FLORA MALESIANA SERIES I - SPERMATOPHYTA Flowering Plants Vol. 10, part 3 Revisions Aceraceae A203 Actinidiaceae s.s. 4: Aizoaceae 4: Alismataceae 5: 317; 6 Alseuosmiaceae 0 Amaranthaceae 4: 69, 593; 6: Anacardiaceae 8: Ancistrocladaceae 4: Aponogetonaceae ae bie'7s Araliaceae-I pt DLs Aristolochiaceae 10: Balanophoraceae a Basellaceae ote Bataceae 5: Betulaceae Sey os Bignoniaceae 8: Bixaceae S.s. 4: Burmanniaceae 555 as. 13.502... 9% Burseraceae 5: 209, Gs 19172 72 2820;9 Butomaceae Ss Byblidaceae a Callitrichaceae 4: Campanulaceae 6: 107,928; 9: Cannabaceae 4: Capparaceae 6: Caprifoliaceae 4: 175, 598; 6: 928; 9: Cardiopteridaceae a Celastraceae 6: 227, 389, Centrolepidaceae 5: Ceratophyllaceae 4: Chenopodiaceae 4: 99, 594; 6: 932; 9: Chloranthaceae 10: Clethraceae * Cochlospermaceae 4: Combretaceae 4: 533; 5: 564; 6: Coniferales 10: Connaraceae 358 - §:-495; 6: 933; 9: Convolvulaceae 4: 388, 5: 5586: 936; 7: | 8235;9: Cornaceae 8: Corynocarpaceae 4: 262; 5: Crassulaceae 4: 197; 9: Cruciferae 10: Crypteroniaceae 8: Ctenolophonaceae 10: Cyperaceae 7: 435; 9: Datiscaceae 4: INDEX TO REVISED FAMILIES Dichapetalaceae5: 305; 6: 941 Dilleniaceae 4: 141; 7: 824 Dioscoreaceae 4: 293 Dipsacaceae 4: 290 Dipterocarpaceae 9: 237 Droseraceae Os? SIs) Oso sie Oe Elaeagnaceae 10: 151 Elatinaceae 4: 203 Epacridaceae 6: 422 Ericaceae 6: 469, 943;9: 562 Erythroxylaceae 5: 543 Fagaceae 73/265; 93.363 Flacourtiaceae 6%) S65 6: 943; 7: 827; 9: 564 Flagellariaceae 4: 245 Geraniaceae 6: 445; 9: 565 Gnetaceae 4: 336; 6: 944 Gonystylaceae 4: 349 Goodeniaceae Be 953,507. 62.949: 7.827 Haemodoraceae Se 1h Haloragaceae 7: 239, 828 Hamamelidaceae 53) 303 Hippocrateaceae 6: 389 Hydrocharitaceae S338); 6:.952;).9: 566 Hydrophyllaceae 4: 207 Hypericaceae ai | Icacinaceae Ti 1:9: 566 Iridaceae ae Ixonanthaceae 10: 621 Juglandaceae 6: 143 Juncaceae 4: 210; 9: 566 Juncaginacae 4: 57 Labiatae 8: 301; 9: 566 Leeaceae yaa iss) Lemnaceae 7: 219 Lentibulariaceae apr ed Liliaceae-I 9: 189 Linaceae 10: 607 Loganiaceae 67 293,.953; 9: 567 Lophopyxidaceae 430e 88 Magnoliaceae 10: 561 Malpighiaceae a PS Martyniaceae 4: 216 Menispermaceae 10: 157 Monimiaceae £03 255 Moringaceae 4: 45 Myoporaceae 4: 265 Myricaceae 4: 277 Najadaceae OE 157 Nyctaginaceae 6: 450 Nyssaceae 4: 29 Ochnaceae (a Olacaceae 163, 4 Onagraceae 8: 98 Opiliaceae 10: "ai Oxalidaceae 72 Wiggs Papaveraceae 5: 114 Passifloraceae 7: 405 Pedaliaceae 4: 216; 7: 829 Pentaphragmataceae 4: 517 Pentaphylaceae 5312) Philydraceae 4; 3 Phytolaccaceae 4: 229 Pittosporaceae 5: 345 Plumbaginaceae 4: 107 Podostemaceae 4: 65;6: 963 Polemoniaceae 4: 195 Polygalaceae 10: 455 Pontederiaceae 4: 255 Portulacaceae Ti aad Primulaceae 6: 173 Proteaceae S214) Punicaceae 4: 226 Restionaceae 5: 416 Rhizophoraceae 5: 429; 6: 965; 8: 550 Salicaceae 5: 107 Salvadoraceae 4: 225 Sarcospermaceae 4: 32 Saururaceae A AT Scyphostegiaceae 5: 297-30 Simaroubaceae 6: 193, 968 Sonneratiaceae 4: 280,513; 6: 973 Sparganiaceae 4: 233 Sphenocleaceae 4: 27 Sphenostemaceae 10: 145 Stackhousiaceae 45 Staphyleaceae 6: 49 Stylidiaceae 4: 529 Styracaceae 4: 49; 9: 568 Symplocaceae 8: 205; 9: 569 Taccaceae 7: 806 Thymelaeaceae 4: 349;'6: 1, 976;.7:\aae Trapaceae 4: 43 Trigoniaceae 4: 59 Trimeniaceae 10: 327 Triuridaceae 10: 109 Turneraceae 4: 235 Typhaceae 4: 243 Ulmaceae S338 Umbelliferae 4: 113, 595 5: 555; 6: 983; 7: 830;9: 569 Valerianaceae 4: 253 Violaceae 73) 1795538 Xyridaceae 4: 366, 598; 9: 571 Zy gophyllaceae 4: 64 ras CONIFERALES (D.J. de Laubenfels, Syracuse, N.Y.) General Introduction In spite of generalized impressions sometimes advanced about the decline and decrease of the Gymnosperms through the enormous development of the Angiosperms in the Cretaceous and their rapidly accelerated development in the Tertiary, it must be realized that this impression is confusing as far as Coniferales are concerned. It is of course a truism that the Gymnosperms are completely outnumbered in genera and spe- cies by the Angiosperms, the latter occupying terrain earlier beset by Gymnosperms. It must be realized, however, that possibly the almost entirely woody Gymnosperms did never have the po- tential for producing such immense numbers of genera and species as now found among the Angiosperms. This statement is also valid for the Coniferales. The Coniferales were only part of the Cretaceous richness in Gymnosperms and whereas many Gymnosperm groups became extinct or lived on with meagre remains, Coniferales — though pro- portionally with few genera and few species — still represent a most essential part of the world’s standing timber and involve a huge biomass through their sociability and their morphology: usual- ly a large size and the little tapering of their cylindric boles. The ‘decline idea’ is thus not valid for the Coniferales and this is further validated by their ex- tremely wide ecological capacity, as they thrive from the Arctic to the Antarctic, in all major parts of the globe, in the lowland, the hills and the mountains, and in the tropics from the seashore almost to the alpine zone, a colossal range, among the Angiosperms shared or approached by only very few families, e.g. Ericaceae and Fagaceae. This universal presence is also due to their most diverse ecological capacities. Coniferales are represented on the permafrost of the taiga as well as in hot semi-deserts, on all sorts of soils, from mineral-rich to mineral-poor, even in peat-swamps, enabling them to stand all sorts of environ- mental conditions. In the forest vegetation they show not seldom a high power of competition, often leading to dominance or codominance, often coupled with longevity. Some are agressive and tend to fill gaps in the vegetation by possessing nomad ecology. Seed is mostly produced in ample quantity; seedlings may be shade-tolerant or -intolerant. A number of species are distinctly fire-resistant. From these facts can be concluded that Coniferales are not just ‘on the decline’, but that they form still a most successful super-order of the Gymnosperms. Distribution. With 12 genera the Coniferales are well represented in the Malesian tropics, while elsewhere in the world only in the rich Sino-Japanese flora is there a substantially larger con- centration of conifer genera; nearby Taiwan for example has 15 genera (only four of which are shared with Malesia, viz. Nageia, Podocarpus, Taxus, and Pinus). Eight wide ranging conifer ge- nera of Antarctic affinities are today well established throughout Malesia up to the westernmost margins and four of these extend well beyond onto the continental part of Asia (Dacrycarpus, Dacrydium, Nageia, and Podocarpus). All are strictly confined to rain-forest habitats. Seven of these have seeds dispersed by birds, the eighth (Agathis) has small seeds with large membraneous wings. Except for the genus Phy/locladus, these genera are well represented at low and medium elevations and it is not necessary to imagine long-range dispersal between isolated mountain peaks as far as altitude is concerned. We have, however, to keep in mind that through the insular physiography of Malesia there may have been the necessity of crossing sea barriers. Unfortunately too little is known in detail about the precise distribution of land and sea in the course of the Tertiary. Two Holarctic conifer genera (Taxus, Pinus) penetrate into western Malesia and two more Ant- arctic genera are at present confined to the eastern half of Malesia (Libocedrus, Araucaria). The fossil record, although incomplete, suggests that the situation was quite different as late as the Miocene. (337) THE LuESTHER T. MERTz LIBRARY THE NEW YORK BOTANICAL GARDEN 338 FLORA MALESIANA [ser. I, vol. 102 Fossil record. Palaeozoic floras of Permo-carboniferous age are known from Sumatra and New Guinea. The Sumatra material shows affinities to Euramerican floras and, by the presence of Gigantopteris especially to the Cathaysian flora of Southeast Asia (JONGMANS & GOTHAN, 1935). In New Guinea fossil floras of Gondwana type as well as with a Cathaysian character are found (JOoNGMANS, 1940; Lacey, 1975), leading HAMILTON (1979) to suspect that at that time New Guinea was situated at mid-latitudes with land connections both to Sumatra and the Australian continent. A Mesozoic flora dated as Neocomian (Lower Cretaceous) has been described by SmiLEy (1970) from the Malay Peninsula. Although considerably poorer in species than the Palaeozoic ones, it is of interest because of the presence of two conifer genera, Frenelopsis of Cupressaceous affinity preserved as macrofossils and the fossil pollen genus Classopollis, derived from the extinct family of Cheirolepidaceae. Frenelopsis ranges from Lower to Upper Cretaceous and was widely distri- buted in North America, Europe and Asia. Classopollis is cosmopolitan and ranges from the Jurassic to the Upper Cretaceous. The general composition of this Lower Cretaceous flora clearly suggests links to Eurasian and North American floras. MULLER (1968) has described a rich and well preserved Upper Cretaceous microflora from Sara- wak, NW. Borneo, in which Caytoniales (Caytonipollenites), Podocarpaceae (Zonalapollenites), Araucariaceae (Araucariacites), Cheirolepidaceae (Classopollis), Cupressaceae or Taxodiaceae (Inaperturopollenites) and a rich assemblage of bisaccate pollen grains of Pinaceous or Podocar- paceous affinity represent the Coniferalean element. Although most of these genera are cosmo- politan, the presence of the bisaccate genus Rugubivesiculites is of considerable interest, since this has been recorded only from the northern hemisphere. Tertiary floras from Malesia, which include both macro- and micro-floras, are mainly restricted to the post-Eocene and, for the Angiosperm component, show a composition not unlike the pres- ent one (KRAUSEL, 1929; PostHuUMUS, 1929, 1931; ANDERSON & MULLER, 1975). However, the reli- ability of identification of many of these records, especially the older leaf remains, is question- able. An exception may be made for wood of Dipterocarpaceae, for which family also fossil pol- len records are available, showing that, at least in the post-Eocene, this family was well established in West Malasia. For Gymnosperms the situation is different and the presence of pollen of Ephedra, Pinus, Picea and 7suga in the Oligocene and Miocene of NW. Borneo, accompanied by pollen of Alnus, its disappearance from the record at the end of the Miocene, followed by immigration of PAyllocla- dus and Dacrycarpus in the Plio-Pleistocene suggests considerable change in the coniferous ele- ments in West Malesia (MULLER, 1966; STEIN, 1978). Dacrydium pollen, which was absent in the Upper Cretaceous-Paleocene assemblages studied by MULLER (1968) from Sarawak is first record- ed for the Oligocene. MULLER (/.c.) has attributed the disappearance of the Asian-montane element to peneplanation, but STEIN (/.c.) believes that they also may have suffered from competition with other forest ele- ments in submontane habitats. Significantly, Pinus has survived longest in NW. Borneo and is still present today in N. Sumatra and the Philippines. KHAN (1976) has established the presence of pollen of the following typical southern conifers in the Upper Miocene and Pliocene of New Guinea: Dacrydium, Dacrycarpus and Microcachrys, the latter disappearing from the record at the end of the Pliocene. Here the southern conifers are accompanied by Nothofagus. This evidence indicates that, in Borneo, boreal conifers have been replaced by Antarctic ones, immigrating from the direction of New Guinea. Dacrydium may have reached western Malesia already in the Oligocene, while PAyllocladus and Dacrycarpus only reached Borneo in the Plio- cene. In New Guinea the latter two were present earlier. This reflects the collision of the northwest moving Australian plate with the Celebes-Borneo area in the mid-Tertiary (HAMILTON, 1979; STEIN, 1978). Before this period Antarctic conifers may have been absent in West Malesia which has remained, at least since the Cretaceous, within reach of the Southeast Asian continent, as indi- 1988] CONIFERALES (de Laubenfels) 339 cated by the presence of boreal conifers, both in the Cretaceous and Tertiary of the Malay Pen- insula and Borneo. New Guinea presumably had been in close contact with Australia since the Palaeozoic. From FLorin’s masterwork (1963) one could deduce that Cryptomeria-like conifers should have occurred in Malesia, since they are found fossil in the Triassic of both Asia and Australia, fading away in the early Cretaceous. Similarly, relatives of Austrotaxus of New Caledonia, A thro- taxis of Tasmania and various Cupressaceae, especially Libocedrus, all genera with Holarctic af- finities, must have occurred in the Cretaceous of Malesia, inasmuch as these genera flourish be- yond to the east and south. There are also reports of fossils belonging to Podocarpaceae and Araucariaceae from Holarctic regions and they could have migrated through Malesia at the same time, but only fossil Araucaria pollen is known from the Upper Cretaceous of Borneo. The main conclusion from the fossil record is that, at least from the Lower Cretaceous onwards till the Oligocene, virtually no southern conifers reached West Malesia where in the Upper Creta- ceous and, more strikingly, in the Oligocene and Miocene a distinct Asian conifer element was present. Only at the mid-Tertiary collision of Australia + New Guinea with West Malesia did an invasion of southern conifers take place, in stages leading to the present-day distribution pattern. However, it is clear that large gaps in our knowledge still exist and it seems most desirable to have reliable records from the Upper Cretaceous and Lower Tertiary of Java, Celebes, the Lesser Sun- da Islands and New Guinea to allow a further confirmation of what is at present still a very tenta- tive picture. I appreciated very much the collaboration of the late Dr. J. MULLER (Leiden) in framing this paragraph on the fossil record. References: ANDERSON & MULLER, Rev. Palaeobot. Palynol. 19 (1975) 291—351; Florin, Acta Horti Berg. 20(4) (1963) 121—312; Hammton, U.S. Geol. Surv. Prof. Pap. 1078 (1979) 1—345; JONGMANS, Meded. Geol. Bur. Mijngebied Heerlen 1938-1939 (1940) 263—274; JoNGMANsS & Go- THAN, Jaarb. Mijnwezen in Ned. Indié 1930, Verh. V.59, pt. 2 (1935) 71—201; KHAN, Austr. J. Bot. 24 (1976) 783—791; KRAUSEL, Verh. Geol. Mijnbouwk. Gen. Ned. Kol., Geol. Serie II (1929) 1—44; Lacey in Campbell (ed.), Gondwana Geology, Austr. Nat. Univ. Press (1975) 125—134; MULLER, Blumea 14 (1966) 231—235; Micropaleontology 14 (1968) 1—37; PostHumus, Bull. Jard. Bot. Btzg III, 10 (1929) 374—384; Leiden Geol. Meded. 5 (1931) 485—508; SmiLey, Geol. Soc. of Malaysia, Bull. n. 3 (1970) 77—113; STEIN, Biogeographica 11 (1978) 1—168. Ecology. As mentioned above, the ecology of conifers shows a considerable variation and a summary may facilitate and stimulate the reader to delve in the text for further details. No main vegetation type, except aquatics and very dry seasonal lowland, is in Malesia devoid of conifers. Though varying in density, they form an essential part of the forest and other vege- tation. In the collecting numberlists of the Indonesian Forestry Service they form from one half to two percent of the total, depending on the area. Biomass of standing timber will attain probably a much higher percentage. Conifers are among the tallest tree species in Malesia. Many possess massive straight boles, mostly cylindric, rising to or over the canopy with a height of 40—50 m. Some reach still larger dimension and tower as emergents over the canopy, equalling the tallest size of the largest Diptero- carpaceae; notably species of the genera Agathis, Araucaria and Pinus may reach 70—75 m in height, the occasional record being a tree of Araucaria hunsteinii of 89 m (B.GrRay, J. Ecol. 63, 1975, 273). Small conifers are also well represented in Malesia. Mature specimens of no more than | m in height are found of Nageia maximus in Bornean swamp forests and of Podocarpus micropedun- culatus on the edges of clearings in and near Brunei. Small conifers are also found in scrub in the mountains, e.g. Dacrydium medium on G. Tahan (Malaya). Colonies of prostrate Podocarpus brassii var. humilis occur on the mountains of New Guinea. Stunted specimens of many other spe- cies are found in poor, rocky habitats in the mountains. Altitude. Lowland species are for example Podocarpus polystachyus which may be locally com- 340 FLORA MALESIANA [ser. I, vol. 102 mon (e.g. in Malaya) on sandy bluffs on the seashore and on low limestone outcrops. Dacrydium pectinatum and D. micropedunculatum can be locally common on low lying sand shoals (e.g. in S. Borneo), while the former along with A gathis borneensis may form nearly solid stands on low- land podsols (kerangas) or lowland peat-forest in Borneo almost at sea-level. The latter occurs sometimes in such quantity as to be worthy of exploitation for timber. A genus ‘descending’ to low altitude is Pinus: in West Luzon and in Mindoro. Pinus merkusii reaches sometimes as low as 50 m altitude as a pioneer in pyrogenous grasslands and up to 150 m on volcanic ash streams (lahars) and lavastreams in N. Sumatra. Descent has also been reported for Araucaria cunninghamii on steep rocky ridges and spurs, occasionally as low down as 75—100 m, the ‘normal’ low parameter being c. 500 m. With increasing elevation conifer populations become more frequent. At high altitudes species become fewer but it is not uncommon to find subalpine forest, whether or not turned into mossy forest, dominated by one or a few species of conifers. For example, the summit forest on Mt Leu- ser (N. Sumatra) is often dominated by a drooping conifer, Dacrycarpus imbricatus var. curvulus ; on Mt Suckling (Papua New Guinea) Araucaria cunninghamii is the dominant conifer. Climate. As mentioned above, in Malesia conifers shun the seasonally very dry /owlands of eastern Java and the Lesser Sunda Islands. They are also rather rare in the mountain rain-forest in this climatically seasonal belt, but Dacrycarpus imbricatus is found as far as Timor as a distinct constituent of the mountain forest; after devastation single trees may even survive as relicts in pyrogenous grassland, adorned with beards of Usnea. Soils. Many conifers prefer nutrient-poor soils, and are often even confined to them, but there are also species which are mostly found on richer latosols, e.g. Dacrycarpus imbricatus which grows excellently on young volcanic soils. As mentioned above, quite a number of conifers grow, sometimes in great quantity, on alluvial sandflats or on podsolized sands and sandstone (kerangas) and in peat-swamps, but they are not always limited to such habitat, as both Dacrydium pectinatum and Agathis borneensis are also commonly met as scattered individuals in middle elevation rain-forest. Some conifers, particularly of the genus Podocarpus, thrive on ultrabasic bedrock in Malesia (as well as in New Caledonia), dense stunted forest with plenty of Podocarpus confertus in Borneo and Podocarpus ridleyi in Malaya are examples. On Mt Soroako (Celebes) scattered specimens of Agathis, Dacrydium, Podocarpus and Nageia are found on ultrabasic bedrock. This may also be true for localized populations of Podocarpus deflexus of Malaya and N. Sumatra. More precise data about possibly specialized soil types are unknown for a number of species with restricted ranges, e.g. Podocarpus levis in Central Malesia, Dacrydium medium in Malaya and Sumatra, Dacrydium ericoides in Borneo, Dacrydium spathoides in New Guinea, and Agathis flavescens in Malaya. A curious conifer taxon is Dacrydium cornwalliana which is found in the mountains of West New Guinea (BW 697) restricted to deep black peat, reminding of the habitat of some Dacrydium and Dacrycarpus species occurring in peat under temperate conditions in New Zealand; both the former species and Dacrycarpus steupii are the only peat-swamp forest trees of Malesia sofar known. Fig. 14. As to limestone, it can in general be said that conifers are rare on this bedrock. In western Ma- lesia Podocarpus polystachyus occurs on low limestone outcrops; in Borneo and Celebes Agathis is also found on soils derived from limestone, and Dacrycarpus imbricatus is found on similar soils on Mt Perdido in Timor. Autecology. In general Malesian conifers are constituents of the rain-forest and as such are dark-germinators, their seedlings growing up under very low light intensity. Germination and up- growth of seedlings in shade is also the rule for high altitudes, but pioneering of conifers on dis- turbed, pyrogenous open land, at 2500—3000 m altitude was observed for Libocedrus and Phyllo- cladus by HOOGLAND. 1988] CONIFERALES (de Laubenfels) 341 A clear exception are the species of Pinus, P. merkusii and P. kesiya, which are light-demanding germinators. They may form permanent, dominant climax stands on very steep, rocky mountain slopes where no litter remains to prohibit germination. One can observe this on the Leuser massif in N. Sumatra at some 2500 m altitude. This occurrence is rare, the main occurrence is of a tem- porary nature, pioneers starting a succession, viz. bare soil of talus, volcanic mudstreams (lahars), lavastreams, earth- and rockslides, places torn open by earthquakes, and further man-made pyro- genous grassland. In the latter they occupy in W. Luzon and N. Sumatra large surfaces which are consequently converted into large, mostly savannah-like stands of Pinus. Under undisturbed con- ditions secondary forest and finally new primary forest will grow up among the pines, the litter and shade of which prohibits their own regeneration. But as Pinus is a long-lived pioneer, under such conditions the pines — which may possibly reach an age of 200—300 years — will remain towering over the later primary forest. Seed of many tropical Malesian conifers rather soon loses its germination power, in line with other rain-forest trees. That of Araucaria hunsteinii is down to zero in ten weeks. According to Wuitmore the viability of seed of Agathis drops rapidly and also that of Araucaria cunninghamii. This is also valid for Pinus merkusii. On the other hand seed in Podocarpaceae may require as much as a year to germinate. To the autecology also belongs the matter of the mycorrhiza, but unfortunately little is known of this relation. It is certain that one of the exomycorrhiza of Pinus merkusii is a Boletus, but it seems that for Pinus and probably other conifers more genera of fungi are involved. BEVEGE (1968) and Honc (Mal. Flor. 41, 1978, 225) have established that in the species of Araucaria an unidentified species of Endogone forms an endotrophic mycorrhiza. Sociology. Mostly conifers occur scattered through the forest, but a number of species have a tendency to occur socially in places. By their large mature size they are then often observed as emergents. It must be remarked, however, that occurrences are often local: A gathis is for instance in Celebes above some 1600—2000 m (where it rather suddenly becomes abundant) and in the Mo- luccas present in most mountain forests and also in the north of West New Guinea, where it is tapped on a commercial scale, but in Papua New Guinea it is very scarce and local. The same can be said of Araucaria in New Guinea, especially A. hunsteinii, which is found as an upper-canopy dominant in the Bulolo area and a few other places, but is elsewhere absent and not found in West New Guinea. In most instances we are ignorant about the underlying cause. As mentioned above, Podocarpus polystachyus is locally common to subdominant in sandy lowland in Malaya and Borneo, and so are Dacrydium pectinatum and Podocarpus micropedun- culatus in Borneo, where the former, sometimes together with A gathis borneensis, may form local stands on lowland podsols (kerangas) or in lowland peat-forest, with very acid soil conditions. Agathis borneensis may also form locally dense stands in the upper canopy. In passing we remark that hardly ever conifers are found in forest dominated by dipterocarps, with the possible ex- ception of the heath-forest in Borneo. In the West Javanese mixed mountain forest (e.g. on Mt Gedeh) there are three large emergents very common: Al/tingia excelsa (Hamam.), together with two conifers, Dacrycarpus imbricatus and Podocarpus bracteatus. But on Mt Tjeremai, a volcano in W. Central Java, there is between c. 1800—2500 m a very large gregarious dominant stand of Dacrycarpus imbricatus only. It remains unclear to what factor in the past this has to be ascribed. In other islands other species of conifers may be very common or gain subdominance in the higher mountains. In Sumatra for example Dacrydium elatum — used for Christmas trees — and on Mt Leuser Dacrycarpus imbricatus var. curvulus, the drooping pine. A similar role in moun- tain forest is found with the dense stunted forests of Podocarpus confertus in Borneo and of Po- docarpus ridleyi in Malaya on ultrabasic bedrock. Localized subdominance is found: Podocarpus deflexus and Agathis flavescens in Malaya, Dacrydium medium in Malaya and Sumatra, and Da- crydium ericoides in Borneo. Mostly conifers occur scattered in the mixed broad-leaved rain-forest, not infrequently repre- sented by several species; e.g. KALKMAN & VINK found on the Doma Peaks in Central New Guinea 342 FLORA MALESIANA [ser. I, vol. 10° Libocedrus papuana as an emergent, accompanied by Dacrycarpus expansus, Phyllocladus hy- pophyllus, and Podocarpus pseudobracteatus (Blumea 18, 1970, 119). Anatomy. The most important wood anatomical surveys of Coniferales are by E.W.J.PHIL- Lips, Identification of softwoods, For. Prod. Res. Bull. London 22 (1948, repr. 1966), and by P.Grecuss, Identification of living Gymnosperms on the basis of xylotomy, Budapest (1955), Xy- lotomy of the living conifers, Budapest (1972). Additional data for Malesia and adjacent regions can be found in R.KANEHIRA, Identification of Philippine woods by anatomical characters, Tai- hoku (1924) 231—244; H.H.JaNssonius, Mikrographie des Holzes der auf Java vorkommenden Baumarten 6 (1936) 469—494; H.Descu, Mal. For. Rec. 15 (1954) 630—632; M.KAEISER, Phyto- morphology 4 (1954) 39—47; J.vAN DER BuRGH, Rev. Palaeobot. Palyn. 15 (1973) 73-275; S.HaAyYASHI c.s., Micrographic atlas of Southeast Asian timber, Kyoto (1973); T.FURUNO, Res. Rep. Foreign Wood 6, Shimane Univ., Matsue (1977); J.H.FUNDTER & J.H.WissE, Meded. Land- bouwhogeschool Wageningen 77-9 (1977); K.OGata, Identification of Southeast Asian timbers (in Japanese), Jap. Ass. Wood Technology (1985). Despite the seemingly homogeneous microscopic structure of the vesselless wood of conifers, there are a number of highly diagnostic wood anatomical differences at various levels of the taxo- nomic hierarchy which can be profitably used for identification and which can help in the recon- struction of a natural classification. In the Malesian representatives of the Coniferales a number of genera can be immediately recognized on unique, single or combined characters: Pinus — Vertical and horizontal resin ducts present, cross field pits (i.e., pits from tracheids to ray parenchyma cells) fenestriform or pinoid, ray tracheids present. (N.B.: resin ducts and ray tracheids are absent from all other Malesian Coniferales.) Agathis and Araucaria — Pits on tracheids alternate and in a closely spaced honeycomb-like pattern (‘araucaroid’). Taxus — Tracheids with distinct spiral thickenings. Libocedrus — Cross field pits strictly cupressoid (i.e., with narrow, included apertures). Podocarpaceae — The distinction of Podocarpaceae from Libocedrus is fairly subtle: cross field pits in Podocarpaceae often include cupressoid types but almost invariably also other types such as taxodioid, pinoid, or piceoid pits. Most Malesian Podocarpaceae and Libocedrus have fairly common to abundant axial parenchyma in their wood, a feature absent from the other coni- fers. However, Phyllocladus and at least some temperate species of Dacrydium lack axial paren- chyma. Presence or absence of parenchyma has been used as an important character for sectional delimitation in Podocarpus sensu lato by KAEIsER, /.c., but some of her observations have been contradicted in a more detailed study by R.N.PATEL, New Zeal. J. Bot. 5 (1967) 307—321. The evidence available from the literature at present suggests that the wood anatomical variation pat- tern within the Podocarpaceae does not coincide with generic delimitation; further studies of well- authenticated samples are needed to assess the taxonomic significance of the wood anatomical va- riation in this family. Leaf anatomy can also play a useful role in identification and classification of the Coniferales, as exemplified in the study by J.T. BucHHOLz & N.E.Gray, J. Arn. Arb. 29 (1948) 49—76 on the systematics of Podocarpus sensu lato and by J.W.LANyon, A card key to Pinus based on needle anatomy, Min. Conservation, N.S.W., Australia (1966), also including the two anatomically dis- tinct Malesian species Pinus merkusii and P. kesiya. The extensive leaf anatomical literature on conifers is summarized in K.NApp-ZINN, Encyclopedia of Plant Anatomy 8 (1), Berlin (1966). — P. BAAS. Palynology. In general gymnospermous pollen is distinguished from angiospermous pollen by the alveolate-granular structure of the sexine, the lamellate structure of the nexine, and the presence of one distal aperture. With the exception of some primitive ranalean groups Angio- sperms have pollen with a columellate sexine, a non-lamellate nexine, and 3 or more equatorial apertures, or have pollen with attributes that can be derived from this basic pattern. The structure of the sexine seems at present the most reliable character. 