg III, 10 (1928) 243, f. 5, incl. ssp. acuminata STEEN. l.c. 247, f. 6; CORNER, Ways. Trees (1940) 168, f. 43 (‘lobbiana’); SANTISUK, Thai For. Bull. Bot. 8 (1974) 29. — R. amoena [non (WALL.) SEEM.] GAMBLE, J. As. Soc. Beng. 74, ii (1905) 381; Rip. Fl. Mal. Pen. 2 (1920) 551. — R. corymbosa STEEN. Bull. Jard. Bot. Btzg III, 10 (1928) 249, f. 7. — R. gigantea [non (BL.) Miqg.] Burk. Dict. (1935). — Fig. 22. Tree, 7-40 m, 60cm @. Leaves biternate or 2-pinnate; leaflets rather coriaceous, obovate to obovate-elliptic, not or short- and blunt-tipped, exceptionally acuminate, basal gland field well- developed, apical one usually distinct, 8-15 by 31/,-71/, cm. Thyrse apical, fairly narrow, 15-25 cm. Calyx 1 cm, rarely to 11/, cm. Basal tube of corolla rather suddenly widened. Distr. S. Peninsular Thailand (Krabi, Pattani); in Malesia: Sumatra (also in Banka; Mentawai Is.: Siberut; Simalur I.), Malaya, Borneo, SW. Philippines (Palawan, Culion). Ecol. Primary and secondary forests, also in open grasslands. In Malaya especially by streams (CORNER), ON granite as well as on limestone; at 0-—400(-800) m. F/. May, July—March, fr. May- Nov. Taxon. Formerly often accepted as conspecific (also by myself, 1927) with R. gigantea cq. amoena, but certainly distinct. Vern. Sumatra: kaju singamba, sindur langit, sundur langit, Batak, kudo kudo pajo, Simalur, mentu, tuih, tuwi(k), Banka; kapung suwi, kéka- 1977] pung, Lampong; bunga pawang, seténgah burong, tangkani, Malaya; Borneo: binutan, kudjuk langit, Dajak-Kapuas; Culion: totancola, Tagb. Note. I must admit that I have somewhat hesitantly kept this apart as a subspecies from true R. pinnata, from which it differs only in shape and texture of leaflets and geographical range. There are some specimens in Borneo which seem transitional. In the Philippines (Palawan and Culion) aber- rant specimens are found with biternate leaves and coriaceous leaflets with recurved margin and rather prominent veins underneath. They were collected in grasslands which may account for their habit. However, in the same islands there are also larger- flowered specimens which I have referred to R. gigantea with similar habit. More field work in these islands is needed to check my tentative conclusions. 4. Radermachera peninsularis STEEN. Blumea 23 (1976) 128, f. la-d. — R. borii (non FiscHER) SANTISUK, Thai For. Bull. Bot. 8 (1974) 30. — Fig. 20a—-d. Tree, 415m. Leaves 3-pinnate, 60-80 cm; leaflets lanceolate-oblong, falcate-caudate, 5-8 by 2-3 cm, on both surfaces microscopically punctate but only with a few scattered, very small ‘larger’ glands; midrib and main nerves (c. 4-5 pairs) puberulous above (as in R. sinica). Inflorescences terminal, similar to those in R. sinica; peduncle firm, 15-35cm long, rachis 4-7 cm; full-grown pedicels 2!/,-7 cm, halfway with 2 decussate linear bracteoles c. 1'/,-3 cm long. Bracts long, linear, exceeding the buds, the lowest up to 5cm long, upper ones 2!/,-3 cm. Calyx campanulate, rather wide and thickish, densely microscopically lepi- dote, 2—2'/, by 1'/, cm, rather irregularly lobed for 1/,/, of its length, with 5 + distinct gland fields. Corolla white or creamy, c. 6-7 cm long (incl. the %entire lobes), with a rather wide (7/,cm) basal tube c. 1'/, cm long concealed in the calyx on top of which the filaments are inserted ; upper tube funnel- shaped widened. Filaments c. 3cm, glabrous; anthers 4'/,mm; connective appendage small. Ovary glabrous. Capsule terete, tortuous, 60-70 cm by 3-5 mm. Distr. Peninsular Thailand (between Phangnga and Krabi); in Malesia: Malay Peninsula (Cameron Highlands), 2 collections. Ecol. In Peninsular Thailand at 8°25’ N, 99°15’ E, in evergreen forest along a stream, on limestone, at 50m altitude, in the Cameron Highlands at 1200 m in mixed rain-forest. Notes. In habit deceptively like R. sinica, but at once different by the shorter (6-7. cm), not salver-shaped corolla and the much wider smooth calyx. The corolla in R. sinica measures c. 8-12 cm including the lobes, with a narrow, very gradually widening tube. R. sinica also has the minute lax bescent nerves and veins above and long narrow SANTISUK /.c. referred the specimens to R. borii, which I refer to R. sinica, a species ranging more northerly in SE. Asia, viz from N. Assam, N. Burma, and Tonkin to S. China, Hainan, Formosa, and the southern Ryu Kyu Is. 5. Radermachera gigantea (BL.) Mig. Ann. Mus. BIGNONIACEAE (van Steenis) 155 Bot. Lugd.-Bat. 3 (1867) 250; SEEM. J. Bot. 8 (1870) 146; Koorp. Atlas Baum. 2 (1914) t. 356 A-K:; Heyne, Nutt. Pl. (1927) 1371; STEEN. Thesis (1927) 983, p.p., excl. syn. lobbii; Bull. Jard. Bot. Btzg III, 10 (1928) 253, f. 9; Blumea 23 (1976) 126. — Bignonia gigantea NORONA, Verh. Bat. Gen. 5 (1790) 70, nomen. — Spathodea gigantea BL. Bijdr. (1826) 761; Mia. Fl. Ind. Bat. 2 (1858) 751. — Bignonia amoena WALL. [Cat. (1832) n. 6512, nomen] Pl. As. Rar. 2 (1831) 78, t. 183; LouDon, Hort. Brit. (1830) 483, err. amara; G. Don, Gen. Hist. 4 (1838) 222. — Bignonia oxyphylla DC. Prod. 9 (1845) 169. — Stereospermum hypostictum Mig. Sum. (1861) 565; CLARKE, Fl. Br. Ind. 4 (1884) 384, excl. syn. S. lobbii; K. & V. Bijdr. Booms. 1 (1894) 72, excl. syn. lobbii; KANNLAL & Das, Fl. Assam 3 (1939) 405. — Lagaropyxis gigantea Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 198, incl. f. sumatrana et f. borneensis Miq. — R. amoena SEEM. J. Bot. 8 (1870) 146; non GAMBLE, J. As. Soc. Beng. 74, ii (1905) 381, quae est R. lobbii. — R. elmeri Merr. Bull. Gov. Lab. Philip. 29 (1905) 48; Philip. J. Sc. 3 (1908) Bot. 334; En. Philip. 3 (1923) 445; STEEN. Thesis (1927) 994. — R. biternata MERR. Philip. J. Sc. 1 (1906) Suppl. 238; ibid. 3 (1908) Bot. 333; En. Philip. 3 (1923) 445; STEEN. Thesis (1927) 970. — R. palawanensis Me_rr. Philip. J. Sc. 3 (1908) Bot. 336; STEEN. Thesis (1927) 977. — R. elliptica MERR. Philip. J. Sc. 3 (1908) Bot. 334; En. Philip. 3 (1923) 445; STEEN. Thesis (1927) 964. — R. sibuyanensis ELMER, Leafi. Philip. Bot. 4 (1912) 1485; Merr. En. Philip. 3 (1923) 447; STEEN. Thesis (1927) 992. — R. elmeri var. fragrans ELMER, Leafl. Philip. Bot. 7 (1915) 2561; Merr. En. Philip. 3 (1923) 446. — R. fragrans STEEN. Thesis (1927) 996. — R. punctata ELMER ex STEEN. Thesis (1927) 982; ELmer, Leafi. 10 (1939) 3709. — R. borneensis STEEN. Bull. Jard. Bot. Btzg III, 10 (1928) 258, f. 10. Shrub or tree, (6—)20-40 m, up to 80 cm @; bark and young leaves bitter. Leaves (1—)2(—3)-pinnate, 12-35(-80) cm, leaflets usually elliptic to oblong, rarely somewhat obovate or lanceolate, shorter or longer acuminate, 4-12(-15) by 2-6(-9) cm, at base underneath mostly with a gland-field, at apex with some scattered glands. Thyrses 8-40 cm, rather open, terminal, ~©-flowered, glabrous. Flowers not rarely fragrant. Calyx (1-)1'/,-2'/,cm long, mostly 2-lobed, sometimes 1- or more-lobed ; glands few or distinct. Corolla 5-6 cm long (incl. lobes), pink or white, usually with yellow streaks in the mouth, above the basal tube rather suddenly widened, campanulate upper tube towards apex almost always short-capitate glandular hairy, and lobes ciliate. Filaments densely capitate-glandular hairy at insertion. Style 2'/,-3 cm. Capsule 15-60 cm long, 5-8 mm @; septum 4mm @. Seeds 8-13 by 2-4 mm. Distr. SE. Asia (Assam: Khasi & Jaintia Hills; Burma: Manipur, Tavoy, WALLICH, CLARKE, /.c., not seen), and Malesia: Sumatra (incl. Billiton and Banka), West to East Java (common in Central and East Java), Lesser Sunda Is. (Bali, Sumbawa, Flores, Alor), Bawean L., SE. Borneo, and Philip- pines. Ecol. Primary and secondary forests, also in areas subject to a dry monsoon, and in East Java in teak forest, very common on Mt Telemojo and near Pringombo (Banjumas) (Koorpers, /.c.), 156 FLORA MALESIANA [ser. I, vol. 8? Ve yaa \ I | Breen Ne CONN — ° ".< ert Ey ET en PR a Sg STRRSah ten era See Se NS = Boss WENA wee) yeaa en ESAT wl : : Wyn Nantes y m (eos wl a, Pm UTUrTTUTyY a oe TT SS a dl flsy Vil LCE Wl {(CUG Ue CULO Y “We (Cec d(C ( PAPAL S nah hg fee Mf (CGC Yyf 2. TD ha Fig. 27. Pajanelia longifolia (WILLD.) K.SCH. a. Fragment of leaf, X */2, b. bud, c. flower, both nat. size, d. capsule, e. ditto in CS, both x 1/,, f. seed, nat. size (a—c after WALLICH, Pl. As. Rar. 1, t. 95/96, e-f Kerr 18547). 1977] BIGNONIACEAE (van Steenis) 163 + constricted above basal tube, zygomorphous; lobes 5, spreading to recurved, nearly equal, 2 lobes connate halfway up, imbricate in bud. Stamens 4, inserted at apex of lower tube, didynamous, subexserted, Sth rudimentary; filaments thick, anther-cells divergent. Disk large, annular, fleshy. Ovary oblong-cylindric; ovules in each cell o4, on 2 placentas, co-seriate. Capsule flat, obovate-lanceolate, tipped, valves broadly winged along the margin. Seeds in several rows on the edges of the septum, compressed, hyaline-winged. Distr. Monotypic, from Malabar eastwards through SE. Asia (E. Bengal: Khasia, Sylhet; Burma: Pegu, Travancore, Chittagong, Tenasserim; Thailand, Andaman Is.) to West Malesia: N. Sumatra (E. Atjeh), Malaya (Perak, N. Kedah, ?Singapore, ?Penang), and the Natuna Is. (Sedanau, Bunguran, Duperré) in the S. China Sea, NW. of Sarawak. Fig. 28. F.-VILLAR (Nov. App. 1880, 150) recorded it erroneously from the Philippines. Ecol. Lowland evergreen mixed forests, in the Andaman Is. also in deciduous forest. I assume bats visit the probably nocturnal flowers, but there are no direct records. 1. Pajanelia longifolia (WILLD.) K.Scu. in E. & P. Nat. Pfl. Fam. 4, 3b (1895) 244; STEEN. Bull. Jard. Bot. Btzg III, 10 (1928) 267; ibid. 12 (1932) 164, f. 2 (map). — Pajaneli RHEEDE, Hort. Mal. 1 (1678) 79, t. 44, sphalma in textu t. “45”. — Bignonia indica var. B LINNE, Sp. Pl. (1753) 625. — Bignonia longifolia WILLD. Sp. Pl. 3 (1800) 306. — Bignonia pajanelia BucH. Hao. Trans. Linn. Soc. 13 (1821) 516, nom illeg., quoting WILLD. — Bignonia multi- ius WALL. [Cat. 6503] Pl. As. Rar. 1 (1830) 81, 9 ; G. Don, Gen. Syst. 4 (1838) 221. — B. multijt uga A.P.DC. Bibl. Univ. Genéve II, 17 (1838) 130: A.DC. Prod. 9 (1845) 227, excl. syn. Lour. et Pers.; Mig. Fl. Ind. Bat. 2 (1858) 758; BurEAU, Mon. (1864) 50, t. 20; Kurz, Prel. Rep. For. Veget. Pegu (1875) App. A: xciii, App. B: 69; Fl. Burma 2 (1877) 237 (‘Payanelia’); GAMBLE, J. As. Soc. Beng. 74, ii (1905) 382; Rip. Fl. Mal. Pen. 2 (1923) 549. — P. rheedii WiGur, Ic. 4 (1848) t. 1343-4; Ill. 2 (1850) t. 161bis; Bepp. FI. Sylv. 3 (1872) clxix, t. 21-5; CLARKE, FI. Br. Ind. 4 (1884) 384; BRANDIS, Ind. Trees (1906) 494; PARKINSON, For. Fl. Andam. (1923) 215. — P. bijuga, lapsus mihi, in syn. Bull. Jard. Bot. Btzg III, 10 (1928) 267. — Fig. 27, 29. Evergreen, small to large tree up to 30 m,1 m@, changing leaves before flowering. Innovations resinous. Twigs terete, thick, lenticellate. Leaves l1-pinnate, 8-12-jugate, 40-120 cm; petiole thick, to 15cm, rachis sharply keeled above, rounded beneath; leaflets entire, oblique, ovate-oblong, speed, chartaceous, 10-20 by 5-8cm, 3-5 mm stalked; nerves anterior side 9-12, posterior 6-9 pairs; beneath with scattered or heaped crateriform glands along the midrib. Thyrses up to 1m, coarse, rachis hollow, lcm @; pedicels 2'/,cm; bracteoles minute. Flowers with a soap smell, se dy tle inside dull-purple shaded. Calyx densely covered with microscopical glands and besides with scattered, rimmed, dish- shaped, large glands; splitting into 5 irregular, crispately-crenulate, acute lobes, tube at base with keel-like folds, persistent. Corolla 5—7'/, cm, thick, + constricted above base, pubescent except at base; lobes imbricate in bud, in anthesis reflexed, broad-obovate, crenulate and crispate; anthers brown to nigrescent, cells divergent. Style lon with a 2-lobed, clavate stigma. Capsule sti 30-45 by 5-9cm (incl. the 2-3cm wide, often splitting wings), with a dorsal ridge, and a corky margin on which the wings; septum quadrangular in CS, contracted in the middle, corky. Seeds in © rows on each margin of the septum, curved, 24/4-3 by 3/,-11/, cm (incl. the hyaline wings). Distr. As the genus. Erroneously recorded for the Philippines by F.-VILLar, Nov. App. (1880) 150. Rarely planted (Atjeh). Fig. 28. Fig. 28. Range of the genus Pajanelia A.P.DC. Ecol. Lowland primary and secondary rain- forest, in the Natuna Is. common, with plenty of seedlings, spared in coconut stands, elsewhere scattered, riverbanks, etc., mostly coastal, from sea-level to 100 m; in the Ghats recorded to 700 m altitude. Fi. Jan.-April, Aug., fr. March-April, ug. Indian authors cite it as being evergreen, but in the Natuna Is. I found it shortly deciduous. Kurz found it in Burma an evergreen tree. He recorded it from the Andaman Is. as ‘very abundant in the leafless jungles’, that is: monsoon forest (Rep. Veg. Andam., Is. 1870, 12, 43, 71). PARKINSON gave no definite clue on the leaf change. 