1988] CONIFERALES (de Laubenfels) 343 The only aperture in pollen of Coniferales is always distal. Mostly it is a thin area (leptoma) in the exine, which is often further distinguishable by a different ornamentation. In Araucariaceae this area is large and circular, in Cupressaceae and Taxaceae small and circular; in Pinaceae and Podocarpaceae it is mostly large and oblong. When large, the thin area may also have a harmome- gathic function beside participating in the germination process. Sometimes an aperture is difficult to trace. A remarkable feature of some Coniferales pollen types is the presence of air bladders (wings, sacci) at the distal pole beside the aperture. Araucariaceae, Cupressaceae and Taxaceae have none, but most Pinaceae (except Larix, Pseudotsuga) and Podocarpaceae (except Saxegothaea) have 2 or 3 of them. Grains without bladders are more or less spherical; those with bladders have a spherical, lens-shaped, or oblong corpus. Most Coniferales pollen is medium-sized (25—50 um). Pinaceous grains measure (40—)50—70 (—80) um; the corpus of the likewise saccate podocarpaceous grains are mostly smaller (up to 50 um). Both cupressaceous and taxaceous grains range from 18 to c. 36 um. Araucariaceous grain size varies from 40—60 um (A gathis) to 60—90 um (Araucaria). In addition Araucaria pollen dif- fers from that of Agathis by the presence of a proximal annular thickening. There is a great deal of variation with regard to wall stratification and structure. Araucariaceae, Cupressaceae, and Taxaceae have a thick intine compared with the exine. In Cupressaceae the in- tine is even very thick, often comprising much more than half of the grain volume. A thick intine has the capacity of swelling after moistening and probably plays an important role in the germina- tion process. In Pinaceae and Podocarpaceae the intine is proportionally less thick. The nexine is lamellate in all Coniferales. The sexine is alveolate-granular in Araucariaceae, Cupressaceae, and Taxaceae, in Larix and Pseudotsuga of the Pinaceae, and in Saxegothaea of the Podocarpa- ceae; in all the rest of the Pinaceae and the Podocarpaceae the sexine offers a columellate-tectate appearance superficially like the situation in Angiosperms, essentially being a variation of the al- veolate structure. In Pinaceae, Podocarpaceae, and in most Araucariaceae pollen the sexine is thicker than the nexine, at least at the proximal side (cappa). In Cupressaceae, Taxaceae, and part of Araucariaceae pollen the nexine is the thickest layer. In the latter three families the surface of the sexine has a perine-like covering, consisting of small (< 1 um) granules (orbicules), which is absent in Pinaceae and Podocarpaceae. Air bladders or sacci form a remarkable aspect of the pollen wall of most Pinaceae and Podo- carpaceae. Sacci develop by proliferation of the alveolate layer of the wall. Probably they func- tion in both flight and harmomegathy of a grain. All Pinaceae have 2 sacci, except Larix and Pseudotsuga which are devoid of them and are fundamentally different from other Pinaceae. Tsuga mostly has pollen with one distal saccus encircling the aperture. In Podocarpaceae the genus Saxegothaea has no sacci and is therefore, and also on account of other features, considered as related to the Araucariaceae. In Podocarpaceae there is more variation in respect to the num- ber, shape, and size of the sacci than in Pinaceae. Pollen grains of Dacrycarpus are provided with 3 sacci, those of Nageia, Podocarpus, and Prumnopitys with 2. Phyllocladus and Falcatifolium pollen also have 2 sacci, but in the first they are very small and in the latter they are narrowly connected around the aperture. In Dacrydium a type occurs which has one fully radiosymmetric saccus around the aperture. Dacrydium has also the Podocarpus-like bisaccate type. Saccate pollen of the extra-Malesian podocarpaceous genera has 2 sacci (Acmopyle, Parasitaxus) or 3 sac- ci (Microcachrys, Pherosphaera). \n the latter two genera and in Dacrycarpus sometimes grains occur which have 4, 5 or 6 sacci due to aberrant tetrad configuration. Pollen of the extra-Malesian Cephalotaxaceae and Taxodiaceae is largely similar to that of Cu- pressaceae and Taxaceae. Coniferales are probably strictly wind-pollinated, the pollen being not sticky, smooth-surfaced, and sometimes provided with sacci. In the former two characters Coniferales pollen resembles some wind-pollinated Angiosperms (e.g. Gramineae, Betulaceae). References: ExDTMAN, Pollen and spore morphology/plant taxonomy, Gymnospermae (1957) 344 FLORA MALESIANA [ser. I, vol. 103 5—44, illus., (1965) 9—82, text; PoCKNALL, New Zeal. J. Bot. 19 (1981) 67—95, 259-266, 267-272; Stvak, Pollen et Spores 17 (1975) 349-421; Stapuin c.s., Rev. Palaeobot. Palyn. 3 (1976) 297-310; TENGNER, Bot. Notis. 118 (1965) 450—452; Van Campo, C. R. Acad. Sc. Paris 272 (1971) 2071—2074; WALKER, The evolutionary significance of the exine, Linn. Soc. Symp. Ser. 1 (1976) 251—308; WEND, J. Inst. Polyt. Osaka City Univ. 11 (1960) 109-136; WoDEHOUsE, Pollen grains (1935). — R.W.J.M. VAN DER Ham. Phytochemistry & Chemotaxonomy. Chemical characters of Coniferales were sum- marized twice in ‘Chemotaxonomie der Pflanzen’ (HEGNAUER, 1962, vol. 1: 293—440, 478—482; 1986, vol. 7: 462—554, 801-802). Here rather extensive bibliographies can be found for all fami- lies of Gymnospermae, Cycadopsida, Coniferopsida, Taxopsida and Chlamydospermae. General characters of Coniferales are: cuticular waxes of the so-called estolide-type; lignin which usually lacks the syringyl component; seeds which store predominantly starch or oils with unusual fatty acids, i.e. bi-tetra-unsaturated Cjs- and C29-acids with an isolated double bond in 5-position; accumulation of cyclitols such as pinitol, sequoyitol and (or) 0-methylmucoinositol in leaves, bark and wood; storage of shikimic and (or) quinic acid in leaves; accumulation of lignans (phenylpropanoid dimers) and (or) agatharesinol-type norlignans in wood, bark, traumatic resins and leaves (here sometimes as glycosides); production and exudation after injury of oleo-resins or gum-resins. Oleo-resins and gum-resins are deposited in schizogenic canals and cavities which seem to be lacking only in some representatives of Taxaceae. Oleo-resins are mixtures of essential oils and resins; turpentine is the essential oil produced by distillation of oleo-resins obtained from several species of Pinus. Gum-resins are mixtures of essential oil, resin and mucilage; Araucaria is the main producer of gum-resins among Coniferales. The predominant constituents of the essential oils of most Coniferales are mono- and sesquiterpenoids; some members of Podocarpaceae and other families produce appreciable amounts of steam-volatile diterpene hydrocarbons and hence yield diterpene-rich essential oils. The resins of Coniferales are mainly composed of diterpenoids; often diterpenic acids predominate. Colophony or rosin is the resin part of pine oleo-resins and Manila copal is the hard oleo-resin from Agathis dammara. Amber or succinite is the fossil resin of pines and Kauri copal is fossilized Agathis resin. Tannins are ubiquitous in Coniferales. They are represented in the taxon exclusively by the so- called condensed tannins and their building stones, the catechins and proanthocyanidins. Galli- and ellagitannins are totally lacking. Other classes of compounds which seem to be totally absent from Coniferales are iridoid com- pounds, cardenolides and steroidal saponins. The same seems to be true of triterpenoids of the ursane, Oleanane and lupane classes, and hence of corresponding saponins. Triterpenoids are represented in Coniferales by lanostane-type tetracyclic, onocerane-type tetra- and pentacyclic, and hopane-type pentacyclic compounds. Steroids are represented by the ubiquitous phytosterols; moreover, the frequent occurrence of phytoecdysones in rather high con- centrations is somewhat typical of the taxon. In the products of steroid and triterpenoid meta- bolism Coniferales strongly resemble Pteridophytes. Polyphenolic compounds other than lignans and tannins are accumulated by all conifers but, besides the general occurrence of flavonoids as a group, most classes of compounds and many individual compounds are restricted to taxa of lower levels such as infrageneric, generic and suprageneric entities. The same is true of alkaloids and several other classes of chemical constit- uents. Some examples to illustrate the situation follow. Agathisflavone-, amentoflavone-, cupressuflavone-, robustaflavone- and hinokiflavone-type biflavones seem to be nearly ubiquitous in leaves of Gymnosperms, but are lacking in Pinaceae which yielded hitherto only one biflavonoid, the flavone-flavonol dimer abiesin. Cephalotaxin-type alkaloids occur in all species of Cephalotaxus. All members of the genus Taxus (but not the other representatives of Taxaceae) produce taxane- type diterpenoids which are often esterified with the so-called Wintersteiner acid, which is a dime- 1988] CONIFERALES (de Laubenfels) 345 thylamino derivative of a hydroxydihydrocinnamic acid; the resulting nitrogen-containing con- stituents, such as the taxines and related compounds, are the ‘Taxus alkaloids’; they are accompa- nied in Taxus by the cyanogenic glucoside taxiphyllin. The latter too seems not to occur in other genera of Taxaceae, but is present in Mefasequoia and in some species of Juniperus. In Podocarpaceae several tendencies concerning secondary metabolism are recognizable: es- sential oils with appreciable amounts of diterpene hydrocarbons, ferruginol- and totarol-type phenolic diterpenes, bitter and biologically highly active mono- and bisnorditerpenoid lactones such as nagilactone, and accumulation of large amounts of phytoecdysones such as the maki- strones and the podecdysones are examples of such family-characteristic tendencies. At the same time the family is the only representative of conifers which makes use of anthocyanins to advert its diaspores: red to pink fleshy parts of Dacrydium, Phyllocladus, and Podocarpus diaspores contain an array of anthocyanins; anthocyanins may also be present in young leaves and strobili; the latter feature is not restricted to Podocarpaceae, however. Antibiotically active carvacrol and thymol derivatives and tropolone-type mono- and sesquiter- penic compounds are present in the wood of many Cupressaceae, including Libocedrus s.1. Exudates of many species of Araucaria contain larger amounts of mucilage than most other conifers; they are true gum-resins; ANDERSON and Munro observed 20—80% of mucilage in Araucaria-exudates with 10—20% of uronic acids, 50—70% galactose and up to 7% of the rather unusual sugar acofriose (3-0-methylrhamnose) as building stones. Acofriose is also present in mucilages of Cycadaceae. An array of low-molecular phenolic compounds, such as hydroxyacetophenones, stilbenes, di- hydrostilbenes and phenylpropanoids (monolignols) is known from Pinaceae ; they occur free and as glycosides and often have a taxon-characteristic distribution, and hence can be useful as taxo- nomic characters. Pinosylvin and its monomethy] ether have been interpreted as phytoalexins of Pinus because their synthesis is induced in the softwood after infection; normally these antifungal compounds are present in Pinus only in hardwoods and in barks. Flavonoid patterns were taxonomically exploited by many phytochemists; flavonoids yielded characters applicable at all levels of the taxonomic hierarchy. Just one example: C-glycoflavones have not yet been traced in Araucariaceae, Cephalotaxaceae, Cupressaceae and Taxaceae, and seem to be restricted in Pinaceae to Abies, Keteleeria, Tsuga and Larix; moreover, they were de- tected in Podocarpaceae in some species of Podocarpus. In recent times detailed analyses of essential oils were performed during biosystematic studies of a number of American conifers; the results proved to be rather promising; in many instances a better understanding of complex population structures was made possible by such inves- tigations. As a whole Coniferales are chemically well characterized by the general presence of several clas- ses of chemical constituents and by the total absence of others. Moreover, secondary metabolites yielded a large number of characters applicable at different levels of the taxonomic hierarchy. — R. HEGNAUER. Systematics. Generic delimitation. Four genera replace earlier broad treatments of the genus Podocarpus, all of which I recognized in 1969 (J. Arn. Arb, 50: 274—369); in part they had for- merly been distinguished as sections of this genus. Certainly there exist substantial relationships between them, but it must be well recognized that this is no sufficient reason for adopting a one- genus concept for the whole. As a matter of fact the morphological differences between these sharply distinct genera are at least of equal taxonomic ‘weight’ as compared to the differences between many other groups of northern hemisphere coniferous genera unequivocally distin- guished. On the other hand | cannot adhere to the recent splitting of the genus Libocedrus. For further argumentation I refer to the text under the genera in the taxonomic part. Cultivation. A fairly large number of exotic conifers are cultivated in Malesia, in part for testing them for forestry purposes, reafforestation, in part as ornamentals in gardens and parks. It falls outside the scope of this Flora to treat the cultivated exotics like the native species, none 346 FLORA MALESIANA [ser. I, vol. 103 of them is naturalized. Quite some have been incorporated in BACKER & BAKHUIZEN VAN DEN BRINK’s Flora of Java (Vol. 1, 1963, 87—95) which may be useful for their identification. A more general work that I can advise for identifying cultivated conifers is W. DALLIMORE & A.B. JACK- son, A handbook for Coniferae and Ginkgoaceae, 4th ed., revised by S.G. HARRISON (1966). Of native conifers several are in cultivation for various purposes. Agathis philippinensis is fre- quently cuitivated as a wayside tree in W. Java and probably elsewhere, and occasionally found in parks. Araucaria cunninghamii is frequently planted in parks and gardens as an ornamental tree. Araucaria hunsteinii, of which very large dominating complexes are found in some places in the Bulolo area in Papua New Guinea, is exploited from native growths, but the cleared ter- ritory is replanted with it on a large scale, because of the valued timber used for plywood. Dacrydi- um elatum is, according to CORNER, widely planted in the hills in Malaya for ornamental purpose. Finally Pinus merkusii is widely used for reafforestation and in addition for timber and the tap- ping of resin for the turpentine industry; notable huge complexes are found at Aek na Uli on the eastern hills around Lake Toba in N. Sumatra; large complexes are also found in S. Celebes; it is also used as an ornamental in parks and gardens. In nurseries, especially of Agathis, a single specimen of an older seedling already provided with mycorrhiza is planted in the centre of the beds in order to speed upgrowth of seedlings. An important point for silviculturists is the fact that, as mentioned above, the seed of many tropical conifers soon loses germination power. For Pinus merkusii, which is distributed on a large scale, very special care must be taken to keep the sundried seed in sealed metal containers with charcoal; even with these precautions a rather rapid loss of germination power takes place. This has been subject to extensive study. Keeping seed storage cooled is also applied. The timber of the large-sized species of all genera is most valuable, but only few are planted for this purpose on a large scale. This is in part due to the fact that of most species growth is slow, with the exception of Pinus species. It depends also for what purpose the timber is used, for pulp, sawn timber or high quality veneer. WHITMORE mentioned for Agathis in Java a rotation period of 30 years for pulp, and 50 years for veneer. For Araucaria in New Guinea a rotation period of 60 years is reckoned for plywood. Economic uses. As mentioned above all larger species of all genera provide excellent timber. In addition, species of A gathis are tapped large-scale, especially in the Moluccas and New Guinea, and to a less extent in Borneo and Celebes, for the resin (‘copal’ or ‘manila copal’, wrongly ‘da- mar’). In addition to the resin obtained from living trees, large bodies of subterranean resin of vanished trees are collected. The market for manila copal declined after the introduction of oil- based synthetics but there is still a use for special purposes. The resin of Pinus merkusii, and to a less extent that of P. kesiya, is collected for the turpentine industry. In N. Sumatra, in the vicinity of Takengon, there was a large factory for this purpose. See C. BRANDTsS Buys c.s. (Meded. Proefstation Boschwezen 19, 1928). References in synonymy. A remark must be made about the references in the synonymy of the species. I have omitted in many cases the mention of names without description or notes which occur in so many local plant lists and casual enumerations. This was made especially urgent by the fact that the names used in these lists, e.g. of Agathis and Podocarpus, are often wrong according to my classification. To account for all these ‘non’ or ‘sensu’ names would have caused an unnecessarily complicated synonymy. If collectors’ numbers were cited in these local lists, proper identity of these records can easily be checked by means of the ‘Identification Lists of Ma- lesian Specimens’ n. 61 (1982), which was issued separately by the Rijksherbarium, Leiden. KEY TO THE FAMILIES based on sexual characters 1. Ovules strictly terminal on short fertile shoots, erect, wingless. Pollen sacs usually several on each micro- SpOLOpHVlleahworsinele tracexcOLyVledOUS sauce eciaa piece eine risa oie ice eee Taxaceae 1. Ovules produced on axillary structures of a fertile shoot. 1988] CONIFERALES (de Laubenfels) 347 2. Seed usually cupped by a fringing epimatium or the inverted seed even completely enclosed by a leathery or fleshy structure, rarely naked and rarely erect, solitary, wingless. Reduced cone bracts often fleshy. Two pollen sacs on each microsporophyll. Cotyledons usually two fused pairs, occasionally more Podocarpaceae 2. Seed produced on an erect, woody, rarely fleshy scale which is often fused with the fertile bract, erect or inverted, occasionally solitary, more often two or more per fertile scale, usually with one or more wing(s). 3. Fertile bract and scale fused, sometimes indistinguishable. Seeds solitary or in variable numbers. More than two pollen sacs on each microsporophyll. Cotyledons usually 2—4. 4. Seed inverted, solitary; large mature seed cones disarticulate. Leaves spirally placed or opposite- decussate and distant. Cotyledons four or two fused pairs ................0-0000: Araucariaceae 4. Seed erect, solitary or in variable numbers; small mature seed cone does not disarticulate. Leaves crowd- ed, opposite-decussate or whorled. Cotyledons two or occasionally more, not fused.. Cupressaceae 3. Fertile bract separate from scale and not woody. Seeds two per scale, inverted, each with a single wing. Two pollen sacs per microsporophyll. Leaves spirally placed. Cotyledons more than two .. Pinaceae ARTIFICIAL KEY TO THE FAMILIES based on vegetative characters 1. Leaves spirally attached, sometimes distichous. 2. Leaves (needles) in bundles of 2—3 with a basal sheath (Pinus) ...............0000000000> Pinaceae 2. Leaves not in bundles with a basal sheath. 3. Leaves needle-like or scale-like. Eeeieeaves scale-ltke (Dacrycarpus, ‘Dacryadium) oo 45 a bees sec ook ese ae ae ee Podocarpaceae 4. Leaves needle-like, triangular or quadrangular in cross-section. 5. Tree with a very dominant primary trunk with regularly placed whorls of lateral branches. Seeds dry, in large bracteate cones (Araucaria cunninghamll) ..........0.0 0c ccc cence cceees Araucariaceae 5. Tree quite irregularly branched. Seeds few, in reduced cones (Dacrycarpus, Dacrydium, Falcatifolium) Podocarpaceae 3. Leaves distinctly flattened and often broad. 6. Leaves bifacially flattened, linear, less than 2.5 mm wide, with a basal constriction (Taxus) Taxaceae 6. Leaves if linear either bilaterally flattened, or more than 2.5 mm wide, or without a basal constriction (Dacrycarpus, Falcatifolium, Phyllocladus, Podocarpus, Prumnopitys) ........... Podocarpaceae 1. Leaves decussate, often distichous. ERNE SCANS (CLI DOCCUTUS) oi 5s sca a's SF oS Pe, MENS Me bye et a Gil ESMaLal ice. 0 REA Cupressaceae 7. Leaves broad and many-veined. 8. Leaves sessile, clasping, lanceate from a broad base (Araucaria hunsteinii)......... Araucariaceae 8. Leaves broad, not with a stem-clasping base. 3. Jerminal bud ‘hemispherical (A gathis) . 2). 5 sis oss | sk ve og soehiele ne wees eee eee Araucariaceae 9-rerminal bud ‘acute (Nageia) 25401. 5092 FT Ta ee Pete ie er ei ale Seen Podocarpaceae TAXACEAE The affinity of Taxaceae has been much debated, with many authors favouring a separate order, 7axales, for it, a position with which I tend to agree. Further questions are raised concerning the grouping of other families with Taxaceae, as against the other conifer families, based on the lack of seed cones, fleshiness of the mature fruit, or lack of a fertile seed scale. Cephalotaxaceae (not in Ma- lesia) has a reduced seed cone structurally organized quite differently from other conifers and vegetatively strongly resembling Taxaceae, so | would group these two together. All other conifer families show seed structures easily derivable from a compound cone with ovules produced on the upper face of a fertile scale which grows in the axil of a bract. Although Taxaceae, perhaps joined by 348 FLORA MALESIANA [ser. I, vol. 103 Cephalotaxaceae, can be set apart from the conifers proper, all can agree that taxads and conifers are more closely related to one another than to any other recognized group. Distribution. Of the five genera recognized for the Taxaceae, only Taxus reaches Malesia. Four are distinctly Holarctic in distribution, including Taxus, which is much the most widespread and reaches into tropical highlands. The fifth, monotypic Austrotaxus, appears on the other side of Malesia in New Caledonia, a distinct fragment of Gondwanaland, obviously a most curious relict on the southern hemisphere (FLoRIN, Acta Horti Berg. 20 (4), 1963, 260, f. 61: map). 1. TAXUS LINNE, Gen. PI. ed. 5 (1754) 462; Sp. Pl. 2 (1753) 1040; Pircrr, Pfl. R. IV, 5, Heft 18 (1903) 110; in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 208; Florin, Acta Horti Berg. 14 (1948) 378; GAussEN, Gymn. Act. & Foss. fasc. 15, ch. 25 (1979) 2. — Fig. 2, 3. Evergreen trees or (prostrate) shrubs. Bark thin, smooth, purple-red, peeling in large thin flakes. Leaves spirally placed (but usually twisted into a single plane), linear to linear-lanceolate, acute, distinctly constricted at the base where the leaf twists into a horizontal position and then widening again in the decur- rent part, penetrated by a single vascular strand marked on the lower surface by a blunt ridge which separates two bands of stomata and on the upper stoma- ta-free surface by a sharp narrow ridge. Foliage and fertile buds small and glob- ular, formed by several small keeled overlapping scales, the lower ones of which remain small while the higher ones expand with growth to become round and membranous. Usually dioecious. The fertile structures produced in the axils of ordinary leaves. Pollen cone solitary above a basal cluster of sterile scales, each peltate microsporophyll with a symmetrical whorl of 6—8 inverted pollen sacs, One microsporophyll in a terminal position and up to a dozen spirally placed la- teral microsporophylls. Seed-bearing structure compound with one or more short ovule-bearing shoots produced subterminally on a very short fertile axis covered by minute keeled spirally arranged scales, each fertile shoot consisting of several decussate pairs of keeled scales which expand as the seed matures into a broad membranous oval shape and together cover the base of the ripened fruit. A small basal aril gradually grows to cup the single erect terminal seed, finally becoming fleshy and bright red. Mature seed flask-shaped, slightly wider than thick with the wider margin slightly keeled. Distr. Seven species on the northern hemisphere middle latitudes and some tropical highlands, almost completely allopatric, but possibly some overlap between two species in the eastern Himalayas; one species in Malesia, and that one more common in subtropical parts of China. The genus has a predominantly northern hemisphere distribution, Central America and S. Celebes being the stations at lowest latitude. Fig. 1. Fossil remains are known from Europe (middle Jurassic to Pliocene) and eastern Asia (Miocene to Plio- cene). Ecol. Understory or canopy plants of moist temperate or tropical mountain forest. From near sea-level in their northernmost occurrence in Norway they reach to nearly 3000 m in subtropical and tropical mountains 1988] CONIFERALES (de Laubenfels) 349 THE WORLD MODIFIED VAN DER GRINTEN. PROJECTION —— © $00 1000 1300 wes \ TS ELT AY EPEES : ED, 60 40 2 oO 20 40 60 80 100 120 140 tast 160 coneiruoe 180 Fig. 1. Range of the genus Taxus L. (solid line) and 7. sumatrana (MiQ.) DE LAvB. (broken line and hatched). and do not go below 1200 m in tropical latitudes. As an undershrub they can be locally quite common, but the trees in this genus rarely take a dominant canopy role. Growth is generally quite slow but dense and speci- mens are often prized as ornamentals. Even the forms which are capable of becoming immense trees usually appear as shrubs or at best as small trees when under cultivation. Pollination is strictly by wind dispersal. Fruits are taken by birds and probably other animals. The seeds are bitter and poisonous when broken into, so that the seeds are normally ingested intact and efficiently dis- persed in animal droppings. Seeds germinate readily in moist shady places. Seedling foliage is essentially similar to that of the adult. Vigorous young plants tend to have leaves larger than those of fully mature specimens, up to two and three times as large and sometimes more lanceolate and falcate. Taxon. All seven species are closely related and some, at least, hybridize readily. As a result some authors such as PitGer prefer to recognize but one species with several subspecies. | would need to know more about the relationships between the taxa before I could take a strong position in this case. Uses. The tough, dense wood has excellent qualities and has been in demand for many uses. Best known is its service for bows and decorative woodwork such as chests and coffins. It is also desirable for fence posts, flooring, and mallots. The well-marked reddish brown heartwood contrasts pleasingly with the pale yellowish sapwood. 1. Taxus sumatrana (Mig.) pe LAus. Kalikasan 7 — nensis Piccer, Pfl. R. 1V, 5, Heft 18 (1903) 112; in (1978) 151. — Cephalotaxus sumatrana Mig. Fl. Ind. —_E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 210. — T. Bat. 2 (1859) 1076. — Podocarpus celebicus Hemst. — baccata ssp. wallichiana (non Zucc.) PicceER, Pfl. R. Kew Bull. (1896) 39. — Cephalotaxus celebica 1V, 5, Heft 18 (1903) 112; BUNNEMEVER, Trop. Na- Wars. Monsunia 1 (1900) 194. — Cephalotaxus tuur 10 (1921) 55, f. 8; STEEN. Bull. Jard. Bot. Btzg mannii (non Hoox./.) Pritzet ex Diets, Bot. Jahrb. Ill, 13 (1934) 194; Sreup, Trop. Natuur Jub. no. 29 (1900) 214; Witson, J. Arn. Arb. 7 (1926) 40. — (1936) 41, f. 1. — 7. baccata var. sinensis HENRY in T. baccata (non L.) Masrers, J. Linn. Soc. Bot. 26 Elwes & Henry, Trees Gr. Brit. & Irel. 1 (1906) 100. (1902) 546, p.p. — T. baccata ssp. cuspidata var. chi- =— T. wallichiana (non Zucc.) Foxw. Philip. J. Se. 350 Fig. 2. Taxus sumatrana (MiqQ.) DE LAvB. on the peak of Mt Bonthain, Celebes, 1700 m alt. (Photogr. J.VAN ZIJLL DE JONG, 1933). 