164 FLORA MALESIANA {[ser. I, vol. 82 Fig. 29. Tree of Pajanelia longifolia (WILLD.) K.Scu. showing the sparingly branched, + pachycaul habit; behind is a limestone hill with dry evergreen forest (photogr. CoRNER, 1935, Sg. Sedili). The calyx contains water in bud, like other coarse-flowered members (Spathodea, Oroxylum, etc.). Uses. GAMBLE (Manual Ind. Timb. ed. 1922, 517) said it is a good timber with close-grained wood. PARKINSON /.c. found it common in the Andaman Is. where it is used for canoes, planking, and boat-building; timber smells like teak and seems to withstand attacks of white ants. Kurz (1870) recorded the largest tree with a trunk of 2m @. Inthe Natuna Is. it is estimated for building boats. In the latter islands a decoction of leaves is used against fever; in Malaya a hot fermentation is applied on the body for stomach disorders (BURKILL, Dict. 1935, 1623). Vern. Békak gunong, bongli, kaju bonglai, M, Malaya, kaju sémua, Natuna Is., abeuéng laut, Meulaboh, Atjeh. Note. For obscure reasons BEDDOME (/.c.) found this anomalous in the family, its flower reminded of Jasmineae; but he also included Schrebera (Oleaceae)! 1977] BIGNONIACEAE (van Steenis) 165 13. NEOSEPICAEA DrELs, Bot. Jahrb. 57 (1922) 500, f. 1; STEEN. Thesis (1927) 899; Bull. Jard. Bot. Btzg III, 10 (1928) 216; Nova Guinea n.s. 8 (1957) 173; ibid. Bot. n. 3 (1960) 15. — Haussmannia F.v.M. Fragm. 4 (1864) 148, non Hausmannia DUNKER, 1846; K.Scu. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 223; STEEN. Thesis (1927) 901. — Nyctocalos subg. Haussmannia SEEM. J. Bot. 8 (1870) 149. — Pandorea sect. Leptophyllae STEEN. Thesis (1927), 841, in clav.; Bull. Jard. Bot. Btzg III, 10 (1928) 200. — Pan- dorea sect. Grandiflores STEEN. Nova Guinea 14 (1927) 301, in clav., pro parte. — Tecomanthe sect. Aurantiacae STEEN. Thesis (1927) 872, in clav.; Bull. Jard. Bot. Btzg II, 10 (1928) 203. — Haussmannianthes STEEN. Proc. R. Soc. Queensl. 41 (1929) 50. — Fig. 30, 37c. Large lianas. Twigs with a distinct gland-field on the nodes. Leaves digitately compound; leaflets 3-5, sessile or short-stalked, only articulated at the base of the petiolule, mostly unequal, terminal one largest, often with a metallic hue above (s.s.), both faces with numerous microscopical glandlets, underneath besides with Fig. 30. Neosepicaea viticoides Die.s. a. Habit, */,, b. LS of flower, showing also disk and staminode, x 2,¢. CS of ovary, x 12. — N. leptophylla (BL.) Stren. d. Just opened capsule and one valve from inside, x 1/,, €. two , * "/,, f. flower, x */, (a-c LEDERMANN 9809, after Diets, d-e Beccari PP 687, f D. BERGMANN 261). 166 FLORA MALESIANA [ser. I, vol. 8? few, flat, scattered, larger, round glands '/,—'/, mm @. Thyrses co-flowered, ter- minal, axillary to ramiflorous. Calyx proportionally small, cupular to campanulate, truncate and minutely toothed, or shallowly 5-lobed by tearing. Corolla short- campanulate or more often + curved, narrow trumpet-shaped, glabrous, or out- side at least the lobes puberulous-papillose ; lobes deltoid, valvate, tomentose on the inner margin, lobes or tube inside sometimes with larger hairs; tube with a dense ring of long hairs at the insertion of the stamens. Stamens 4, didynamous, exserted, 5th rudimentary; anther-cells divaricate. Disk cupular, enveloping the ovary base. Ovary glabrous; style exserted; stigma 2-lamellate, 2-celled, with 2 placentas in each cell and c ovules. Capsules stipitate, c. 10-20 cm long, narrow oblong, terete to broad-ellipsoid in section, beaked; valves boat-shaped. Seeds co, thin-winged, rectangular. Distr. Queensland (1 sp.) and Malesia: New Guinea (3 spp.). Fig. 31. Ecol. Rain-forests, from the lowland up to c. 2000 m. Note. In the Campsis-alliance possibly closest related to Pandorea, different by the almost regular flowers, the valvate corolla lobes, the exserted stamens and style, digitate leaves, and cupular disk clasping the base of the ovary. KEY TO THE SPECIES 1. Corolla brown, basal tube, upper tube and lobes all about '/, cm long; lobes inside bearded, tube on one side so 1. N. viticoides 1. Corolla red or orange, at least 3 cm long, the tube many times as long as the lobes. 2 Gorolasorances | Oem longuncl: the lobese a ac . 2 h° Bs a ze Pliocene Symplocos subg. Hopea AY) - : a oO) MIGCEne teense Symplocos subg. Symplocos e Oligocene + Eocene es 1. ———— Fig. 1. Range of the genus Symplocos, recent and fossil. The fossil localities in Europe, Japan, and E. North America are all belonging to species of subg. Hopea. There is no species common to the Old and New World, but the E. Asian S. lucida is closest allied to the N. American S. tinctoria. Taxonomy. BRAND (1901) has made an intricate subdivision of the genus, partly based on former generic names. I believe we cannot go further than a subdivision into two subgenera, in which macromorphology is supported by chemotaxonomy and palynology, viz subg. Symplocos and subg. Hopea. Subdivisions could be based on one important single character: straight versus curved embryo, spiral versus distichous phyllotaxis, continuous versus flushwise growth from scaly buds, but it appears that such subdivisions do not coincide. This leads to the view that there is a block of species with reticulate affinities. This view also emerges from the palynological results. 1977] SYMPLOCACEAE (Nooteboom) 207 Both subgenera occur in the New and the Old World; subg. Symplocos, which is almost strictly tropical, possesses only 2 spp. in Indo-Malesia, but probably many more in America. In this revision 58 spp. are distinguished in Malesia; there are more new species, but I have refrained from describing them as the material is incomplete; I have enumerated them in my revision /.c. 296. Fossils. Before the Glacial Epoch Symplocos occurred also in Europe in the mixed mesophytic subtropical to warm-temperate forest, onwards of the Eocene, obviously as a common con- stituent of the Tertiary mixed mesophytic forest, as shown from fossil stones. Cf. KIRCHHEIMER, Palaeontographica 90B (1949) 1-52, t. 1-2. These stones are very similar to endocarps of recent species; obviously no major changes did occur in the genus during this era. The three fossil Pliocene species in Japan are almost certainly the same as those that are living there today. One fossil species is known from the Eocene in the eastern U.S.A. Fig. 1. Ecology. All species are evergreen, except a single deciduous one, S. paniculata (THUNB.) Mia. from Kashmir to Manchuria and Japan. They grow under tropical to temperate conditions in mixed evergreen rain-forest, not under arid conditions. Their stature is mostly small and they make part of the undergrowth and lower storeys, in exceptional cases attaining a maximum height of c. 30m and 60cm @. In Malesia they are found from sea-level up to the alpine zone at c. 4000 m (Mt Kinabalu; New Guinea), where they are represented by mostly microphyllous (fig. 12) dwarf shrubs in the dense elfin and mossy forest on slopes, summits and ridges where they may be common; but they are almost nowhere recorded as a dominant. A few species, e.g. S. polyandra, are restricted to the lowland, but most species have a fair altitudinal range, and are most commonly collected in the hill and mountain forest. A few are restricted to high altitude, e.g. S. buxifolia, S. deflexa, S. johniana, S. zizyphoides, and several varieties of S. cochinchinensis. A fair number seem to be rare and have been seldom collected, others are common and widely distributed in the archipelago, notably S. cochinchinensis, S. celastrifolia, S. fasciculata, S. laeteviridis, S. ophirensis, and S. odoratissima. Especially these species, several of which are variable, grow on a variety of soils, including young-volcanic; they are scarce on limestone and generally prefer more acid, humous soils, e.g. S. celastrifolia is common in coastal forests, especially in the transition between mangroves and freshwater swamps, but it occurs also on kerangas, along river banks, and even in peat swamp forest. S. cochinchinensis var. sessifolia is very resistant against poisonous crater gases and acid soil conditions and can act as a pioneer in crater fields in Java, sometimes dwarfing down to very small size, although still producing flower and fruit; in the surrounding closed elfin forest it is a common small tree, growing together with Vaccinium, Myrica, Myrsine, Leptospermum, etc. Density of species. In fig. 2 the density of species has been indicated for each province and island (group). The richest areas are those of continental SE. Asia and West Malesia, while the number of species tapers out towards East Malesia and the SW. Pacific. The greatest number of endemic species is found in West Malesia, notably (as usually) in Borneo and the Philippines. However, in East Malesia New Guinea has a fair number of endemic species. The high number of endemics in New Caledonia is a bit exaggerating the situation as all are certainly derivatives of S. cochin- chinensis. The same holds for the endemics of New Guinea (with the exception of S. cylindracea) and for Australia (with the exception of S. cyanocarpa C. T. Wuire). Flower biology. \n all Symplocos spp. the flowers of an inflorescence open almost simultaneously and on one tree almost all inflorescences are open at the same time, so that the whole crown is for a short time gay with the blossoms (fig. 3). Of S. cochinchinensis var. sessifolia flowers are deliciously scented, as hawthorn, but field records mention other species as scentless or faintly scented. This varies obviously with the species. Pollination. Docrers VAN LEEUWEN (Verh. Kon. Ak. Wet. A’dam sect. 2, 31, 1933, 218) reported of S. cochinchinensis var. sessifolia, on the summit of Mt Pangrango, West Java, at c. 3000 m, that flowers expand in the morning but open only halfway, the corolla remaining bent over the sexual organs; at 8 h. anthers are open and often touch the stigma on which the sticky 208 FLORA MALESIANA [ser. I, vol. 82 Fig. 2. Density of species in Old World Symplocos; above the hyphen the endemic species for each island (group) or country, below the hyphen the non-endemic species. Fig. 3. Symplocos laeteviridis STAPF var. laeteviridis in full flower, showing also alternate phyllotaxis.— Sabah (Nooresoom 1017). Photogr. NooTEBOOM, Febr. 1969. 1977] SYMPLOCACEAE (Nooteboom) 209 pollen readily falls; on the 2nd flowering day the corolla is widely open, anthers are empty, and the stigma is always pollinated. This means self-pollination. DocCTERS VAN LEEUWEN found, however, also the flowers frequented by various insects, among them bees and bumble-bees. They are not so much attracted by the little nectar, but are in search of pollen. Some species may have locally a strict flowering time; e.g. S. cochinchinensis var. sessifolia flowers, according to DOCTERS VAN LEEUWEN (i.c., fig. 52), from October to January, in the rainy season, on the summit of Mt Pangrango, West Java. Hybridization. Though there are in a few instances indications (by high sterile pollen °%) that hybridization may occur, no clear cases are recorded. It is, however, clear that cross-fertilization must occur in the polygamous species in East Malesia. Galls. DOCTERS VAN LEEUWEN (Zoocecidia, 1926, 460) found in S. cochinchinensis var. sessi- folia small leaf galls, caused by psyllids by which the two halves of the leaf curve upwards till margins touch and a narrow cavity is formed. He recorded similar galls also from other forms of this species. In S. fasciculata he found a stem gall caused by a gall-midge and in S. brandisii a flower gall caused by a gall-midge. Dispersal. RipLey (Disp. 1930) assumed that bats may be fond of the hard-fleshed drupes (l.c. 347). He mentioned that in North America tyrant birds (Sayornis phoebe) eat amongst others fruit of S. tinctoria (1.c. 483) and that in South America a curassow, a sort of turkey, would feed on the fruit of S. cernua. DOCTERS VAN LEEUWEN (Verh. Kon. Ak. Wet. A’dam sect. 2, 31, 1933, 220) believed Symplocos to be dispersed by birds but did not find endocarps in the stomach of fruit-eating birds. VAN STEENIS found fruit of S. henschelii abundant on the ground below trees at Tjibodas, although this species has a fairly thick, hard-fleshed exocarp, in contrast to most species in which the exocarp is thin. Also in fossils sometimes immense quantities of stones are found together, about which KIRCHHEIMER reported (Palaeontographica 90B, 1949, 1-52): ina total mass of c. 3500 m? he estimated the number of endocarps at some 2?/, billions. He assumed that these were deposited within one century in a site of forest dominated by Symplocos. However, he added that the layers in which the endocarps were deposited gave no evidence of rivers which could have transported and accumulated the seeds and he concluded that they have dropped to the soil in situ. For these reasons abundant dispersal by birds or bats is in Symplocos not very likely. Dispersal by water takes place in species in which some fruit cells are barren and remain empty, e.g. S. celastrifolia. Morphology. The phyllotaxis is variable but constant for the species; it is either spiral or alternate (distichous) in which latter case the twigs are often zigzag (fig. 3). In most species leaves are more or less equally dispersed along the twigs, but in other species there is a tendency that the leaves are becoming crowded towards the end of the year’s growth, e.g. in S. macrocarpa, as noted by TRIMEN (Handb. FI. Ceyl. 3, 1895, 103). In Malesian spp. this occurs also in S. herzogii and S. gigantifolia where the large leaves occur crowded at the end of the year’s growth. There is a single species in which all the leaves are in real pseudo-whorls, viz S. verticillifolia from the Philippine Is. (fig. 20). The leader-shoots in Symplocos, e.g. S. fasciculata, have spiral phyllotaxis; such shoots may, however, also carry flowers. Rejuvenation is in certain species by continuous growth of the twig apex, as is e.g. charac- teristic in S. fasciculata. In other species, however, there are clear buds with conspicuous bud- scales, indicating that the growth mode is flushwise and discontinuous, as e.g. in S. costata and S. lucida (fig. 15). This might be a good character of subdividing subg. Hopea. It can, however, only be used if one has accurate knowledge of the rejuvenation process of each species. This is sometimes difficult to ascertain from herbarium material as the bud-scales do not always leave traces of distinct scars, field data hardly ever mention the character, and material is seldom collected in the stage of flush. If the growth mode were well examined in all species I believe it would represent a good key character. Flushwise, discontinuous growth, with scaly buds could be assumed to be an adaptation to seasonally cold climates. It is a life form intermediate halfway evergreen and deciduous. It is rare in the Malesian tropics where it is known e.g. from Acer, some genera of Lauraceae, Fagaceae, 210 FLORA MALESIANA [ser. I, vol. 8? which also in the tropics are found in the cool, tropical-montane climate, which is however hardly seasonal. It still could be viewed as an indication of former immigration of taxa of higher latitude. Once acquired this growth mode must then have been conserved, as it occurs also in S. barring- toniifolia which is restricted to the tropical lowland. The inflorescence is either a panicle or a raceme or spike. Morphologically it is cymose, the flower always being sustained by two bracteoles which may at times carry abortive buds in their axil (fig. 11b). In some cases the inflorescence is condensed to a fascicle or cluster of flowers (fig. 20a) or even be reduced to a single flower (fig. 19a). In a few species flowers occur on old wood, as e.g. in S. polyandra, S. wikstroemifolia (p.p.), S. rubiginosa, and S. tricoccata. The flowers are bisexual but functionally unisexual flowers are found in several taxa, especially in New Guinea. Such taxa are either dioecious or polygamous. In male flowers the style is small and without a stigma, in female flowers the number of stamens is reduced (even to less than 10) and anthers are sterile. In subg. Symplocos the stamens are monadelphous with a long tube (fig. 6a, d); in subg. Hopea they are only connate at the base for at most 2 mm (fig. llc), and intergrading from strictly monadelphous to strictly pentadelphous, the phalanges being alterni- petalous. In my revision it has been explained that, in contrast with former opinion, the ovary is initially 1-celled, with the ovules attached close to the centre on the induplicate part of the carpels, each of the 2-5 compartments having usually 4 ovules; in fruit these appear as cells. In each developed cell there is usually one seed. The latter and the embryo it contains may be curved or straight. See fig. 4b, c, g, h, j, k. Fig. 4. Symplocos ophirensis CLARKE ssp. perakensis (K. & G.) Noor. var. perakensis. a. LS of fruit, out of centre, b. seed, c. curved embryo, with 2 short apical cotyledons, all x 4. — S. ophirensis CLARKE ssp. cumingiana (BRAND) Noor. var. cumingiana. d. LS of fruit, seed cavity empty, e. stone, x 4. — S. macro- phylla WALL. ex DC. ssp. cordifolia (THw.) Noor. var. apicalis (THw.) Noort. f. Ribbed stone, with fold, g. seed, the curved embryo enveloped by the albumen, x 2. — S. paniculata (THUNB.) Mia. h. LS of seed showing curved embryo, h’. ditto in CS, showing how such seed may appear deceptively as 2 seeds, x 11/5. — S. glauca (THUNB.) Koipz. i. Fruit, j. seed in LS showing straight embryo, x 3. — S. paniculata (THunB.) Mig. k. Germinating seedling with LS of endocarp and seed, showing mode of exist of embryo, x 11/3, 1. seedling, x 2/,. — S. celastrifolia GRIFF. ex CLARKE. m. U-shaped seed, x 6 (a-c BuRKILL 1013, d-e NooTeBooM 2229, f-g ASHTON 2480, k—/ after LuBspock). — alb albumen, ec stony endocarp, mc mesocarp, os outer surface of fruit, sc seed cavity. 1977] SYMPLOCACEAE (Nooteboom) 211 The fruit is a drupe, with a fleshy, corky or woody mesocarp and a very hard stone (endocarp). The endocarp may be smooth (fig. 10c, 19d) or show outside ridges or irregularities (fig. 4e, f, 9c, 10e, 14d); the same holds for the inside of the endocarp. In the centre of the copious endosperm the embryo is embedded. It is slender and may be straight or curved. In the tropical subg. Symplocos it is always straight. In subg. Hopea it is straight in all American spp. and in 80% of the living species in the Old World and also in all fossil species in Europe. From this it is con- cluded that a straight embryo seems to be the primitive state in the genus. Only the three Pliocene fossil species of Japan, which can be matched with living species, have curved seeds and conse- quently curved embryos. Curved seeds occur in degree, they may be hook-shaped or U-shaped or even be twice curved (S-shaped in S. brachybotrys). See fig. 4. This may give some difficulty in studying sections of the stones to count the number of seeds in a fruit (e.g. fig. 4h-h’). Although of the living species only 20% have curved seeds the vast majority of the individual living plants have curved seeds; so it seems that this probably recent trend in the evolution of the genus was successful although the reason for its origin and advantage of its function remains obscure. Seedlings. Few observations are made. LussBock (Contr. Knowl. Seedlings, 1892, 206-208, fig. 509) noted for S. paniculata (sect. Hopea): the endocarp does not burst during germination; the radicle emerges by a small hole at the apical narrow end; the hypocotyle elongates, becoming curved, finally straightening, carrying up the endocarp containing the embryo. As the cotyledons elongate, they push out at the small hole in the endocarp (so to say throw the latter off), and finally get free and spread out to the light; they enlarge but remain narrow. The first two leaves are opposite, hairy on both sides and serrulate which may persist in leaves of saplings (fig. 4l). Spot-characters. In the herbarium a Symplocos of subg. Hopea can mostly easily be spotted by spiral, exstipulate, eglandular, serrate or crenate leaves discolouring pale greenish or yellowish or greenish-brown, a feature connected with a high Al-content of the tissues. At a very young stage, the just expanding leaves have proportionally conspicuous gland-like teeth on the margin. A significant character is that in the herbarium the midrib is always sulcate above, with the exception of 4 spp. in which it is prominent: S. anomala, S. lancifolia, S. lucida, and S. wikstroemifolia. The cup-like 3 bracts (of which 2 bracteoles) below the flower (fig. 11b) is also characteristic as is the inferior ovary and fruit. Innovations and newly expanded leaves are in many species a beautiful violet, afterwards changing into violet-brown while the drupes are often blue to black-violet, features found in many aluminium-accumulating plants (Eurya, Helicia, etc.). Anatomy. For general surveys also covering the older literature, see SOLEREDER, Syst. Anat. Dicot. Stuttgart (1899) 587-589 (under Styracaceae) and ibid. (1908) 208-210; METCALFE & CHALK, Anat. Dicot. Oxford (1950) 890-893. Selected references: JANssONIUS, Mikr. 4 (1925) 471-498 (wood anatomy); DEN BerGeR, Determinatietabel Malesié, Veenman, Wageningen (1949) (wood identification); JANssonius, Blumea 6 (1950) 422-423 & 424 (wood anatomical affinities); Descu, Mal. For. Rec. 15 (1954) 591-593 (wood); ZAHUR, Mem. Cornell Univ. Agric. Exp. Stn. 358 (1959) 35 (bark anatomy); Huser, Mitt. Bot. Staatssamml. Miinchen 5 (1963) 1-48; Baas, Blumea 21 (1973) 201-216 (ecological wood anatomy); Nootesoom, Leid. Bot. Ser. 1 (1975) 20-22 (leaf and wood anatomy). The wood is characterized by the following primitive set of characters: Vessels solitary and with many-barred scalariform perforations. Fibre-tracheids with conspicuously bordered pits on both radial and tangential walls. Parenchyma diffuse or diffuse-in-aggregates. Rays hetero- geneous, usually of two distinct sizes. The bark is also of a primitive type with compound sieve plates. Mechanical bark tissue is poorly developed and composed of groups of sclereids (ZAHUR, Lc.). The leaf anatomy exhibits few constant characters such as paracytic stomata, clustered crystals and dorsiventral mesophyll. Presence or absence of a hypodermis, of idioblastic leaf sclereids, of a complex vasculation pattern in the midrib, and of an indumentum varies. The diagnostic and systematic value of these characters remains to be assessed. The anatomical evidence is inconclusive with respect to a positive indication of the closest affinities of Symplocaceae. The traditional treatment of the family as a member of the Ebenales 212 FLORA MALESIANA [ser. I, vol. 8? close to Styracaceae must, however, be refuted. The anatomy is more compatible with suggestions of a Cornalean or a Thealean alliance as advocated by NOOTEBOOM I.c. Palynology. The palynology of the Old World spp. was examined by R. VAN DER MEIJDEN (Pollen et Spores 12, 1970, 513-551, 1971, suppl. in my Monograph, 1975, 9-15). The essential results are the following: the two main pollen types coincide with the distinction of the two sub- genera. In subg. Symplocos there are two minor types, one belonging to the Old World spp., the other to those of America. In subg. Hopea there are 9 subtypes, but none is apparently peculiar to American spp. The distribution of these subtypes is rather complicated and leads to the view of reticulate relation- ship, which agrees with the impression gained from macromorphology. Another feature is that within the variable species several subtypes are represented, and furthermore that a number of subtypes are found in species which are taxonomically not closely related. There is no agreement between the shape of the embryo, straight or curved, and pollen subtypes. Echinate pollen is found in the Philippine S. whitfordii and in the East Malesian and Pacific varieties of S. cochin- chinensis ssp. leptophylla; also the 9 endemic spp. of Symplocos in New Caledonia which are all related to this subspecies have echinate pollen. In several taxa a certain amount of pollen is sterile and I have ascribed this to hybridization. Phytochemistry. Many species of Symplocos, especially from subg. Hopea, contain alu- minium compounds, a feature which manifestates itself in the yellow colour of dried leaves. Especially when the plants are dried after having been conserved in alcohol vapour according to the Schweinfurth method, the yellow colour becomes very intense. The yellow colour is the result of a reaction of aluminium compounds with flavonols in the drying leaf. The amounts recorded in literature vary between 0.05 and 4.2% of dry weight of the leaves; barks may contain similar amounts of aluminium (CHENERY, Kew Bull. 1948, 173-183; Analyst, 1948, 501; NooTEBOoM, Leid. Bot. Ser. 1, 1975, 19). RADLKOFER (Ber. Deut. Bot. Ges. 22, 1904, 216-224) already men- tioned that the ash of Symplocos leaves contains c. 50% aluminium oxide. He also described the so-called ‘‘Tonerdek6rper”’ in the leaves of Symplocos. These are masses of colourless material filling often large parts of the cells, predominantly in the palissade parenchyma. According to RADLKOFER these masses consist mainly of aluminium compounds. KRATZMANN (Sitz. Ber. Ak. Wiss. Wien, 1913, 311-336) found that these aluminium bodies also contain much other material, for instance silicates, and that the aluminium is also accumulated in other parts of the leaf. NEGER (Flora N.F. 16, 1923, 326-330) observed that the development of plants of Symplocos lucida (THuNB.) S. & Z. depends on the amount of aluminium compound in the solution they are culti- vated on. Plants grew best on a solution containing 1 promille aluminium. Besides aluminium many other compounds are found (HEGNAUER, Chemotaxonomy der Pflanzen 6, 1973). The more important are: 1) Phenolic compounds (see also BATE SmiTH, J. Linn. Soc. Bot. 58, 1952, 95-173). Gallic and ellagic acid seem to be rather common. Leucoanthocyanins occur in varying amount. BATE SMITH /.c. also found quercetin, and caffeic acid. In the bark of S. /ucida (THUNB.) S. & Z. the lignan glycoside symplocosin has been found, and traces of methylsalicilate were demonstrated in the bark of several species. True tannins were not yet found in Symplocos. 