6 (1911) Bot. 166; Merr. En. Philip. Fl. Pl. 1 (1923) 5; YAMAMOTO, J. Soc. Trop. Agric. 10 (1938) 182, f. 42. — Tsuga mairei LEMEE & LEVEILLE, Le Monde des Plantes, année 16 (2me sér.) n. 88 (May 1914) 20; Bull. Acad. Int. Geog. Bot. 16 (1914) 20. — T. cuspidata var. chinensis (PILGER) REHDER & WILSON in Sargent, Pl. Wils. 2 (1914) 8, p.p. — T. cuspidata (non Step. & Succ.) KANEH. Formos. Trees (1917) 616. — T. chinensis (PILGER) REHDER, J. Arn. Arb. 1 (1919) 51; ibid. 4 (1923) 119; DALLIMORE & JACK- son, Handb. Conif. (1923) 71; Witson, J. Arn. Arb. 7 (1926) 41; ibid. 8 (1927) 88; HAND.-Mazz. Symb. Sin. 7 (1929) 2; Hu & Cuwn, Icon. PI. Sin. 2, 8 (1929) pl. 53; BEAN, Trees & Shrubs Brit. Is. 3 (1933) 476; Orr, Not. R. Bot. Gard. Edinb. 18 (1933) 124; ibid. 19 (1937) 261; KANEH. Formos. Trees rev. ed. (1936) 31; REHDER, Man. Cult. Trees & Shrubs 2 (1940) 3; MetcaLF, Fl. Fukien | (1942) 23; FaNa, Icon. PI. Omeiens. II, 2 (1946) t. 190; Law, Bot. Bull. Acad. Sin. I, 2 (1947) 143. — T. wallichiana var. chinensis (PILGER) FLorin, Acta Horti Berg. 14 (1948) 378, pl. 5; GAUSSEN, Gymn. Act. & Foss. fasc. 15, ch. 25 FLORA MALESIANA [ser. I, vol. 103 (1979) 16. — T. speciosa Ftorin, Acta Horti Berg. 14 (1948) 382, pl. 6; L1 & KENG, Taiwania 1 (1954) 29, pl. 2. — T. mairei (LEMEE & LEVEILLE) Hu & Liv, II- lus. Nat. & Introd. Lign. Pl. Taiwan 1 (1960) 16; GAUSSEN, Gymn. Act. & Foss. fasc. 15, ch. 25 (1979) 16, f. 858. — T. celebica (WALL.) L1, Woody FI. Taiwan (1963) 34; Harrison, Handb. Conif. & Ginkgo (1967) 598. — T. yunnanensis CHENG, CHENG & Fu, Acta Phytotax. Sin. 13 (4) (1975) 86. — Fig. 2, 3. Fig. 3. Taxus sumatrana (MiQ.) DE LAuB. Habit, fe- male, with fruit, x 1 (DE LAUBENFELS P668). Large, slow-growing tree to as much as 45 m high and over | m diam., but considerably shorter on ex- posed ridges. Leaves on juvenile specimens or on vig- orous shoots linear lanceolate and often falcate with a prominent bend near the base and also sometimes with a slight reverse curve near the apex which is nar- rowly acute and often slightly spiculate, 2—4 cm long by 2—2.5 mm wide at the widest part below the centre of the leaf. Leaves on older specimens or from ex- posed positions more nearly linear and straight and abruptly narrowed at the apex, 1.5—2.5 cm by 1.5—2 mm, with slightly recurved margins. Pollen cones 1988] globular on a short stalk, about 4 mm in diameter. Mature seed c. 6 by 5 mm, and 4 mm thick. Distr. Eastern Himalayas, N. Burma, SE. Chi- CONIFERALES (de Laubenfels) 351 yond their early stages of growth in lowland China, while highland trees, as is usual elsewhere, are of much reduced stature. na, Taiwan, South Vietnam; in Malesia: Sumatra (from Karoland southwards to Benkulen), Philip- pines (Luzon: Lepanto, Benguet, Laguna, Tayabas; Mindanao; Davao), Celebes (Central and SW.: Bon- thain Peak). Fig. 1, 4. Ecol. Moist subtropical forests and tropical high- land ridges and mossy forests in the canopy and locally dominant; 1400—2300 m. Uses. A magnificent timber tree, but occurring too locally to be of importance and too slow-growing for cultivation. Vern. Tampinur batu, Karo, kaju tadji, Mt Dempo. Note. The extensive synonymy partly stems from the discontinuous distribution, but several authors insist that two types exist in China. Whereas im- mense trees are seen in undisturbed forests of Taiwan and in Malesia, on the mainland only smaller trees are normally seen with one type reported mostly at lower elevation and another at higher elevation. Both types, however, often appear from the same collec- tion area and | was able to collect both from a single large Formosan tree, part from low on the tree and part from high up. It appears that trees rarely get be- Fig. 4. Range of Taxus sumatrana (M1Q.) DE LAUB. PODOCARPACEAE Monoecious or dioecious trees and shrubs, some prostrate (and one parasitic on another member of the family, Parasitaxus, in New Caledonia). Each cotyle- don, of which there are usually two but in a few cases more, a fused pair with a corresponding bifid tip. Foliage buds ranging from a loose cluster of reduced leaves to a complex specialized structure (in Podocarpus). Leaves of many shapes and sizes. Pollen produced in small cones with many microsporophylls, each of which have two inverted dorsal pollen sacs above which is a small sterile tip. Male cones may be solitary in the axils of ordinary leaves, sometimes many adjacent cone subtending leaves, or they may be terminal or clustered on special structures involving sterile scales, or in a few cases they may arise in the axils of scales at the base of a new foliage shoot. The po//en for all genera (except extra-Mal. Saxegothaea) is provided with two or more bladders or ‘wings’, a trait shared with many genera of Pinaceae. More than two are found only in Dacrycarpus (and extra-Mal. Microcarpus and Microcachrys). The basic seed producing structure in Podocarpaceae is a compound terminal or lateral cone in which fertile scales arise in the axils of cone bracts. The cone is further often subtended by a specialized shoot with scales or modified leaves or even a naked peduncle. The bract is usually a small scale but may be larger and in some cases 352 FLORA MALESIANA [ser. I, vol. 103 hardly differs from foliage leaves. The fertile scale or epimatium is a highly modified shoot and in this family bears a single naked ovule on its upper sur- face. In one genus there is no scale while this and one other genus have erect ovules, but in the great majority of genera the ovule is inverted. The fertile scale cups the developing seed and may even completely surround it with only the micropyle protruding at the time of pollination. Fleshiness, either of the cone bracts or of the fertile scale (or both) is common and the whole female structure may be reduced to only one or a few fertile units and a few sterile units. This can yield plum-like fruits or, in other cases, structures resembling those of the cashew (Anacardium occidentale). Distribution. There is a strong Antarctic relationship with a broad extension into Malesia for the 172 known species in 13 genera (in Malesia7 genera with 61 spp.). Of six local and generally primitive genera, four are in the Antarctic zone and two are in New Caledonia. Three wide- ranging genera extend also into the tropical American highlands and two of these further range across the tropical African highlands. All seven of the wide-ranging genera are common in Male- sia, only one of which (Falcatifolium), however, is confined to the Asian tropics; five are in New Zealand. A few species reach into moist subtropical forests of eastern Asia. Fossils. The early fossil record is quite limited unless one includes forms that merely resemble the Podocarpaceae and whose relationships range from uncertain to highly doubtful. Suggestive macrofossils and pollen of the Jurassic indicate that the family was probably already present in India as well as in the far southern latitudes including New Zealand and W. Antarctica, that is to say Gondwanaland. Similar finds continue through the Cretaceous except for India, where Po- docarpaceous fossils no longer occur. In the Eocene and Oligocene, fossils of this family become more abundant in the higher southern latitudes and can often be assigned to modern genera. Fos- sils, particularly pollen, show that some of the presently endemic genera such as (extra-Mal.) Ac- mopyle and Microcachrys were formerly of much greater distribution. Some recent authors insist that fossils of Podocarpaceae occur in boreal regions. FERGUSON (1967) identified Podocarpus among Cenozoic fossils from Europe, suggesting a recent wider pre- glacial expansion of the genus. REYMANOWNA (1975) recently recognized Dacrydium and Stachy- carpus (Prumnopitys) from the Jurassic of Europe. Furthermore, I have been told by palynologists that typical Podocarpaceae type pollen is well represented in northern latitudes. On the other hand, FLorin (1963), after having examined both such alleged macrofossils and pollen fossils, expressed strong scepticism concerning their relationship to Podocarpaceae while pointing out that southern hemisphere strata are rich in Podocarpaceous remains. Certainly if any such plants ever existed beyond subtropical China and India, they have since disappeared completely while primitive forms of the family now survive only in the far south where they also have a res- pectable fossil history. Fossil and present-day distributions suggest that the Podocarpaceae, as we know it, developed in cool moist Antarctic forests early in the Mesozoic period. Modern genera were already differen- tiated before the parts of Gondwanaland became isolated. Several genera had reached India and Kerguelen where they eventually disappeared, but two genera still survive in Africa. All of the im- portant genera were included in the South American landmass where one interesting endemic ge- nus, Saxegothaea, is also still found, but some of the other genera have since become extinct there. Probably all the recognized living genera had already differentiated before the Tertiary as a major element of the Antarctic flora. Only in Malesia and probably only in late Tertiary times have mem- bers of this family descended into the lowlands of the tropics to any significant extent. They are still a major element not only in the Antarctic forests and in the Malesian highlands, but also in the tropical highlands of Africa and America. 1988] CONIFERALES (de Laubenfels) 353 Maps of fossil distribution are given by CouPER (1960) and FLORIN (1963). References: COUPER, Proc. R. Soc. Lond. ser. B, 152 (1960) 491—500, maps; FERGUSON, Palaeogeogr., Palaeoclimatol., Palaeoecol. 3 (1967) 73—110; FLorin, Acta Horti Berg. 20 (4) (1963) 121-312, 68 maps; REYMANowNna, XIIth Intern. Bot. Congr., Leningrad; Abstracts (1975) 99. Ecology. Mostly trees of moist forests at all elevations and well into the middle latitudes both as major canopy trees and as understory plants, rarely in areas with a marked dry season. A few species are scrubby or even prostrate and as such may be found beyond the tree line, on rocky outcrops or other specialized habitats. The family is well represented in, but not confined to dif- ficult soils such as sand and ultrabasics as well as in mossy forests. There are nodules regularly present on the roots, but their function is unclear. FURMAN (Amer. J. Bot. 57, 1979, 910) showed that they contained endotrophic mycorrhizae and that nitrogen fixa- tion did not occur. BECKING (Ann. Inst. Pasteur. 111, 1966, 295) indicated that the mycorrhizae were Phycomycetes. Growth is possible in sterile soil without mycorrhizae. Most genera are dioecious and pollination is by wind. Individuals are usually scattered but lo- cally common and the large quantities of pollen that are often produced seem to be able to reach effectively across considerable distances. Seedlings are found scattered and even quite isolated from seed sources due no doubt to dispersal by birds or fruit bats which eat the fleshy fruit. Co- ordination of fruiting times is for many species in tropical regions not well developed because at any given time it is often possible to find examples at any and all stages of reproduction and the collection of ripe fruit is variously reported for a given species at disparate dates. I have even seen two stages on the same tree. Growth is distinctly cyclic and in some genera there are elaborate terminal buds and similar buds for pollen cones. Seed-bearing structures are usually produced on the latest shoots while pollen cones frequently emerge from shoots of the previous cycle. Sometimes leaves of only the last cycle persist on a tree but more common is the display of three or four cycles of growth. Seeds germinate on or near the surface of the forest floor. The cotyledons remain at least partly inside the seed coat absorbing nourishment from the endosperm while the radicle penetrates the soil and begins forming a root system. Eventually the linear cotyledons shed the emptied seed coat and persist at the base of the growing shoot for a variable length of time. When functioning leaves are established, the cotyledons will be shed. Even if the adult leaves have some other form, the first foliage leaves in almost all taxa are bifacially flattened, often with an abrupt transition where the adult foliage is distinct. Various parasites are known for this family. Members of Podocarpaceae are the exclusive hosts of three genera of fungus in the family Coryneliales, in Malesia recorded for Podocarpus crassi- gemmis. Their fruiting bodies can often be seen erupting from leaves or stems in Podocarpus or Nageia, but this does not seem to be particularly harmful. One species of Korthalsella (Viscaceae), a dwarf mistletoe, is also parasitic in Podocarpaceae: K. dacrydii has been reported both on Dacrycarpus and on Dacrydium in various parts of Malesia (WASSCHER, Blumea 4, 1941, 320, 1 map). Embryology. The fertilized egg undergoes four or five mitoses resulting in up to 16 to 32 free nuclei. Most of these are then walled off and cluster at the base of the archaegonium forming a pro-embryo of several tiers of cells. Those in the lowest tier are embryonic and divide to form binucleate cells of which there may be but one to in some genera as many as 16. The next tier of cells elongates into a ‘prosuspensor’ consisting of from 3 to 25 cells, the number of cells being roughly proportional to the size of the seed and therefore the length needed to reach the centre of the female gametophyte (later to become endosperm). A third tier of cells is not completely walled off and is left behind to degenerate as the embryonic mass is projected away. At the apex of the embryo there may be one or a few cells forming a ‘cap’. Unless there are five mitoses (i.e. Nageia and Prumnopitys — both with large seeds) a larger number of suspensor cells means fewer embryonic cells. In the majority of cases the embryonic mass divides, along with the secondary 354 FLORA MALESIANA [ser. I, vol. 10° suspensor which it generates, into several competing units, the common conifer condition known as cleavage polyembryony. Simple polyembryony resulting from more than one fertilized archae- gonium also occurs. Growth of the embryo begins when the nuclei of the binucleate cells divide and then form groups of four cells. Probably an actual developed embryo derives from but a single binucleate cell so that, when there are more, they are competitive. The reduction of the number of embryonic cells often to a single cell in the genus Podocarpus appears to be a derived character. The binucleate embryo stage itself is unique in Podocarpaceae, while the number of mitoses leading to the pro-embryo is intermediate between a large number for Araucariaceae and non-coniferous Gymnosperms on the one hand and a smaller number for most other conifers on the other. Sciadopitys in the Taxodiaceae has five, while Cephalotaxus and most of the Taxaceae also have four. Chromosomes. According to Hair & BEUZENBERG (Nature 181, 1958, 1584) the chromo- somes in Podocarpaceae are remarkable. Basically the number for the great majority is in effect n=10 while for Phyllocladus it is n=9 (and for extra-Mal. Halocarpus n= 8). For a great many species in most genera, however, there are two kinds of chromosomes. One type, always present, is median to submedian, while the other, sometimes present, is subterminal to subtelocentric. Two of the latter always correspond to one of the former indicating either a progressive splitting of some of the chromosomes or less likely a progressive pairwise fusion of some or all of the chromo- somes. Phyllocladus and the genera with bilaterally flattened leaves (Dacrycarpus, Falcatifolium, and Acmopyle) have only the one kind of chromosome. The large genera Dacrydium, Nageia, and Podocarpus are partly with one kind and partly mixed. The other six (mostly small) genera always have mixed chromosome types. The result is a wide range of actual chromosome numbers from n=8 ton=19. Occasionally hybrids have been noted or suspected. Many species occur side by side in nature without any apparent hybridization. Taxonomy. Two recent works have treated all of what is recognized as a single family here. GAUSSEN (Les Gymnosperms actuelles et Fossiles, fasc. 13 & 14, 1974 & 1976) separates each of the three most distinct genera into families of their own, viz. Saxegothaeaceae, Phyllocladaceae, and Pherosphaeraceae. He recognizes one section of Nageia (Afrocarpus) as a distinct genus, while grouping the rest of this genus and Parasitaxus with Podocarpus. There are eight genera in Podocarpaceae as he envisions it. In my taxonomic revision (J. Arn. Arb. 50, 1969, 274—369) I recognize a single family and 13 genera (including the recently published Halocarpus by implica- tion only). Uses. The wood of trees in this family is light coloured, usually yellowish, is durable, easy to work, and generally similar to pine though rather harder. It is extensively used for lumber where sufficiently dense stands of good-sized trees occur, mostly outside of Malesia. In Borneo wood of Nageia is sometimes mixed with Agathis (‘dammar’) in commercial cuttings. Specimens of many genera are selected for planting around native settlements although the specimens seen in urban areas within Malesia usually come from China or Japan. In fact, natives in many areas so prize the wood for construction that, as I have been told on several occasions and have confirmed through experience, it is often necessary to go some distance from the nearest village to find mature wild trees. In some species the fruits are edible and I have found a few in tropical America that were locally appreciated but I have not discovered any such example in Malesia. Note. Conifers lack flowers and even where brightly coloured fruit occurs it tends to be very transitory, thus conifers tend to be bypassed by collectors. Most genera are dioecious and separate collections of male and female are necessary. It is often desirable to have a juvenile specimen (low branches in the shade usually have the juvenile form) to appreciate the range of foliage form. Sometimes immense numbers of recently shed pollen cones are encountered on the forest floor and these are worth collecting. 1988] CONIFERALES (de Laubenfels) as VEGETATIVE KEY TO THE GENERA veltacein the,form,of = = = CONIFERALES (de Laubenfels) Fig. 21. Range of Dacrydium comosum CORNER (dots) and D. ericoides DE LAuB. (triangles). 14. Dacrydium ericoides DE LAUB., nov. sp. Arbor ad 17 malta. Folia linearia recta, paginis su- peris planis, apicis abruptis apiculatis, 5—10 mm longa, 0.7—1 mm lata, 0.2 mm crassa, dorsis carina- tis. Strobili fere laterali, masculi 7—10 mm longi, 2—2.5 mm diametri, apicis microsporophyllorum calcaria 1 mm longa. Type: BRUNIG S 8722 (L, holo), Merurong Plateau, Sarawak. Tree 10—17 m tall, 25—30 cm diam., with drooping twigs. Leaves linear, straight, spread out more or less perpendicular to the shoot except on new growth, narrowing abruptly at the apex to an apiculate tip, flat on the upper surface but becoming slightly con- cave towards the apex, stomata on the upper surface in two bands separated over the midvein, sharply keeled on the lower surface, 5—10 mm long, 0.7—1 mm wide, 0.2 mm thick. Fertile structures usually lateral, subtended by a cluster of reduced leaves which are 2—3 mm long. Pollen cones 7-10 mm long and 2—2.5 mm diam. Apex of the microsporophyll a lanceolate spur c. 1 mm long and 0.7 mm wide. Seed bracts 3—4 mm long with sometimes two fertile. Mature seed unknown. 371 Distr. Malesia: Borneo (Sarawak, known only from Mt Dulit and the Merurong Plateau in N. Sara- wak). Fig. 21. Ecol. Locally common in primary forest on ex- posed mossy ridges at 1000 to 1500 m. Vern. Sempilor, Bintulu. Note. The spreading straight linear leaves con- trast rather strikingly with other members of the ge- nus and rather resemble the juvenile foliage of Cupressaceae. Earlier | had included it in D. spathoides where the shorter leaves are also more or less linear and much wider than thick, but in this spe- cies the fertile structures are usually terminal and are subtended by nearly typical leaves not greatly re- duced as in D. ericoides, while the leaves are distinct- ly bent forward and not straight. 15. Dacrydium leptophyllum (WASSCHER) DE LAUB., nov. comb. — Podocarpus leptophylla WaAssCHER, Blumea 4 (1941) 414, t. 4, f. 9. — Dacrycarpus lepto- Phylla (WASSCHER) GAUSSEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 150. Leaves diverging widely from the stem but sharply bent forward parallel to the stem or even directed in- ward, lanceolate, pungent, 1—1.5 mm long, 0.2—0.3 mm wide, 0.1 mm thick, strongly keeled on the dor- sal side, flat or slightly concave on the axial side. Leaves on vigorous branches larger, up to 3 mm long and 0.6 mm wide. Fertile material unknown. Distr. Malesia: West New Guinea (known only from the top of Mt Goliath), at 3000—3600 m. Note. The original description expressed uncer- tainty between Dacrydium and Podocarpus sect. Dacrycarpus for this unique taxon, but unfortunate- ly settled for the latter. The tiny leaves are typical for Dacrydium and the primary branches show no sign of the dimorphism which characterizes Dacrycarpus. 3. FALCATIFOLIUM DE Laus. J. Arn. Arb. 50 (1969) 308; Fl. Nouv. Caléd. et Dép. 4 (1972) 30; GAUSSEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 67. — Fig. 22. Dioecious shrubs to large trees to 36 m tall with thin more or less smooth brownish bark with scattered lenticels, reddish and somewhat fibrous within, breaking off in occasional flakes on larger specimens. Loosely and irregularly branched. Leaves spirally placed, single veined, and alternating with elongated appressed scales which are loosely clustered at the shoot apices to form foliar buds between episodes of growth. Seedling leaves narrowly lenticular, apiculate, bifacially flattened, giving way abruptly to distinct juvenile leaves in about the second year of growth. Juvenile and adult leaves distichous, bilater- ally flattened and falcately curved away from the branch with the apex in most cases oppositely curved in the direction of shoot growth. Reproductive struc- 372 FLORA MALESIANA [ser. I, vol. 10° tures on short scaly shoots which are either axillary or terminal and may bear a few reduced leaves. Pollen cones cylindrical, solitary or clustered; microsporo- phyll a small acuminate spur above the two pollen sacs. Seed-bearing structures solitary, consisting of up to about a dozen large acuminate scales which become greatly swollen, red, and fleshy when mature; normally one subapical scale fer- tile with a cup-shaped epimatium which has a distinct hump opposite the base of the included seed positioned well beyond the subtending fleshy scale so that the solitary seed and its basal humped epimatium are fully exposed; the inverted ovule gradually turning upward as it matures into a nearly erect seed; the mature seed with two lateral weak ridges along its wider sides which come together in an apical ridge, otherwise the seed is more or less egg-shaped. Distr. New Caledonia (1 sp.); in Malesia: New Guinea, Moluccas (Obi I.), N. & Central Celebes, Philip- pines (Mindoro), Borneo, Riouw-Lingga Arch. (Lingga), and Malaya. Note. Obviously related to Dacrydium but differing in the dimorphic foliage with specialized fertile shoots and the exposed hump of the epimatium opposite the base of the seed. In Dacrydium the base of the seed lies close to its attachment and is always well covered by the subtending bract. KEY LO THE SPECTE'S 1. Adult leaves normally bent at least slightly forward at the apex, tapering from at least the centre of the leaf; pollen cones at least 17 mm long. 2. Adult leaves linear-lanceolate, sun leaves at least 20 mm long, not glaucous; pollen cone 2.5—3.5 mm ONZE Mis Jove Beg c 4 6 cuo.c. 0 6.61 HO Iad AROS ERE tc Sea NEE ORS Cn SM ERs Par SE Rei 1. F. falciforme 2. Adult leaves mostly lanceolate, sun leaves 13—20 mm long, glaucous; pollen cone 1.5—3 mm diam. 2. F. gruezoi 1. Adult leaves often not bent forward at the apex, the sides parallel for most of their length; pollen cones no more than 13 mm long. 3. Adult leaves weakly keeled if at all, 2—3.5 mm wide, 12—20 mm long ............ 3. F. papuanum 3. Adult leaves distinctly keeled on their broader surfaces, 1—2.5 mm wide, 18—35 mm long 4. F. angustum 1. Falcatifolium falciforme (PARL.) DE LAus. J. Arn. Arb. 50 (1969) 309; GaussEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 73. — Podocarpus falciformis ParL. in DC. Prod. 16, 2 (1868) 685. — Nageia falciformis (PARL.) O.K. Rev. Gen. Pl. 2 (1891) 800. — Dacrydium falciforme (PARL.) PILGER, Pfl. R. IV, 5, Heft 18 (1903) 45; Foxw. Philip. J. Sc. 6 (1911) Bot. 153; RrpLey, J. Str. Br. R. As. Soc. n. 60 (1911) 56; PitGER, Bot. Jahrb. 54 (1916) 35; Stapr, J. Linn. Soc. Bot. 42 (1914) 191, f. 8; BurKiLL & HoLTTum, Gard. Bull. S. S. 3 (1923) 76; RipLey, Fl. Mal. Pen. 5 (1925) 280; CorRNER, Wayside Trees (1940) 722; KENG in Whitmore, Tree Fl. Mal. 1 (1972) 46, f. 2. — Fig. 22. Large shrub from 1.5 m to occasionally a large tree as much as 36 m tall, more commonly 5—12 m, 4—40 cm diam. Seedling leaves widening gradually from a petiole several mm long to margins parallel in the middle of the leaf, apex more abrupt, acute, apiculate, midrib a low blunt ridge above and a nar- row sharp ridge below, slightly revolute, 4—9 by 2—3.5 mm. Juvenile leaves on the first branches only slightly longer and wider than the seedling but soon becoming as much as 12 cm long and more gradually becoming as much as 12 mm wide, the lanceolate apex strongly curved so as to become parallel to the shoot, midribs on either side a weak ridge. Adult shade leaves spreading at a large angle with more or less parallel margins in the centre of the leaves and broadly lanceolate apex which curves strongly for- ward but still at an angle from the shoot, 4—7 cm by 5—9 mm. Adult sun leaves much more abrupt at both ends so as to form a broad lens shape to almost a par- allelogram with rounded corners, 2—4 cm long by 5—7 mm wide, the apex sometimes not bent forward. Pollen cones 2—4 cm long by 2.5—3.5 mm diam. Re- ceptacle of seed-bearing structure 4—5 mm long; ma- ture seed 6—7 mm long, 5 mm wide, and 3.5—4 mm thick, becoming black. Distr. Malesia: Malaya, Riouw-Lingga Arch. (Lingga: P. Tanda) and Borneo (mainly Sarawak and Sabah). Fig. 23. Ecol. Locally common along ridges as a bushy tree or in the subcanopy of primary rain-forest, often 1988] Fig. 22. Falcatifolium falciforme (PARL.) DE LAvuB. Twig with male cone, x | (after Wyatt-SMITH KEP 93115). on podsol sands and kerangas, but occasionally on deeper fertile soils a somewhat emergent forest giant, from 400—2100 m. Vern. Kayu china, Sabah, Lahad Datu, iguh gawah, \ban, Merurong Plateau. Notes. Inthe forests of Mt Kinabalu the juvenile plants have smaller leaves than elsewhere, but other- wise there do not appear to be any differences. Sever- al collections of more or less juvenile material have been made in Celebes and Central Moluccas (Obi), but these resemble more F. gruezoi of the Philip- pines. A single specimen from high kerangas on the Usan Apan Plateau in Sarawak has leaves in the form of adult shade leaves, but these are only 6 by 2 mm. CONIFERALES (de Laubenfels) 373 It is called a ‘young tree’ but given as 24 m tall. This may well be a new species. 2. Falcatifolium gruezoi DE LAUB., nov. sp. — Dacrydium falciforme [non (PARL.) PILGER] Foxw. ex MERR. Philip J. Sc. 2 (1907) Bot. 257; Foxw. ibid. 6 (1911) Bot. 153, t. 28, f. 1; MERR. En. Philip. 1 (1922) 4. Arbor 4—12 m alta. Folia juvenilia ad 7.5 cm longa, 7 mm lata, falcata et apice versus apex ramo- rum curvo, lanceolata; folia adulta umbrae minora, 3.5 cm longa, 6—7 mm lata; folia solis plus minora, 13—20 mm longa, 3.5—6 mm lata, acuta, apiculata, glauca. Strobili masculi 1.7—6 cm longi, 1.5—3 mm diametri. Strobili feminei receptaculo 2 mm, semina 7 mm longo. Type: GRUEZO WM 4052 (L, holo; CALP, iso), Naujan, Paitan access, Paitaraan (Mt Halcon area), Mindoro Oriental, Philippines. Tree 4—12 m tall. Juvenile leaves to 7.5 cm by 7 mm, falcate with the apex curved forward more or less parallel with the branch; lanceolate. Adult leaves in the shade smaller, 3.5 cm by 6—7 mm; sun leaves even smaller, 13—20 by 3.5—6 mm, acute, apiculate, glaucous. Pollen cones 1.7—6 cm long and 1.5—3 mm diam. Receptacle of the seed-bearing structure 2 mm long; seed 7 mm long. Distr. Malesia: Philippines (Luzon: Tayabas, Nueva Ecija; Mindoro; Panay; Mindanao: Davao, Surigao); Celebes: Manado (Poso; Gorontalo; Palu); Moluccas (Obi). Fig. 23. Veet 4 Se R\ Sra aes OS @ Falcatifolium falciforme (Pax..) or Lau. & Falcatifolium gruezoi 08 LAUD. 4 Falcatifolium papuanum oe Laun ane a se © & Falcatifolium angustum 0® LAUD. Fig. 23. Range of four species-of the genus Falcatifo= lium. Ecol. In exposed locations along ridges or on the borders of open areas, 1600-2200 m in the Philip- pines, 1200-1400 m in Celebes, 700 m in Obi. Note. See note under F. falciforme. 