2) Alkaloids. Only for two species structurally known alkaloids were described. More research is needed. 3) Saponins. In several species saponin-like compounds were found, as well in the bark as in the leaves. — R. HEGNAUER. Chromosomes. In my monograph I have given an account of chromosome numbers, which are unfortunately too few. However, the majority is n = 11, with some deviations; rarely 2n = 24, and one count of the North American S. tinctoria of 1n = 14, all in subg. Hopea. The one count known of subg. Symplocos in Malesia yielded 2n = c. 90 (2n = 88 would fit an octoploid). It would be too rash to conclude that polyploidy would be normal in that subgenus. On the other hand it may tentatively be concluded that species in subg. Hopea are diploid, with x = 11. This does not fit the numbers found in other Ebenales families, nor in Theaceae, but it does agree with Cornaceae sens. lat. Uses. As timber Symplocos has no great value, according to HEYNE (Nutt. Pl. 1927, 1262). Leaves and bark of Symplocos contain a fair amount of alum, both in Asian and American spp. (cf. Ber. Deut. Bot. Ges. 22, 1904, 126). This was commonly used, mostly from decoctions of the bark, in dyeing processes (red and brown), e.g. in the batik industry in Java. RuMpPuius already 1977] SYMPLOCACEAE (Nooteboom) 213 mentioned this use from the Moluccas. Several species were used for this purpose, e.g. S. cochinchinensis, S. fasciculata, S. odoratissima (HEYNE, /.c.). The same compound is probably also the constituent active in medicinal uses against so-called sprue (‘thrush’) as ‘obat seriawan’. Notes. Identification of material of Symplocos is for several reasons far from easy. Because of simultaneous flowering flowers and fruits are practically never found together and both stages are properly needed. Only few species possess well definable vegetative characters. Moreover, a few widespread species have proved to be rather variable, to a fair degree by racial differentiation. These are the reasons that besides a general key in which all characters are used, I have found it useful to add a number of partial keys for islands or island groups in a double series, either for fruiting or for flowering material. In fig. 5 a scheme is given elucidating the way in which for this genus descriptive terminology is used in the keys and descriptions. Fig. 5. Schemes elucidating descriptive terminology used in the text. — A: a length of leaf, 6 width of leaf, c divided by b is leaf index, d length of acumen. — B: way of expressing base angle «. — C: deflorated flower; br bract, bra bracteole, c length of calyx, c/ length of calyx lobes, ov height of ovary. KEY TO THE SUBGENERA 1. Petals connate at least halfway up. Leaves usually not becoming yellow when drying, not discolouring, spirally arranged, entire. Flowers very fragrant. Seeds and embryo straight. Spp. 1-2 1. subg. Symplocos 1. Petals connate only at the very base. Leaves usually becoming more or less yellow or greenish yellow when dried. Leaves spirally arranged or distichous, exceptionally in pseudowhorls. Flowers not or mostly only faintly fragrant. Seeds and embryo straight or curved. Spp.3-58 . . . . 2. subg. Hopea 1. Subgenus Symplocos Cf. Noor. Leid. Bot. Ser. 1 (1975) 36. — Cordyloblaste Mor. Bot. Zeit. 6 (1848) 606; RipL. Fl. Mal. Pen. 2 (1923) 307; ALston, Handb. FI. Ceyl. 6 (Suppl.) (1931) 186. — Symplocos sect. Cordyloblaste B. & H. Gen. Pl. 2 (1876) 669; BRAND, Pfl. R. Heft 6 (1901) 88; Steen. Bull. Bot. Gard. Btzg III, 17 (1948) 429. — Symplocos subg. Cordyloblaste GAMBLE, J. As. Soc. Beng. 74, ii (1906) 248. — Fig. 6. Leaves usually not becoming yellow when dry. Corolla tubular, erect, often to above the middle adherent to the staminal tube and then suddenly expanded; 214 FLORA MALESIANA [ser. I, vol. 8? margins of the petals free, thus sometimes obscuring the coalescence. Stamens monadelphous; free part of filaments ribbon-shaped, in several whorls, in the outer whorl often very short, always suddenly attenuate below the anther. Fruits 2-5- celled, usually none of the cells aborted. Seeds straight, cylindrical. Distr. Tropics of Indo-Malesia and South America, largely within 30° N and S, more than 100 spp. described from the New World, in Malesia 2 spp. Fig. 1. Ecol. Rain-forest, from the lowland up to c. 3300 m (Mt Kinabalu). KEY TO THE SPECIES 1. Calyx c. 6(-10) mm long. Corolla 21/,-5 cm long. Fruits 3-Scmlong. ..... . 1. Calyx 3-5 mm long. Corolla */,-13/, cm long. Fruits 1-1'/, cm long 1. Symplocos henschelii (Mor.) BrH. ex CLARKE, FI. Br. Ind. 3 (1882) 588, quoad nomen et basionym, excl. stirp.; BRAND, Pfl. R. Heft 6 (1901) 89; Bull. Herb. Boiss. II, 6 (1906) 750; Koorp. Atlas 2 (1914) t. 390; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 440, f. 2a—l; Nova Guinea n.s. 10 (1959) 210; Back. & BAKH. f. Fl. Java 2 (1965) 204; STEEN. Mt. FI. Java (1972) pl. 52-3; Noor. Leid. Bot. Ser. 1 (1975) 37, pl. 1g. — Cordyloblaste henscheli Mor. Bot. Zeit. 6 (1848) 606. — Eugeniodes henscheli O. K. Rev. Gen. Pl. 2 (1891) 975. — S. nageli K. & V. Bijdr. 7 (1900) 159. — S. scortechinii KING & GAMBLE, J. As. Soc. Beng. 74, ii (1906) 250. — Cordyloblaste scortechinii Rip. Fl. Mal. Pen. 2 (1923) 309. — S. dolichantha MeErR. Sar. Mus. J. 3 (1928) 545. — S. stenosepala STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 444, f. 2 m—n. — Fig. 6a—c. For further synonyms see under the variety. Shrub, or mostly a tree, to 30 m; innovations glabrous to grey or rusty velvety. Leaves glabrous, sometimes the midrib above and underside hairy, 7-17(-22) by 3-7!/,cm; petiole */,-11/,(—-2) cm. Racemes up to 10 cm, incl. bracts and flowers grey or rusty tomentose, short-peduncled, 1-12- flowered. Bracts narrow-triangular; pedicels 0-6 mm, with 2(-3) tiny bracteoles. Calyx lobes rounded to triangular, mostly erect, 1—4'/, by 2-3 mm, persistent. Corolla sericeous (in Mal.), club-shaped in bud, 2!/,-5 cm, connate for 3/,—%/,, tube 3-4 mm @, lobes spathulate. Staminal tube 1/,cm shorter than corolla, adnate to the corolla tube except towards apex, free part 1/,-1'/, cm; anthers 20-110, filaments unequal. Ovary 3-4- celled; ovules 2-4 per cell, usually only 1 develop- ing. Fruit obovoid to spindle-shaped, 3-5 by 2-3 cm; mesocarp thick, hard-fleshy to + woody. Distr. Continental SE. Asia (Burma, Thailand, Indo-China) and West Malesia (Sumatra, Malay Peninsula, W. Java, Borneo), a distinct subspecies in Thailand. Note. Additional material has shown that S. stenosepala STEEN. cannot be upheld and, more- over, that S. maingayi CLARKE deserves only varietal rank. KEY TO THE VARIETIES 1. Leaves and twig ends usually glabrous. Free part of staminal tube 7-15 mm. a. var. henschelii 1. Twig ends and leaves underneath hairy. Free part of staminal tube 5-7 mm b. var. maingayi 1. S. henschelii 2. S. pendula a. var. henschelii. — Fig. 6a—b. Shrub or tree, up to 25 m, 45 cm @. Twigs glab- rous, the youngest ones sometimes more or less grey or rufescent appressedly pubescent to velvety or tomentose. Leaves glabrous, or the midrib beneath sparsely short fine-hairy, rarely with same indument as var. maingayi. Free part of staminal tube 7-15 mm; anthers (40—)55—-75(—110), in the upper 5-10mm, ascendent and nearly sessile above to descendent on a slender filament below, the lowest ones hanging from a 2-5 mm long fila- ment. Fruit with + fleshy mesocarp. Distr. As the species. Ecol. Below 1100 min mixed dipterocarp forest, also once in swamp forest, and on podsol (Kalabit), at higher altitude in oak-chestnut mountain forest, also on ridges and in mossy forest, 600-2000 m (in continental SE. Asia at 130-800 m). F/. Jan.—Dec., fr. Febr.—Sept. Vern. Sumatra: kayu djaram-djaram bosi, Batak; Borneo: té baradang, Sarawak, Kalabit, yum, Kenyah lang., /amau-lamau, Brunei. b. var. maingayi (CLARKE) Noor. Leid. Bot. Ser. 1 (1975) 39. — S. maingayi BTH. ex CLARKE, FI. Br. Ind. 3 (1882) 588; BRAND, Pfl. R. Heft 6 (1901) 90; K. & G. J. As. Soc. Beng. 72, ii (1906) 249; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 445. — Euge- niodes maingayi O. K. Rev. Gen. Pl. 2 (1891) 975. — Cordyloblaste maingayi Rip. Fl. Mal. Pen. 2 (1923) 309. — Fig. 6c. Tree up to 21 m, 40cm @. Twigs densely rusty tomentose or velvety, glabrescent. Leaves sparsely fine-hairy beneath, especially on midrib and nerves, to greyish tomentose or velvety. Free part of staminal tube 5-7 mm; anthers 20-60, in the upper 5 mm, ona very short (!/,—1/, mm) thin free part of the filaments. Fruit with -- woody mesocarp. Distr. Malesia: Malay Peninsula and Borneo (Sarawak, Brunei). Ecol. Evergreen primary and depleted lowland forest, 15-150 m; in Borneo often on low sandy ridges, raised beaches, and large sandy podsols (kerangas). F/. April-May, fr. Jan. 2. Symplocos pendula WiGuHT, Ic. 4 (1848) 10, t. 1237; Ill. Ind. Bot. 2 (1850) t. 151-b, 7-12; CLARKE, FI. Br. Ind. 3 (1882) 587; BRAND, Pfl. R. Heft 6 (1901) 88; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 437; Noor. Leid. Bot. Ser. 1 (1975) 40, pl. lh. — S. scortechinii (non K. & G.) RIDL. J. Linn. Soc. Bot. 38 (1908) 315. — S. pulcherrima 1977] SYMPLOCACEAE (Nooteboom) Fig. 6. Symplocos henschelii (Mor.) BTH. ex CLARKE var. henschelii. a. Flower, b. fruit, exocarp halved. — S. henschelii var. maingayi (CLARKE) Noor. c. Fruit. — S. pendula WiGut var. pendula. d. Flower. — S. pendula var. hirtistylis (CLARKE) Noor. e. Fruit. All nat. size (@ WILSON 2547, 6 after STEEN. 1972, pl. 52-3b, c KOSTERMANS 9328, d father ANGLADE s.n., e MEER 3618). Ript. J. Fed. Mal. St. Mus. 6 (1915) 160. — Cordyloblaste pulcherrima Rip. Fl. Mal. Pen. 2 (1923) 308. — Fig. 6d-e. For further synonyms see under the variety. var. pendula. — Fig. 6d. Small shrub '/,—-3 m or tree up to 27 mand 50 cm @. Twigs glabrous to rusty tomentose. Leaves glabrous or nearly so, elliptic to obovate or orbicu- lar, entire to crenate, apex rounded to acuminate, (1-)2#/,-12'/, by (1-)1'/.-6cm; nerves 4-8(-11) pairs; petiole (1-)5-15 mm. Racemes very short, sometimes flowers solitary. Bracts to 1 mm. Brac- teoles 2-4, narrow-triangular. Pedicels 0-5 mm, longer in solitary flowers. Calyx lobes very short and rounded, ciliate. Corolla tubular-trumpet- shaped, (5-)10-17 mm, fleshy, silver-white to creamy, fragrant, the petals connate halfway up, spathulate, rounded at apex, glabrous to tomentose. Staminal tube adnate to corolla except for upper 3-5 mm, hairy to glabrous inside; anthers 30-50 (-80). Ovary semi-inferior, glabrous, the apex semi-globose, c. 1'/, mm high, densely grey-hairy; style c. 1 cm, more or less hairy at the base to brous at the apex. Fruits spindle-shaped, 10-15 y 3-6cm, green pinkish red, the enlarged calyx lobes surrounding the hairy, conical, persistent style-base. Distr. Continental SE. Asia (Ceylon, Deccan, Hainan), in Malesia: Malay Peninsula. Ecol. Mountain forests and open heath and scrub, often on ridges, 600-1750 m. Fl. Febr.- March, /r. Sept. var. hirtistylis (CLARKE) Noor. Leid. Bot. Ser. | (1975) 42, f. 2a, with full synonymy. — S. henschelii (non BTH.) CLARKE, FI. Br. Ind. 3 (1882) 588, pro stirp., incl. var. hirtistylis CLARKE. — S. confusa BRAND, Pfl. R. Heft 6 (1901) 88; Bull. Herb. Boiss. II, 6 (1906) 750; K. & G. J. As. Soc. Beng. 74, ii (1906) 248; BRAND, Philip. J. Sc. 3 (190%) Bot. 3; Me_rr. En. Philip. 3 (1923) 297; Steen. J. Arn. Arb. 28 (1947) 423; Bull. Bot. Gard. Btzg III, 17 (1948) 432. — S. albifrons BRAND, Pfi. R. Heft 6 (1901) 88; Bull. Herb. Boiss. II, 6 (1906) 750; Nova Guinea 14 (1924) 189. — S. capitellata BRAND, Pfi. R. Heft 6 (1901) 88; Bull. Herb. Boiss. II, 6 (1906) 750; Nova Guinea 14 (1924) 188. — S. fox- worthyi BRAND, Philip. J. Sc. 3 (1908) Bot. 3; Merr. En. Philip. 3 (1923) 299. — Styrax obovatus Rip. J. Str. Br. R. As. Soc. m. 61 (1912) 8. — S. obovata Rio. J. Fed. Mal. St. Mus. 6 (1915) 51. — §. crenulata Riwv. l.c. — S. novoguineensis Gisss, Arfak (1917) 176. — Cordyloblaste obovata Rip. Fl. Mal. Pen. 2 (1923) 308. — Cordyloblaste crenulata Riwi. lc. 309. — S. atrata BRAND, Nova Guinea 14 (1924) 188. — S. topica BRAND, l.c. 189. — Fig. 6e. Ovary hairy. Distr. Continental SE. Asia (N. Burma, Indo- China, China, Japan, Formosa), throughout Malesia, except Java and Lesser Sunda Is. Ecol. Primary and secondary montane and sub- alpine forest, mossy forest, often common on ridges, or in open fern thickets (Tamrau), on sand or clay, 1500-3300 m, but in kerangas forest in Sarawak at 800 m. Fi. March—Aug. (Sept.—Febr.), Sr. Febr.—April, July-Sept. At higher altitude often a dwarf shrub with small leaves, but sometimes also a dwarf shrub with large leaves in high forest. 2. Subgenus Hopea CLARKE, FI. Br. Ind. 3 (1882) 572; BRAND, Pfl. R. Heft 6 (1901) 25; Noor. Leid. Bot. Ser. 1 (1975) 43, with full synonymy. — Hopea Linné, Mant. (1767) 105, nom. 216 FLORA MALESIANA [ser. I, vol. 8? a 000 0486000 3 3 4 8 9 10 10 1a 12 a \ ‘ 65° 0000086 14 16-1a 16-1b 16-1C 16-1d 16-2e 16-3f 16-4g 16—4g OHo00008 16-49 16-49 16-49 16-4h 116-41 16-41 16-4m 16-4mM 16-4nN 16-40 605608 0 4 6 8 o 16-4p 16-4Q 16-4rF 16-4F 16—4S 16-4t 16-4U 16-4U 16-4W 0 9 00.0()¢ 20C 24 27 28 30 31 32 33a 33eE 34 37 40 41a 41b 42-1a 42-3f 42-3f 42-3g 42-2Cc 43 0000500 Fig. 7. Fruits in outline, in the dried state. Of each fruit the voucher specimen is cited by the number of the taxon. If for showing variability more fruits of the same taxon are drawn, read from left to right corres- ponding with the voucher numbers. All drawings natural size. — 3 CHEw WEE LEK 938 — 3 CF 104879 — 4 SAN 56690 — 5 DE Wipe 13773 — 6 A. Ernst 736 — 7 KING’s Coll. 6179 — 8 MEUER 7581 — 9 45 eos 1977] SYMPLOCACEAE (Nooteboom) 217 rejic. — Dicalix Lour. Fl. Coch. 1 (1790) 663; BL. Bijdr. (1826) 1116 (‘Dicalyx’). — Sariava REINW. Syll. Ratisb. 2 (1825) 12. — Carlea Pr. Epim. Bot. (1851) 216. — Baranda Luianos, Mem. Ac. Cienz. Madrid 3, 2 (1857) 502. — Eugeniodes O. K. Rev. Gen. Pl. 2 (1891) 409, 975, nom. illeg. — Fig. 7-20. Leaves usually becoming more or less yellow when drying. Petals glabrous, or hairy in only few species, connate only at the very base, mostly expanded. Stamens monadelphous to pentadelphous, only connate at the very base (for at most 2 mm); filaments cylindrical, slender to rather stiff, often gradually attenuate towards the anther. Fruits 2—3(—5?)-celled, often 1-celled by abortion. Seeds either straight or curved, and then with curved embryo. Distr. About 150 spp., as for the genus. Fig. 1. Note. As explained in the note under the genus, a general overall key is given to all species, as much as possible based on vegetative characters and on flowering material. To facilitate identification additional local keys are given for the main Malesian islands or island groups, one each for flowering and for fruiting material. In addition in fig. 7 fruits are drawn of all species as far as available in the dried state. They have been ate thor according to the number of the taxa. The following terminology has been adopted for fruit- shapes: globose 34, 41b ampulliform 23, 42c, 43 ellipsoid 4, 19, 33e, 46 spindle-shaped 38: fig. 19d. ovoid 48 cylindrical 20c, 21 obovoid la, 1b: fig. 6c It should be observed that the shape of the stone may differ from the shape of the fruit and that for instance ovoid fruits may possess an ampulliform stone. There is no strict relation between the shape of the seed and the shape of the fruit or stone, but ampulli- form fruits have always a curved seed and curved embryo and spindle-shaped and cylindrical fruits have always a straight seed and embryo. Besides the overall-shape of the drupe, the shape of the stone can be important: sometimes it bears lower or higher ridges, which ornamentation provides good characters. KEY TO THE SPECIES 1. Leaves (pseudo-)verticillate. Zaeerpes Mae OF Jeaves glabrous, Twigs hirsute. . .. . % \ ©. °WeU ee 55. S. verticillifolia eee mice Of leaves'hairy. Twigs tomentose ... . . » » » «wemplue eeuuele 30. S. herzogii 1. Leaves not verticillate. 3. Midrib prominent on the upper surface. 4. Twigs glabrous. 5. Leaves crowded towards the end of the twigs, minutely appressedly hairy beneath 37. S. wikstroemifolia in menves evenly distributed, glabrous... . «).6) sss 50) eye ee 35. S. lucida 4. Twigs hairy. HALuieR f. 2197 — 10 CLemens 32525 — 10 CLEMENS 32478 — lla bb 23324 — 12 SAN 46543 — 14 KoOSsTERMANS 9158 — 15a KOSTERMANS & ANTA 527 — 16—la Forbes 861 — 16—1b MEWER 1690 — 16-Ic CLEMENS 17224 — 16-1d BS 4476 — 16-2e LARSEN c.s. 887 — 16-3f NGF 33643 — 16-4g ANU 2027 — 16-4g A. C. Smitu 1054 — 16-4g BW 4970 — 16—4g GILLespie 3918 — 16—-4g NGF 28481 — 16—4h VINK 17308 — 16—4i Brass 28343 — 16-41 Brass 29919 — 16-4m LEDERMANN 8946 — 16-4m T. G. HARTLEY 13135 — 16-4n PULLEN 479 — 16-40 NGF 49168 — 16-4p PULLEN 7783 — 16-4q KosTERMANS & WIRAWAN 878 — 16—4r NICOLAS 19 — 16—4r VAN BALGOoy 862 — 16-4s KALKMAN 5128 — 16-4t VINK 16079 — 16-4u KosterMAns 2375 — 16~—4u Forses P. P. 652 — 16-4w @rass 28191 — 16-4w CLEMENS 1661 — 16-4x NGF 23728 — 17 CLemens 33706 — 19 Koorpers 15596 — 20c VAN BEUSEKOM c.s. 837— 20d ENpert 2580 — 20f CF 97832 — 21 bb 22503 — 22 Jacoss 5766 — 23 SAN A2240 — 24 MERRILL 6148 — 27 BS 45592 — 28 KEP/FRI 8236 — 30 T. G. Hart.Ley 12509 — 31 Noorrenoom & ABAN 1500 — 32 HILDEBRAND 55 — 33a SAN 65017 — 33e SAN 44386 — 34 PNH 18483 — 35 BURGER s.n. — 36 Cel. 11-374 — 37 F. C. How 73506 — 40 BS 26447 — 4la Ja 7723 — 41b SAN 57045 — 42-la BurRN Murpocnu 340 — 42-3f Dinc Hou 274 — 42-3f Nooresoom 2229 — 42-3g BS 83753 — 42-2c CF 98890 — 43 Kasewsx1 1208 — 44 § 17287 — 45 Carr 12782 — 46 Rip_ey 16102 — 47a ROBINSON & KLoss 199 — 47b Mever 7665 — 48 Ismaet 9 — 50 Beccari P. S. 106 — 51 Koeiz 29538 — 52 CLemMeNns 32559 —53 S 26305 — 55 PNH 14397 — 56 BS 45675 or 45775 — 57 Jacons 7484 — 58 Nooresoom 1491, 218 FLORA MALESIANA [ser. I, vol. 8? 6. Leaves crowded towards the end of the twigs, minutely appressedly hairy beneath 37. S. wikstroemifolia 6. Leaves evenly distributed, glabrous or sparsely fine hairy beneath. 7. Underside of leaves glabrous. Corolla:4—6 nm: j x5... - ee ea 4. S. anomala 7, Underside o1 leaves hairy: ‘Corolla 2*/-4 mm iF 2 ek kt ls . 34. S. lancifolia 3. Midrib sulcate above. Pe Gea len flee la ieee gwen eee Oo ee ee re oe 41. S. odoratissima 8. Corolla glabrous. 9. Twigs hairy. 10. Underside of leaves glabrous. (When petiole and leaf margin beset with closely spaced vesicular glands: 3. S. adenophylla). fiaideaves:disuchonus Viluitioee fie. C2. 2S SL WeDo A ZEB 33. S. laeteviridis 11. Leaves spirally arranged. 12. Calyx and ovary glabrous. 13. Petole:0-5:tmim:, Fae. ser oie a. ca) ON a 16-4. S. cochinchinensis ssp. leptophylla 13. Petiole more than 5 mm. 14. Leaves shorter thanScm .......... 16-4. S. cochinchinensis ssp. leptophylla 14. Leaves longer than 5 cm. 1 DEY. Ss ck © de rentaien eee be 16-4. S. cochinchinensis ssp. leptophylla 15. Disk glabrous. 16. Twigs (appressedly) pubescent, puberulous or pilose. Seeds not straight. 16. S. cochinchinensis 16. Twigs tomentose or tomentellous. 17. Petiole 12-17 mm. Acumen 2-7 mm long. Nerves 8-12 pairs. Fruits more than 10 mm long ede cme 8A cantina fhoue) 2 os cece > gh ee 5. S. atjehensis 17. Petiole 5-12 mm. Acumen longer than 7 mm. Nerves 10-16 pairs. Fruits to c. 10mm long. 28. S. glomerata 12. Calyx and/or ovary hairy. 18. Leaves crowded towards the end of the twigs, the latter tapering off towards the apex. S. polyandra 18. Leaves evenly distributed, twigs not obviously tapering off. 19. Ovary glabrous. 20-“Inflorescence only A=flowered”) = 65 FP SONS ee: 38. S. multibracteata 20. Inflorescence more-flowered. PPebiskinaryer is ot ete GO, oR 16-4. S. cochinchinensis ssp. leptophylla 21. Disk glabrous. 22. Seed and embryo uncinately curved towards the base 16-4. S. cochinchinensis ssp. leptophylla 22. Seed and embryo twice curved .. . . 16-1. S. cochinchinensis ssp. cochinchinensis 19. Ovary hairy. DACA E SLA DTOUS gy ets 0d eS) sg) ihe ae 16-4. S. cochinchinensis ssp. leptophylla 23. Calyx hairy. 24. Bracts caducous. 25. Inflorescence an often branched raceme to 4 cm. Calyx 1-2 mm long 47. S. robinsonii 25. Inflorescence a 1—3-flowered short spike. Calyxc.3mm ... . 10. S. brachybotrys 24. Bracts persistent. 262 Penole 055 maven. 2 OR) ara. 16-4. S. cochinchinensis ssp. leptophylla 26. Petiole more than 5 mm. 213: BECKS SUBICNE SS 5c FRO a Bok ee ee ee 24. S. filipes D272 Seeds riot 'straignt: | ors ee a. ee 16-4. S. cochinchinensis ssp. leptophylla 10. Underside of leaves hairy. 28. Leaves distichous. 29. Nerves up to 6 pairs. 30; Argie:of leaf base more than 00? 0) 29% £00 2s Ae, Bee © 31. S. johniana 30. Angle of leaf base less than 90°. 31. Dek Gatiious 25-612. 2,06 wench) ghd TER Wa ate 52. S. trichomarginalis 31 Diskhatty——- £28 aged fe! SS eee IS no Ee 33. S. laeteviridis 29. Nerves (5-)6 pairs or more. 32. Leaves longer than 5 cm (mean length). 33. Flowers c. 3 in an up to 3 cm long lax raceme. Fruits 10-14 mm long. Stamens c. 90 or more. 17. S. colombonensis 33. Inflorescence usually different. Fruits to c. 12 mm long. Stamens c. 70 or less. 34. Inflorescence a fascicle. Bracts to c. 1 mm long, persistent, bracteoles persistent. Ovary c. 1 mm high, calyx c. 1 mm long, lobes not becoming longer by tearing. Corolla c. 2-4"/, min. Style base hairy. Fruits: ampulliform” 2A +. 0 See 23. S. fasciculata 34. Inflorescence not a fascicle. Bracts longer than 1 mm, caducous, bracteoles caducous. Ovary more than 1 mm high, calyx longer than 1 mm, lobes becoming longer by tearing. Corolla more than 4 mm long. Style base glabrous. Fruits ovoid to ellipsoid 33. S. laeteviridis 1977] SYMPLOCACEAE (Nooteboom) 219 32. Leaves shorter than 5 cm. 35. Inflorescence only 1-flowered. 36. Reticulation not prominent. Ovary c. 1 mm high, calyx longer than 2 mm, lobes c. 3 mm. Bracts several. Corolla c.4 mm. Fruits 8-9 mm long . . 252. S. trichomarginalis 36. Reticulation present beneath. Ovary 1—11/, mm high, calyx c. 2mm long, lobes 1—1!/, mm long. Bract 1. Corolla 4-6 mm long. Fruits 10-12mmlong .... 58. S. zizyphoides 35. Inflorescence more-flowered. Beeracts.and bracteoles caducous . .. . . «asgte Bodeeeee avo: 33. S. laeteviridis 37. Bracts and bracteoles persistent. ae narys hrmsisc:..10. by 5 mm ..< \3 eae. Boe Pee ee eee se 22. S. deflexa 38. Disk glabrous. Fruits 10-12 by 5-6mm. .........+... 58. S. zizyphoides 28. Leaves spirally arranged. 39. Upper side of leaves hairy. feerenele of leaf base more than 90°. . . 26. 3/3 sche ieee 13. S. calycodactylos 40. Angle of leaf base less than 90°. 41. Leaf margin (and petiole) beset with closely spaced glands. ..... 3. S. adenophylla 41. Leaf margin (and petiole) often glandular but glands not closely spaced. 42. Ovary hairy. Fr. cylindrical, 13-18 by 3-5 mm. Embryo straight. . . . 20. S. crassipes 42. Ovary glabrous. Fr. ampulliform, 6 by 4 mm. Embryo twice curved 43. S. paucistaminea 39. Upper side of leaves glabrous. 43. Calyx and ovary glabrous. 44. Leaves crowded towards the end of the twigs, minutely appressedly hairy beneath 37. S. wikstroemifolia 44. Leaves evenly distributed, glabrous or longer hairs beneath. 45. Seeds straight 46. Leaf index 2-3. Fruits less than20mmlong............ 5. S. atjehensis 46. Leaf index more than 3. Fruits longer than 20 mm. Ari ereesiicss than 1@ipans -. . . .-ey oe De eee 15. S. cerasifolia a9." Nexves more‘than 10 pairs ~~... . . Se 15b. S. cerasifolia var. grandifolia meemerais tie Stone ee 16-4. S. cochinchinensis ssp. leptophylla 43. Calyx and/or ovary hairy. 48. Leaves crowded towards the end of the twigs ......... 37. S. wikstroemifolia 48. Leaves evenly distributed. 49. Ovary glabrous. 50. Disk hairy. 51. Inflorescence only 1-flowered. Seeds straight. ........ 38. S. multibracteata 51. Inflorescence more-flowered. Seeds not straight. we, Pee O-) WIT ss ck aur pk ee 34. S. lancifolia 52. Petiole morethanSmm......... 16—4. S. cochinchinensis ssp. leptophylla 50. Disk glabrous. BE emy toc, tenon =... ws 16-1. S. cochinchinensis ssp. cochinchinensis 53. Ovary more than | mm high. 54. Twigs (appressedly) pubescent, puberulous or pilose. 16-4. S. cochinchinensis ssp. leptophylla 54. Twigs not appressedly pubescent or puberulous. 55. Leaf index 2-3. Bracts persistent, shorter than 3 mm, bracteoles persistent. Calyx lobes not becoming longer by tearing. Corolla shorter than c. 4 mm. Stamens less than 30. Fruits to c. 10 mm long. Mesocarp fleshy (shrivelled when dry). Seeds not straight. 16-4. S. cochinchinensis ssp. leptophylla 55. Leaf index more than 3. Bracts caducous, longer than 3 mm, bracteoles caducous. Calyx lobes becoming longer by tearing. Corolla c. 5mm long. Stamens more than 30. Fruits more than 20 mm long. Mesocarp woody or corky. Seeds straight 15. S. cerasifolia 49. Ovary hairy 56. Calyx ibrous 6. te.0\6. of tense Tele 16-4. S. cochinchinensis ssp. leptophylla 56. Calyx hairy. 57. Bracts caducous. 58. Leaves longer than 15 cm. 59. Fruits more than 10 mm long, 2-5-celled. Mesocarp woody or corky. Stone with high lengthwise not interrupted ridges. Seeds straight . 15b. S. cerasifolia var. grandifolia 59. Fruits to c. 10 mm long, 1-celled. Mesocarp thin, friable in dry state. Stone with a transverse constriction at one side. Seeds not straight .... . 