3. Faleatifolium papuanum pe LAus. J. Arn. Arb. 50 (1969) 312, f. 6; Blumea 17 (1969) 274; GAuUSsEN, Gymn. Act. & Foss. fase. 13, ch. 20 (1974) 73, f. 698. — Dacrydium falciforme {non (Part.) PILGerR] Laut. Bot. Jahrb. 68 (1937) 247. 374 Tree 6—22 m tall, 8—40 cm diam. Seedling leaves 6—18 by 0.6—0.8 mm. Juvenile leaves the same as the smaller adult leaves, glaucous beneath. Adult leaves falcate and then more or less linear in the distal part of the leaf or tapering slightly, narrowing almost ab- ruptly to an apiculate apex, the apex occasionally bent slightly forward, 10-20 by 2—4 mm. Pollen cones 5—13 mm long and 2—2.5 mm diam. Recep- tacle and seed each 6—7 mm long. Distr. Malesia: New Guinea. Fig. 23. Ecol. Understory tree of moist mountain forests, often associated with Nothofagus spp., Myrtaceae and other Podocarpaceae, 1500—2400 m. Vern. Mungag, Hagen Togoba, tug/, Wahgi, Min}. Note. Anentire plant scarcely 20 cm tall with tiny leaves mentioned and illustrated in the type descrip- tion from the Vogelkop either represents perhaps a FLORA MALESIANA [ser. I, vol. 103 reduced form of exposed ridges or a distinct new spe- cies. 4. Falcatifolium angustum be Laus. J. Arn. Arb. 50 (1969) 312, f. 7a; GAUSsSEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 73, f. 699. Tree to 20 m tall, 6—25 cm diam. Juvenile leaves narrowly lanceolate and gradually curved slightly forward towards the apex, c. 7 cm long and 1.2 mm near the base. Adult leaves less curved or straight, pungent, keeled on each side, 18—35 by 1—2.5 mm. Somewhat immature pollen cones 8 mm long and 2 mm diam. Seed-bearing structures unknown. Distr. Malesia: Borneo (known from two loca- tions near the coast of Sarawak). Fig. 23. Ecol. In forests, 90—240 m, on podsolized sands and kerangas, associated with Parastemon, Shorea albida, and Gymnostoma sp. 4. DACRYCARPUS (ENDL.) DE Laus. J. Arn. Arb. 50 (1969) 315; Fl. Nouv. Caléd. et Dép. 4 (1972) 34; GAUSSEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 133; DE LAUB. Kalikasan 7 (1978) 125; vAN RoyeNn, Alpine Fl. New Guinea 2 (1979) 11. — Podocarpus sect. Dacrycarpus ENDL. Syn. Conif. (1847) 221; CARRIERE, Traité Gen. Conif. ed. 1 (1855) 477; ed. 2 (1867) 676; GorDOoN, Pinetum ed. 1 (1858) 289; ed. 2 (1875) 356; Mia. Fl. Ind. Bat. 2 (1859) 1074; HENKEL & HOCHSTETTER, Synop. Nadelholz. (1865) 403; DE Borer, Conif. Arch. Ind. (1866) 25; PARL. in DC. Prod. 16, 2 (1868) 520; DE Kirwan, Conif. 2 (1868) 224; EICHLER in E. & P. Nat. Pfl. Fam. II, 1 (1889) 105; BrissNER, Nadelholzkunde (1891) 17; Pricer, Pfl. R. IV, 5, Heft 18 (1903) 55; in E. & P. Nat. Pfl. Fam., Nachtr. 3 (1908) 3; Foxw. Philip J. Sc. 6 (1911) Bot. 156; Stites, Ann. Bot. 26 (1912) 448; Grpss, Ann. Bot. 26 (1912) 525; PILGER in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 242; HicKEL, Fl. Gén. I.-C. 5 (1931) 1066; WasscHER, Blumea 4 (1941) 386; BUCHHOLZ & Gray, J. Arn. Arb. 29 (1948) 56. — Podocarpus sect. Dacrydioi- deae BENNETT in Bennett & R.Br. Pl. Jav. Rar. 1 (1838) 41. — Podocarpus sect. Dacrydium BERTRAND, Ann. Sc. Nat. V, 20 (1874) 67. — Fig. 26, 28, 31—33. Dioecious shrubs or trees, to 41 m tall. Bark hard, dark brown or blackish but weathering to gray, surface rough with occasional lenticels, inside pink to reddish brown and granular or slightly fibrous, on older trees breaking off in small thick, somewhat vertically elongated plates or sometimes short strips. Leaves amphistomatic, spirally placed, broadly decurrent, apiculate. Leaves on primary shoots as well as on the basis of foliage shoots and fertile structures bi- facially flattened, keeled on the dorsal side, lanceolate or sometimes triangular, often nearly appressed, mostly 1.5—3 by 0.5—0.8 mm, but wider on the decur- rent part, up to at least 4 mm long on young plants and mostly 1—2 mm long at the base of foliage shoots or fertile structures. The apex of resting shoots a 1988] CONIFERALES (de Laubenfels) 375 loose cluster of reduced leaves. Juvenile type of leaves on special shoots that generally do not continue growth after reaching a resting stage, bilaterally flat- tened and usually slightly keeled on both faces, falcate and then curved forward towards the apex so that the apiculate tip is oriented more or less parallel with the shoot, otherwise linear or less often lanceolate, spreading at about a 60° angle, the centre of the leaf either straight or gradually curving to the tip, dis- tinctly shorter towards either end of the shoot so that the whole shoot has a feather-like appearance, mostly 6—12 mm long and c. 1 mm wide, usually distichous, gradually changing to an adult form but often fertile at intermediate stages of this transition. Final adult forms not distichous, generally shorter and more robust than the juvenile leaves, more or less uniform along the shoot, the apiculate tip still bent forward parallel to the shoot, in some species nearly iden- tical with the leaves of primary shoots. Fertile structures terminal on short, mostly lateral shoots, the seed-bearing structures usually on a considerably longer shoot than that of the pollen cone. Immature pollen cones at first some- times nearly spherical, then becoming somewhat elongated but finally elongat- ing abruptly with a slight decrease in diameter at maturity, then mostly c. 6—10 mm long and 2—3 mm diam., sometimes longer. Apex of microsporophyll trian- gular, acute to apiculate, c. 1.2 by 0.8 mm. Shoots for female structures 3—17 mm long. Leaves at the base of the seed-bearing structure sharply elongated to form an involucre which often surrounds the immature seed-bearing structure but which in the shorter examples becomes spreading as the structure grows. Seed-bearing structure composed of a small warty receptacle, 2.5—4 mm long, 2.5mm diam., which becomes greatly enlarged, fleshy, and first orange then red or in other species purple when ripe, later turning brown, bearing one or two protruding short sterile leaf-like bracts and one or two subterminal fertile bracts. The inverted ovules completely surrounded by the epimatium and fused as a rib along one side with the fertile bract whose short free tip forms a small off-centre crest over the mature structure. Mature seed nearly spherical (or oval) but remaining covered by the leathery epimatium and scale, forming an erect or somewhat oblique structure which in most species is c. 5-6 mm long and 4.5—5.5 mm diam., dark in colour. Distr. Ranging from northern Burma and southernmost China to Fiji and New Zealand 9 spp.; in Malesia abundant with 7 spp., reaching their greatest variety in New Guinea with 5 spp. Fig. 24. Fossils are known from N. Antarctica (Graham Land) and New Zealand (Middle Jurassic) and S. Patagonia (Upper Cretaceous to Oligocene) where the genus is now extinct. In the Eocene found in New Zealand, and since the Oligocene also in SE. Australia, where it became extinct (FLoRIN, Kongl. Svensk. Vet. Ak. Handl. Ill, 19, n. 2, 1940, 70; Acta Horti Berg. 20 (4), 1963, 188, f. 19: map). Note. Sterile specimens strongly resemble Dacrydium and hence the generic name. The fusion of the fertile scale with the epimatium is a unique trait of the genus while the seeds of Dacrydium are furthermore naked. In most cases sterile specimens can be readily distinguished from Dacrydium by the distinctly dimorphic fo- liage. 376 FLORA MALESIANA [ser. I, vol. 10° Fig. 24. Range of the genus Dacrycarpus (ENDL.) DE LAus. Figures above the hyphen indicate the number of endemic species, that below the hyphen the total number of species. REY TO, DME SSPE GLEs 1. Involucral leaves short (2.5—5 mm long) and mostly spreading so that the immature receptacle becomes or more frequently is always exposed; adult leaves scale-like or equally keeled on four sides, 1—3 mm long. 2. Adult leaves less than 2 mm long, more or less in the form of scales .............. 1. D. imbricatus DaeAduitieavesrateledst 2mm long, in the fonm of short needles.) san. .se ss] le ees 2. D. steupii 1. Involucral leaves curved to surround the young fertile structure (3—13 mm long) and still covering at least the entire immature receptacle before it enlarges when ripe; adult leaves not scale-like nor equally keeled on four sides, 1—6 mm long. 3. Adult leaves bilaterally flattened. 4. Involucral leaves surrounding receptacle and mature seed (7-13 mm long); foliage leaves slender (OLS =—O23 inavon Kistolas Werhiteelhy eo lealy 2 oa mia oats c Rico Meola atte omomanonadca es oor 3. D. cumingii 4. Involucral leaves reaching the base only of the mature seed (5—8 mm long); foliage leaves robust (0.8—1 MIM Wide) PStLONS VA Keeled brs sister. oe Cite th Aces aise Slemicsa gcts sehesenstnustg tort atten 4. D. kinabaluensis 3. Adult leaves bifacially flattened. 5. Involucral leaves less than 5 mm long, not reaching the mature seed; foliage leaves 0.6—1 mm wide. 6, Seed not large/(6—6 mm) long) foliage! leaves spreading’. 12.056. 2s. 4.-60 42-6. se 5. D. expansus Gy iSeed larce!(/—8 mmi long); sfoliaselleavesiimbnicate! .3 5 anes dees ee eeele ee 6. D. compactus 5. Involucral leaves 6—10 mm long, mostly covering the mature seed; foliage leaves 0.4—0.6 mm wide 7. D. cinctus 1. Dacrycarpus imbricatus (BLUME) DE LAus. J. Arn. Kalikasan 7 (1978) 126; vAN RoyeEN, Alpine Fl. New Arb. 50 (1969) 320, f. 8a; GAussEN, Gymn. Act. & Guinea 2 (1979) 13. — Podocarpus imbricatus Foss. fasc. 13, ch. 20 (1974) 152, f. 724; pE Laus. BiumE, En. Pl. Jav. 1 (1827) 89; Pitcer, Pfl. R. IV, 1988] CONIFERALES (de Laubenfels) 377 5, Heft 18 (1903) 56; Koorpb. Exk. Fl. Java 1 (1911) 64, f. 2; Atlas 3 (1915) f. 585, 586; Fl. Tjibodas (1918) 3; PrcGer in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 245, f. 124E; HicKkEL, Fl. Gén. I.-C. 5 (1931) 1068; STEEN. Trop. Natuur 29 (1940) 75, 1 fig.; Was- SCHER, Blumea 4 (1941) 388, t. III, f. 2; BACKER & Baku./. Fl. Java 1 (1963) 89; STEEN. Mount. Fl. Java (1972) t. 13, f. 2. — Podocarpus cupressina R.BR. ex MIRBEL, Mém. Mus. Hist. Nat. Paris 13 (1925) 75, nomen; BENNETT in Bennett & R.Br. Pl. Jav. Rar. 1 (1838) 35, f. 10; ENpL. Syn. Conif. (1847) 222; BiumeE, Rumphia 3 (1847) 218, t. 172, f.2 & t. 172B, f. 2; Miq. Pl. Jungh. 1 (1851) 3; Fl. Ind. Bat. 2 (1859) 1074; HENKEL & HOcHSTETTER, Synop. Nadelhdolz. (1865) 403; pE Boer, Conif. Arch. Ind. (1866) 25; Par. in DC. Prod. 16, 2 (1868) 521; EICHLER in E. & P. Nat. Pfl. Fam. 2, 1 (1887) 106; Hook.f. FI. Br. Ind. 5 (1896) 650; Wars. Monsunia | (1900) 191; K. & V. Bijdr. Booms. Java 10 (1904) 262. — Podocar- pus javanicus (non Burm.f.) MERR. Philip J. Sc. 19 (1921) 338; En. Philip. 1 (1922) 3. — Fig. 26. KEY TO THE VARIETIES 1. Leaves slender (0.4—0.6 mm wide); involucral leaves always spreading. » Beaves imbricate .< ¢ ss ..«..-)- a. var. imbricatus . Beaves'spreading ...:...)....... b. var. patulus 1. Leaves robust (0.6—1 mm wide); involucral leaves sometimes clasping the receptacle. . Leaves spreading ........... c. var. robustus Nm NM w EER ewe aca erg rod Fo oe 3. eavesumibricates 2. 52..0..5% d. var. curvulus a. var. imbricatus Majestic columnar tree to 50 m tall, up to 2 m diam., crown large, often dome-shaped. Leaves of primary shoots imbricate; leaves of juvenile foliage shoots distichous, nearly linear, up to 10-17 mm long by 1.2—2.2 mm wide at the centre of a shoot, gradually losing the distichous habit as the tree ma- tures, but shoots with more or less bilaterally flat- tened leaves distinctly longer in the middle of the shoot almost always present on even the oldest trees. Terminal shoots on young plants often elongated whip-like up to 20 cm. Leaves on older trees even- tually becoming mostly scale-like, imbricate, dis- tinctly keeled on the dorsal side, long-triangular, 1—1.8 by 0.4—0.6 mm. Involucral leaves becoming spreading, acicular, 2.5—4 mm long, rarely to 5 mm. Ripe receptacle red. Distr. Malesia: Java, all Lesser Sunda Islands (Bali—Timor) and SW. & Central Celebes. Fig. 25. Ecol. Mostly scattered and common in primary and secondary rain-forest, not rarely as an emergent, and co-dominant in West Java with Podocarpus ne- riifolius and Altingia noronhae, on the south slope of Mt Tjeremai volcano characterizing the zone be- tween 2400—2700 m without other co-dominants, a situation not yet explained (VAN STEENIS, 1972), in Timor found under more or less seasonal conditions in isolated specimens laden with Usnea in grassland after deforestation, mostly between 1000—2500 m, ~ \ * \ ‘ A a Fig. 25. Range of Dacrycarpus imbricatus (BLUME) be LauB. var, imbricatus (triangles) and var, patulus DE Laus. (dots). 378 FLORA MALESIANA [ser. I, vol. 103 Fig. 26. Dacrycarpus imbricatus (BLUME) DE Lau. var. patulus DE LAUB. (drawing by R.S.KENG in Whit- more, Tree Flora of Malaya 1, 1972, 51, f. 4). SS 1988] but in Lombok reported as low as 200 m and in Cele- bes ascending to 3000 m. Probably exterminated at lower elevations in Java by deforestation. Male flow- ers at Tjibodas in Aug.—Sept. Sometimes the stem of full-grown trees produces sprouts at the base (VAN STEENIS, 1940). Uses. A most valuable timber tree. Vern. Java: djamudju, ki hadji, ki putri, tjemoro (tukung), Tjidadap, S; kadju pakis, tjemara binéh, Md.; SW. Celebes (Bonthain): kayu angin, k. pa- rang; Lesser Sunda Islands: Bali: tarupanda; Sum- ba: kKayu awama, Lairondja, kadju uamang; Lom- bok: majangmekar; Flores: oh-ru, Ruteng; Timor: haae tuni, W. Timor, Nenas, ai-caqueu fuie, E. Timor, Tetun lang. b. var. patulus DE Laus. J. Arn. Arb. 50 (1969) 320, f. 8b. — Podocarpus cupressina Riw.ey, J. Str. Br. R. As. Soc. n. 60 (1911) 58. — Podocarpus imbrica- tus sensu Gipss, Ann. Bot. 26 (1912) 525, t. 49, f. 1—8; Stapr, J. Linn. Soc. Bot. 42 (1914) 193; MERR. En. Born. Pl. (1921) 31; Rrpiey, Fl. Mal. Pen. 5 (1925) 283; Merr. Contr. Arn. Arb. 8 (1934) 14; CoRNER, Wayside Trees (1940) 723; KENG in Whit- more, Tree Fl. Malaya 1 (1972) 51, f. 4. — Podocar- pus kawaii HayatTa, Bull. Econ. Indochine 20 (1917) 439. — D. kawaii (HAYATA) GAUSSEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 140, f. 726. — Fig. 26. Tree, 5—40 m tall, 10—100 cm diam. Adult foliage leaves not imbricate, spreading sharply, acicular, dis- tinctly keeled on four sides, 0.8—1.5 by 0.4—0.6 mm. Involucral leaves to 3 mm long, spreading. Distr. Northern Burma and _ southernmost China, through Southeast Asia to Malesia: Sumatra, Malaya, Borneo, Philippines (Luzon, Mindanao), Central Celebes, along the N. coast of New Guinea (incl. New Britain and New Ireland), and New Hebrides to Fiji. Common, but not in Java. Fig. 25. Ecol. Scattered and common in primary and sec- ondary rain-forest, mostly between 700 and 2500 m, in N. Sumatra at c. 400 m on sinterlimestone near sulphur springs near Tinggi Radja, up to c. 3000 m in Borneo, and occasionally to near sea-level in Fiji. Vern. Malaya: ru bukit, Kedah; Sumatra: am- bun, Solok, W. Coast, balanidju, Kerintji, beru, Karo Lands, ki mérak, marak, Bencoolen, damanik, Simelungun, sampinur bunga, Karo-Toba, falas, Kroé, Bencoolen; Borneo: menjilu, Dusun lang., Kp. Tindai, Sabah, fampilas, Sensurun, Sabah, Dusun lang.; Celebes: siozi, Mt Wuka Tampai, Palu. Note. Only in Celebes is there an overlap with var. imbricatus with possibly transitional forms. In Borneo, Mindanao, and along the northern coast of New Guinea specimens approach var. robustus in form. Specimens from the western and eastern parts of the range are identical and easily distinguishable from other varieties. CONIFERALES (de Laubenfels) 379 c. var. robustus DE LAus. J. Arn. Arb. 50 (1969) 323, f. 8c; VAN RoyeEN, Alpine Fl. New Guinea 2 (1979) 16, f. 35d—g. — Podocarpus imbricatus sensu Foxw. Philip J. Sc. 6 (1911) Bot. 157. — Podocarpus pa- puanus RIDLEY, Trans. Linn. Soc. London II, 9 (1916) 158; GrspBs, Contr. Phytogeogr. Arfak Mts (1917) 80, f. 4; PiLGER in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 245; Bot. Jahrb. 68 (1937) 244; WASSCHER, Blumea 4 (1941) 402, t. 4, f. 3. — Podo- carpus javanica sensu MERR. Philip J. Sc. 19 (1921) 338; En. Philip. 1 (1922) 3, pro specimina. — Podocarpus cupressina sensu LANE-POOLE, For. Res. Terr. Papua & New Guinea (1925) 73. — D. papuana (RIDLEY) GAuSSEN, Gymn. Act. & Foss. fase. 13, ch. 20 (1974) 142, f. 731. — D. steupii (non DE LAUB.) DE Laus. Kalikasan 7 (1978) 127. Tree, 5—45 m tall, S—130 cm diam. Adult foliage leaves like var. patulus but distinctly more robust, 1.2—1.8 by 0.6—0.8 mm. Involucral leaves to 3 mm long and spreading or in some areas to 5 mm and more or less clasping the receptacle. Distr. Malesia: Borneo (Sarawak, once), Philip- pines (Luzon, Mindanao), Moluccas (Morotai, Ce- ram), and throughout New Guinea. Fig. 27. Fig. 27. Range of Dacrycarpus imbricatus (BLUME) DE LAuB. var. robustus DE LAuB. (dots) and var. cur- vulus (M1Q.) DE LAuB. (triangles). Ecol. Scattered and, in New Guinea, very com- mon in primary and secondary rain-forest, canopy tree, or sometimes emergent, often co-dominant, in mossy forest associated with Nothofagus and Phyllo- cladus, also in Lithocarpus-Castanopsis mixed forest co-dominant, (S00—)700—3000 m, a specimen from Borneo reported from 240 m. Vern. Borneo: pierur, Sarawak, Kelabit lang.; Philippines: ‘upi, Cotabato, Mindanao; West New Guinea: apé, Wissel Lakes, Kapauko lang., betjhiea, Arfak, Hattam lang., jamari, Wondiwoi Mts, Wandammen lang., kKaowie, kowi(e), Ransiki, Manikiong lang., nijoop, Kebar lang., toromoai, Dojodial, Cycloop Mts; East New Guinea: gubin, kubin, Hagen Togoba, iljo, Enga lang., kaibelpiti, Waghi, Minj, /ou, Wabag lang., pau, Lake Inim, 380 FLORA MALESIANA [ser. I, vol. 10° Enga lang., tibuidi, Anga Valley, Mendi lang., wba, Chimbu, Masul, umba, Waimambuno, paupeepeen, Mt Ne. Note. Specimens from Borneo and the Philip- pines have been identified as D. steupii where the fo- liage leaves fall within the size range of this species because in these areas the distichous habit is lost rath- er early causing the foliage to resemble D. steupii closely. Unlike D. steupii, however, the leaves range to the smaller sizes of D. imbricatus. The involucral leaves in these areas, unlike elsewhere in D. im- bricatus, are at least 5 mm long and curved upward. Perhaps this material represents a distinct variety. d. var. curvulus (Mig.) DE LAus. J. Arn. Arb. 50 (1969) 326, f. 8d; GAUssEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 154. — Podocarpus cupressina var. curvula Miq. Pl. Jungh. 1 (1851) 4; Fl. Ind. Bat. 2 (1859) 1074. — Podocarpus imbricatus var. curvula (MiQ.) WASSCHER, Blumea 4 (1941) 398. — Fig. 28. Shrubby pyramidal tree to 8 m tall and sometimes procumbent. Foliage shoots curved downwards but main branches curved upwards. Adult foliage leaves like var. imbricatus but distinctly more robust, 1.2—2 by 0.8—1 mm. Involucral leaves 2.5—4.5 mm long and more or less clasping the receptacle. Distr. Malesia: North Sumatra (Atjeh: Leuser complex & G. Bandahara) and western half of Java (Priangan; Diéng). Fig. 27. Ecol. In N. Sumatra pure stands on exposed mossy mountain peaks, on blangs and steep slopes, between 2000 and 3420 m. Vern. Tjamarah, J, at variance for Casuarina which does not occur so far west in Java. 2. Dacrycarpus steupii (WASSCHER) DE LAuB. J. Arn. Arb. 50 (1969) 328; GaussEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 158, f. 728; vAN ROYEN, Alpine Fl. New Guinea 2 (1979) 16, f. 35a—c. — Podocarpus steupii WASSCHER, Blumea 4 (1941) 405, t. 4, f. 4. — Podocarpus papuanus (non RIDLEY) STEUP, Trop. Natuur 27 (1938) 145. Conical tree, 4—36 m tall, 15—100 cm diam. Leaves of primary shoots spreading slightly. Leaves of juvenile foliage shoots distichous, nearly linear, soon losing the distichous habit as the tree matures. Leaves on older trees eventually becoming nearly quadrangular in cross section, widely spreading, tapering slightly, uniform in size along a shoot, 2—3 by 0.4-0.6 mm. Involucral leaves becoming spreading, 3—4 mm long. Distr. Malesia: Central E. Borneo (G. Beratus, Fig. 28. Dacrycarpus imbricatus (BLUME) DE LAUB. var.. curvulus (MIQ.) DE LAus. on Mt Bandahara, Sumatra, 2500 m. Habit of branch with female cones (Photogr. W.J.J.O.DE WILDE, 1972). 1988] near Balikpapan, once), Central Celebes (Latimod- jong Mts) and throughout New Guinea. Fig. 29. h PF ad 7 a . A L = a. | } Ln J Oh J ~ ioe 4 te ais ogee UV od a } ’ ar y me 4 | tld AY >= ~ 7 oe % ne = — a Z = +> \ r Bs ~ 4 Dacrycorpus steupii (WasSCHER) Dz LAUB. % Se @ Decrycarpus cumingii (PaRL_) DE Laus. \ @ Dacrycarpus kinabaluensis (WASSCHER) DE LAUB. \ » Sa | Fig. 29. Range of three species of the genus Dacry- carpus. Ecol. Locally common, particularly in disturbed forests, or in poorly drained areas where it may form nearly pure stands, in boggy grasslands and reed- swamps, on sandy clay, once on a rocky riverbank, once on a limestone hillock in mossy forest (Mt Bera- tus), 860—3420 m, but mostly c. 1500—2000 m. Vern. New Guinea: miejoop, nak, Kebar lang., ape, Wissel Lakes, Kapauko lang., pau, Wabag, Enga lang. Note. The spreading needles give this species a rather distinct appearance from D. imbricatus var. robustus which it otherwise strongly resembles. In New Guinea it has a markedly distinct ecology. 3. Dacrycarpus cumingii (PARL.) DE Laus. J. Arn. Arb. 50 (1969) 329; GaussEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 156, f. 727; DE Laus. Kalika- san 7 (1978) 128. — Podocarpus cumingii Part. in DC. Prod. 16, 2 (1868) 521; WasscHER, Blumea 4 (1941) 407, t. 4, f. 5. — Nageia cumingii (PARL.) O. K. Rev. Gen. PI. 2 (1891) 800. — Podocarpus im- bricatus var. cumingii (PARL.) PiLGerR, Pfl. R. IV, 5, Heft 18 (1903) 56. Tree, 8—25 m tall, up to 18—75 cm diam. Leaves of primary shoots spreading slightly, often curved so that the apex is directed inward slightly towards the axis. Leaves of juvenile foliage shoots distichous, nearly linear. Leaves of older trees similar but mostly not distichous, often more robust and scarcely keeled on the lateral faces, 3—6 by 0.6—0.8 mm. In addition to primary shoots and purely foliage shoots there are intermediate shoots with leaves 2—3 mm long, strongly keeled on four sides, but distinctly bilateral- ly flattened. The intermediate shoots bear foliage shoots and fertile shoots but are caducous like the fo- liage shoots and unlike the primary shoots. Pollen cones usually normal but on one specimen from low elevation up to 4 cm long. /nvolucral leaves greatly elongated resembling the foliage leaves but curved, not straight, 7~13 mm long, completely surrounding CONIFERALES (de Laubenfels) 381 the developing fertile structure which scarcely sur- passes them when fully mature. Ripe receptacle red- dish. Distr. Malesia: N. Sumatra (Leuser complex, 2 coll.), Borneo (Sarawak, rare), Philippines (Luzon, Negros, Panay, Mindanao). Fig. 29. Ecol. Locally common from 1000 to 3314 m, but mostly between 1850 and 2650 m in mossy primary forest. Locally it occurs above D. imbricatus most of whose varieties do not enter the mossy forest. Vern. Sumatra: sangu, Gajo, Mt Leuser; Philip- pines: igem, Davao, Mindanao. 4. Dacrycarpus kinabaluensis (WASSCHER) DE LAUB. J. Arn. Arb. 50 (1969) 330; GAUSsEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 154, f. 725. — Podocar- pus imbricatus var. kinabaluensis \WASSCHER, Blumea 4 (1941) 400, t. 4, f. 2. — Podocarpus im- bricatus (non BLUME) GiBBs, Ann. Bot. 26 (1912) 525, p.p., t. 49, f. 1-9. Shrub or small, sometimes gnarled tree, 2-13 m tall, 1S—30 cm diam. Leaves of primary shoots near- ly imbricate with the apex often curved slightly in- ward. Leaves of juvenile foliage shoots distichous, nearly linear. Leaves on older trees similar but soon becoming not distichous and more robust, distinctly keeled on the lateral faces, 3—6 by 0.8—1 mm. Together with primary shoots and foliage shoots are intermediate shoots with leaves 2—5 mm long and tri- angular or quadrangular in cross section. The inter- mediate shoots bear foliage shoots and fertile shoots but are also deciduous. /nvolucral leaves greatly elongated resembling the foliage leaves but more dis- tinctly curved, 5—8 mm long, reaching only the lower part of the seed when it is mature. Ripe receptacle blue or purple. Seed with its covering 6—7 mm long and 5—6 mm diam. Distr. Malesia: Borneo (Sabah: Mt Kinabalu). Fig. 29. Ecol. Common, sometimes in nearly pure stands in dwarf mountain scrub from 2700 m to the tree line at c. 4000 m. Note. On Mt Kinabalu D. imbricatus does not occur above c. 2000 m, leaving a considerable gap be- fore D. kinabaluensis is seen, which represents the mossy forest zone. In fact, D. kinabaluensis rather strongly resembles D. cumingii, differing particular- ly in the distinctly shorter involucral leaves. Foliage leaves of D. cumingii when collected from exposed parts of the tree approach this species in robust form. The receptacle colour of D. cumingii is poorly docu- mented and may well become purple also. 5. Dacrycarpus expansus DE LAus, J. Arn, Arb, 50 (1969) 334, f. 7b; GAUssen, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 162, f. 733. Tree 9~25(—30) m tall, 22—58 cm diam. Leaves of 382 FLORA MALESIANA [ser. I, vol. 103 primary shoots on young plants nearly imbricate but on older plants spreading and then curved forwards. Leaves of juvenile foliage shoots distichous, nearly linear, soon losing the distichous habit as the tree ma- tures. Leaves on older trees eventually becoming wider than thick but distinctly keeled on the upper and lower surfaces, spreading but the upper half curving forwards, uniform in size along the shoot, tapering slightly, 1.5—3 by 0.4—0.8 mm or a little lar- ger on younger trees. /nvolucral leaves loosely sur- rounding the young fertile structure but covering on- ly the receptacle of the mature seed with its covering, which is c. 3—3.5 mm long. Distr. Malesia: Central Highlands of Papua New Guinea. Fig. 30. Ecol. Locally common or even in pure stands (e.g. at Wabag), or co-dominant, sometimes emergent, often in disturbed situations, e.g. on edges of treefern grassland, 1300—2750 m. Vern. Pa’u, pau, Kepilan, Enga lang. Note. Not associated with moist habitats like D. steupii, a species which is also associated with dis- ae ha t Fig. 30. Range of Dacrycarpus expansus DE LAUB. (squares) and D. compactus (WASSCHER) DE LAUB. (dots). turbed habitats. This latter species differs in the form of the foliage leaves and the involucral leaves. 6. Dacrycarpus compactus (WASSCHER) DE LAUB. J. Arn. Arb. 50 (1969) 336, f. 9c; GAUSSEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 158, f. 730; VAN Fig. 31. A boggy hollow, filled with the tall grass Deschampsia klossii RDLEy (c. 1 m high) with on the edge tall Dacrycarpus compactus (WASSCHER) DE LAvB. in dense, very mossy shrubberies of mixed composition. About | km north of Lake Habbema, West New Guinea, 3300 m (Photogr. L.J.Brass, 1938). 1988] CONIFERALES (de Laubenfels) 383 RoyYEN, Alpine Fl. New Guinea 2 (1979) 20, f. 36, t. 80. — Podocarpus compacta WASSCHER, Blumea 4 (1941) 411, t. 4, f. 8a, b. — Podocarpus papuanus (non RIDLEY) PitGER, Bot. Jahrb. 68 (1936) 244. — Fig. 31, 32. Irregular tree 2—20 m tall, up to 25—60 cm diam. Leaves of primary shoots spreading slightly, often curved so that the apex is directed inward towards the axis. Leaves of juvenile foliage shoots not distichous, lanceolate, strongly keeled laterally, 2—2.5 by 0.6 mm. Leaves on older trees becoming similar to the leaves on primary shoots, spreading but curved through most of their length, strongly keeled on the dorsal side, 1—2.5 by 0.6—1 mm. Pollen cones on a 3 mm or more often longer shoot. /nvolucral leaves robust, curving to surround the receptacle, strongly keeled on the dorsal side and more or less triangular in cross section, 4—5 by 0.8—1.2 mm. Ripe receptacle purple to black. Seed with its covering 7—8.5 mm long and 7—8 mm diam. Distr. Malesia: New Guinea. Common inE., but rare in W. New Guinea (Habbema Lake, Quarles Lake, 2 coll.). Fig. 30. Fig. 32. Dacrycarpus compactus (WASSCHER) DE Laus. Detail of female branch with cones. Mt Amungwiwa, New Guinea, 3050 m (Photogr. P.VAN ROYEN 11072, June 1976). Ecol. Common on the higher peaks near the tree line, sometimes forming pure stands or emerging above a subalpine shrubbery, or scattered in alpine grassland, often in isolated specimens and obviously fire-resistant, in Podocarpus-Libocedrus forest, tarely on wet peaty soil (Lake Aunde), 2800—3950 m, but mostly above 3400 m. Vern. Kadzinam, kaibigl, Minj, kaipik, Kubur, Minj, umbwa, Lake Aunde, Chimbu, Waimam- bano, umba-nifiogo, Chimbu, Lake Aunde. 7. Dacrycarpus cinctus (PILGER) DE LAus. J. Arn. Arb. 50 (1969) 332, f. 9a; GAUSSEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974), 160, f. 732; VAN ROYEN, Alpine Fl. New Guinea 2 (1979) 17, f. 35h, t. 79. — Podocarpus cinctus P1LGER, Bot. Jahrb. 69 (1938) 253; WASSCHER, Blumea 4 (1941) 409, t. 4, f. 6. — Podocarpus dacrydiifolia WASSCHER, /.c. 410, t. 4, f. 7. — D. dacrydiifolia (WASSCHER) GAUSSEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 148, f. 729. — Fig. 33. Often flat-crowned tree up to 33 m, 20—90 cm diam., or a shrub 2—4 m tall. Leaves of primary shoots spreading slightly, straight or more often curved forward, 5—6 mm long on young plants, be- coming 3—4 mm long on adult plants and 2-3 mm long at the base of foliage shoots and fertile struc- tures. Leaves of juvenile foliage shoots not disti- LAUB. Detail of female branch with cones. Papua New Guinea, Southern Highlands District, 27 miles from Mendi, 2743 m (Photogr. Womerstey LAE 55322, Sept. 1972). Fig. 33. Dacrycarpus cinctus (PILGER) DE 384 FLORA MALESIANA [ser. I, vol. 103 chous or perhaps slightly so, linear-lanceolate, the upper half curved forward, 0.5—0.8 mm _ wide. Leaves on older trees eventually becoming similar to the leaves of primary shoots but somewhat narrower and curved like the juvenile leaves, 2—5S by 0.4—0.6 mm, uniform along a shoot, often glaucous. Pollen cone sometimes on long shoots. /nvolucral leaves re- sembling the foliage leaves but curved throughout their length, completely surrounding the developing seed with its covering which rises slightly above them when mature, 6—10 mm long. Receptacle bright red when ripe. Seed with its covering 7 mm long and 6—7 mm diam. Distr. Malesia: Central Celebes, Moluccas (Cen- tral Ceram: G. Binaja), and throughout New Gui- Ecol. In New Guinea extremely common and often dominant, or co-dominant with Nothofagus, Libocedrus, Elaeocarpus and Podocarpus, in moun- tain forest and mossy forest, on Mt Binaja in orchard-like pure stands with a mossy ground cover, rarely in muddy parts of swamps (lowasi swamp near Woitape), a canopy tree or sometimes emergent, often thick-trunked, the foliage glaucous or not, 1800—2850 m, occasionally as high as 3600 m, in Ce- ram from 1300—3000 m, in Celebes reported as low as 900 m. Vern. Celebes: sareh, Upper Binuang, Ulu Sahu; New Guinea: djasiha, Asaro, Kefamo, gu-gra-goin, Goroka, Hagen lang., jumbiri, Mt Giluwe, Mendi lang., kaiwilpitti, Waghi, Minj, kubil-kaibigl, ku- buk-kajbek, Kubor Ra., Minj, kubin, Hagen, Togoba, ma-u, Finisterre Ra., Naho lang., pau, Hagen-Wabag, Enga lang., piepienie, Mt Ne, Tari, Hula lang., u(m)ba, Chimbu lang. at various places. Note. Collectors complain that this species grades into D. compactus and indeed in the zone of overlap between these two species specimens of D. cinctus have shorter and more robust leaves resemb- ling D. compactus, while the common glaucousness of D. cinctus has been observed to disappear above 2950 m. The two species are substantially different, however, and both become much less common in the elevations where they overlap (2900—3400 m). Per- haps hybridization occurs where they overlap. nea. Fig. 34. Fig. 34. Range of Dacrycarpus cinctus (PILGER) DE LAUB. 5. PRUMNOPITYS PHILIPPI, Linnaea 30 (1860) 731; DE Laus. Fl. Nouv. Caléd. et Dép. 4 (1972) 55; Blumea 24 (1978) 189. — Stachycarpus (ENDL.) TIEGH. Bull. Soc. Bot. Fr. 38 (1891) 162; GaussEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 81. — Podo- carpus sect. Taxoideae BENNETT in Bennett & R.Br. Pl. Jav. Rar. 1 (1838) 40. — Podocarpus sect. Stachycarpus ENDL. Syn. Conif. (1847) 218; HENKEL & HOCHSTETTER, Synop. Nadelholz. (1865) 399; Par. in DC. Prod. 16, 2 (1868) 518; EICHLER in E. & P. Nat. Pfl. Fam. 2, 1 (1889) 105; Pircrr, Pfl. R. IV, 5, Heft 18 (1903) 63; Gress, Ann. Bot. 26 (1912) 537; BuCHHOLZ & Gray, J. Arn. Arb. 29 (1948) 58. — Podocarpus sect. Prumnopitys (PHILLIPPI) BERTRAND, Ann. Sc. Nat. V, 20 (1874) 65. — Podocarpus subg. Stachycarpus (ENDL.) ENGLER in E. & P. Nat. Pfl. Fam., Nachtr. 1 (1897) 21; PitGer in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 242; WasscHER, Blumea 4 (1941) 380. — Fig. 36, 38, 39. For further synonyms see under section Sundacarpus. Densely branched dioecious trees to 60 m tall. Bark smooth, fibrous, and red- dish to yellowish brown, often darker on the surface but weathering to gray, on older trees breaking off in irregular more or less quadrangular plates 3—5 mm thick and 3—10 cm across, with scattered lenticel-like mounds. Foliage buds 1988] CONIFERALES (de Laubenfels) 385 small and inconspicuous with overlapping triangular scales. Leaves spirally placed, bifacially flattened, linear, uninerved, without hypoderm, hypostomat- ic, narrowed at the decurrent base with a twist where the leaf leaves the stem so that the leaves appear distichous. Pollen cones axillary and solitary or grouped on scaly spike (or even compound structures). Seed with its covering solitary and subterminal or grouped along a scaly or leafy shoot, inverted and completely covered by a fleshy epimatium with an apical crest; the seed with a slightly asym- metrical ridge at the micropylar end. Distr. 10 spp. in two slightly geographically overlapping sections, with the type section extending from Australia and New Caledonia to New Zealand and from Chile to Venezuela and Costa Rica. The monospecific section Sundacarpus is confined to Malesia and NE. Queensland. Fig. 35. Uses. Several species are important timber trees. a _ — —t 1 ———— 4 4 din re 4 Fig. 35. Range of the genus Prumnopitys Puiippl. Figures above the hyphen indicate the number of endemic species, that below the hyphen the total number of species. 1. Section Sundacarpus (BUCHHOLZ & GRAY) DE LAuB. Blumea 24 (1978) 190. — Podocarpus sect. Sun- dacarpus BucHHOLZ & Gray, J. Arn. Arb. 29 (1948) 57; Frorin, Acta Horti Berg. 20 (4) (1963) 190, f. 20 (map). — Stachycarpus sect. Sundacarpus (BUCH- HOLZ & GRAY) GAUSSEN, Gymn. Act. & Foss. fasc. 13, ch. 20 (1974) 81. 1. Prumnopitys amara (BLUME) DE LAus. Blumea 24 (1978) 190. — Podocarpus amara BiumE, En. PI. Java | (1827) 88; BENNETT in Bennett & R.Br. PI. Jay. Rar. 1 (1838) 40; ENDL. Syn. Conif. (1847) 217; BLume, Rumphia 3 (1849) 213, t. 170; Mia. FI. Ind. Bat. 6 (1859) 1073; Henket & Hocustetrer, Synop. Nadelhdélz. (1865) 395; pe Boer, Conif. Arch. Ind. (1866) 20; Part. in DC. Prod. 16, 2 (1868) 516; Ber- TRAND, Ann. Sc. Nat. V, 20 (1874) 67; TieGu. Bull. Soc. Bot. Fr. 38 (1891) 38; Wars. Monsunia | (1900) 192; Picer, Pfl. R. IV, 5, Heft 18 (1903) 68, f. 13A—D; K. & V. Bijdr. 10 (1904) 263; Foxw. Philip. J. Sc. 6 (1911) Bot. 159; Koorp. Exk. Fl. Java 1 (1911) 64, f. 1; Hatt. Meded. Rijksherb. n. 14 (1912) 34; Koorp. Atlas 3 (1915) t. 590, 591; MErRR. En. Philip. 1 (1922) 2; Koorp., Fl. Tjibodas (1922) 2; LANE-POOLE, For. Res. Terr. Papua & New Guinea (1925) 73; Pitcer in E. & P. Nat. Pfl. Fam, ed. 2, 13 (1926) 245, f. 131A—D; Wasscuer, Blumea 4 (1941) 381; Orr, Trans. Bot. Soc. Edinb. 34 (1944) 11; Gray & Bucunoiz, J. Arn. Arb, 32 (1951) 93; Backer & Baku. Fl. Java | (1963) 89. — Podocar- pus eurhyncha Mig. FI, Ind. Bat. 2 (1856) 1074; De Borer, Conif. Arch. Ind. (1866) 24, t. 3, f. 2; PARL. 386 FLORA MALESIANA [ser. I, vol. 103 Fig. 36. Prumnopitys amara (BLUME) DE Laus. A. Habit of tree; B, C. twigs with pollen cones; D—F. micro- sporophylls (different views) (from Koorp. Atlas 3, 1915, t. 590). 1988] CONIFERALES (de Laubenfels) in DC. Prod. 16, 2 (1868) 518; Wars. Monsunia 1 (1900) 193. — Podocarpus dulcamara SEEM. Bonplandia 9 (1861) 253; ibid. 10 (1862) 365. — Na- geia amara (BLUME) O. K. Rev. Gen. PI. 2 (1891) 800. — Nageia eurhyncha (Miq.) O. K. /.c. 800. — Podo- carpus pedunculata Batley, Queensl. Agr. J. 5 (1899) 390, t. 144; Queensl. Fl. 5 (1902) 1498; BAKER & Situ, Res. Pines Austr. (1910) 441; Hardwoods Austr. (1919) 429. — Stachycarpus amara GAUSSEN, Gymn. Act. & Foss. fasc. 12 (1973) f. 635, 646; ibid. 13, ch. 20 (1974) 105, f. 715 (map). — Fig. 36, 38, 39. Tree 10—60 m high, 12—140 cm diam. Bark sur- face checkered by numerous cracks. Cotyledons 3 fused pairs. Foliage buds small, globose, with over- lapping, rounded, keeled scales up to 2 mm long. Juvenile leaves oblong, 4—12 by up to 2 cm, more or less abruptly narrowing at the base to a 3—5 mm long petiole, narrowing abruptly at the apex to an elongat- ed tapering ‘drip tip’ which is c. 2 mm wide and up to 20 mm long, apex rounded to acute. Mature leaves becoming longer and narrower, linear, narrowed at the base to a c. 5 mm long petiole, usually slightly acuminate and acute, a distinct groove over the mid- vein above, broadly raised below, 5—15 cm by 6—14 mm. Pollen cones 15—35 by 2.5—3.5 mm, solitary and terminal or grouped to at least seven on an auxil- lary 1-7 mm peduncle with several sterile basal scales. Apex of microsporophyll acute, triangular, keeled, c. 0.8 mm long. One to several ovules scat- tered laterally along a 3—5 mm scaly shoot, the scales triangular to rounded, decurrent, spreading, 1.5—2 mm long, the sterile scales deciduous. Ovule and its covering oval, longer than its bract (fertile scale) and distinctly crested at its apex, dark blue and glaucous. Growing seed and its covering elongated at both the micropylar end and the forwardly bent apex; ma- ture seed and its covering nearly spherical, with a small obtuse crest, c. 25 mm diam., becoming red- dish and then dark purple and glaucous. Seed c. 20 mm diam., with an indistinct ridge and minute apicu- lus formed from the micropyle, the smooth outer hard shell c. 1 mm thick, the fleshy covering c. 3 mm thick becoming wrinkled as it dries and often falling off. Distr. NE. coastal Queensland; in Malesia: through and very common in New Guinea (incl. New Britain & New Ireland), Moluccas (Buru, Halma- heira, Morotai), Lesser Sunda Islands (Timor, Flores, West Sumbawa, Lombok), throughout Java, Central and SW. Celebes (Bonthain), Philippines (Mindanao, Luzon), Borneo (only in Sabah!), and Sumatra (Central—N., Batak region, rare in S. Pa- lembang). Fig. 37. Ecol. Scattered and often common in primary and secondary rain-forest, in New Guinea very com- mon, often in Fagaceous forest, sometimes in mossy forest, in submontane forest at c. 900 m with Dyso- YD» \. I} a | ~A\ \—__—__—— —L —————— =. r We J WX, ss i se ~ SEE =. Bare: Pe € =e oe eee | “.% ‘Xg j La wis te >, - . —{/ oe em Wee ae i Le ee al r a et VIN, ay —) \ Pe = eer eT yw -— Fig. 37. Range of Prumnopitys amara (BLUME) DE LAUB. xylum, Macaranga, Ficus, sometimes emergent as a colossal tree, often on latosols, rarely on sandy soils or on marshy ground, (sea-level—)500—2000(—2300) m, according to SMYTHIEs (in sched.) to 3000 m in Sabah. A few times it is mentioned that the tree is but- tressed (Pinosok Plateau and Mt Cyclops, New Gui- nea), or spurred, a rare feature in Podocarpaceae. Uses. A fine timber tree, often of large dimen- sion. In New Guinea mentioned to be used for joinery and furniture. Vern. Sumatra: kuta-béa, sitoba, sitobu, Toba- Batak, besi, Karo-Batak, buluh, Pasemah, Palem- bang; Java: ki bima, ki manang pait, ki marak, ki merak, ki pait, ki putri (confusion with Podocarpus neriifolius), S; tadji, Tengger, tjemoro, Madiun (confusion with Casuarina), J; Borneo: sempilau, Kinabatangan; Lesser Sunda Islands: pinis, Flores, kayu santen, Batulante, Sumbawa; West New Guinea: bublugie, Wissel Lakes, Kapauko lang., ir- bouska, Manikiong, Ransiki, du-on, nepa, niepa, nipah, Kebar lang., niwup, Kebar Valley, Andjai dial., (jermes, Ransiki, Hattam lang.; East New Gui- nea: kaibiltugl, Wahgi, Minj, kepim, Pokaris, Wa- bag, Enga lang., kumburum, Mt Entaldam, Telefo- min, /a-u, Poget logging area, Wabag, /evekus, Tele- fomin, migimi, Ogeramnang, powa, Pomagos R.C. Mission, Weng lang., singai, Bulolo, Garaina dial., taso, Wonatabe, til, Hagen, Togoba, fsula, Chimbu, Masul; New Britain: neli/, Mt Talawe. Black pine, Queensland. Note. The leaves are variously reported as bitter (‘amara’), to which also the Sundanese name ‘pait’ refers, bittersweet (‘dulcamara’), or sweet tasting. This and the groove over the midvein most readily distinguishes it from similar-leaved associated Podo- carpus species while the lack of hypoderm also gives a distinct texture to the leaves. The striking form of the juvenile leaves led to the description of Podocar- pus eurhyncha. Gray & BucHHoLz (1951) report that the leaves occasionally have a lateral pair of vascular resin canals in addition to the conspicuous central canal beneath the vascular bundle. Two collectors 388 FLORA MALESIANA [ser. I, vol. 10° Fig. 38. Prumnopitys amara (BLUME) DE Lavus. G, H. Twigs with pollen cones; I. twig with seeds; K, L. twig and leaf of a young tree; M. leaf of an adult tree; N—P. pollen cones with details; Q, R. seed, also in cross section (from Koorp. Atlas 3, 1915, t. 591). 1988] CONIFERALES (de Laubenfels) 389 Fig. 39. Prumnopitys amara (BLUME) DE LAUvB. Ripe seed red (Photogr. BW 2318). report seeds with distinctive sculpturing on their sur- _ two fused pairs of cotyledons is unique. The limited face but this is not evident in the corresponding pre- _ occurrence in Borneo is curious. served specimens. The normally three rather than 6. NAGEIA GAERTN. De Fruct. & Sem. (1788) 191; Gorpon, Pinetum (1858) 134; O. K. Rev. Gen. Pl. 2 (1891) 798; Florin, Acta Horti Berg. 20 (4) (1963) 190, f. 21 (map); DE Laus. Blumea 32 (1987) 209. — Decussocarpus DE LaAuB. J. Arn. Arb. 50 (1969) 340; Fl. Nouv. Caléd. et Dép. 4 (1972) 48; Kalikasan 7 (1978) 130. — Fig. 41. Dioecious, erect shrubs or trees, 1—54 m tall. Bark tan to brown within and dark brown to black on the surface but often weathering to gray, peeling in ir- regular shaped plates to short vertical strips. Leaves opposite-decussate (or mixed with some leaves spirally placed), distinctly narrowed to a decurrent base. Juvenile leaves mostly larger than the adult leaves which are otherwise similar, twisted at the base so as to appear distichous, in most cases amphistomatic with 390 FLORA MALESIANA [ser. I, vol. 103 the abaxial face uppermost on the left side of the shoot and the axial face upper- most on the right side (in a few species the leaves are hypostomatic and without this unique orientation). Pollen cones sessile or terminal, solitary or grouped, cylindrical (or oval). Each inverted seed completely covered by the fertile scale (epimatium), one or occasionally two subterminal on a scaly shoot, the usually persistent leathery covering becoming more or less fleshy when ripe. Distr. There are 12 spp. in the three sections through most of the tropical forests of the world, throughout Malesia, where two sections overlap each other. Fig. 40. KEY TO THE SECTIONS 1. Leaves large, broad, ovate to broad-lanceolate, with many longitudinal veins......... 1. Sect. Nageia | Mbeavesssmallecwithkarsinglesntopeercs mete esse ios nee ara wt «ae ee ees 2. Sect. Polypodiopsis 1. Section Nageia DE LausB. Blumea 32 (1987) 209. — Podocarpus sect. Dammaroides BENNETT in Bennett & R.Br. Pl. Jav. Rar. 1 (1838) 41. — Podocarpus sect. Nageia ENDL. Syn. Conif. (1847) 207; Mia. FI. Ind. Bat. 2 (1859) 1071; HENKEL & HOcHSTET- TER, Synop. Nadelholz. (1865) 378; DE BorEr, Conif. Arch. Ind. (1866) 12; PaRL. in DC. Prod. 16, 2 (1868) 507; BERTRAND, Ann. Sc. Nat. V, 20 (1874) 66; Pitcer, Pfl. R. IV, 5, Heft 18 (1903) 59; Grpss, Ann. Bot. 26 (1912) 533; PILGER in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 242; WasscHER, Blumea 4 (1941) 415; BUCHHOLZ & Gray, J. Arn. Arb. 29 (1948) 56; GAussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 11. — Decussocarpus sect. Dammaroides (BENNETT) DE Laus. J. Arn. Arb. 50 (1969) 348; Kalikasan 7 (1978) 130. Erect shrubs or trees, 1—48 m tall. Bark smooth, on trees peeling in large thin irregular shaped plates with scattered lenticels. Foliage buds a compact cluster of lanceolate deciduous scales abruptly wider than the shoot and distinctly acute, when terminal often 2—3 mm beyond the last leaf bases, 3—6 mm long, lateral buds sessile. Primary shoots not differentiated from ultimate shoots. Leaves with many parallel vascular bundles (one of which is medial) converging towards the acute to acuminate apex, more or less elliptic but juvenile leaves very acuminate and adult leaves sometimes more or less blunt, distichous, amphistomatic with equal basal twists (or hypo- stomatic with opposite basal twists), narrowed to a short broad petiole. Fertile shoots arising in the axils of leaves from sessile buds essentially the same as foliage buds, one to several pairs of deciduous scales below the first pollen cone or the female receptacle. Pollen cones cylindrical, solitary or grouped with one terminal and others forming sessile decussate pairs about 5 mm apart, each cone in the axil of a sterile scale but the terminal three often fused at their bases. In some species a part of the fruit-bearing shoot becoming enlarged and eventually fleshy forming a receptacle, otherwise a part of the shoot often remaining attached to the seed when it falls. Seed smooth, nearly spherical but elongated on the micropylar end into a small asymmetrical beak. Distr. There are 5 spp. from southern India and Bangladesh across Indochina and Malesia to New Britain and through southern China to southern Japan; in Malesia 3 spp. Fig. 40. | SSS) re eee 1988] CONIFERALES (de Laubenfels) 391 Notes. Sterile specimens often confused with A gathis with very similar leaves, but in Agathis the parallel vascular bundles do not markedly converge towards the leaf apex and the terminal bud is globular, not acute, but very blunt-rounded. The key to the three species is inadequate for identification of sterile material, but the leaf-size and length of the petiole may be helpful, though sizes overlap. sect. AFROCARPUS Fig. 40. Range of the genus Nageia GAERTN. and its three sections, with the number of species. KEY TO THE SPECIES 1. Fruit with fleshy receptacle. 2. Pollen cones grouped on a peduncle. Leaves at least 6cm long .................. 1. N. wallichiana 2. Pollen cones solitary, sessile. Leaves mostly less than 5 cm long...............-.2+05- 2. N. motleyi 1. Fruit lacking a fleshy receptacle. Large-leaved .... 1. Nageia wallichiana (PResL) O. K. Rev. Gen. Pl. 2 (1891) 800; pe Laus. Blumea 32 (1987) 210. — Podo- carpus latifolius (non THUNB. 1794) BLuME, En. PI. Jav. 1 (1827) 89; Watt. Pl. As. Rar. (1830) 26; HorsFieé_p in Bennett & R.Br. Pl. Jav. Rar. | (1838) 41; Loupon, Arb. & Fruct. Brit. 4 (1838) 2100, f. 1995; Hassk. Cat. Hort. Bog. (1844) 70; ENDL. Syn. Conif. (1847) 208; Mig. Pl. Jungh. 1 (1851) 1; FI. Ind. Bat. 2 (1859) 1071; Henket & HOCHSTETTER, Synop. Nadelhdélz. (1865) 379; pe Boer, Conif. Arch. Ind. (1866) 12; Part. in DC. Prod. 16, 2 (1868) 508; Hook.f. Fl. Brit. India 5 (1896) 649. — Podocarpus wallichianus Prest, Bot. Bemerk. (1844) 110; Purcer, Pfl. R. [V, 5, Heft 18 (1903) 59; Riptey, J. Str. Br. R. As. Soc. n. 60 (1911) 57; Hicket, Fl. Gén. I.-C. 5 (1931) 1068; Keno in Whit- more, Tree Fl. Malaya 1 (1972) 53, f. Sd—f; STA. de ae CEES 3. N. maximus PHENGKLAI, Thai For. Bull. Bot. 7 (1973) 17, f. 11; GaussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 18. — Podocarpus blumei ENDL. Syn. Conif. (1847) 208; HENKEL & HOCHSTETTER, Synop. Nadelhdlz. (1865) 380; Part. in DC. Prod. 16, 2 (1868) 508; Becc. Malesia 1 (1877) 178; Wars. Monsunia | (1900) 193; Pircer, Pfl. R. IV, 5, Heft 18 (1903) 60, t. 9; K. & V. Bijdr. 10 (1904) 261; Koorp. Exk. FI. Java | (1911) 67; Foxw. Philip. J. Sc. 6 (1911) Bot. 158, t. 28, f. 2; Koorp. Atlas 3 (1915) f. 588; RipLey, Fl. Mal. Pen. 5 (1925) 281; Pircer in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 245, t. 134B; Bot. Jahrb. 68 (1937) 245; Corner, Wayside Trees (1940) 717; Wasscuer, Blumea 4 (1941) 416; BACKER & BAKu//. Fl. Java 1 (1963) 89; Gaussen, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 20, f. 736. — Podocarpus agathifolia Blume, Rumphia 3 (1849) 217, t. 173. — 392 FLORA MALESIANA [ser. I, vol. 103 Fig. 41. Nageia wullichiana (PRESL) O.K. A. Twig with seed; B, C. twig portion with young pollen cones; D. ditto with mature pollen cones; E. longitudinal section of seed; F. leaf of a mature tree; G. leaf of a young tree; H, I. terminal foliage buds; K. habit of tree (from Koorp. Atlas 3, 1915, t. 588). 1988] N. blumei (ENDL.) GoRDON, Pinetum (1858) 138. — Podocarpus latifolia f. ternatensis DE BoER, Conif. Arch. Ind. (1866) 14. — Decussocarpus wallichianus (PRESL) DE Laus. J. Arn. Arb. 50 (1969) 349; Kalikasan 7 (1978) 131. — Fig. 41. Tree, 10—54 m high, 7—60 cm diam., clear bole to 30 m. Leaves quite variable both on juvenile and adult plants with the larger leaves growing in the shade, 6—14(—23) by 2—5(—9) cm, 2 to at least 6 times as long as broad; petiole S—10 mm. Pollen cones in groups of up to at least 7 on a 2—10 mm ped- uncle; mature cone 8—18 by 3—4 mm. Apex of the microsporophyll lanceolate, 2-3 mm long. Seea- bearing structure solitary on a 8 to at least 20 mm peduncle with caducous scales or occasionally re- duced leaves. Receptacle with 4—7 sterile, deflexed, slightly enlarged bracts, 7-18 mm long before be- coming ripe and fleshy, dark purple or black when ripe. Seed with its covering 15—18 mm diam. Distr. Southeast Asia (S. extremity of the Dec- can Peninsula, Assam, Burma, Thailand, Indochina, Yunnan); in Malesia: Sumatra, Malaya, Banka I., W. Java, Lesser Sunda Islands (Flores), Borneo (incl. Karimata I.), Philippines (Luzon, Sibuyan, Mindoro, Panay, Samar), N. & Central Celebes, Moluccas (Obi, Ternate, Morotai, Ceram), New Guinea (incl. Meos Num, Biak, Japen, and Norman- by Is.). Fig. 42. : “ . ‘ x gph \ ; \ eye cress leit ieee 7. Sect. Rumphius 7. Outer budscales erect and free. Midrib narrow and in most species prominently raised. Ripe receptacle usuallytpunplenSppr 28 —SOMs Yay. et ae Soe Aceye eee atktey. ec abte 2.5, tales 8. Sect. Polystachyus VEGETATIVE, KEY TO,THE SECTIONS 1. Primary budscales imbricate, forming a ball or pyramid with few or none of their apices raised, bud no longer than diameter. 2. Primary budscales overlapping and forming a ball or in some cases meeting pyramid-like at the bud apex but the scale apices never recurved outward; leaf hypoderm absent or occasionally slightly developed; leaf midvein generally more than half a mm wide but never a fullmm ............... 3. Sect. Globulus 2. Primary budscales mostly in the form of a pyramid with usually one or more scales recurved outward at the apex; leaves with continuous upper hypoderm and well developed lower hypoderm; leaf midvein more GiplessTone ymin WIdee re Atte Peete ete Ree + Mor ae oe en) eee LEE 7. Sect. Rumphius 1. Primary budscales with completely free apices, bud usually longer than diameter. 3. Adult leaves distinctly acuminate; primary budscales mostly more than four times as long as wide 2. Sect. Acuminatus 3. Adult leaves not acuminate but where transitional leaves may be slightly acuminate, the primary budscales less than four times as long as wide. 4. Foliage buds at least twice as long and generally much longer than basal diameter (leaves not at all acuminate). 5. Leaves lanceolate, widest near the base. 6. Midrib on upper side of leaf at least 0.2 mm high, often prominent ..... 4. Sect. Longifoliolatus 6. Midrib on upper side of leaf less than 0.2 mm high, indistinct... 8. Sect. Polystachyus (P. ridleyi) 5. Leaves more or less linear, not widest near the base (buds only sometimes more than twice as long as basal diameter). 7. Leaves less than 7 mm wide, 5 cm long, midrib on the upper side less than 0.2 mm high 4. Sect. Longifoliolatus (P. gibbsii) 7. Leaves at least 8 mm wide, 4 cm long, midrib on upper side c. 0.3 mm high 1. Sect. Foliolatus (P. borneensis) 4. Foliage buds less than twice as long as basal diameter. 8. Basal diameter of foliage bud 1—2 mm (midrib on upper side of leaf 0.2 mm high and less than 0.5 mm wide). SeMidribron upper side‘of leaf 02 mnt wide. -. «eee Set oo tele see eee 5. Sect. Gracilis Oy Midribsoniupper side of leafiiat leasti0:3 mmijwide 29-22: 6.20.25 eee 8. Sect. Polystachyus 8. Basal diameter of foliage bud at least 2 mm, usually considerably more. 10. Midrib on upper side of leaf less than 0.2 mm high and 0.7 mm wide. ie eat maidmb.0:2—O0Simmiwidey aan. esc ee see asset Cie eee ier. Ge 6. Sect. Macrostachyus 11. Leaf midrib at least 0.5 mm wide. 12. Leaf less than 7 mm wide, 5 cm long; basal diameter of foliage bud less than 3 mm; leaf midrib OSS Oe mmm Widera GISTINICE g 4..cxs castor eee. tee Seed 4. Sect. Longifoliolatus (P. gibbsii) 12. Leaf at least 8 mm wide, 5 cm long; basal diameter of foliage bud 3—4 mm (P. /evis) or leaf midrib broadeandomdistinet (24 SDGINOIGES)\ cia ae 2 ou ete aici versus uetoe a crietele ee 1. Sect. Foliolatus 1988] CONIFERALES (de Laubenfels) 399 10. Midrib on upper side of leaf at least 0.2 mm high and 0.3 mm wide but where less than 0.5 mm wide it is at least 0.3 mm high. feleat midrib: at Jeast.0:3.mm high) 2.02 -ceret- relat Seecisy eee ae 1. Sect. Foliolatus 13. Leaf midrib 0.2—0.3 mm high. i4eLeaf midrib 0.5—0.6 mm wide .... 2.0.02... c0.0000% 8. Sect. Polystachyus (P. macrocarpus) Baers midrib.0.8—1.2 mm wide cas ccstk:.aacs Stansenles 7. Sect. Rumphius (P. /aubenfelsii) 1. Section Foliolatus DE LAuB. Blumea 30 (1985) 264. Primary foliage budscales erect, triangular to shortly lanceolate; secondary scales acute, the secondary bud when it first appears generally a pyramid whose apex is formed of a cluster of crowded scale tips. Leaves mostly with more or less parallel sides and mostly acute, only in a few cases acuminate; upper midrib a blunt ridge at least 0.3 mm wide, broader but less prominent below; usually three vascular resin canals, sometimes more. Buds for pollen cones mostly 2—3 mm long, either sessile or on a short peduncle, the secondary pollen cone buds when they first appear around ball of overlapping imbricate scales. Pollen cones mostly solitary or in groups of no more than 3. Apex of the microsporophyll a small triangular spur c. 0.3—0.5 mm long over a base c. 1 mm wide. Foliola of the female structure mostly c. 2 mm long. Receptacle normally formed of two bracts, one of which is fertile and longer than the other or both fertile and equal, the fertile bracts 8—10(—12) mm long before becoming fleshy, in most species the ripe receptacle is known to be red. Seed with its covering 7—12 mm long by 5—8 mm diam., in most species more particu- larly 9-10 mm long and 6—7 mm diam., globular, apex mostly blunt. Peduncle mostly 5—10(—15) mm long but shorter in some species. Distr. From Nepal throughout Malesia to the Solomon Islands, Fiji and Tonga 10 spp. The range is greatly dominated by the impressive, wide distribution of P. neriifolius, the most widespread species of the genus, which covers virtually the entire Asian—Pacific wet tropical forest zone. The one highland species, P. rubens, is also widely distributed, otherwise the species are more local but combine to form a nearly continuous slight- ly overlapping series from Sumatra to Tonga. All but 2 spp. lie within Malesia. Ecol. Scattered either in primary rain-forest (but rarely encountered in the dipterocarp forest: P. nerii- folius), generally prominent in ridgetop mossy forests at moderate elevations (most species), or widespread in mountain forests at higher elevations (P. rubens). One species is confined to riverbanks. Note. Sect. Foliolatus consists of a group of closely related species which lack the specialized character- istics which distinguish each of the remaining sections of the genus. KEY TO THE SPECIES 1. Midrib on the upper side of the leaf prominent, at least 0.3 mm high and less than 1 mm wide (primary budscales no more than 5 mm long). 2. Juvenile leaves acuminate, midrib bold with vertical sides. Primary budscales 1—1.5 mm wide. 3. Leaf at least 7 cm by 10 mm, not apiculate, new leaves green to reddish, midrib 0.4—0.8 mm wide (where only 0.4—0.5 mm wide the leaf usually acuminate). Pollen cones normally sessile. Ripe receptacle red. 1. P. neriifolius 3. Adult leaf no more than 6 cm by 10 mm, often apiculate, new leaves bright red, midrib 0.3 mm wide. Pollen cones often on a short peduncle. Ripe receptacle often purple ..............+. 2. P. rubens 2. Juvenile leaves never acuminate, midrib with sloping sides. Primary budscales 2 mm wide (pollen cones usually pedunculate). 4. Leaf at least 7 cm by 10 mm, linear-oblong (midrib 0.5 mm wide). Pollen cone usually solitary. Re- GCemacie often with three fleshy Dracts + «0.00.00 co Wielele hisitle « Adee A n:o wile Rb oka e eles 3. P. archboldii 4. Adult leaf no more than 9 cm by 9 mm, elliptic. Pollen cones usually in clusters of 2—3. Receptacle of OD TUB CER OTUY: oi snie wininia! the bid Chl eh «, Ceol, ove.e vind Ly kbiblels all odtteh dh elie hae 4. P. insularis 400 FLORA MALESIANA [ser. I, vol. 103 1. Midrib on the upper side of the leaf at least 1 mm wide or indistinctly raised or both. 5. Midrib prominently raised above, 0.3 mm high, often drying to a channel below (leaves distinctly thicker than usual for the genus). Primary budscales may reach at least 1 cm long. 6. Leaves more than 10 cm long. Female peduncle 9-15 mm. Primary budscales and leaves deflexed 5. P. deflexus 6. Leaves less than 10cm long. Female peduncle 2 mm. Primary budscales and leaves erect 6. P. borneensis 5. Upper surface of leaf flat or nearly so, lower surface generally lacking a channel (leaves not thicker than usual for the genus). Budscales never as long as 1 cm. 7. Leaves lanceolate or narrowly linear.......... 