48. S. rubiginosa 58. Leaves shorter than c. 15 cm. 60. Calyx lobes longer than 1'/, mm. Style base hairy. 61.) Leaves aaprter than 5 ott «a: 05) aoe ees 10. S. brachybotrys 61. Leaves longer than 5 cm. Gz. Tloreacenes @ Teeth os Seve: eR we 2 50. S. sumatrana 62. Inflorescence a 1-3-flowered short spike or a spike to 4.cm. eS 220 FLORA MALESIANA [ser. I, vol. 8? 63. Petiole 3-4 mm. Ovary c. 1 mm high, calyx c. 3 mm, lobes longer than 2!/, mm. Ovary (appressedly) pubescent. Disk inconspicuous. Fruits c. 5mm _ broad, (-celled?Scedsinotistraisht] Were 2) aes OS CO: 10. S. brachybotrys 63. Petiole 5-7 mm. Ovary c. 1!/, mm high, calyx 2 mm long, lobes 1'/,-21/, mm long. Ovary sericeous. Disk clearly present. Fruits more than 5mm broad, 3-celled. Seeds isttaiehias 51-341. 4 BO Se eee era . 1. AA 50. S. sumatrana 60. Calyx lobes */,-1*/, mm long. Style base ele 64. Nerves more than 10 pairs ..... 1... 6-2. 5 5. 47. S. robinsonii 64. Nerves less that 10 pairs. 65. Ovary c. 1 mm high, lobes triangular. . 42-1b. S. ophirensis var. densireticulata 65. Ovary 1-2 mm high, lobes not triangular. ......... 47. S. robinsonii 57. Bracts persistent. 66. Inflorescence only 1-flowered. Gi-wAngle ofleafibase.more than\90°° >... 33. ts SO See ee. ee 31. S. johniana 67. Angle of leaf base less than 90°. 68. Leaf index 4-7. Ovary c. 1'/,mm high. Stamens less than 30. Disk hairy, incon- spictousastone:smoothiad tstabsale nehOoie toe ee 49. S. salicioides 68. Leaf index 1.3-4. Ovary to c. 1 mm high. Stamens more than 30. Disk glabrous, clearly present. Stone with ridges or grooves ........ 20. S. crassipes 66. Inflorescence more-flowered. 69. Seeds straight. 70. Bracts to c. 1 mm long. 71. Ovary to c. 1 mm high. 72. Reticulation fine. Calyx longer than 1 mm. Inflorescence a much reduced often clustered spike. Fruits not ampulliform, 13-18 mmlong . . . 20. S. crassipes 72. Reticulation coarse. Calyx c. 1 mm long. Inflorescence a fascicle. Fruits ampulli- form; 5—7'mmlongee sash wean aed «fnew: scien aaa 23. S. fasciculata 71. Ovary more than 1 mm high. 73. Leaf index more than 3. Calyx lobes longer than!/,mm. . . 3. S. adenophylla 13- seat index. 2-3 Calyx lobes toc: 4/,mm jong, -.5)2.. . 3 cee 24. S. filipes 70. Bracts longer than 1 mm. (4-eAnelejof leaf base more than90>, 405 teeter sa Ga ee 20. S. crassipes 74. Angle of leaf base less than 90°. 75. Underside of leaves especially hairy onthe margin. .... . 20. S. crassipes 75. Underside of leaves not especially hairy on the margin. 76. Calyx lobes longer than 1!/, mm. Disk hairy. Stamens more wie Pat Style base hairy: Eruits’more than (Ommilons ~ . 3). . . . . se . S. crassipes 76. Calyx lobes 1-1!/, mm long. Disk glabrous. Stamens less than 0. Style base glabrous se ruts.cs1 0mm Jong). |. .2 Sy tay ee ee 56. S. vidalii 69. Seeds not straight. Ue ISkeClADTOUS. /~ an ope h od Gees 2 os bw ae 16-4. S. cochinchinensis ssp. leptophylla 77. Disk hairy. Peer secdtandlembryOsU-snapeG a a seg spe, es 34. S. lancifolia 78. Seed and embryo uncinately curved towards the base. 16-4. S. cochinchinensis ssp. leptophylla 9. Twigs glabrous. 79. Underside of leaves hairy. 80. Leaves crowded towards the end of the twigs, minutely appressedly hairy beneath 37. S. wikstroemifolia 80. Leaves evenly distributed. 81. Calyx and ovary glabrous. 2) Diskhiaye ss. £0 fo BOTS, BALES NO Wom, 16-4. S. cochinchinensis ssp. leptophylla 82. Disk glabrous. 83. Seed and embryo uncinately curved towards the base. 16-4. S. cochinchinensis ssp. leptophylla 83. Seed and embryo not uncinately curved towards the base ...... 5. S. atjehensis 81. Calyx and/or ovary hairy. 84." Weaves. distichotis < Pain, «ag oT ahs YS Pia. a. 5 ee 33. S. laeteviridis 84. Leaves spirally arranged. $5. Leaves shorter than Ss cat “4 4a oe ee. dun 2 eel sie ee 10. S. brachybotrys 85. Leaves longer than 5 cm. 86. Calyx glabrous. 87. Calyx lobes becoming longer by tearing. Seeds straight ....... 20. S. crassipes 87. Calyx lobes not becoming longer by tearing. Seeds not straight. 16—4. S. cochinchinensis ssp. leptophylla 86. Calyx hairy. 88. Petiolé 3=4. mm. 2 82.22 Se ee ee ee 10. S. brachybotrys 88. Petiole more than 5 mm. 89. Ovary glabrous 00 < wo ote hese 16—4. S. cochinchinensis ssp. leptophylla 1977] SYMPLOCACEAE (Nooteboom) 221 89. Ovary hairy. nt Bracts ane bracteoles caducous«.) .yc of 4. Wetted paces 48. S. rubiginosa 90. Bracts and bracteoles persistent. Sieeeacwsite.c. tm long. Seeds straight .. . .. . . seecwsles ok wets 24. S. filipes 91. Bracts longer than 1 mm. Seeds not straight. 16-4. S. cochinchinensis ssp. leptophylla 79. Underside of leaves glabrous. RS CISTICHOUS MU Sr are kite SE) op a 5 sy! op hayh sadaeen oy eee 33. S. laeteviridis 92. Leaves spirally arranged. 93. Calyx and/or ovary hairy. 94. Ovary glabrous. 95. Bracts caducous. ERA RO CE LRA CM... 6). 0s: asa nies Coed Se ee 12. S. buxifolia SP OAVES (OL GHN Ls PSF a! s/s.) sx. La ces 6 SR eee 21. S. cylindracea PP CANCSNONCEr HAN MO CMs) ys. «woes sue © eee 7. S. barringtoniifolia 95. Bracts persistent. me senorescence Only I-flowered . . . . . . « + seisudlp ue eee 12. S. buxifolia 97. Inflorescence more-flowered. Op DISET gS So a oor 16-4. S. cochinchinensis ssp. leptophylla 98. Disk glabrous. 99. Calyx lobes toc.*/,mmlong....... 16-4. S. cochinchinensis ssp. leptophylla 99. Calyx lobes longer than /, mm. foe Ovary toiemymmobighe soo ooo. ace Cl cl Re, ee 16. S. cochinchinensis 100. Ovary more than 1 mm high. Pe IIE RITEADDS ile oe eos: «deus wt ole eee 42. S. ophirensis 101. Seeds not straight. 102. Seed and embryo uncinately curved towards the base. 16-4. S. cochinchinensis ssp. leptophylla 102. Seed-and embryo different .. . «6.» volte 42. S. ophirensis 94. Ovary hairy. 103. Leaves shorter than 5cm. Petiole3-4mm ........... 10. S. brachybotrys 103. Leaves longer than 5 cm. 104. Calyx glabrous. 105. Disk glabrous. 106. Calyx lobes not becoming longer by tearing. Seeds not straight. 16-4. S. cochinchinensis ssp. leptophylla 106. Calyx lobes becoming longer by tearing. Seeds straight .... . 20. S. crassipes 105. Disk hairy. 107. Seeds not straight. Bracts and bracteoles persistent. 16-4. S. cochinchinensis ssp. leptophylla 107. Seeds straight. Bracts and bracteoles caducous. Ess SOOTONAL SOMME sys) Kee a eh» set wt Re 21. S. cylindracea A ee oe er 32. S. junghuhnii 104. Calyx hairy. 109. Bracts caducous. 110. Calyx longer than 1 mm. Style base hairy. 111. Petiole 3-4 mm. Inflorescence a (basally branched) spike. Ovary c. 1 mm high. Disk inconspicuous. Fruits c. 10 mm long, I-celled. Seeds not straight .10. S. brachybotrys 111. Petiole more than 5 mm. Inflorescence a panicle. Ovary 1-1'/, mm high. Disk clearly present. Fruits 15 mm long, 3-celled. Seeds straight .... . 21. S. cylindracea 110. Calyx to c. 1 mm long. Style base glabrous .......... 42. S. ophirensis 109. Bracts persistent. RAMEN P SITING Guy te 8 8 eo» 4) ew 42. S. ophirensis 112. Petiole more than 5 mm. 113. Leaves crowded towards the end of the twigs, the latter tapering off, at least 5 mm @ IER. 5g x01 che nilesun > ip eee eee 44. S. polyandra 113. These characters not combined. 114. Nerves 13-20 pairs. Intramarginal vein absent. Leaves 21-62 cm 26. S. gigantifolia 114. Nerves 4-13 pairs. Leaves 4-23 cm. 115. Disk glabrous. 116. Bractslongerthanimm ...... 16-4. S. cochinchinensis ssp. leptophylla 116. Bracts to c. 1 mm long. 117. Calyx lobes to c. '/, mm long. Stone with low ridges .... . 24. S. filipes 117. Calyx lobes longer than '/, mm. Stone with high lengthwise interrupted ridges. 42. S. ophirensis 115. Disk hairy. 118. Nerves 4-7 pairs. Fruits ampulliform with long beak, c. 7 by 5mm 42. S. ophirensis 118. These characters not combined. . . . 16-4. S. cochinchinensis ssp. leptophylla ——- © = 222 FLORA MALESIANA [ser. I, vol. 8? 93. Calyx and ovary glabrous. 119" Inflorescence fetmindl. nn... te Pe eee oe ees 46. S. pyriflora 119. Inflorescence axillary. 120. Bracts caducous. 121. Nerves more than 10 pairs. 122. Inflorescence a (basally branched) spike, forming a cone in bud. Fruits more than 20 mm [Steere ere ye a ae Get oi, ee nines LS) 5) OPS ee es 122. Inflorescence not a spike. Fruits less than 20 mm long. 123) Beactssana bracteoles plabrous 2. 92... ss use ... . 46. S. pyrifiora 123. Bracts and bracteoles hairy. 124. Leaf margin entire. Disk inconspicuous ........... 36. S. maliliensis 124. Leaf margin not entire. Disk clearly present. 125. Bracts shorter than 3 mm. Stamens less than 100. Corolla c. 4mm 11. S. brandisii 125. Bracts longer than 3 mm. Stamens more than 100. Corolla c. 5 mm long. 11b. S. brandisii var. pseudoclethra 121. Nerves less than 10 pairs. 126. Disk hairy. I27= Stamens 15-40 betiole | —sstittie ses aot. so es ee 34. S. lancifolia 127. Stamens more than 40. Petiole more than 5 mm. 128. Ovary to c. 1 mm high. 129. Inflorescence a (basally branched) lax spike. Bracts to c. 1 mm long 25. S. gambliana 129. Inflorescence not a spike. Bracts longer than 1 mm. 130. Inflorescence a (basally branched) raceme. Stamens 40-c. 60. Calyx lobes becoming longer by tearing. Style base glabrous .......... 14. S. celastrifolia 130. Inflorescence a panicle of racemes. Stamens more than 100. Calyx lobes not becom- me longer by tearing. style base hairy... - =. + «. - =. «eee 39. S. nivea 128. Ovary more than 1 mm high. SI Petiolers Arie re te ke ser eae as toe ’.. . . 9. S. borneensis 131. Petiole more than 5 mm. 132. Terminal buds glabrous. 133. Inflorescence a (basally branched) raceme. Calyx 11/, mm long. Style base glabrous. EUUISEG ol OMIM ONPG cue usec yn) «, cdaaice camseees ee 8. S. batakensis 133. Inflorescence a fascicle or very short spike. Calyx c. 2 mm long. Style base hairy. Bris NGO E, ee fee, ees, ce, ek, peo sie 53. S. tricoccata 132. Terminal buds hairy. Inflorescence a panicle. Calyx longer than 2 mm, lobes longer than 1!/, mm. Style base hairy. Fruits morethanl10 mmlong . 21. S. cylindracea 126. Disk glabrous. 134. Fruits 2—5-celled. 135. Inflorescence a fascicle or a very short spike. Ovary more than 1 mm high. 53. S. tricoccata 135. Inflorescence a (basally branched) raceme. Ovary to c. 1 mm high. 136. Inflorescence axis hairy. Corolla more than 4mm long. Calyx lobes becoming longer by tearing. Stone smooth. Seeds not straight .... . 14. S. celastrifolia 136. Inflorescence axis glabrous. Corolla shorter than c. 4 mm. Calyx lobes not becoming longer by tearing. Stone with ridges or grooves. Seeds straight 27. S. glabriramifera 134. Fruits 1-celled. 137. Reticulation fine. Ovary 2-3 mm high ............ 2. S. buxifolia 137. Reticulation coarse. Ovary 1-2 mm high. 138. Inflorescence much branched ..... 16-3. S. cochinchinensis ssp. thwaitesii USE... Itloresrence Semple ais Fe. fe A ae fen oo Se Ce 51. S. sumuntia 120. Bracts persistent. 139. Leaves shorter than 5 cm. 140. Inflorescence only 1-flowered. Bracts several. 141. Bracts shorter than 3 mm. Corolla shorter than c. 4 mm. Ovary 1-2 mm high. Stamens less than 30. Stone smooth. Seed and embryo uncinately curved towards the base. 16-4. S. cochinchinensis ssp. leptophylla 141. Bracts longer than 3 mm. Corolla more than 4 mm long. Ovary more than 2 mm high. Stamens more than 50. Stone with ridges or grooves. Seed and embryo not uncinately curved: towardsithe: base:. s.0.°5. *. eae, eee censor tee ties 12. S. buxifoli 140. Inflorescence more-flowered. Bract 1. 142. Petiole 0-5 mm. 143; Corolla’ S=fimm eo ee eee ee ee 57. S. whitfordii 143. Corolla shorter. 144. Leaf index less than 2. Acumen shorter than 5 mm. Bracts longer than 1 mm. 144. Leaf index more than 2. Acumen longer thanSmm...... 34. S. lancifolia 142. Petiole more than 5 mm. 1977} SYMPLOCACEAE (Nooteboom) 223 145. Inflorescence a basally branched raceme. Corolla 5-7 mmlong . . 57. S. whitfordii 145. Inflorescence a (basally branched) spike. Corolla shorter than c. 4 mm. 16-4. S. cochinchinensis ssp. leptophylla 139. Leaves longer than 5 cm. are PIO Mer SATII ea fc. wy Sud. we us 5. ag Sk Sh 00 choap hace oe ee ee 34. S. lancifolia 146. Petiole more than 5 mm. 147. Inflorescence not a spike. 148. Inflorescence not a fascicle. 149. Reticulation fine. Ovaryc. 1 mmhigh. ........ 18. S. composiracemosa 149. Reticulation coarse. Ovary more than I mm high 16-3. S. cochinchinensis ssp. thwaitesii 148. Inflorescence a fascicle. 150. Disk glabrous. 151. Petiole 12-17 mm. Ovary more than | mm high. Nerves 8-12 pairs. Fruits 10-12 mm MBTIP elie a eile eo es 8 ew od eee 5. S. atjehensis 151. Petiole 5-12 mm. Ovary c. 1 mm high. Nerves 10-16 pairs. Fruits 7-10 mm long. 28. S. glomerata 150. Disk hairy. 152. Leaves obovate, longer than 10 cm. Acumen longer than 5 mm. Inflorescence axis hairy. Calyx regularly 5-lobed. Fruits ovoid or obovoid, 1I-celled. Seeds 1, not REERINNE pers Sire Be. cin cis wok). (Sear& 16-4. S. cochinchinensis ssp. leptophylla 152. Leaves elliptic or circular, shorter than c. 10 cm. Acumen shorter than 5 mm. In- florescence axis glabrous. Calyx 2-4-lobed or symmetrically cleft. Fruits cylindric or ellipsoid, 2—S-celled. Seeds more than 1, straight. . ... . 