7. Woeaves oblong; not tapering. . 45. ..e-s- eee 1. Podocarpus neriifolius D.Don in Lambert, Gen. Pinus ed. 1 (1824) 21; Prod. Fl. Nep. (1825) 55; SPRENG. Syst. Veg. 3 (1826) 889; BENNETT in Bennett & R.Br. Pl. Jav. Rar. 1 (1838) 40; ENpL. Syn. Conif. (1847) 215; Part. Bot. Mag. 78 (1852) t. 4655; HENKEL & HOCHSTETTER, Synop. Nadelholz. (1865) 381; Par. in DC. Prod. 16, 2 (1868) 514; EICHLER in E. & P. Nat. Pfl. Fam. 2, 1 (1887) 104; Hook.f. Fl. Br. Ind. 5 (1888) 649; Srapr, Trans. Linn. Soc. II, Bot. 4 (1894) 249; Wars. Monsunia | (1900) 193; Pricer, Pfl. R. IV, 5, Heft 18 (1903) 80; K. & V. Bijdr. 10 (1904) 265; Merr. Philip. J. Sc. 1 (1906) Suppl. 24; Foxw. Philip. J. Sc. 2 (1907) Bot. 258; Gisss, J. Linn. Soc. 39 (1909) 183; Foxw. Philip. J. Sc. 6 (1911) Bot. 162; RmpLey, J. Str. Br. R. As. Soc. n. 60 (1911) 57; Koorp. Exk. Fl. Java 1 (1911) 65, f. 3; Gipss, Ann. Bot. 26 (1912) 549, t. 51, f. 48—51, t. 53, f. 78; Koorp. Atlas 2 (1915) t. 589; Fl. Tjibodas I CL92a) eS: RIptny.. El Male Rents (1925). 281: Pricer in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 247; HickEL, Fl. Gén. I.-C. 5 (1931) 1069; WaAsscHER, Blumea 4 (1941) 437, incl. var. polyantha WASSCHER, l.c. 455; Gray, J. Arn. Arb. 39 (1958) 460, 467; BACKER & BaKH./f. Fl. Java 1 (1963) 90; GAUSSEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 187, f. 805; DE Laus. Kalikasan 7 (1978) 139. — Nageia neriifolia (D.Don) O.K. Rev. Gen. Pl. 2 (1891) 800. — P. neglecta BLtumME, Rumphia 3 (1849) 213; HENKEL & HOcuSTETTER, Synop. Nadelholz. (1865) 396; DE Borer, Conif. Arch. Ind. (1866) 21, t. II, 2; Part. in DC. Prod. 16, 2 (1868) 516; Wars. Mon- suna | (1900) 193. — Nageia neglecta (BLUME) O.K. Rev. Gen. Pl. 2 (1891) 800. — P. discolor BLUME, Rumphia 3 (1847) 213; Mia. Fl. Ind. Bat. 2 (1859) 1074; HENKEL & HocHSTETTER, Synop. Nadelholz. (1865) 396; pE BoER, Conif. Arch. Ind. (1866) 23, t. III, 1; PARL. in DC. Prod. 16, 2 (1868) 518; E1cHLER in E. & P. Nat. Pfl. Fam. 2, 1 (1887) 104; Wars. Monsunia 1| (1900) 193. — Nageia discolor (BLUME) O.K. Rev. Gen. Pl. 2 (1891) 800. — P. leptostachya BiumeE, Rumphia 3 (1849) 214; Mia. Fl. Ind. Bat. 2 (1859) 1073; HENKEL & HOCHSTETTER, Synop. Nadelholz. (1865) 392; pE Borer, Conif. Arch. Ind. (1866) 19, t. II, 1; PARL. in DC. Prod. 16, 2 (1868) 515; WarB. Monsunia | (1900) 193. — Nageia lepto- stachya (BLUME) O.K. Rev. Gen. PI. 2 (1891) 800. — dss, eek alco eetec Me hole arlene 7. P. levis P. junghuhniana Mig. Pl. Jungh. 1 (1851) 2; Fl. Ind. Bat. 2 (1859) 1073. — P. polyantha (WASSCHER) GaussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 191, f. 811. — P. decipiens Gray, J. Arn. Arb. 36 (1955) 204; GaussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 225, f. 844. Tree (3—)5—30(—45) m tall, 10—100 cm diam., clear bole up to 20 m; crown often dome-shaped. Fo- liage buds 2—5 mm long, occasionally longer, the pri- mary scales often spreading. Juvenile leaves 15—24 by up to 2.4 cm, acuminate and acute, narrowing ab- ruptly at the base to a short petiole, becoming linear- lanceolate and c. 1.6 cm wide on older fast-growing saplings in open forest situations. Leaves of mature trees similar with a short petiole up to 6 mm. Shade leaves acuminate, 8—12 by 1.2—1.8 cm. More ex- posed leaves linear-lanceolate, 12—18 by 1.1—1.5 cm or on particularly large trees more nearly linear and 7—10 by 1—1.1 cm. Midrib abruptly raised on the up- per side of the leaf at least 0.3 mm high and usually 0.6—0.8 mm wide but as little as 0.4 mm wide on less vigorous leaves or in the Borneo region on most leaves. Pollen cones solitary or in groups of two or most commonly three, sessile. Receptacile bright red when mature. Distr. From Nepal, Sikkim, Assam (Khasya), Thailand and Indochina through Malesia to the Solomon and Fiji Islands; in Malesia: Malaya, Sumatra (incl. Simalur I.), throughout Java and Bor- neo (incl. Karimata I.), the Philippines (Mindanao), Celebes, the Lesser Sunda Islands (Bali, Flores), the Moluccas (Obi, Ceram, Halmaheira), and New Guinea (incl. New Britain, New Ireland, Rossel I., Fig. 45. Range of Podocarpus neriifolius D.Don. 1988] Manus I., Biak, Job I., and Numfoor), common in many islands. Fig. 45. Ecol. Scattered and locally common in primary rain-forests from near sea-level to c. 2100 m. In most areas it appears as an understory tree with occasional much larger, emergent specimens in the canopy but in other areas, such as Java, Fiji, efc. it is normally a canopy tree. Habitats vary: rarely riverine, often on rocky hill- tops, in mossy forest, twice recorded from limestone, and twice from swampy forest. Also as to soils there is diversity: in Sarawak it is found on kerangas in heath forest and on sandstone ridges, but also on an- desitic laterites, which is the common latosol in Java, and sandy clay. In the Morobe District (New Guinea) it is recorded from ultrabasic. As to associates it is recorded from pelawan (Tris- tania) forest on sandstone ridges in S. Borneo; in the Javanese mountain forest its codominants are Dacry- carpus imbricatus and Altingia noronhae; in New Britain it occurs in the hills with Pometia and Calophyllum, in the montane forest in New Ireland with Fagaceae, Eugenia and Schizomeria; in the Morobe District (New Guinea) it is associated with Anisoptera and Flindersia in the canopy. Field notes. The bole is columnar, as usual; very rarely buttresses were recorded on field labels, 120 by 60 cm and 200 by 40 cm; sometimes the base was slightly spurred. Uses. A valuable timber tree, used for construc- tion. Vern. Brown pine, E; Malaya: jati bukit, Pahang; Sumatra: ambai ayam, Indragiri, hatang, Tapanuli, kayu tadji, Palembang, minangkas, Ben- coolen, naru dotan, Simalur |., sito bu hotang, Karo- Batak; Java: antoh, J, Japara, ki bima, ki merak, ki pantjar, ki putri, S; Borneo: belah buloh, Lawas, Sa- rawak, djadjaruman, Sampit, ki beling, Sabah; Celebes: kurniah, Nokilalaki; Lesser Sunda Islands: hadjo ketong, hadju pinis rona, Flores; Moluccas: bitao, Halmaheira; West New Guinea: aiwimunuwa- mee, Biak lang., bung, Mayu lang., buskagid/ji, buts- gagyi, Andai, Manikiong lang., gedorra, Fakfak, Esania lang., kayu tjina merah, Kp. Baros, kor- rikain, Tehid lang., wwa, Amberbaken lang., wajar, Mandobo lang., wasabraren, Numfoor |.; East New Guinea: isimberi, Nindewari, sipiri, Kikori R., sirau, Bulolo, Gairana dial. Notes. Much the most widespread species of the genus, but other species are very commonly identi- fied under this name causing confusion as to its exact character and retarding the recognition of the other species. In spite of the great geographic range, only slight variation exists within the species. The most distinct element is in the substantially isolated occurrence in Fiji where primary foliage budscales can reach 5 mm CONIFERALES (de Laubenfels) 401 and where extra vascular resin canals are often found in the leaves (P. decipiens). In Borneo the midrib on the upper surface of the leaf is weak while from India to southeastern Asia the leaves are more commonly lanceolate and the foliage buds are usually no more than 2 mm long with barely spreading primary scales. 2. Podocarpus rubens DE LAus. Blumea 30 (1985) 266. — P. neriifolius var. timorensis W ASSCHER, Blu- mea 4 (1941) 451. — P. pilgeri (non Foxw.) VAN Royen, Alpine Fl. New Guinea 2 (1979) 29, f. 39A. — Fig. 46. Fig. 46. Podocarpus rubens DE LAuB. Twig with seed-bearing structure showing two fruits on their re- ceptacles (after NGF 6980). Tree 2—30 m tall, 4—36 cm diam. Foliage buds 2—3 mm long, the primary scales erect or slightly spread- ing. Developing leaves bright red. Juvenile leaves linear, narrowly acuminate, becoming apiculate, 6—8 by 1.1—1.4 cm, narrowing at the base to a 2—3 mm petiole. Adu/t leaves linear to elliptic, 3—6 by 0.6—1 cm, apex acute to rounded and often apiculate, base same as juvenile leaves, margin revo- lute, midrib above 0.3 mm high with vertical sides, 0.3 mm wide. Pollen cones solitary or in threes, ses- sile or on a short peduncle. Receptacle becoming first red then purple. Distr. Malesia: Central to S. Sumatra, Celebes (Central: Latimodjong & Kambuno Mts; SW: G. Bonthain), Lesser Sunda Islands (Timor: G. Mutis), and New Guinea (incl. Normanby & New Britain). Fig. 47. Ecol. Scattered as a medium-sized, substage tree in primary rain-forest mostly above 1500 m but as 402 FLORA MALESIANA [ser. I, vol. 103 aa « as. z . ? ° Fig. 47. Range of Podocarpus rubens DE LAUvuB. low as 800 m on smaller islands or locally common to dominant as a small tree on ridges between 2000 and 3000 m or occasionally higher. Mostly on lato- sols, in New Guinea in Fagaceous mossy forest, rare- ly in swampy forest on peaty soils with Dacrydium. Vern. West New Guinea: bebi-€, Wissel Lakes, Kapauko lang., ungpop, Arfak, Hattam lang.; East New Guinea: bin, Ingembit village, W. Distr., kaip, Wabag, Enga lang., ne/il, Mt Tangis, New Britain, sukou, Wigote, Wapi lang. Notes. The bright red colour of a new flush of leaves which has been seen in this species can make the trees quite conspicuous. Herbarium specimens can resemble P. pil/geri but the upper midrib of the leaf rises c. 0.3 mm with vertical sides, while that of P. pilgeri is hardly 0.2 mm high with sloping sides. Leaves of P. pilgeri which approach P. rubens in size are from shady situations and are distinctly thin and delicate while those whose texture resembles the coarse rigid leaves of P. rubens are no more than 4—6 mm wide. The always solitary pollen cone of P. pilgeri is distinctly more slender and the ripe re- ceptacle is red. Unique in the section is that most specimens of P. rubens have some leaves with a nar- row-acuminate or more or less apiculate apex. The specimens of P. rubens from Ambon have a continuous upper hypoderm while elsewhere there are only scattered upper hypoderm cells and vascular sclereids. Specimens assigned to P. rubens from Normanby 1. and New Britain have quite blunt leaves with a less prominent midrib. In all other ways they conform to the species but it is possible that they represent a dif- ferent taxon. The specimen IcHLAs 166 from a high elevation in Sumatra seems too long and narrow to be placed comfortably here, even though more typical speci- mens exist nearby at lower elevation. The foliage bud is rather robust and the lateral resin canals are quite large. It resembles P. brevifolius and P. crassigem- mis and could represent a new allied species. 3. Podocarpus archboldii N.E.Gray, J. Arn. Arb. 39 (1958) 452; GaussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 177; pE Laus. Blumea 26 (1980) 140. Tree 10—40 m tall, 30—100 cm diam., clear bole up to 12—24 m, sometimes slightly fluted. Foliage buds 2—4 mm long, the primary scales slightly spreading. Juvenile leaves to at least 18 by 1.6 cm, otherwise like adult leaves. Adu/t leaves linear to linear-lanceolate, 7—12 by 1—1.4 cm, apex broadly acute, base ab- ruptly narrowed to a petiole up to 5 mm long, midrib above blunt, 0.3 mm high, with sloping sides, 0.5 mm wide. Pollen cones solitary, sessile or on a peduncle to at least 4mm. Receptacle often with a third lateral bract smaller than the second bract, red when ma- ture. Distr. Malesia: New Guinea. Fig. 48. Ecol. Scattered and locally common in the can- opy of mixed mid-mountain rain-forest with Casta- nopsis from 720 to 2200 m. Vern. West New Guinea: mu, soa, Kebar lang.; East: sarau, Bulolo, Geraina. Note. Sterile specimens strongly resemble P. ne- riifolius but strictly lack acuminate leaf tips and the upper midrib has sloping, not vertical sides. Unlike P. neriifolius the pollen cones are sometimes pedun- culate and the fruit receptacle sometimes has an extra lateral bract. 4. Podocarpus insularis DE LAuB. Blumea 30 (1985) 266. Small to medium-sized tree, 3—39 m tall, up to 20-60 cm diam., bole up to 24 m. Foliage buds 2.5—3 mm long and up to 3.5 mm diam., the primary budscales strongly spreading. Juvenile leaves linear, to 15 by 1.4 cm. Adult leaves elliptic, 5.5S—9 cm by 7—9 mm, apex acute, narrowed at the base to a 3—4 mm petiole; midrib above 0.3 mm high with vertical sides, 0.3 mm wide. Pollen cones solitary or in threes, sessile or with a short peduncle. Receptacle red when mature. Distr. New Hebrides and all Solomon Islands; in Malesia: New Guinea and adjacent islands: Rossel, Sudest, Misima, Woodlark, Fergusson, and New Britain. Fig. 49. Ecol. A good-sized lesser canopy tree, scattered and locally common in wet rain-forest, also in Nothofagus forest with undergrowth of Nastus, from near sea-level to 1680 m, and smaller trees from low ridge habitats. Vern. East New Guinea: dala, tunum, Milne Bay, Daga lang., ida-ayebo, Kutubu lang. Note. This species is distinguished by its rather small and narrow elliptic leaves with narrow but prominent midrib with nearly vertical slides and by the robust compact foliage buds with outward curl- ing scales. Dried specimens sometimes develop a rich red-brown colour on the underside. 1988] 5. Podocarpus deflexus Riptey, Fl. Mal. Pen. 5 (1925) 283; WaAssCHER, Blumea 4 (1941) 427; Gray, J. Arn. Arb. 39 (1958) 450; GaussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 175, t. 802. Small tree, 5—10 m tall, to 10 cm diam. Foliage buds c. 3 mm long but with the primary budscales up to 12 mm long and curling sharply outward with bluntly rounded apices, the secondary scales much shorter, broader, and rounded. Juvenile leaves up to 26 by 1.2—1.5 cm. Adult leaves linear to linear- lanceolate, 11—22 cm by 8—10(—13) mm, acute or slightly rounded at the apex, narrowing gradually more or less to a petiole, cuticle thick, often with five vascular resin canals, sharply bent at the base so as to hang downward, midrib above 0.4 mm high and 1 mm broad with indistinct margins, midrib beneath drying to a channel below. Pollen cones in groups of three, sessile or with a 2 mm peduncle. Primary pol- len cone budscales 2 mm wide. Receptacle formed by two fertile bracts and a third shorter sterile bract crowded more or less to one side between the other two. Seed 11—12 by 6-8 mm. Distr. Malesia: N. Sumatra (Gajo Lands: G. Ketambe) and Malaya (G. Tahan). Fig. 48. ‘> - sae a, YA / 8 es te / SN f Anne / Pe a Sy) forthe rv f - ae Seer = = 4 = = ae 2 al oo aS ° “Ys CR lo a git ee: = iy Ue / et Re 8 > ae ! a, HE es “2 ran £> Ee is Khe —] f\ Ee a — QL 4 ma yiabk Fig. 48. Range of Podocarpus archboldii N.E.Gray (A), P. deflexus Ripiey (D), and P. borneensis DE Laus. (B). Ecol. Rising above and locally dominant in dwarf mountain scrub, 1500-2100 m. Note. A rather distinctive species, its large smooth leaves and large fruit resembling among others P. rumphii and P. solomoniensis in other sec- tions of the genus but differing in the key characters of those sections. This species, unlike most of the genus, appears to be monoecious inasmuch as a ran- dom field inspection of about a dozen trees showed tvery single one to bear male buds (fruiting speci- mens have been collected at a different time of year). CONIFERALES (de Laubenfels) 403 6. Podocarpus borneensis DE LAus. Blumea 30 (1985) 266. — P. polystachus var. rigidus WASSCHER, Blumea 4 (1941) 460; Gray, J. Arn. Arb. 39 (1958) 471. Small to medium-sized tree, (2—)5—12(—23) m tall, 2.5—21 cm diam. Foliage buds 4—10 mm long, the primary scales erect. Juvenile leaves linear, to 16 by 1.4 cm. Adult leaves linear to ovate, (2.5—)4—7.5 (—9) cm by 8—13 mm, narrowing at the base to a 3—5 mm petiole, more or less acute but usually slightly rounded at the apex, thick, with abundant auxillary sclereids, midrib above prominent, 1 mm wide and 0.3 mm high, often drying to a channel below, often crowded around the foliage bud. Pollen cones soli- tary or in threes, sessile or on a short peduncle. Seed- bearing structure on a2 mm peduncle, receptacle red when mature. Seed with a small crest. Distr. Malesia: Borneo (incl. Karimata Is.). Fig. 48. Ecol. Locally common or even dominant (Meru- rong Plateau) on mossy rocky ridges, or scattered in nearby forest, in high kerangas forest and on white, sandy soils, 700—2070 m, one collection from a swamp at 360 m. Vern. Bisit, Sarawak, Kenya lang., bubung, Iban lang., buloh, Merurong Plateau. Note. As in P. deflexus the leaves are thicker than usual for the genus and the lower midrib dries to a channel. The abundant foliar sclereids is a char- acter shared apparently with P. novaecaledoniae in this section. 7. Podocarpus levis DE Laus. Blumea 24 (1978) 496. Tree, 8—25(—35) m tall, 16—40 cm diam. Foliage buds 3—9 mm long, the primary scales spreading. Ju- venile leaves up to 20 by 1.5 cm, narrowly acute. Adult leaves \inear-lanceolate, 8—14 by 1—1.4 cm, acute or narrowly rounded at the apex, narrowing at the base to a 4—9 mm petiole, stiff, upper hypoder- mal fibres in a continuous layer, upper midrib a low blunt ridge 0.3—0.5 mm wide and 0.1—0.2 mm high which furthermore often collapses upon drying so that the leaf appears smooth or even channeled, often with five vascular resin canals. Pollen cones typical of the section or longer, up to 8 cm long, soli- tary or in groups of up to three. Receptacle often with two bracts fertile and then sometimes with a third shorter sterile bract on one side between the other two, red when mature; foliola 2—4 mm long. Seed with its covering 12—13 by c. 8 mm. Distr. Malesia: E. Borneo (once; Berau), Cele- bes, Moluccas (Talaud Is.: Karakelong, Mt Piapi; Ambon: Mt Salhutu & Mt Hori), and West New Gui- nea (Meos Noom & Japan Is.). Fig. 49. Ecol. Scattered and locally common in primary rain-forest, from sea-level to 1650 m. In E. Borneo on limestone. FLORA MALESIANA [ser. I, vol. 103 Fig. 49. Range of Podocarpus insularis DE LAUB. (east of line) and P. levis DE LAuB. (west of line). Vern. Celebes: marisa, Donggala, sanru, Malili; New Guinea: kayu tjina, Meos Noom, wasiwarare, Japen I. 8. Podocarpus spathoides DE LAusB. Blumea 30 (1985) 267. Tree, 3—20 m tall, to 30 cm diam. Foliage buds 2—6 mm long, the primary scales spreading. Juvenile leaves up to 23 mm wide. Adult leaves linear, 5—13 by 0.8—2 cm, apex acute or rounded, narrowing abruptly at the base to a 3—5 mm petiole, midrib above broad and obtuse. Pollen cones in threes on a short peduncle. Distr. Solomon Islands; in Malesia: E. New Guinea (Louisiades: Rossel I.), N. Moluccas (Moro- tai: G. Pare), and Malaya (G. Ledang = Mt Ophir; type), all isolated occurrences. Fig. SO. \ (| SKE of ~e\ Ip a Na ra S ome ere pe LY es one ee oe oh ee ti ike Si : > co | Be a a seat : : a goer get Me Fig. 50. Range of Podocarpus spathoides DE LAUB. Ecol. Scattered and locally common at 1000— 1200 m in the western two stands and near sea-level in the east. Note. In Malaya and Rossel I. the foliage buds are no more than 3 mm long, the others are twice as long. In Malaya the trees grow in a summit scrub and are only 3—4 m high, elsewhere collectors report 12—20 m high trees. Perhaps intermediates exist or alternatively more than one similar taxon may be in- volved here. 2. Section Acuminatus DE LAuB. Blumea 30 (1985) 267. Primary foliage buds of variable diameters, at least 4 mm long, often much longer; primary scales erect, lanceolate, with free tips, as long as the bud itself, up to 3 mm wide at the base in vigorous buds but c. 1.5 mm wide in weaker buds; secondary budscales acuminate, the secondary bud when it first appears a loose cluster of free scales whose tips resemble the surrounding primary scales. Leaves with parallel margins, distinctly acuminate, narrowing more or less abruptly at the base, midrib above at least 0.5 mm wide and 0.3 mm high, broader and less prominent beneath, usually three vascular resin canals, occasionally more. Buds for pollen cones c. 3—5 mm long, either sessile or on a short peduncle, secondary scales acuminate, the secondary pollen cone bud when it first appears is briefly a round ball of overlapping imbricate scales. Pollen cones 3—5 cm long or longer, occasionally solitary but mostly in groups of three. Apex of the microsporophyll a triangular spur 0.5—0.7 mm long over a broader base. Peduncle of seed-bearing structure vari- able. Receptacle of the seed formed of one or two fertile bracts of variable sizes and one shorter sterile bract, the ripe receptacle red. Seed with its covering of variable size, globular, apex blunt or a slight crest. Distr. From Borneo to Queensland 3 spp. , of which 2 in Malesia: in Borneo and New Guinea respectively. Ecol. Low elevation primary rain-forest understory trees, in one case also common along disturbed forest margins. Note. The only section with both acuminate secondary budscales and acuminate leaves. 1988] CONIFERALES (de Laubenfels) 405 KEY TO THE SPECIES 1. Seed and its covering at least 11 by 8 cm, not crested. Adult leaves mostly more than 17 mm wide. Female peduncle at least 4 mm long. Plant lacking rhizomes ...................2.2--000- 9. P. ledermannii 1. Seed and its covering less than 11 by 8 mn, slightly crested. Adult leaves less than 17 mm wide. Female peduncle c. | mm long. Plant with rhizomes ..... 9. Podocarpus ledermannii PirGer, Bot. Jahrb. 54 (1916) 210; in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 248; WASSCHER, Blumea 4 (1941) 456; N.E.Gray, J. Arn. Arb. 39 (1958) 447; GaussenN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 173, t. 827; DE Laus. Blumea 26 (1980) 140. — P. idenburgensis N.E.Gray, J. Arn. Arb. 39 (1958) 447; GAUSSEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 171, t. 826. — P. ridleyi N.E.Gray, J. Arn. Arb. 39 (1958) 426, f. 1. Tree, 4—26(—33) m tall, 8—60 cm diam. Foliage buds 4—8 mm long by 3—4 mm in diam., the primary budscales more or less spreading. Juvenile and adult shade leaves \inear, 11—22 by 2—2.4 cm, narrowing abruptly to an acuminate apex and at the base toa 4—10 mm petiole; /eaves more exposed to the sun no more than 16—18 mm wide, weakly acuminate if at all and narrowing less abruptly at the base; midrib above broad and rounded, 1.5—2 mm wide and c. 0.6 mm high, sometimes collapsing on dried specimens to a small narrow ridge. Pollen cones c. 4.5 cm long, grouped on a 3—4 mm peduncle. Seed-bearing struc- ture on a 4—15 mm peduncle; foliola 2 mm long; re- ceptacle 9— 16 mm long. Seed with its covering 11—13 by 9—10 mn, slightly crested. Distr. Malesia: New Guinea and New Britain. Fig. 51. Ecol. Scattered and locally common in primary rain-forest understory from low elevation to at least 1800 m. Vern. West New Guinea: sua, Kebar, bébieai, Kapauko lang.; East New Guinea: babako, Aijura, Anona; New Britain: ne/eel, Cape Gloucester, nelil, Mt Talawe. Note. Leaves in the type collection are at the short and broad extremes for the species. Leaves in two examples were found to have two large addition- al vascular resin canals and one of these also had a continuous upper hypoderm. 10. Podocarpus micropedunculatus pe LaAus. Blumea 30 (1985) 268. SE Ek LE eh o> 10. P. micropedunculatus Shrub or small tree, 1—7(—13) m tall, 5—20 cm diam., with rhizomes which spread under the forest litter. Foliage buds 6—15 mm long by 1.5—3 mm diam. Juvenile leaves 14—18 by 1.5—2.1 cm. Adult leaves 8—17 by 1—1.5 cm; petiole 3—5 mm; midrib above 0.5—0.8 mm wide and 0.4 mm high, drying to a channel below. Pollen cones 3.5—7.5 cm, sessile or with a short peduncle. Primary pollen cone budscales 4--5 mm long. Seed-bearing structure on a 1 mm ped- uncle; foliola 3—4 mm; receptacle 8—10 mm long. Seed with its covering 8—10 by 6 mm with a small crest. Distr. Malesia: Borneo (Sarawak: Marudi For. Res.; Brunei; Sabah: Papar, Tawao). Fig. 51. Fig. 51. Range of Podocarpus ledermannii PILGER (L) and P. micropedunculatus DE LAuB. (M). Ecol. Scattered in understory of A gathis forest or forming a major element in thickets along the mar- gins of clearings, mostly on sandy, podzolic soils, ke- rangas, sandstone, humic peaty podsols of raised beaches, and peat-swamp forest, from sea-level to oc- casionally as much as 500 m. Sometimes associated with Dryobalanops rappa or Shorea albida. Vern. Sarawak: kayu china; Sabah: kayu tjina. Note. Propagation by rhizomes is a remarkable character otherwise unknown among erect conifers and obviously facilitates rapid appropriation of dis- turbed situations. 3. Section Globulus DE Laus. Blumea 30 (1985) 268. Primary foliage bud a globular head of completely imbricate scales, gradually expanding with 406 FLORA MALESIANA [ser. I, vol. 103 the growth of a new shoot until the secondary bud, also globular, erupts; primary budscales acute to more or less rounded, c. 1.5—2 mm long; secondary scales oval and blunt. Leaves with parallel sides or somewhat lanceolate, slightly rounded at a narrow apex to broadly rounded and sometimes acuminate, midrib above blunt, 0.2 mm high and at least 0.5 mm broad, broader and indistinct below, 3 vascular resin canals. Buds for pollen cones sessile or on a short peduncle, the primary scales c. 1.5 mm long; the secondary pollen cone bud similar to the secondary foliage bud. Pollen cones mostly 2.5—4.5 cm long (smaller in some species), solitary or in groups of three. Apex of the microsporophyll mostly a small triangle less than 0.5 mm long (longer in P. nakaii). Seed-bearing structure on a peduncle of variable length; receptacle formed of two bracts, one of which is longer and mostly 7—9 mm long (longer in P. /ucienii) or both are fertile and equal, where known the ripe receptacle red. Seed with its covering 8—10 mm long by 5.5—6 mm diam. (or larger in some species), globular, apex blunt or in some species with crest. Distr. From Vietnam and Formosa to Borneo and New Caledonia 6 spp.; in Malesia: 2 spp., in Sumatra/Malaya and Borneo respectively. Ecol. Mostly short trees of low (to 2000 m) elevation tropical forest either in the understory of primary or secondary rain-forest (occasionally larger and in the canopy), on exposed mossy ridges, or in short open forest. Note. The gap between Borneo and New Caledonia is largely filled by the related sect. Rumphius which overlaps this section geographically only slightly. The primary budscales in sect. Rumphius rather than being completely imbricate and overlapping more or less meet at their apices with usually one or more actually curl- ing outward at their apex while, in addition, the pollen cones in sect. Rumphius are normally in clusters of more than three and all the species of sect. Rumphius occur as large canopy trees. KEY TO THE SPECIES 1. Seed and its covering with a small crest; foliola 2 mm long. Adult leaves not acuminate, 9-15 mm wide 11. P. globulus 1. Seed and its covering without a crest; foliola less than 1 mm long. Adult leaves distinctly acuminate, 14—21 ANTE WIE sre Ae ePRe eo os eee I~. Meitey. aihic kolaioe ces © nS rewiele = Se eae 12. P. teysmannii 11. Podocarpus globulus DE Laus. Blumea 30 (1985) 269. Tree, 3.5—27 m tall, to 18 and more cm diam. Ju- venile leaves 7.5—16 by 1.5—2.4 cm, abruptly acu- minate. Adult leaves linear, narrowing more or less abruptly at the base to a 2—3 mm petiole, acute to rounded apex, 3.5—8 by 0.9—1.5 cm, midrib above 0.5—0.7 mm wide. Pollen cones 2.5—4.5 cm, solitary or occasionally in threes, sessile or on a short (1 mm) peduncle. Seed-bearing structure on a 3—4 mm ped- uncle; foliola 2 mm and early deciduous. Seed with its covering with a brief crest. Distr. Malesia: northern part of Borneo (Sara- wak; Sabah: Mt Silam). Fig. 52. Ecol. In primary rain-forest or mossy forest on ridges and peaks from 300 to 1500 m apparently where the forest is not dominated by dipterocarps. In some cases at least, an ultrabasic soil is indicated. Vern. Sapiro, Lawas, Murut lang. 12. Podocarpus teysmannii Mig. Fl. Ind. Bat. 2 (1859) 1072; bE Boer, Conif. Arch. Ind. (1866) 14, t. 1; Pari. in DC. Prod. 16, 2 (1868) 516; WaRB. Monsunia | (1900) 193; Pircer, Pfl. R. IV, 5, Heft Fig. 52. Range ot Podocarpus globulus DE LAUB. (G) and P. teysmannii Mia. (T). 18 (1903) 81. — Nageia teysmannii (Miq.) O. K. Rev. Gen. Pl]. 2 (1891) 800. — P. neriifolius var. teysman- nii (MiQ.) WaAsscHER, Blumea 4 (1941) 453; N.E.Gray, J. Arn. Arb. 39 (1958) 468; GAUsSEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 189. Tree, 4—12 m tall, up to 30 cm diam. Juvenile leaves to 16 by 2.7 cm. Adult leaves linear, 9—13 by 1.4—2.1 cm, acuminate, narrowing more or less ab- ruptly at the base to 4—7 mm petiole, midrib above 0.6—0.8 mm wide. Pollen cones 2.5 cm, solitary or occasionally paired, sessile. Seed-bearing structure 1988] CONIFERALES (de Laubenfels) 407 on a 6—11 mm peduncle; foliola 1 mm and early Ecol. Understory tree of primary or secondary deciduous. rain-forest from sea-level to occasionally as high as Distr. Malesia: Malaya and Sumatra (Westcoast 1140 m, in Banka on granite sand. Res.), incl. Riouw-Lingga and Banka Is. Fig. 52. Vern. Sumatra: kalek rotan, Westcoast. 