40. S. obovatifolia 147. Inflorescence a (basally branched) spike. 153. Twigs (exceptionally) thick. 154. Terminal buds hairy. Disk hairy . . . . 16-4. S. cochinchinensis ssp. leptophylla 154. Terminal buds glabrous. Disk glabrous. 155. Inflorescence axis hairy. Bracts 2-3 mm, hairy. Bracteoles hairy. Calyx 11/,-2 mm long. Prots:c210'mm longs. . .c.1 2s So eee 6. S. barisanica 155. Inflorescence axis glabrous. Bracts 5-7 mm, glabrous. Bracteoles glabrous. Calyx longer than;2'mm. Fruits c:-13.mm long: .. . os nis kee 45. S. pulvinata 153. Twigs not (exceptionally) thick. 156. Calyx 2-4-lobed or symmetrically cleft. Baie REOOIEAD SAAN sk ke wt oe 54. S. trisepala DP PCHOIG: =-e-Ots Gok Gd ea ee kw ce 40. S. obovatifolia 156. Calyx regularly 5-lobed. 158. Base angle to 20—30°. Leaf index 3'/,—5. Nerves 11-13 pairs. . 29. S. goodeniacea 158. Base angle more than 30°. Leaf index less than 3'/,. Nerves at most 11 pairs. 159. Seed and embryo uncinately curved towards the base. 16-4. S. cochinchinensis ssp. leptophylla 159. Seed and embryo twice curved. ..... 16-2. S. cochinchinensis ssp. laurina KEYS TO FLOWERING MATERIAL ARRANGED BY ISLANDS AND ISLAND GROUPS Sumatra 1. Midrib prominent on the upper surface. hes SitenckGiniviens 7.15) . + « 2 + 4 0.6, #, © ¥, Wee ee 4. S. anomala 5. Twigs hairy. ’ ; 6. Leaves distichous. Calyx 2—4-lobed or symmetrically cleft, calyx lobes becoming longer by tearing. 33. S. laeteviri 6. Leaves spirally arranged. : 7. Leaves crowded towards the end of the twigs. Twigs thick, tapering towards apex. Petiole more than 20 mm. Corolla more than 7 mm long. Apex of leaves rounded or acute. 44. S. poly. 7. Leaves evenly distributed. - Colvx end ovmry. giabrous .. . . . «+> ‘es ion ote ee eee ee 5. S. atjehensis 8. Calyx and/or ovary hairy. : 9. Leaves longer than 10 cm. Nerves more than 10 pairs. Inflorescence a (basally branched) spike. Bracts persistent. Ovary glabrous, to c. | mm high, calyx lobes longer than 1'/, mm. Disk glabrous, clearly present. Fruits ampulliform, —— P . S. cochinchinensis ssp. cochinchinensis 224 FLORA MALESIANA [ser. I, vol. 8? 9. Leaves shorter than c. 10cm. Nerves less than 10 pairs. Inflorescence a raceme. Bracts caducous. Ovary hairy, 1-2 mm Heh ates lobes iiilinia mm es Disk hairy, inconspicuous. Fruits ellipsoid, 3-celled . . : . . . . 47. §. robinsonii 5. Twigs glabrous. POMINET Ves 4-5, DAMS 0 yes x Mey segieme, so we 8) es eee oes ae vie as oA OL 10. Nerves more than 5 pairs. 11. Calyx and/or ovary hairy. 12: dkeavesiGistichous 12 «cele .Shariccles ack BE Ae. Deol SSS eiserceruiis 12. Leaves spirally arranged. 13. Leaves crowded towards the end of the sean -off me: ..... . 44. §. polyandra 134 Plantidiiierentes.... Bad SE a alk thal - oo oe « « AZ SS ODMIeNSIS 11. Calyx and ovary glabrous. 14. Inflorescence not a spike. 15 inflorescence. a‘fascicle. Stamens'c./50!' sou view. ..eo. si. 2S Se eiensis 15. Inflorescence a raceme. 16s Stamensi40=60; 211-40 eecw gels eam 1 oe ter) wie) 4) 31-2 e 16. Stamens c. 100 . ee Peres) fe Lec Sie 14. Inflorescence a (basally branched) spike. ijemuivwies thick. betminaljbudsilarce, sa.) ie) Seales . 6. S. barisanica 17. Twigs not thick. Terminal buds small ....... 16-2. S. cochinchinensis ssp. laurina 4. Underside of leaves hairy. 18. Twigs glabrous. 19. Leaves distichous. Petiole 1-4mm. Corolla3-5 mm .......... . 33. S. laeteviridis 19. Leaves spirally arranged. Petiole more than 5 mm. 20. Inflorescence a fascicle. Bracts persistent, c. 2mm. Calyx glabrous. Calyx lobes not becoming longer DY: tearing ty.c Sieecticg ‘aya ks bcc oa ee Cee EP ie? Be 5. S. atjehensis 20. Inflorescence a spike forming a cone in bud. Bracts caducous, 3-5 mm. Calyx hairy. Calyx lobes becoming longer by tearing. ..... » Uae) ceereielg.cnu lanl? Aa Sarbioinss 18. Twigs hairy. 21. Leaves distichous. 22. Calyx usually hairy. Inflorescence a fascicle. Bracts persistent. Ovary c. 1 mm long. Calyx c. 1 mm long. Calyx lobes not becoming longer by tearing. Style base hairy . 23. S. fasciculata 22. Calyx often glabrous. Inflorescence a raceme or panicle of racemes. Bracts caducous. Ovary 1-1'/, mm high. Calyx 2-3 mm. Calyx lobes becoming longer by tearing. Style base glabrous 33. S. laeteviridis 21. Leaves spirally arranged. 23. Calyx and ovary glabrous. D4niLeaf index. 2.to.32 Nerves!8=12 pairs, ...¢) ies) a 0 Si Soak ae. APE Se Seaehensis 24. Leaf index more than 3. Nerves 6-9 Dae 25. Nerves less than 10 pairs. . . . artnet se : . S. cerasifolia 255 INetvesumoreithan: 10) pairs? ., .. « sch aves en _ 15b. S. poe var. grandifolia 23. Calyx and/or ovary hairy. 26. Upper side of leaves hairy (pulverulent). Leaf margin and petiole beset with many closely SpacedavesiculanmclandSin-t.a ee ceiiia cucees Moen sees ete e . . 3. S. adenophylla 26. Upper side of leaves glabrous. Leaf margin and petiole different. 27. Ovary glabrous. 28. Nerves less than 10 pairs. Bracts caducous. Ovary 1—1!/, mm high. Calyx ee mm. Calyx lpbes/becominelonger by teaming . ete ee oe ee . S. cerasifolia 28. Nerves more than 10 pairs. Bracts persistent. Ovary to c. 1 mm long. Guise t to 2 mm long. Calyx lobes not becoming longer by tearing. 16-1. S. cochinchinensis ssp. cochinchinensis 27. Ovary hairy. 29. Leaves longer than 15 cm. BO MMICHVESHIOSISsDAITS us Sais 54.0 oat one eee eek aes} 15b. S. cerasifolia var. grandifolia SOMINGHVESEL2—IVAPallSs Fe Naaincuk: Piecet Seeerals «os ss) 6 Fo ee ee 48. S. rubiginosa 29. Leaves shorter than c. 15 cm. 31. Bracts to c. 1 mm long. 32. Leaf margin (and petiole) beset with closely spaced glands. . . . . 3. S. adenophylla 32. Leaf margin (and petiole) often glandular but glands not closely spaced. 33. Nerves less than 10 pairs. Reticulation coarse. Inflorescence a fascicle. Bracts persistent. Ovaryie.f muni high. Stylebase hairy’) 3 sao. o9) ep) et DS 23. S. fasciculata 33. Nerves more than 10 pairs. Reticulation fine. Inflorescence a (basally branched) raceme. Bracts caducous. Ovary more than 1 mm high. Style base glabrous . 47. S. robinsonii 31. Bracts longer than 1 mm. 34. Angle oljleanbase.c. 90° 1 sa oer es es 3 OS Pe 50. S. sumatrana 4 Angleton leat base less'than'607).Gai)) 6. Ra ae eee 47. S. robinsonii 1977] SYMPLOCACEAE (Nooteboom) 225 Malay Peninsula 1. Midrib prominent on the upper surface. 2. Twigs hairy. 3. Leaves evenly distributed, underside glabrous ........2.2.2.2.2.4.2. 4. S. anomala 3. Leaves crowded towards the end of the twigs, minutely sparsely appressedly hairy beneath. 37. S. wikstroemifolia 2. Twigs glabrous. 4. Leaves crowded towards the end of the twigs ............. 37. S. wikstroemifolia EaAUCuen IY Cistripnted (bows) «ke se oe eee SD ee ee 35. S. lucida 1. Midrib impressed on the upper surface. Ty. ec SOS ee Se. 8 eS eae 41. S. odoratissima 5. Corolla glabrous. 6. Twigs hairy. 7. Leaves distichous. eemeorsice Of leaves SlabrouS . . . . «ics ay ce © « ommcgeinelne Wee 33. S. laeteviridis 8. Underside of leaves hairy. 9. Bracts persistent. 10. Inflorescence a true fascicle. Ovary c. 1 mm high. Calyx 1 mm long . . . 23. S. fasciculata 10. Inflorescence a short, often clustered spike. Ovary 1-2 mm high. Calyx 1'/,-4 mm long 20. S. crassipes SMP EAUICOUS ba > si ERAT. Wins) Se <5) oh os dee Se ee ee 33. S. laeteviridis 7. Leaves spirally arranged. 11. Upper side of leaves hairy. 12. Angle of leaf base more than 90°. Bracts and bracteoles caducous. Hairs on twigs more than PenarMREYTTI HF tg stats ede ie bree eee 13. S. calycodactylos 12. Angle of leaf base less than 90°. 13. Leaf margin (and petiole) beset with closely spaced glands. Bracts to c. 1 mm long. Calyx to c. 1 mm long, calyx lobes */,-1 mm long. Disk glabrous. Style base not conical. Fruits to c. meemiong PP Mae a. 3S re 3. S. adenophylla 13. Leaf margin (and petiole) often glandular but glands not closely spaced. Bracts longer than 1 mm. Calyx 11/,4 mm, calyx lobes longer than 1'/, mm. Disk hairy. Style base conical. Bee ts- 1S mint long Oe 20. S. crassipes 11. Upper side of leaves glabrous. 14. Leaves crowded towards the end of the twigs... ......... 37. S. wikstroemifolia 14. Leaves evenly distributed. 15. Underside of leaves glabrous. 16. Calyx glabrous. Inflorescence a fascicle. Disk pulvinate or cylindric. Fruits cylindrical. Seeds nee 2. UP Webbe wood «les eater ae 28. S. glomerata 16. Calyx hairy. Inflorescence a (basally branched) spike. Disk annular. Fruits ampulliform. Becas notetraicnt-; Foo. Ss . eeG@adls 16-1. S. cochinchinensis ssp. cochinchinensis 15. Underside of leaves hairy (pulverulent, nearly glabrous, in S. adenophylla). 17. Calyx and ovary glabrous. ae pecmoreacence’a spiké.. . . . . . mie ee ee A 15. S. cerasifolia Deameeenrenee a igeticle. 2. nis « caite eee eee 28. S. glomerata 17. Calyx and/or ovary hairy. 19. Ovary glabrous. 20. Nerves less than 10 pairs. Bracts and bracteoles caducous. Ovary 1-1'/, mm high, calyx longer than 2 mm, calyx lobes 1-1'/, mm long, becoming longer by tearing. Fruits 22-40 ge ee aera ee 15. S. cerasifolia 20. Nerves more than 10 pairs. Bracts and bracteoles persistent. Ovary to c. 1 mm high, calyx c. 2mm long, calyx lobes longer than 1'/, mm, not becoming longer by tearing. Fruits sya some, A-colled «5. + 2 hm 2 16-1. S. cochinchinensis ssp. cochinchinensis 19. Ovary hairy. 21. Ovary more than 1 mm high. 22. Leaf margin (and petiole) beset with closely spaced glands. Bracts to c. 1 oc long. Calyx teccbmmlong 3 es! oh. eels Ce SET ee Pee 3. S. adenophylla 22. Leaf margin (and petiole) often glandular but glands not closely spaced. Bracts longer than 1 mm. 23. Nerves 6-11 pairs. Reticulation faintly prominent. Bracts and bracteoles persistent. Calyx lobes not becoming longer by tearing. Fruits 13-18 mm long . 20. S. crassipes 23. Nerves 12-17 pairs. Reticulation much prominent. Bracts and bracteoles caducous. Calyx lobes becoming longer by tearing. Fruits toc. 10 mmlong . . 48. S. rubiginosa 21. Ovary to c. 1 mm high. 24.\ heaves longer than:i3.0m. «sis a2 «nan elt 6 nod? re ah 20. S. crassipes 24. Leaves shorter than c. 15 cm. 25. (Reticulation fine.) Bracts and bracteoles caducous ....... 42. S. ophirensis 25. (Reticulation coarse.) Bracts persistent. 226 FLORA MALESIANA [ser. I, vol. 8? 26. Inflorescence an often clustered short spike. Bracts 1-4 mm. Calyx 2!/.-3 mm. Stamens more than 60. Fruits not ampulliform, 13-18 mm long ... .. 20. S. crassipes 26. Inflorescence a fascicle. Bracts to c. 1 mm long. pes c. 1 mm long. Stamens 12-35. Fruits ampulliform, 5-7 mmlong ............. . 23. S. fasciculata 6. Twigs glabrous. 27. Nerves more than 10 pairs. 28. Underside of leaves hairy. S90 inflorescence aspike. .. 6 2645 4 ede ey a 99. Inflorescence afascicle .. 2.0.0 62 © 6 1 oe es hw ow «ogee eA RE ONIMCIE TSE 28. Underside of leaves glabrous. 30. Petiole more than 20 mm. Inflorescence a spike or a cone. Fruits 3-celled 7. S. barringtoniifolia 30. Petiole less than 20 mm. 31 Inflorescence terminal =") =). es ie. we es es OUT AGS aor 31. Inflorescence axillary. 32. Calyx andjor ovary hairy. Corolla. 2-5mm . . . ws ww. CS 42. S. ophirensis 32. Calyx and ovary glabrous. 33. Intramarginal vein far from margin. Inflorescence a fascicle. Bracts persistent, hairy, shorter than 3 mm. Ovary c. 1 mm high, calyx 1-2 mm long, calyx lobes not becoming longer by tearing. Corolla 4-5 mm. Stamens less than 50. Fruits 7-10 mm long. . 28. S. glomerata 33. Intramarginal vein close to margin. Inflorescence a raceme or panicle of racemes. Bracts caducous, glabrous, longer than 3 mm. Ovary 1'/,-2 mm high, calyx 3-5 mm, calyx lobes becoming longer by Eee Corolla 8-10 mm long. Stamens c. 100 or more. Fruits c. 15 mm lONS Vie ae eo a ae ete Sen ee Se a 27. Nerves less than 10 pairs. 34. Underside of leaves hairy. 35. Leaves crowded towards the end of the twigs .. ..... . . . . 37. S. wikstroemifolia 35. Leaves evenly distributed. 36. Bracts persistent. Disk glabrous. Fruits 13-18 mm long Jw) 70) ZO Saerassines 36. Bracts caducous. Disk hairy. Fruits 7-12mmlong ....... . . . 33. S. laeteviridis 34. Underside of leaves glabrous. 37. Calyx and/or ovary hairy. 38. Disk hairy. 39. Leaves distichous . . fy eee PE 8 Gren) GG 39. Leaves spirally arranged Sh oto gg eee ab ae. oa: th 42. S. ophirensis 38. Disk glabrous. 40. Inflorescence a short, often clustered spike. Bracts persistent. . . . . . 20. S. crassipes AQ: Inflorescence’a raceme? Bractsicaducous . . . . /S0.0) ee Cee 51. S. sumuntia 37. Calyx and ovary glabrous. 41. Inflorescence a (basally branched) spike. Bracts persistent 16-2. S. cochinchinensis ssp. laurina 41. Inflorescence not a spike. Bracts caducous. 42. Inflorescence a panicle of racemes. Stamens more than 100 ........ 39. S. nivea 42. Inflorescence a (basally branched) raceme. Stamens 25-60. 43. Calyx lobes becoming longer by tearing. ............ 414. S. celastrifolia 43. Calyx lobes not becoming longer by tearing .... . &! & =o eS Sy snes Java & The Lesser Sunda Islands PrietoachnaIty oe Sono te es Boe pea de A A eee 41. S. odoratissima 1. Corolla glabrous. 2) Midrib proniinent on theiupper suriace?;\"> 202". pe, ey) es RD) 35. S. lucida 2. Midrib impressed in the upper surface. 3. Twigs hairy. 4° Leaves distichous) 5.7) Deco meee Wey Gi) co gs SR OE 23. S. fasciculata 4. Leaves spirally arranged. 5. Underside of leaves glabrous. (If leaf ene and Hg he beset with closely spaced vesicular glands: 35 Sy adenopnylia:) ss.) ee 16-1. S. cochinchinensis ssp. cochinchinensis 5. Underside of leaves hairy. 6.“ Upper side of leaves hairy (pulverulent).5" s We. GOkA. 2st. i228 See 3. S. adenophylla 6. Upper side of leaves glabrous. 7. Ovary glabrous, calyx longer than 1 mm. Bracts longer than 1 mm 16-1. S. cochirichinensis ssp. cochinchinensis 7. Ovary hairy. 8. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands. Ovary 12mm high . 2. se ts Spee oe Se nee aoe eee 3. S. adenophylla 8. Leaf ndee 2-3. Leaf pena (and petiole) often glandular but glands not closely spaced. Ovary’c. 1! murvhigh . . . . . . UR eee eee eee 23. S. fasciculata 1977] SYMPLOCACEAE (Nooteboom) 227 3. Twigs glabrous. 9. Calyx and/or ovary hairy. 10. Ovary hairy. Inflorescence a raceme. Calyx glabrous .......... 32. S. junghuhnii 10. Ovary glabrous. Inflorescence a spike. Calyx hairy. 16-1. S. cochinchinensis ssp. cochinchinensis 9. Calyx and ovary glabrous. 11. Inflorescence axis glabrous. Pree IesSitnans2ONM aie i SG See.) TR 16-2. S. cochinchinensis ssp. laurina i@ereuoic more than 20mm... .........-. 16-4. S. cochinchinensis ssp. leptophylla 11. Inflorescence axis hairy. mecemesoess than iO.pairs 6. is. 6 ed SI ee 16. S. cochinchinensis 13. Nerves more than 10 pairs. 14. Angle of leaf base 25—40°. Inflorescence a (basally branched) spike, forming a cone in bud. 19. S. costata 14. Angle of leaf base more than 60°. Inflorescencea raceme ........ 11. S. brandisii Borneo SS a a co a a rr mri eS Re 41. S. odoratissima 1. Corolla glabrous. Sessa fiat ‘or prominent on the upper surface .......:..2.082..-. 4. S. anomala 2. Midrib impressed in the upper surface. 3. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular glands: 3. S. adenophylla). 4. Twigs hairy. ReEIPEPECISUICHOUS, . ceo sss wes ww ws vee 33. S. laeteviridis 5. Leaves spirally arranged. femeeaves 4-6.em. Petiole 3-4mm. . . . . .. «5 a os el ee 10. S. brachybotrys 6. Leaves longer than 6 cm. Petiole more than 5 mm. 7. Leaves evenly distributed. Twigs not thick, cylindrical. Leaf margin not entire. Petiole less than 20 mm. Ovary glabrous, to c. 1 mm high, calyx 1-2 mm long. Corolla 3-5 mm. Fruits ampulli- form, 1-celled. Seed 1, not straight. Apex of leaves acuminate. 16-1. S. cochinchinensis ssp. cochinchinensis 7. Leaves crowded towards the end of the twigs. Twigs thick, tapering towards apex. Leaf margin entire. Petiole more than 20 mm. Ovary hairy, c. 2mm high, calyx 2-3 mm long. Corolla 8-10 mm long. Fruits ellipsoid, 3-celled. Seeds more than 1, straight. Apex of leaves rounded or MS ke fe ee 44. S. polyandra 4. Twigs glabrous. 8. Calyx and/or ovary hairy. ees cisticnous,..Calyx SIabrOUS. «. «= «. ++. «lels) ll) an 33. S. laeteviridis 9. Leaves spirally arranged. 10. Inflorescence a 1—3-flowered spike. Bracts caducous. Stamens c. 100. Petiole 3—-4 mm. Leaves ae CON o 50 (tI PRIN 58 O.5 5. 4) i) et SR 10. S. brachybotrys 10. These characters not combined. eaves 15—SO.mmilong . 0... 2 os os 1 Wt eae 12. S. buxifolia 11. Leaves longer than 5 cm. 12. Petiole more than 20 mm. Twigs thick. Inflorescence a spike. Calyx 3-3!/, mm 7. S. barringtoniifolia 12. Petiole less than 20 mm. Inflorescence a raceme or a spike. Calyx to c. 1 mm long 42. S. ophirensis 8. Calyx and ovary glabrous. a peeves more wen TO paite.. . ww tt nw + ke 29. S. goodeniacea 13. Nerves less than 10 pairs. 14. Bracts persistent. Seer waeawee suorier than 5 Cm... «i «4s « «©, peel eee 12. S. buxifolia SRR CM IMIG TROT OC ws) gk ie oe 16-2. S. cochinchinensis ssp. laurina 14. Bracts caducous. 16. Ovary to c. 1 mm high. Apex of leaf mostly abruptly acuminate. 17. Leaf margin entire. Inflorescence axis glabrous. Petiole 5-10 mm . . . 25. S. gambliana 17. Leaf margin not entire. Inflorescence axis hairy. Petiole 3-I15mm_ . 14. S. celastrifolia 16. Ovary more than | mm high. Leaf apex usually not or faintly acuminate. 18. Leaf margin entire. Calyx */,-1 mm long. Disk hairy. Petiole 3-4mm . 9. S. borneensis 18. Leaf margin not entire. Calyx longer than 1 mm. Disk glabrous or the style base hairy. 19. Leaves shorter than 5 cm and acumen shorter than 5 mm. Reticulation fine. Inflorescence a few-flowered raceme. Bracts longer than 3 mm. Ovary 2-3 mm high, calyx 2—5 mm long. Style base glabrous. Fruits 1-celled 9) o/5)4 Gla ake 12. S. buxifolia 19. Leaves longer than 5 cm and acumen longer than 5 mm. Reticulation coarse. Inflorescence a fascicle or very short spike. Bracts c. 1'/, mm. Ovary c. 2 mm high, calyx c. 2 mm long. Style base hairy. Fruits 3-celled. Stone smooth. Seeds more than! . . 53. S. tricoccata 228 FLORA MALESIANA [ser. I, vol. 8? 3. Underside of leaves hairy. 20. Leaves distichous. 21...\Calyx symmetrically teared whenvolder ...). Grace 6 4s ee he . . 33. §. laeteviridis 21. Calyx regular. 22. Stamens 25-35. Inflorescence a true fascicle. Petiole 2-8 mm. Leaves 5-18 cm. Ovary c. 1 mm High ave 6.0 MN IONS te isles) ws «0 we cf ook 2) 3 se 23. S. fasciculata 22. Stamens more than 40. pipers solitary or ina raceme or panicle. Leaves 2-9 cm. Calyx 1-3 mm. 23. Nerves 7-11 pairs. Petiole 3-4 mm. Leaves 4-9 cm. Stamens more than 90. Calyx 13/,-3 mm 17. S. colombonensis 23. Nerves 3-8 pairs. Stamens 40-100, but when more than 90 Bonde 1-2 mm. 24. Petiole 2-4 mm. Leaves 2- 31/5 cm. Calyxcc.Sitiim) .o. eee . . 52. S. trichomarginalis 24. Petiole 1-2 mm. Leaves 21/,—7 cm. aes 1-2 mm. 25 4Galyaccuommilonewicaiset. vies) 4 sarktovotie |. OR -oner. ae. 58. S. zizyphoides 25. Calyx 1-1"/, mm long. 26 Ovary, limm nich) weer & oe. Sanit, “te wat) eae. ead Leask 31. S. johniana DOmOV ALY ales —2 meh Nun. ei A ea sensin Silos 8s (ce es hele cate ee 22. S. deflexa 20. Leaves spirally arranged. 27. Twigs glabrous. PR MIBCAVESTA—OIGIT) wa ciultt we gies) clog curts. Pets nisi! aseecs: oumblenite Tacit re 10. S. brachybotrys 28. Leaves longer than 6 cm. 29e Teaves: /—16cm., Nerves 6-9) palTS... «. -, cusses Mtr Mucmacnel. bereaaleieain. 15. S. cerasifolia DO eaves 1 5—45/ CIN NEVES I 2-7, Dalts 0.) 5, oasems Deen een) Otten 48. S. rubiginosa 27. Twigs hairy. 30. Upper side of leaves hairy (pulverulent). .........2.2.2.... 3. S. adenophylla 30. Upper side of leaves glabrous. S@Alwe AUG OVALY .SIADTOUS: 1.4. 4 soa « & 5 a 2 » GS = = CRE 15. S. cerasifolia 31. Calyx and/or ovary hairy. 32. Ovary glabrous. 33. Nerves less than 10 pairs. Bracts caducous. Ovary 1—1*/, mm high, calyx longer than 2 mm. Rrutsiellipsoida22—40immilones3-celled) 7) 4.5 suis eerie ariel) Saleuee 15. S. cerasifolia 33. Nerves more than 10 pairs. Bracts persistent. Ovary to c. 1 mm high, calyx 1-2 mm long. Fruits ampulliform, 5—7 mm long, 1-celled. 16-1. S. cochinchinensis ssp. cochinchinensis 32. Ovary hairy. Sdseeavesslongenthanilisycom: ©. side: “erly ..mta dS. cout) Suse: ome 48. S. rubiginosa 34. Leaves shorter than c. 15 cm. 35. Inflorescence only 1-flowered. 36, Angleton leaf base more than OO kkk se eo a 31. S. johniana 36. Angle of leaf base less than 90°. Sie bracts. le 'Calyxsl—Z2 mmMVONg, cto , « c: ahs... %. « 1.1 cee 10. S. brachybotrys 35. Inflorescence more-flowered. 38. Bracts caducous. Petiole 3-4 mm. Nerves 6-9. Stamens c. 100 . . 10. S. brachybotrys 38. Bracts persistent. Petiole 2-12 mm. Nerves 3-12. Stamens 12-more than 100. 39. Calyx to c. 1 mm long. Stamens 12-50. 40. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands. Ovanyal=2 mmhighiot Aosta as Boies wae. -ees UE eel Eee oe 3. S. adenophylla 40. Leaf index 2-3. Leaf margin (and petiole) often glandular but glands not closely spaced. Ovary scalimmihish: ps wenene. A eres tek ose) Bt oe 23. S. fasciculata 39) Galyx longer than 1mm. Stamens 25-100: 7 >. 5). we ne 20. S. crassipes Philippines ieelbeavesverticiilate... . . « «hy 4 . uw aha a in" od ey) GR Se oe 55. S. verticillifolia 1. Leaves not verticillate. 2. Midrib prominent in the upper surface. 5; ULWwigs glabrous) Petiolesmorethanpormim) eiemes see ee Sens este 35. S. lucida Sihwies hairy. Petiole:l!—Simm ainiss: sels ate tye Geta nt ee 34. S. lancifolia 2. Midrib impressed on the upper surface. 4. Corolla hairy (in the Philippines sometimes nearly glabrous!) ...... . 41. S. odoratissima 4. Corolla glabrous. 5. Twigs hairy. Ovwleeaves distichous ls» ..1.....08 ain 4 hele eee ee Do Eee 23. S. fasciculata 6. Leaves spirally arranged. 74 Galyx divided into: three 23/,,-mm long lobes) asus o-aesiee ee 54. S. trisepala 7. Calyx not so. 8. Leaves crowded towards the end of the twigs. Twigs thick, tapering towards the apex. Fruits 3-celled. Apex of leaves rounded or acute... 5. «1.5.8 os 44. S. polyandra 1977] SYMPLOCACEAE (Nooteboom) 229 8. Leaves evenly distributed. 9. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular glands: 3. S. adenophylla). 10. Ovary hairy, c. 1'/, mm high. Inflorescence a lax raceme. Bracts to c. 1 mm long. Calyx lobes omni lone: stamens ¢.25 . . . . ks fs so 5 ne” Ree 24. S. filipes 10. Ovary glabrous (hidden between bracts!). Inflorescence a spike. 16-1. S. cochinchinensis ssp. cochinchinensis 9. Underside of leaves hairy. 11. Upper side of leaves hairy (pulverulent) ..........2... 3. S. adenophylla 11. Upper side of leaves glabrous. 12. Ovary glabrous. 13. Leaves longer than 10cm. Petiole more than 5 mm. Nerves more than 10 pairs. Calyx lobes longer than 11/, mm. Stamens more than 30. Disk glabrous. Style base glabrous. enusranipuitorn 2s! rk) i 16-1. S. cochinchinensis ssp. cochinchinensis 13. Leaves shorter than c. 10 cm. Petiole 1-5 mm. Nerves 4-11 pairs. Calyx lobes 1/,-11/, mm long. Stamens 15-40. Disk hairy. Style base hairy. Fruits not ampulliform 34. S. lancifolia 12. Ovary hairy. 14. Leaf margin (and petiole) beset with closely spaced glands .. . . 3. S. adenophylla 14. Leaf margin (and petiole) often glandular but glands not closely spaced. 15. Style base glabrous. 16. Bracts to c. 1 mm long. Calyx lobes c. '!/, mm long, not triangular . . 24. S. filipes 16. Bracts 2-3 mm. Calyx lobes 1-1!/, mm, triangular ......... 56. S. vidalii 15. Style base hairy. 17. Intramarginal vein present. Inflorescence a fascicle. Fruits ampulliform 23. S. fasciculata 17. Intramarginal vein absent. Inflorescence a (basally branched) spike. Fruits ellipsoid to BEcHiatiM: walutsitis matic (fic sc Jee, (a Se ee ee 34. S. lancifolia 5. Twigs glabrous. 18. Calyx and ovary glabrous. 19. Inflorescence a (basally branched) spike. en eran Sring? )sis) oS. lec! oro, ye, a 34. S. lancifolia 20. Acumen shorter than 5 mm. 21. Angle of leaf base less than 60°. Bracts 2-3 mm. Calyx lobes c. 2 mm long. Style base hairy. 40. S. obovatifolia 21. Angls of leaf base c. 90°. Bracts 3-5 mm. Calyx lobes longer than 2'/, mm. Disk glabrous. 54. S. trisepala 19. Inflorescence not a spike. 22. Bracts and bracteoles persistent. 23. Leaves 2-5?/, cm. Inflorescence a (basally branched) raceme. Bracts longer than 3 mm. Calyx regularly 5-lobed, calyx lobes semi-ovate. Fruits ovoid, 5-7 mm long, 1-celled. 57. S. whitfordii 23. Leaves 7!/,-11 cm. Inflorescence a fascicle. Bracts 2-3 mm long. Calyx 3-lobed, the lobes semi-elliptic. Fruits ellipsoid, 11 mm long, 3-celled ........ 40. S. obovatifolia 22. Bracts and bracteoles caducous. 24. Inflorescence axis glabrous. Corolla 3-4 mm. Calyx lobes not becoming longer by tearing. 27. S. glabriramifera 24. Inflorescence axis hairy. Corolla4-6mm...........+.-. 14. S. celastrifolia 18. Calyx and/or ovary hairy. 25. Inflorescence a spike. Ovary glabrous. Seevaty*/—immbhigh .......+..-s 16-1. S. cochinchinensis ssp. cochinchinensis Teawvere 2 iste mem kk we tw te ee eg 8 ee ee cee 42. S. ophirensis 25. Inflorescence a (sometimes compound) raceme. Ovary hairy. 27. Inflorescence a very lax raceme of 4-10 cm. Pedicels slender, 2-15 mm. Axis of raceme sparsely pulverulent-puberulous. Stamensc.25 ..... + ee eee eee es 24. S. filipes 27. These characters not combined. ...... 5.5626 + sseeerevee 42. S. ophirensis Celebes & The Moluccas CE EE eer Me Pe a 41. S. odoratissima 1. Corolla glabrous. 2. Midrib prominent on the upper surface ©. 6 6 ee ee 35. S. lucida 2. Midrib impressed in the upper surface. 3. Twigs hairy. 4. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular glands: 3. S. adenophylla). 5. Leaves distichous. Bracts caducous .......+ + ss ee etree 33. S. laeteviridis 5. Leaves spirally arranged. 230 FLORA MALESIANA [ser. I, vol. 8? 6. Calyx and ovary slabrous. ... <2 )