4. Section Longifoliolatus DE LAuB. Blumea 30 (1985) 269. Primary foliage budscales erect, narrowly lanceolate with free tips, occasionally as little as 4 mm long on weak shoots but mostly at least 6 mm long and often much longer; secondary bud- scales acuminate, the secondary bud when it first becomes visible a cluster of erect scale tips shor- ter than the primary scales. Leaves either linear and no more than 11 mm wide or narrowly lan- ceolate, narrowing gradually at the base, midrib above variable in width and mostly c. 0.3 mm high but lower in some species, broader and less prominent beneath (and in some species drying into a channel), in most species more than 3 vascular resin canals, frequently a well-developed or continuous upper hypoderm, especially where there are no extra resin canals. Buds for pollen cones 3—4 mm long, mostly sessile but with a short peduncle in a few species, the primary scales mostly 1.5 mm wide at the base, occasionally wider; secondary scales mostly acuminate, the sec- ondary pollen cone bud when it first appears sometimes still a ball of overlapping scales but more often with the scale tips starting to separate. Pollen cones variable in length, slightly greater in diameter than usual for the genus where the apex of the microsporophyll is longer than usual, soli- tary or sporadically in groups of 2 or 3. Apex of the microsporophyll usually a spur c. 0.5 mm long on a wider base but longer in some species and shorter in others. Seed-bearing structure on a peduncle (2—)7—15(—20) mm long; foliola in all species to over 3 mm and only sometimes as little as 2mm in any species. Receptacle formed of at least two bracts at least one of which is fertile and longer, while often, or in some species usually, two are equal, with one to three smaller bracts crowded between them and sometimes one or two of these smaller bracts fertile, the larger bracts variable in size between the species but at least 9 mm long; where known the ripe receptacles are red to dark red. Seed with its covering also variable in size, at least 8 mm long by 6 mm diam., globular, in some species with a weak crest. Distr. Mostly in highly localized stands from Sumatra to Fiji, 10 spp., 5 of which are in Malesia. Ecol. Mostly small or stunted trees (or even decumbent) in more or less specialized habitats, particularly over ultrabasic soils. Notes. This section can be divided roughly into two parts, one with more or less parallel-margined leaves with extra resin canals and normally at least two fruits on each fertile structure as well as small pollen cones 10—28 mm long and a strong relationship to ultrabasic soils; the other part with lanceolate leaves, which only in some cases have extra resin canals and normally only one fruit in each fertile structure as well as larger pollen cones 3—4.5(—6) cm long and a relationship to a variety of habitats. In all species the foliage buds on vigorous shoots surpass 8 mm, a condition found in the subgenus only elsewhere in sect. Acuminatus. The female foliola usually or always at least 3 mm long are rare elsewhere in the genus. KEY TO THE SPECIES 1. Pollen cones less than 3 cm long; leaf elliptic, somewhat revolute, adult leaves less than 5 cm long 13. P. gibbsii 1. Pollen cones more than 3 cm long; leaf mostly lanceolate or linear-lanceolate, not revolute, adult leaves at least 5 cm long. 2. Leaf linear-lanceolate or linear, midrib broad and blunt on the upper side (no crest on fruit) 14. P. confertus 2. Leaf lanceolate, midrib sharp and narrow on the upper side. 408 FLORA MALESIANA [ser. I, vol. 103 3. Leaves more than 7 mm wide; scales at base of pollen cone long triangular; fruit with a crest. 4. Leaves mostly over 10 mm wide; foliola at base of receptacle more than 3 mm long; bud for pollen cone 4—5 mm in diameter; pollen cone initially sessile, when mature with an elongated scaly base 15. P. bracteatus 4. Leaves mostly less than 10 mm wide; foliola at base of receptacle up to 3.4 mm long; bud for pollen cone c. 2 mm in diameter; pollen cone with a short peduncle to 2 mm.... 16. P. pseudobracteatus 3. Leaves less than 8 mm wide; scales at base of pollen cone broad and rounded; fruit blunt 13. Podocarpus gibbsii N.E.Gray, J. Arn. Arb. 39 (1958) 429; GaussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 155, t. 803. Tree 2.5—20 m tall. Foliage buds 4—9 mm long. Juvenile leaves linear, acute, 4—9 cm by 5—9 mm. Adult leaves linear to ovate, 2—5 cm by 4—7 mm, acute to almost rounded at the apex, narrowed at the base to a 1—3 mm petiole; midrib above weakly de- veloped, 0.5—0.7 mm wide and 0.1—0.2 mm high and often collapsing upon drying leaving a weak depres- sion with a narrow ridge in the centre, upper hypo- derm continuous. Pollen cones 1—1.5 cm, sessile, solitary or occasionally a pair; apex of microsporo- phyll triangular, up to 0.8 mm long. Seed-bearing structure on a peduncle at least 3 mm long; foliola 4 mm long; receptacle formed of two unequal bracts; mature seed unknown. Distr. Malesia: Borneo (Sabah: Mt Kinabalu). Fig.953:. . 8) x | \Y ; Fig. 53. Range of Podocarpus gibbsii N.E.GRay (G) and P. atjehensis (WASSCHER) DE LAUB. (A). Ecol. Mossy ridges between 1200 and 2400 m, mostly or always on ultrabasic soil. Note. The leaves generally resemble those of P. pilgeri, a species which is not otherwise similar at all. 14. Podocarpus confertus DE Laus. Blumea 30 (1985) 271. Tree 1—36 m tall. Foliage buds 6—10 mm long. Juvenile leaves up to 20 by 1.2 cm. Adult leaves linear to linear-lanceolate, 5-12 cm by 7-11 mm, acute, narrowing at the base to a 3—6 mm petiole; midrib above obtuse, 1 mm wide and 0.2—0.3 mm high, collapsing when dry to form a narrow irregular ridge or becoming completely flat, upper hypoderm 17. P. atjehensis continuous. Pollen cones 3—4.5 cm, sessile, solitary or occasionally in pairs; apex of microsporophy]ll tri- angular, 0.3—0.5 mm long. Seed-bearing structure ona5—13mm peduncle; foliola S—6 mm long; recep- tacle 8—12 mm, formed of two unequal bracts. Seed with its covering 10—11 by 6—6.5 mm. Distr. Malesia: Borneo (Sabah: Mt Silam). Fig. 54. Ecol. In dense local populations on various poor soils some or most of which are ultrabasic; subdomi- nant in somewhat open and sometimes stunted forest from 90 to 1200 m. 15. Podocarpus bracteatus BLUME, En. Pl. Jav. 1 (1827) 88; BENNETT in Bennett & R.Br. Pl. Jav. Rar. 1 (1838) 40; ENDL. Syn. Conif. (1847) 216; BLUME, Rumphia 3 (1849) 214, t. 172, f. 1; Mig. Pl. Jungh. 1 (1851) 2; Fl. Ind. Bat. 2 (1859) 1072; HENKEL & HOcHSTETTER, Synop. Nadelholz. (1865) 391; DE Boer, Conif. Arch. Ind. (1866) 16; Pari. in DC. Prod. 16, 2 (1868) 515; EICHLER in E. & P. Nat. Pfl. Fam. 2, | (1887) 104; Wars. Monsunia | (1900) 192. — P. nertifolius var. bracteatus (BLUME) WASSCHER, Blumea 4 (1941) 449; GaussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 189, t. 806. — P. bracteatus var. brevipes BLUME, Rumphia 3 (1849) 214; Mra. Pl. Jungh. 1 (1851) 2; HENKEL & HOCHSTETTER, Synop. Nadelholz. (1865) 392; Pari. in DC. Prod. 16, 2 (1868) 515. — P. neriifolius var. brevipes (BLUME) Pitcer, Pfl. R. [V, 5, Heft 18 (1903) 81; GaussENn, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 187. — P. neriifolius (non D.Don) STEEN. Mt. Fl. Java (1972) Les eafe ale Tree 10—40 m tall, 15—100 cm diam. Foliage buds 5—12 mm long. Juvenile leaves up to 23 by 2 cm. Adult leaves distinctly lanceolate, 6—14 cm by 9—14 mm, narrowly acute, narrowing at the base to a 2—4 mm petiole, sometimes with 5 vascular resin canals, midrib above a sharp ridge 0.4 mm wide and 0.3 mm high, sometimes collapsing when dried into a trough. Pollen cones 3.5—6 cm by 3—4 mn, sessile but elon- gating when mature through the scaly base as well as the zone of microsporophylls and the scales then fol- lowing, the elongated scaly base up to 8 mm long, solitary or occasionally in pairs; apex of microsporo- phyll c. 1 mm long. Seed-bearing structure on a 10-20 mm peduncle; foliola 4-5 mm long; re- ceptacle 10-14 mm long, often with two fertile 1988] CONIFERALES (de Laubenfels) 409 bracts and additional bracts between them. Seed with its covering including a small crest, 11—14 by 7 mm. Distr. Malesia: N. & Central Sumatra (rare), throughout Java and the Lesser Sunda Islands (Flo- res: Mt Ranaka). Fig. 54. i \ e eg / t a Fe » p~ ef _ \( “a = —— of As 7 = WT bx Xs \ / S = le a P D SS slo oe q < ae “See > ~ ~--—- 7 . —t 2 1 h — “NS a ae | \ 4 ~ be . + \ ¢ . 8 > \ ‘SY Fig. 54. Range of Podocarpus confertus DE LAUB. (C), P. bracteatus BLUME (B), and P. pseudobractea- tus DE LAUB. (P). Ecol. Scattered in the canopy of moist mountain rain-forest from 1000 to 2600 m or occasionally as low as 400 m. Uses. An excellent timber tree. Vern. Sumatra: kayu unung unung, Toba Batak; Java: bima, J (Pekalongan), ki marak, ki pantjar, ki putri, S. Note. The most common Podocarpus of the mountain forests of Java. Podocarpus neriifolius also occurs there, but rarely above 1600 m, while P. bracteatus is common to over 2000 m. Sterile speci- mens of the two are sometimes similar, but P. bracteatus has a narrower midrib, longer budscales, and a more distinctly lanceolate shape. 16. Podocarpus pseudobracteatus pe LAuB. Blumea 26 (1980) 142. — P. archboldii var. crassiramosus N.E.Gray, J. Arn. Arb. 39 (1958) 453. Tree 1-15 m tall, 5-20 cm diam. Foliage buds 5—14 mm long. Juvenile leaves linear-lanceolate, up to 22 by 1.7 cm. Adult leaves linear-lanceolate to lan- ceolate or exceptionally elliptic, 6-15 cm by 7—12 mm, narrowly acute, tapering more or less abruptly at the base to a 2—4 mm petiole; midrib above a prominent narrow ridge with nearly vertical sides, 0.4—0.5 mm wide and 0.3—0.4 mm high. Pollen cones 4—4.5 cm, on a short peduncle up to 2 mm long, solitary; apex of the microsporophyll 0.5—0.7 mm long. Seed-bearing structure on a 2—5 mm ped- uncle; foliola 2.5—3 mm long, often thick and lan- ceolate; receptacle 7-11 mm long, formed of two bracts and becoming first orange, then red, then al- most black when ripe. Seed with its covering 10-11 by 8—9 mm, blunt. Distr. Malesia: New Guinea. Fig. 54. Ecol. Scattered and locally common in the understory of mossy Castanopsis-Nothofagus forest and Dacrydium swamp forest, sometimes entering the alpine shrubbery, from 1740 to 2850 m. Vern. East New Guinea: kaip, Wabag, Enga lang., kebu, Tari Gap, puling, Togoba, Chimbu. 17. Podocarpus atjehensis (WASSCHER) DE LAUB. Blumea 30 (1985) 271. — P. neriifolius var. atjehen- sis WASSCHER, Blumea 4 (1941) 450; N.E.Gray, J. Arn. Arb. 39 (1958) 466; GAaussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 189. Tree 8—15 m tall, 20 cm diam. Foliage buds 6—14 mm long. Adult leaves linear-lanceolate to lanceo- late, 7-11 cm by 6—8 mm, narrowly acute, narrow- ing somewhat gradually at the base to a 3—4 mm petiole, cuticle thick, sometimes with lateral vascular resin canals distinctly larger than the median canal, midrib above a prominent ridge with nearly vertical sides, 0.2—0.3 mm wide, 0.2 mm high. Pollen cones 3.5 cm, sessile, solitary; apex of the microsporophyll a small triangular spur c. 0.2 mm long. Seed-bearing structure on a 8—16 mm peduncle; foliola 2-4 mm long; receptacle 10—11 mm long, formed of two un- equal bracts, becoming red when mature. Seed with its covering, including a blunt apex, 9-11 by 7-8 mm. Distr. Malesia: N. Sumatra (Gajo Lands: Kemiri & Bandahara Mts) and West New Guinea (Wissel Lakes). Fig. 53. Ecol. In local forest populations, probably on poor soils, in N. Sumatra at 2500-3300 m, near Wis- sel Lakes at 1800 m. 5. Section Gracilis pE Laus. Blumea 30 (1985) 272. Primary foliage bud small, up to 2 mm diam., but smaller in some species; primary scates erect or slightly spreading, triangular to lanceolate, up to 3 mm long and | mm wide at the base, only 0.6 mm wide in P. glaucus; secondary scales also acute, the secondary bud when it first appears a cluster of free bud tips. Leaves linear to ovate, acute to rounded at the apex, particularly delicate with remarkably blunt apices when growing in the shade, generally distinctly rigid when growing in exposed situations; midrib above distinct but gracile with sloping sides and about the size of FLORA MALESIANA [ser. I, vol. 103 410 an ordinary pencil line (up to 0.3 mm wide), broader and blunt to nearly flat below, three vascular resin canals. Buds for pollen cones sessile and quite small, the primary scales no more than 1.5 mm long with free tips; secondary scales about twice as long as broad, the secondary pollen cone bud when it first appears a spherical ball cupped by the primary scales. Pollen cones slender, most- ly 2—3.5 cm long and c. 3 mm diam. before shedding pollen but 2—2.5 mm diam. after shedding, solitary, often elongating in the scaly base as well as in the zone of microsporophylls when mature. Apex of the microsporophyll a small triangular spur 0.3—0.5 mm long over a base c. 1 mm wide. Seed-bearing structure on a peduncle (3—)5—10(—16) mm long. Foliola 1.5 mm long. Receptacle formed of two bracts, the fertile bracts 7—8(—10) mm long, sterile bracts shorter, becoming red in some species and purple in others when mature. Seed with its covering globular, 7—8 mm long without a small crest and 8—9 mm long in species with a crest, mostly 5.5—6 mm diam. (less in P. affinis from Fiji). Distr. From southern China through Malesia to Fiji, but not in Malaya, Sumatra, and Java, 5 spp. Two species widespread and the other three quite rare and outside the range of the other two. Four of the five spe- cies occur in the Philippines. Ecol. Mossy mountain forests either on isolated peaks or at high elevation. Note. A group of closely related species, some of which could conceivably be considered varieties in- asmuch as certain variations of a like nature are also known within P. pilgeri. For example, P. lophatus has crowded leaves and a crest on the fruit, characters shared with P. glaucus but otherwise it corresponds with P. pilgeri. On the other hand, P. wangii, here included in P. pilgeri, has very small foliage buds and a purple ripe receptacle, characters also shared with P. glaucus. KEY TO THE SPECIES 1. Leaves over 2 cm long; foliage bud at least 1.5 mm long. 2. Leaves less than 8 mm wide, not always blunt (sun growth leaves acute). de seaves dispersed: Eruitnot crested) .=.42..2c-5 - - 3. Leaves crowded (overlapping). Fruit crested ... A. Sh. co eas es ee 18. P. pilgeri «gird a Cele Sea. Bebe eee 19. P. lophatus 2. Leaves more than 8 mm wide, always blunt (dispersed; fruit not crested)........... 20. P. rotundus 1. Leaves less than 2 cm long (less than 6 mm wide, always blunt, crowded); foliage bud less than 1.5 mm lonea(irnitgenested) ieee 44? sienna. sea. : 18. Podocarpus pilgeri Foxw. Philip. J. Sc. 2 (1907) Bot. 259; ibid. 6 (1911) Bot. 160; PitGeEr, Bot. Jahrb. 54 (1916) 38; in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 248; N.E.Gray, J. Arn. Arb. 39 (1958) 459; GAUSSEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 185, t. 810; bE Laus. Kalikasan 7 (1978) 135. — P. celebica (non HEMSL.) Wars. Monsunia 1 (1900) 192; Pitcer, Pfl. R. IV, 5, Heft 18 (1903) 78. — P. costalis (non PRESL) Foxw. Philip. J. Sc. 6 (1911) Bot. 161. — P. schlechteri Pitcer, Bot. Jahrb. 54 (1916) 209; in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 248; Laut. Bot. Jahrb. 63 (1930) 474; Pmtcer, Bot. Jahrb. 68 (1936) 246; GaussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 187. — P. wangii CHANG, Sun- yatsenia 6 (1941) 26. Tree 1—25 m tall, 8—60 cm diam.; bole to 12 m, rarely fluted. Foliage buds 2.5—3 mm long (or some- times shorter). Juvenile leaves linear, up to 7 cm by 9 mm, broadly acute and apiculate. Adult shade leaves ovate, widest part closer to the apex, 2—4 cm by 5—8 mm, abruptly rounded at the apex, nar- rowing more gradually at the base to a short 2-3 mm innate feud. £ Bel eek: dees 21. P. glaucus petiole; exposed leaves elliptic, 2.5—4 cm by 4—6 mm, acute, revolute, with intermediate forms to- wards shade leaves common; midrib above on all leaves 0.2 mm wide and high. Pollen cones often elongating in the basal scaly part to 3 mm. Female re- ceptacle becoming red when mature (dark violet has also been reported). Seed with its covering without a crest. Distr. S. China; in Malesia: Philippines (Negros Occidental, Mindanao), Central Celebes, and com- mon in New Guinea. Fig. 55. Ecol. Scattered and locally common in moist and often mossy forest, (700—)1200—1300 m, as a me- dium-sized tree, but dwarfed on ridges and at high elevation. Mostly in the understory, in beech forest with mossy undergrowth, associated with Phyllocla- dus and Myrsine, in New Guinea in elfin woodland on Mt Hunstein and on Mt Nettoti in low Xantho- myrtus-Podocarpus crest forest. Vern. West New Guinea: bempop, Hattam lang.; East New Guinea: gihura, Asaro, Kefamo, iamu- gang, Goroka, Togoba, jamega, Hagen, Togoba, CONIFERALES (de Laubenfels) Fig. 55. Range of Podocarpus pilgeri Foxw. kaibeltugl, Wahgi, Minj, kebu, Mt Ne, monopana, Mairi, Natabung, puling, Hagen, Wankl, sosumehi, Dunantina, sula, Chimbu, Masul, yamga, Hagen, Minj, yazib, Nondugl, Minj. Note. The great variability of the leaves can be confusing and the position on the tree of leaf collec- tions should be noted. 19. Podocarpus lophatus pE Laus. Kalikasan 7 (1978) 137. — P. brevifolius (non STAPF) Foxw. Philip. J. Sc. 6 (1911) Bot. 160, t. 29, f. 2; GausSsEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) t. 801 p.p. Small tree. Foliage buds 3 mm long. Adult leaves densely crowded, thick, elliptic, 3 cm by 5 mm, slightly revolute, acute, narrowed at the base to a short 2 mm petiole; midrib above a distinct ridge c. 0.2 mm wide. Pollen cones unknown. Seed with its covering with a distinct crest. Distr. Malesia: Philippines (Luzon: Mt Tapu- lao), one locality only. Fig. 56. Ecol. Mossy forest at 1800 m. 20. Podocarpus rotundus pe Laus. Kalikasan 7 (1978) 136. Tree 5—15 m tall. Foliage buds 2—3 mm long. Juvenile leaves \inear, up to 10 cm long. Adult leaves oval to slightly linear, 2-5 cm by 8—11 mm, very rounded at the apex even when growing in exposed situations, sometimes apiculate, narrowed at the base to a 2 mm petiole; midrib above a small ridge 0.2 mm wide and high. Pollen cone elongating through the base to produce a scaly section 3-10 mm long. Female receptacle becoming red when mature. Seed with its covering without a crest. Distr. Malesia: E. Borneo (Mt Beratus near Ba- likpapan) and Philippines (Luzon: Mt Banajao in Laguna Prov. and Lucban in Tayabas Prov.), 3 col- lections. Fig. 56. Ecol. Dwarf mossy forest, on Mt Beratus at c. 1000 m, in Luzon up to 2200 m. 411 os | V Sy y, Ea Se of = : ae é e ap nn ae yes densa ee ini: D “DS Fig. 56. Range of Podocarpus lophatus DE LAvB. (L) and P. rotundus DE LauB. (R). 21. Podocarpus glaucus Foxw. Philip. J. Sc. 2 (1907) Bot. 258; ibid. 6 (1911) Bot. 159, t. 29, f. 1; PmLGER in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 248; Was- SCHER, Blumea 4 (1941) 468; N.E.Gray, J. Arn. Arb. 39 (1958) 440; GaussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 163, t. 825; DE Laus. Kalika- san 7 (1978) 138. Decumbent shrub to small or medium-sized tree, 2-15 m tall, up to 20—25 cm diam. Foliage buds 1—1.5 mm long and 1.5 mm in diam.; primary bud- scales 0.6 mm wide. Juvenile leaves oval, 2—3.5 cm by 5—7 mm. Adult leaves ovate, the widest part somewhat beyond the centre, 1—2 cm by 3—6 mm, round and very blunt at the apex, narrowing at the base to a 2—3 mm petiole, revolute, crowded, flush- ing red, at least sometimes glaucous; midrib above a distinct ridge 0.2 mm wide. Pollen cones 1-2 cm long, often elongating in the basal scaly part to 1—2 mm. Female receptacle becoming purple when ma- ture. Seed with its covering with a distinct crest. Distr. Solomon Islands; in Malesia: New Guinea Fig. 57. Range of Podocarpus glaucus Foxw. 412 FLORA MALESIANA [ser. I, vol. 103 (also on Arfak Mts, and incl. Manus in Admiralty Is.), Moluccas (W. Ceram), and Philippines (Min- doro). Fig. 57. Ecol. A medium-sized tree in the forest or more often dwarfed or even decumbent on mountain crests in stunted mossy forests, often locally common, (500—)1000—2800 m. Recorded from stony, sandy clay and from a limestone ridge associated with Gymnostoma and Rhododendron, near Kiunga and Wissel Lakes (New Guinea) on peaty soil. Vern. West New Guinea: nipa, Kebar, bébiai, Kapauko lang. Note. Sterile young plants resemble P. pilgeri, but with crowded leaves. 6. Section Macrostachyus DE LAus. Blumea 30 (1985) 273. Primary foliage bud mostly at least 4 mm diam. on vigorous shoots, but 3—3.5 mm diam. on weaker shoots, and generally somewhat smaller in some species, about as long as or somewhat longer than the diameter, none more than 5 mm long; primary scales triangular and erect or lan- ceolate and often with outwardly curling tips and thus longer than the bud itself; secondary scales acute or apiculate, the secondary bud when it first appears a nest of scale tips which elongates into a loose pyramid. Mature leaves only c. 5—8 mm wide and mostly less than 6 cm long, acute to rounded at the apex, narrowed gradually at the base, mostly revolute, tough, midrib above nar- row but distinct, 0.2—0.4 mm wide and 0.1—0.2 mm high, broader and blunt below, sometimes drying to a channel, three vascular resin canals. Buds for pollen cones sessile (or in P. crassigem- mis on a short peduncle), the secondary pollen cone bud when it first appears more exposed and still a globular ball. Pollen cones 2.5—5.5 cm long and up to 7 mm diam. before shedding pollen, but 4—5 mm diam. after shedding and even smaller in one variety which does not have the usual lanceolate 2—3(—4) mm apex of the microsporophyll, solitary or occasionally in pairs. Seed- bearing structure of various sizes, usually with a short peduncle 2—6 mm long and foliola c. 2mm long; where known the fully ripe receptacles are dark purplish black passing through red in ripen- ing. Distr. In isolated populations from southern Cambodia and islands south of Taiwan to New Guinea, 5 spp., of which 4 in Malesia (two widespread examples in New Guinea). Ecol. Locally common at high elevations or on mossy ridges. One species, P. costalis, largely confined to a group of small islands at low elevation. Note. A similar habitat and general distribution to sect. Gracilis but members of sect. Macrostachyus have robust leaves where those of sect. Gracilis are delicate. One sterile specimen from Sumatra (ICHLAS 166, Mt Singgalang, 2800 m) placed with P. bracteatus (sect. Longifoliatus), resembles P. brevifolius of the present section also, but more information is needed. KE Ye LO THE SEROLES 1. Foliage bud at least 3 mm in diameter. Leaves elliptic. 2. Seed with its covering not crested, more than 9 mm diameter. 3. Leaf at least 7 times as long as wide. Pollen cone pedunculate ............... 22. P. crassigemmis Sltcat less than’ 4 times as: long as)wide. Pollen conersessile’ Saye.hys- n a0. set ee 23. P. brassii 2. Seed with its covering crested, less than 8 mm diameter. (Leaf less than 7 mm wide) 24. P. brevifolius 1. Foliage bud less than 3 mm in diameter. Leaves more or less linear blunt. (Fruit crested, less than 7 mm (QUEINEKS Cl\in:53 bra -stgasees Rees a Sot OrG cc OER CO arr eR PE acl eR Ra Pere enle esol Sc 25. P. costalis 22. Podocarpus crassigemmis DE Laus. Blumea 26 (1980) 141. — P. archboldii (non N.E.GRay) GaAussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 177, t. 828; vAN RoyeEN, Alpine Fl. New Guinea 2 (1979)30; t. 41. Tree (3—)8—38 m tall, 10—75 cm diam., bole occa- sionally fluted. Sometimes pagoda habit. Foliage buds 3—5 mm long, the primary scales up to 8 mm long, on juvenile plants to over 10 mm, strongly curl- ing outward. Juvenile leaves linear-lanceolate, nar- 1988] CONIFERALES (de Laubenfels) 413 rowly acute, up to 20 by 1.4cm. Adult leaves elliptic, 3-11 cm by 4.5—12 mn, acute to narrowly acute, narrowing at the base to a 2—5 mm petiole, revolute, midrib above a sharp ridge 0.2—0.4 mm wide and 0.2 mm high. Buds for pollen cones on a 1—7 mm pedun- cle, primary scales up to 4.5 mm long. Pollen cones occasionally in pairs. Seed-bearing structure on a 5—14 mm peduncle; receptacle 10—15 mm long. Seed with its covering 11—14 by 9-10 mm. Distr. Malesia: New Guinea (except Vogelkop Peninsula). Fig. 58. Fig. 58. Range of Podocarpus crassigemmis DE LAUB. Ecol. Common or subdominant in the canopy of high mountain mossy forest, or emergent, often in Nothofagus and Phyllocladus forest, rarely in sec- ondary forest and occasionally in grassland, (1800—)2100—3400 m. Vern. East New Guinea: a-pul, kaboga, morum- ba, Mt Giluwe, Mendi lang., baula, Kundiawa, Chimbu, iamuka, Tomba, jamekang, Hagen, Tom- ba, juba, kamga, puling, Hagen, Togoba, kabor, Anga Valley, Mendi lang., kabiltugl, kaibelparu, kaibig, Wahgi, Minj, kaip, Kepilam, Enga lang., no- nofan, Mairi, Watabung, ronohanini, Asaro, Kefamo, ((/sula, Chimbu, Masul. Note. Leaves from lower parts of trees substan- . tially larger than those from higher or more exposed parts of the same tree can at the same time bear fertile material. In the Goroka Subdistrict SrEvENs found two trees (LAE 51011) of which the leaves were infected with Corynelia uberata Fries, widely distributed in the Old World. 23. Podocarpus brassii Pitcer, Bot. Jahrb. 68 (1937) 246; Wasscuer, Blumea 4 (1941) 469; N.E.Gray, J. Arn. Arb. 39 (1958) 440; Gaussen, Gymn. Act. & Foss. fase. 14, ch. 21 (1976) 163, t. 824; VAN ROYEN, Alpine Fl. New Guinea 2 (1979) 24, t. 37, 38. KEY LO; THE YiARIE TIES 1. Apex of microsporophyll lanceolate, at least 2mm Kel cfs OL Ng farina te SenGA Mas oF a. var. brassii 1. Apex of microsporophyll triangular, less than 1 mm long. Small tree to prostrate shrub b. var. humilis a. var. brassii Tree 3—30 m tall, up to 75 cm diam. Foliage buds 4—5 mm long, primary scales erect and up to 5 mm (or more) long. Juvenile leaves linear, 2.5—4 cm by 5—7.5 mm, acute or even apiculate, narrowing ab- ruptly at the base. Adult leaves oval, 1—2.5 cm by 4—7 mm, broadly acute, narrowing more or less ab- ruptly at the base to a 1-2 mm petiole, revolute, glaucous on the underside; midrib above a sometimes indistinct ridge 0.2 mm wide and 0.1 mm high. Pollen cone 6—7 mm diam.; apex of the micro- sporophyll lanceolate or slightly rounded at the tip, 3—4 mm long. Seed-bearing structure on a 1-9 mm peduncle; receptacle 6—9 mm with the apex of the otherwise adnate bracts spreading. Seed with its cov- ering 10-13 by 9 mm. Distr. Malesia: New Guinea (except Vogelkop Peninsula). Fig. 59. Fig. 59. Range of Podocarpus brassii P1LGER (east of line), P. brevifolius (STAPF) Foxw. (B), and P. costalis PREsL (C). Ecol. Common and sometimes dominant near the tree line, often an emergent in alpine scrub, often flat-topped, also a gnarled treelet in fire-induced grasslands or in coppices on edge of grassland, on limestone fields, rarely in mossy forest, commonest between 3000 and at least 3750 m, but also occasion- ally lower: Mt Ambua 2600 m, Ibiware 2700 m, Wissel Lakes 2000 m. Vern. East New Guinea: bacela, Kundiawa, 414 FLORA MALESIANA [ser. I, vol. 10° Chimbu, baugwa, Waimambano, Chimbu, chuga, Chimbu dial., kaibig/-tuga, ra, Kubor, Minj, kaipil, Wahgi, Minj, maja, Mairi, Mondo, tsu/o, Masul, Chimbu. b. var. humilis DE LAuB. Blumea 30 (1985) 274. Decumbent shrub to small tree up to 5 m high. Pollen cones 3.5—4.5 mm diam.; apex of the micro- sporophyll triangular, | mm long. Distr. Malesia: New Guinea (except Vogelkop Peninsula). Fig. 59. Ecol. High elevation scrub, 2600—3600 m, pros- trate in open areas, sometimes with pools, on Mt Capella forming dense mats 3 m square, 15—30 cm high. Note. Definitely not a stunted alpine form, being found at lower elevations than the other (larger) va- riety which actually occurs as a tree in the alpine scrub at or near the tree line. Only the pollen cone and the prostrate habit that often develops distin- guish the two varieties so that herbarium specimens cannot always be assigned with confidence to their proper variety. 24. Podocarpus brevifolius (STAPF) Foxw. Philip J. Sc. 6 (1911) Bot. 160, t. 29, f. 2; PrtcEr, Bot. Jahrb. 54 (1916) 40; WasscHER, Blumea 4 (1941) 466; N.E.Gray, J. Arn. Arb. 39 (1958) 441; GAussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 165, t. 801, p.p. — P. neriifolius var. brevifolius STAPF, Trans. Linn. Soc. Bot. II, 4 (1894) 249. Small, often gnarled, sometimes conical tree 2—7.5 m tall. Foliage bud 3.4—4 mm long, primary scales erect, spreading at their tips, occasionally somewhat longer than the bud itself. Juvenile leaves 5—8 cm by 6—9 mm, narrowly acute. Adult leaves elliptical, 2—5 cm by 4—6.5 mm, acute, narrowed at the base to a broad 1—2 mm petiole, slightly revolute; midrib above a narrow ridge 0.2 mm wide and 0.1 mm high. Seed-bearing structure on a 2—4 mm peduncle; foliola c. 3 mm long; receptacle 6—8 mm long. Seed with its covering 10—12 (including a small crest) by 6.5 mm. Distr. Malesia: N. Borneo (Sabah: Mt Kinaba- lu). Fig. 59. Ecol. Scattered and locally common in or under dwarf forest, (2100—)2650—3750 m, on granite rocks. Note. Reports of occurrences in various other places have all proven to be distinctly different spe- cies. 25. Podocarpus costalis PREsL, Epim. Bot. (1849) 236; Pitcer, Pfl. R. 1V, 5, Heft 18 (1903) 78; Foxw. Philip J. Sc. 6 (1911) Bot. 161, p.p.; N.E.Gray, J. Arn. Arb. 39 (1958) 456; GaussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 183, t. 831. — P. polystachyus (non R.Br.) Li & KENG, Taiwania 5 (1954) 34, t. 5; L1, Woody Fl. Taiwan (1963) 41, f. 5; GAUSSEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) (ated PE VaR op Small tree c. 1—5 m high, possibly higher. Foliage buds 2—4 mm long, primary scales erect. Juvenile leaves up to at least 9 by 1.3 cm, acute or more or less rounded at the apex. Adult leaves linear, 4—7 cm (or as little as 2.5 cm on short side branches) by 5—10 mm, broadly acute or more usually rounded at the apex, sometimes with a small blunt apiculus, narrow- ing more or less abruptly at the base to a 2—3 mm petiole, slightly revolute; midrib above a distinct ridge 0.3 mm wide and 0.2 mm high. Seed-bearing structure on a 4—6 mm peduncle; foliola c. 1.5 mm long and early caducous; receptacle 12—15 mm long, reported to be red when mature. Seed with its cover- ing 9—10 (including a small crest) by 6—7 mm. Distr. S. Taiwan (Orchid I.); in Malesia: Philip- pines (N. Luzon, on Bucas and other isles between Luzon and Taiwan and possibly on the northcoast of Luzon). Fig. 59. Ecol. Coastal bluffs near sea-level to at least 300 m. Note. Popular in cultivation in the Philippines and often confused with P. polystachyus because of a similar habitat and similar sized leaves. The leaves of P. costalis are slightly revolute while those of P. polystachyus are not. 7. Section Rumphius DE LaAvuB. Blumea 30 (1985) 274. Primary foliage bud globular, up to 4 mm long; primary scales as long as the bud, triangular, crowded together more or less into a pyramid usually with the very tip of some or all of the scales bent away from the apex of the pyramid; secondary scales acute to slightly acuminate, the second- ary bud when it first appears in most cases a spherical ball as in sect. Globulus but in P. lauben- felsii the scale tips may already be free. Leaves linear, sometimes larger than average for the genus, acute or on juvenile specimens sometimes slightly acuminate, narrowing more or less abruptly at 1988] CONIFERALES (de Laubenfels) 415 the base to a 4—16 mm petiole, stiff, mostly with a continuous upper hypoderm, midrib blunt above, at least 0.7 mm wide, broader and sometimes almost flat below, usually three vascular resin canals. Buds for pollen cones 1.5—2 mm long, either sessile or on a short peduncle, the pri- mary scales 1—1.5 mm wide at the base; secondary scales rounded, the secondary pollen cone buds when they first appear a round ball of overlapping imbricate scales. Pollen cones 2 to at least 4 cm long, usually in groups of more than three. Apex of the microsporophyll a small triangular spur 0.2 mm long over a wider base. Seed-bearing structure on a (2.5—)6—16 mm peduncle; foliola 1—1.5 mm long; receptacle mostly formed of two bracts, the longer fertile one 9-15 mm long or both fertile and equal, in P. rumphii a third lateral smaller bract often found; where known the ripe receptacle becomes red. Seed with its covering globular, at least 8 mm long, larger in P. rumphii. Distr. One very widespread species from the fringes of Asia through New Guinea and two localized spe- cies, one in northern Borneo and the other in northern Queensland. In Malesia 2 spp. Ecol. Scattered, often widely separated stands involving large, primary rain-forest canopy trees at general- ly low elevation. Note. A transitional section which shares multiple clustered pollen cones with sect. Polystachyus, but lacks the purple fruit of that section and shares the spherical developing pollen cone buds with sect. Globulus along with generally broad and blunt upperside of the midribs. The ecology and large linear leaves are some- what distinct. KEY TO THE SPECIES 1. Pollen cones sessile; foliage budscales often completely adpressed; receptacle often with a third lateral a CECMMIEN TIT ARTY. x. is. Sonat in Bisisic oak ae Oe ore wie ile Seta. vice) afore lave ualete, scale alctantetsie 26. P. rumphii 1. Pollen cone clusters usually on a small peduncle; foliage budscales always spreading at the tips; receptacle never with a third lateral sterile bract........... 26. Podocarpus rumphii BLUME, Rumphia 3 (1849) 214; Mig. FI. Ind. Bat. 2 (1859) 1073; HENKEL & HocHsTETTER, Synop. Nadelhdlz. (1865) 393; DE Boer, Conif. Arch. Ind. (1866) 15; Pari. in DC. Prod. 16, 2 (1868) 515; Becc. Malesia 1 (1877) 179; EICHLER in E. & P. Nat. Pfl. Fam. 2, 1 (1887) 104; Wars. Monsunia | (1900) 192; PmGer, Pfl. R. IV, 5, Heft 18 (1903) 81; Foxw. Philip. J. Sc. 2 (1907) Bot. 258; Merr. Interpr. Rumph. (1917) 75; Foxw. Philip. J. Sc. 6 (1911) Bot. 164; WasscHER, Blumea 4 (1941) 432; N.E.Gray, J. Arn. Arb. 39 (1958) 455; GausseNn, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 179, t. 815; pe Laus. Blumea 24 (1978) 496; Kalika- san 7 (1978) 141. — Nageia rumphii (BLUME) F.v.M. Descr. Not. | (1877) 93. — P. koordersii P1LGER ex K. & V. Bijdr. Booms. 10 (1904) 268; Koorp. Exk. Fl. Java 1 (1911) 66; K. & V. Atlas Baumart. Java 3 (1915) t. 587; Wasscuer, Blumea 4 (1941) 431; N.E.Gray, J. Arn. Arb. 39 (1958) 433; BACKER & Baku. f. Fl. Java | (1963) 90; Gaussen, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 159, t. 804. — P. phi- lippinensis Foxw. Philip. J. Sc. 6 (1911) Bot. 163, t. 30; N.E.Gray, J. Arn. Arb. 39 (1958) 434; GAUSSEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 159. Tree 12—45 m tall, up to 35—75 cm diam, Foliage buds 2.5—4 mm long, the tips of the primary scales mecting at the apex with one or two often bent out- ie) ceauccauch tear tAe hei mekeke eee 27. P. laubenfelsii ward at the tip. Juvenile leaves 19—26 by 1.9—2.6cm, acute or slightly acuminate. Adult leaves on smaller trees and lower branches of taller trees linear, 12—22 by 1.1—1.9 cm, acute or slightly acuminate, narrow- ing distinctly at the base to a 4—16 mm petiole, stiff; leaves from the exposed parts of taller trees 9-14 cm by 9-14 mm, acute, with a 4—10 mm petiole; upper side of midrib of all leaves a blunt ridge 0.7—1.2 mm wide and 0.3 mm high, often collapsing when dried to a flat surface or a small irregular ridge. Buds for pollen cones sessile. Pollen cones in groups of up to at least five, 3.5—4.5 cm long; microsporophylls somewhat elongated and tightly crowded. Receptacle frequently with a third lateral bract smaller than the other two. Seed with its covering glaucous, 12—15 by 10-12 mm. Distr. Hainan; in Malesia: Malaya (Genting Highl.), S. Central Java (Nusa Kambangan), Ba- wean |. (Java Sea), Borneo (Sabah, incl. Selangan Is.), Philippines (Luzon: Agusan del Norte), Cele- bes, Lesser Sunda Islands (Flores, Timor) Moluccas (Obi, Weda, Aru Is.), New Guinea (incl. Misool & Numfoor Is.). Fig. 60. Ecol. Locally common in primary rain-forest but rather in widely separate localities, frequently on is- lands, in Java on limestone, 5—200(—600—1550) m. Uses. Reported to be a good timber tree. 416 FLORA MALESIANA [ser. I, vol. 10° 2.5—4 mm long, the tips of the primary scales gen- erally spreading. Juvenile leaves 11—24 by 1.7—2.4 ei — cm. Adult leaves linear to linear-lanceolate, 7—19 by 1—1.8 cm, narrowly acute to acuminate, narrowing more or less abruptly at the base to a 6-14 mm * 3\~ petiole, upper side of midrib a blunt ridge 0.8—1.2 : s 0 mm wide and 0.2—0.3 mm high. Buds for pollen J : — AeU Ta cones on a peduncle to 6 mm long or less commonly f v ? 2 at FE sale sessile. Pollen cones grouped (3) 4 (5), 2—4 cm long; . ee ee On SA. microsporophylls more or less elongated. Seed with hi 3 i" its covering at least 8 mm long but fully mature ex- Fig. 60. Range of Podocarpus rumphii BLUME. amples unknown. Distr. Malesia: Borneo (Sarawak: Lawas; Sabah: Trusmadi & Kinabalu; E. Kalimantan: Vern. Borneo: kayu china, Sabah, Selangan; Kutei). Fig. 61. Philippines: malakanayan, Luzon, Agusan; Celebes: sandu, Malili; Lesser Sunda Islands: mermolas, moak, Flores, e-tama, Timor, Bunaq lang.; Moluc- cas: kayu china, rangundjela, Aru Is., P. Wokam; West New Guinea: aibemmunowame, aiwimun- wame, Fakfak, Biak lang., eswasa, weswaze, Arguni Bay, Irahutu lang., manulit, Misool I., onen, War- samson R., Mooi lang., osien, Sorong, Mooi lang., djera, si-era, Mimika, wasabraran, Numfoor, Biak lang. Note. Generally homogeneous throughout its range but the transition from large juvenile to small adult leaves mostly takes the form of long slender leaves towards the west and shorter broader leaves further east, especially in the Aru region. The Aru Fig. 61. Range of Podocarpus laubenfelsii TionG. specimens apparently lack the usual continuous up- per hypoderm as well. Ecol. Scattered in primary rain-forest, in mossy 27. Podocarpus laubenfelsii TioNnc, Blumea 29 forest, a large emergent on rocky ridge on kerangas, (1984) 523. dominant in heath forest, also in waterlogged acid Tree, 14—35 m tall, 20—60 cm diam. Foliage buds soil of Agathis forest. 600—1600 m. 8. Section Polystachyus DE LausB. Blumea 30 (1985) 275. Foliage bud on vigorous shoots mostly 2—2.5 mm diam. and on weaker shoots 1—1.5 mm diam., larger in P. macrocarpus, up to 4.5 mm long in species with more or less linear leaves and 4—9 mm long in species with distinctly lanceolate leaves; primary foliage budscales erect, some- times slightly spreading, triangular to lanceolate, up to 1.5 mm wide at the base in vigorous buds and | mm wide in weaker buds; secondary budscales more or less rounded or slightly acuminate, the secondary bud when it first appears a cluster of scale tips or when with particularly short pri- mary scales may still be more or less in a ball. Leaves linear and almost blunt to lanceolate, some- times in the same species at different stages of growth; midrib prominent above, usually less than 0.6 mm wide, broader and blunter below, usually three vascular resin canals but occasionally more (or less). Buds for pollen cones up to 3 mm long, either sessile or on a short peduncle; the secondary pollen cone buds when they first appear a round ball of overlapping scales. Pollen cones variable in length among the species but most commonly 2—4 cm long, normally in clusters of up to five; apex of the microsporophyll similarly variable in length but most commonly a small 1988] CONIFERALES (de Laubenfels) 417 spur 0.2—0.3 mm long over a wider base. Seed-bearing structure on a variable-sized peduncle, mostly 3—12 mm long but shorter in some species; foliola mostly 1.5—2 mm long; receptacle nor- mally formed of two unequal bracts with one fertile but occasionally both are fertile and equal, the fertile bracts mostly 7—12 mm long (longer in P. fasciculus), in most species known to become purple after passing through red when ripening. Seed with its covering of various sizes, globular and blunt. Distr. Central China and southern Japan through Malesia to eastern Australia, 9 spp. Most of these species occur in or near China. In Malesia 3 spp. Ecol. Scattered in (subtropical and) highland tropical forests but one species occurs on tropical sandy coastal bluffs or low elevation limestone outcrops. Note. In two species only pink ripe receptacles have been reported but in this section just as in familiar Rubus fruits, a crop of pink or red fruit is later seen to become purple or in other cases fruit bats strip off the fully ripe fruit leaving only the less ripe red, so reports of red or pink may not be conclusive. Isolated specimens may well have pollen cones in clusters of no more than 3 but a normal flush of pollen cone produc- tion will have larger clusters well represented and collectors should note this. KEY TO THE SPECIES 1. Adult leaves linear and more or less rounded at the apex, not lanceolate, upper midrib prominent. 2. Seed and its covering less than 8 mm in diameter. Midrib above sharp.......... 28. P. polystachyus 2. Seed and its covering more than 10 mm in diameter. Midrib above blunt ....... 29. P. macrocarpus 1. Adult leaves lanceolate, narrowly acute at the apex, upper midrib indistinct............ 30. P. ridleyi 28. Podocarpus polystachyus R.Br. ex ENDL. Syn. Conif. (1847) 215; Mia. Fl. Ind. Bat. 2 (1859) 1072; HENKEL & HOCHSTETTER, Synop. Nadelholz. (1865) 392; PARL. in DC. Prod. 16, 2 (1868) 515; Wars. Monsunia | (1900) 192; Pitcer, Pfl. R. IV, 5, Heft 18 (1903) 79; Merr. Philip J. Sc. 3 (1908) Bot. 394; Foxw. Philip. J. Sc. 6 (1911) Bot. 161; RipLey, FI. Mal. Pen. 5 (1925) 282, t. 228; WasscHER, Blumea 4 (1941) 456, t. 5, f. l4a—c; N.E.Gray, J. Arn. Arb. 39 (1958) 469, f. 4; KENG in Whitmore, Tree FI. Mal. 1 (1972) 49, f. 3, p.p.; GAUSSEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 191, t. 812, p.p.; DE LaAus. Kalikasan 7 (1978) 142. — P. neriifolius D.Don in Lamb., Pinus | (1824) 21, p.p.; Hook. /. Fl. Brit. In- dia 5 (1888) 649, p.p. — P. thevetiifolia BLUME, Rumphia 3 (1849) 213; Mig. Fl. Ind. Bat. 2 (1859) 1074; Henket & HocusTeTTeR, Synop. Nadelholz. (1865) 397; pe Borer, Conif. Arch. Ind. (1866) 22, t. 2; Part. in DC. Prod. 16, 2 (1868) 518; Becc. Malesia | (1877) 180; Wars. Monsunia | (1900) 192; Pitcer, Pfl. R. 1V, 5, Heft 18 (1903) 79; WaAsscHER, Blumea 4 (1941) 462; N.E.Gray, J. Arn. Arb. 39 (1958) 457; Backer & Baku. /. Fl. Java 1 (1963) 90; Gaussen, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 183, t. 829. — Nageia thevetiaefolia (BLUME) F.v.M. Descr. Not. Pap. Pl. 1 (1877) 93. — Nageia polystachyus (R.Br. ex ENpL.) O. K. Rev. Gen. PI. 2 (1891) 800. — Fig. 62. Tree 1-20 m tall, 3-45 cm diam., most commonly c. 6m. Foliage buds 1.5—3 mm long. Juvenile leaves generally within the upper range of adult leaf size, Fig. 62. Podocarpus polystachyus R.BR. ex ENDL. linear to linear-lanceolate, acute and almost Pollen cones, * 1.4 (Photogr. A.ELSENER, 1965). 418 FLORA MALESIANA [ser. I, vol. 103 apiculate, sometimes mixed with more typical adult leaves. Adult leaves linear to oval, 3—10 cm by 6—13 mm, more or less acute to rounded at the apex, nar- rowed abruptly at the base to a 1—3 mm peduncle, margin flat or nearly so; midrib above a sharp ridge 0.3—0.4 mm wide and 0.2 mm high. Buds for pollen cones sessile. Pollen cones 2—4 cm long, clustered in groups of up to at least five. Seed-bearing structure on a 1—6 mm peduncle; foliola 1—1.5 mm long, fall- ing; receptacle 7-10 mm long. Seed with its covering 7-9 by 5—7 mm. Distr. Southernmost Peninsular Thailand; in Malesia: Malay Peninsula, Riouw-Lingga & Banka Is., Borneo (W. Kalimantan: Pasir Pandjang & Karimata I.; Sarawak; Brunei; Sabah), Philippines (Palawan; Luzon: Tayabas & Ilocos Norte Prov.), Moluccas (Obi, Waigeu), West New Guinea (Vogel- kop Peninsula). Fig. 63. Fig. 63. Range of Podocarpus polystachyus R.BR. ex ENDL. Ecol. The main occurrences are at low altitudes and fall apart for the major part into three ecologies. First, the principal habitat is sandy beaches, often gregariously bordering the sea at hightide mark and sandy coastal bluffs and low outcrops, also mention- ed for sandy ridges in the mangrove. On coastal gran- ite and limestone rocks the trees are gnarled. Second, it is often frequent on lowland coastal kerangas and sandy ‘pandangs’ (degraded heath forest) and sandy heath forest (Menchali For. Res., Pahang). These two habitats are typical in the Sunda Land. Third, on limestone hills inland, for instance in Malaya and the Philippines, in Obi, Waigeu, and the Vogelkop Pen- insula in New Guinea at 180, 280 and 550 m, at 1000 m in Palawan. In East Malesia these occurrences are scattered. The bole is sometimes recorded to be fluted. The tree is found in Obi exceptionally tall, 40 m, with a clear bole of 25 m and 38 cm diam., and buttresses of 1 by 1.5 m. A most interesting ecology. Vern. Malaya: jati bukit, Selangor; Lingga: kayu karamat; Borneo: W. Kalimantan: mayu serai, Bt. Besar, tentada, Matan; Sarawak: /andin, Bintulu; Brunei: anggeriting; Sabah: kandabang, Bajau I’tan, kKayu china, Sibuboh For. Res., saumah, Manadahan; New Guinea: Vogelkop: arbudjin, Maibrat lang., rabudien, Lake Ajamaru. Note. Often cultivated (e.g. in Medan in gardens and parks) and remarkably similar to the also widely cultivated P. macrophyllus whose native range and ecology nevertheless is quite distinct. The leaves of P. macrophyllus usually have narrow but definitely revolute margins which narrow gradually towards the base while the leaves of P. polystachyus are not revolute and narrow abruptly at the base. Gray (1958) reported that the leaves of P. polystachyus have upper hypodermal fibres 70 um diam., while in P. macrophyllus these are less than 20 um. 29. Podocarpus macrocarpus DE LAuB. Kalikasan 7 (1978) 140. Tree 10—20 m tall, up to 30 cm diam. Foliage buds 2—4 mm long and the same in diameter; the longer buds with distinctly spreading upper parts of the primary scales; the secondary bud when it first ap- pears may still be a globular ball with the shorter ex- amples of primary budscales. Juvenile leaves linear- lanceolate, 8—15 by 1—1.4 cm, acute. Adult leaves linear to linear-lanceolate, 6-10 cm by 8—13 mm, acute but often slightly rounded at the apex, nar- rowed at the base more or less to a 2—4 mm petiole, thick with nearly continuous upper hypoderm and more or less shiny on the upper surface; midrib above a blunt ridge 0.5—0.6 mm wide and 0.2—0.3 mm high. Buds for pollen cones sessile. Pollen cones 2.5 cm long, grouped in clusters of up to at least four. Seed-bearing structure on a 3—12 mm peduncle; re- ceptacle 10-12 mm long. Seed with its covering 15—17 by 10—13 mm. Distr. Malesia: Philippines (northern Luzon). Fig. 64. Fig. 64. Range of Podocarpus macrocarpus DE Laus. (M) and P. ridleyi (WASSCHER) N.E.GrRay (R). Ecol. Scattered and sometimes common in cloud forests, c. 2000—2100 m. One collection reported at 700 m is doubtful. 1988] 30. Podocarpus ridleyi (WASSCHER) N.E.Gray, J. Arn. Arb. 39 (1958) 435; GaussEN, Gymn. Act. & Foss. fasc. 14, ch. 21 (1976) 159, t. 814. — P. nerii- folius var. ridleyi WASSCHER, Blumea 4 (1941) 453; KENG in Whitmore, Tree Fl. Mal. 1 (1972) 49, f. 3, p.p. Tree 4—24 m tall, 20—30 cm diam. Foliage buds 4-8 mm long. Juvenile leaves linear-lanceolate, 11—20 by 1.1—1.6 cm, narrowly acute. Adult leaves linear-lanceolate to lanceolate, 5—12 cm by 7-14 mm, acute, narrowed at the base more or less to a 2—3 mm petiole, slightly revolute, with a continuous upper hypoderm, sometimes with five vascular resin CONIFERALES (de Laubenfels) 419 canals; midrib above a low ridge 0.2—0.5 mm wide and 0.1—0.2 mm high. Buds for pollen cones sessile or on a 1 mm peduncle. Pollen cones 1.5—2 cm by 2 mm, clustered to at least four; apex of the micro- sporophyll a tiny triangular spur 0.1 mm long. Seed- bearing structure on a 3—12 mm peduncle; receptacle 8—9 mm long, known to turn pink. Seed with its covering 7 by at least 4 mm. Distr. Malesia: Malay Peninsula. Fig. 64. Ecol. Localized and more or less dominant on several isolated peaks with poor soils in a somewhat stunted rain-forest, 480—1300 m. On ridge in Panti For. Res. over standstone, on Mt Ophir on granite. ARAUCARIACEAE Monoecious, medium-sized to very large trees (rarely shrubby in very exposed situations). Either four independent cotyledons or two fused pairs (which may be retained in the seed after germination). The growing point of foliage shoots quite distinct between the two genera, being just a few highly reduced leaves in Araucaria and a highly organized bud formed of overlapping scales in A gathis. The leaves vary from scales or needles to broad leathery forms with many paral- lel veins sometimes on the same plant at different stages of growth. Pollen pro- duced in cylindrical cones from one to as much as twenty cm long with numer- ous pedunculate spirally placed microsporophylls each with several to many pendent elongated pollen sacs attached to the lower side of an enlarged shield- like apex which also projects apically more or less overlapping the adjacent microsporophylls. Pollen cones solitary, terminal or lateral, on branches separ- ate from those bearing seed cones, subtended by a cluster of more or less modi- fied leaves in the form of scales, deciduous when mature. Pollen globular, with- out ‘wings’. Seeds produced in large, well-formed cones which disintegrate when mature, dispensing the seeds in most cases with the help of wing-like struc- tures; the seed cone terminal on a robust shoot or peduncle with more or less modified leaves that change in a brief transition zone at the base of the cone into cone bracts, formed of numerous spirally-placed bract complexes, usually ma- turing in the second year. Individual seed cone bract leathery or woody and fus- ed with the fertile scale which bears one large inverted seed on its upper surface. Distribution. The 40 species in two genera are well represented in Malesia (13 spp.) and ex- tend eastward and southward into Fiji, New Caledonia (18 spp.), Australia, and New Zealand, with 2 spp. also in the cooler parts of South America, giving the family a distinct Antarctic rela- tionship. Only one species of Araucaria (in South America) occurs completely outside of the tropics, while the majority of the species in the family belong in the lowland tropics and others grow in the tropical highlands. Fossils. Early coniferous fossils are often characterized as ‘Araucarioid’ because of the mor- phological resemblance of fossil foliage shoots to certain well-known juvenile forms of Araucaria, 420 FLORA MALESIANA [ser. I, vol. 10° but there is no reason to conclude that these actually belong in Araucariaceae. Acceptable fossils of Araucariaceae, however, from Jurassic and Cretaceous age are well represented in the general areas of their modern occurrences, often at higher latitudes to be sure, and also in India and South Africa. More surprising is the apparent occurrence of fossils belonging to the family during the same time span but far away in North America and Europe (FLoRIN, 1963; GAussEN, 1970). A close relationship of English Jurassic fossils specifically to Araucaria bidwillii is indicated by StocKEY (1981). Wherever the family may have originated, it became well established in the southern hemisphere in Mesozoic times and has since disappeared from whatever northern oc- currences it may have had. The genus A gathis is first recorded in the Oligocene of Australia and New Zealand and today extends into the Asian tropics, while no convincing fossils have ever been found in any other part of the world for this genus. Tertiary records of the two modern genera of Araucariaceae are all well south of the equator across all of the southernmost land areas including the Palmer Peninsula of Antarctica. Specula- tion about tropical origins or early penetration of the tropics is not supported by any evidence and it is at least as likely that the occupation of the Malesian region took place during Plio- Pleistocene times alongside advancing members of Podocarpaceae. Maps of fossil distributions are given in FLORIN (1963). References: FLorin, Acta Horti Berg. 20 (4) (1963) 121—312, 68 maps; GAUSSEN, Gymn. Act. & Foss. fase. 11, ch. 14 (1970) 42—56, f. 555—558; MACARTHUR, The genus Araucaria in its geo- graphical aspects, Univ. W. Austr., Geogr. Lab. Econ. Dept., Research Rep. 5 (1949); StockeEy, Canad. J. Bot. 59 (1981) 1932—1940. Ecology. Canopy trees or emergents of moist forests at a wide range of elevations starting at sea-level in the tropics and extending to tropical highlands and to intermediate elevations in the middle latitudes. (One species in New Caledonia grows in drier forests and open places.) Some species clearly follow disturbances and others just as clearly do not. Many contrasting soil types are associated with different species. Most or probably all species are monoecious and pollination is by wind. Frequent reports of dioeciousness result from a single sex stage, inasmuch as many species produce seed cones well before pollen cones appear. Dense stands are common for many but by no means all species and the large amounts of pollen are adequate to reach substantial distances effectively. Seeds are car- ried only short distances by wind in most cases and germinate in large numbers near their parent tree. The occurrence of isolated specimens shows that seeds sometimes are carried across substan- tial distances. Growth occurs in distinct episodes and where distinct terminal buds are not evident whole units of growth tend eventually to be shed as a unit. Leaves normally persist for several years and may be shed separately from branch units in some cases. Cyclic growth results in false whorls of branches and a tendency for highly formal tree architecture. A number of pests and diseases specific and otherwise of Agathis have been reported by WHIT- MORE (1977) mostly outside of Malesia. In particular a moth genus, Agathiphaga, specifically at- tacks cones, while a coccid, Coniferococcus agathidis, causes defoliation. Problems mostly arise when normally dispersed trees are concentrated in plantations. GAUSSEN (1970: 62 & 66) lists a variety of pests of Araucaria, mostly associated with cultivated examples. Notable are a ‘pine bark weevil’ (Aesiotes notabilis) and the ‘hoop pine borer’ (Ca/ymmaderus). Termites of the genus Coptotermes do serious damage to A. Aunsteinii. Embryology. The fertilized egg undergoes at least five mitoses resulting in 32 or more free nuclei before cell walls form. The resulting cluster of cells deep inside the egg (proembryo) is then organized into three parts. The cells closest to the archaegonium elongate to form a massive ‘pro- suspensor’ while those on the opposite side form a temporary ‘cap’. The cells at the centre of the proembryo become the embryo proper, which does not undergo cleavage as in many other coni- fers. Simple polyembryony resulting from more than one fertilized archaegonium, however, may occur. The large number of proembryo cells and the massive embryonic ‘cap’ are distinct for 1988] CONIFERALES (de Laubenfels) 421 Araucariaceae within the conifers. The chromosome number is n= 13. No hybridization is sus- pected. Taxonomy. Two well marked genera are recognized. Uses. The large size of individuals in many species in this family along with the excellent qual- ity of the wood has made them prime candidates for lumber production where sufficiently dense stands occur. The wood is light coloured, yellowish or brownish, straight grained, easily worked, durable, and generally similar to pine but somewhat harder than the more familiar types. The wood is sometimes intermingled and hardly distinguishable from material of Podocarpaceae. Im- portant stands of Agathis have been exploited in Borneo and stands of Araucaria in New Guinea (Ismail, 1964; Gray, 1975; HAVEL, 1971), as well as many locations outside of Malesia. Heavy exploitation has reduced the economic importance of this family. Some attempts have been made to establish tree plantations, but this effort is in the early stages of development (WHITMORE, 1977, 1980; BowEN & WHITMORE, 1980). Large quantities of pitch have been gathered, particularly from certain species of Agathis where it is known as ‘dammar’. Both fossil pitch with darker colours and fresh pitch which is much lighter have been produced. Immense dammar trees sometimes have some form of steps built into their trunks to enable collectors to reach the accumulating pitch. Specimens of various species make handsome ornamentals and are widely used in landscaping in the warmer parts of the world. (The seeds of several Araucaria species are in great demand as food.) References: BOWEN & WHITMORE, Commonw. For. Rev. 59 (1980) 307; B.Gray, J. Ecol. 63 (1975) 273-289; HaveEL, J. Ecol. 59 (1971) 203-213; IsmamL BIN Hayi ALI, Mal. For. 27 (1964) 354—360; Wuitmore, A first look at Agathis, Oxford (1977); Econ. Bot. 34 (1980) 1. Note. The great size of trees in this family has led to an emphasis on collection of juvenile foliage specimens and immature cones and when this is not admitted by the collectors the result can be misleading. Because the seed cones shatter on maturity and the pollen cones are deciduous, it is next to impossible to collect attached mature fertile material. Fallen cone scales and pollen cones abound below mature trees and should be collected. KEY TO THE GENERA 1. Seed coat fused with cone-scale complex, apex of fertile scale forming a ‘ligule’, bract usually with membra- nous wings. Leaves spirally placed, crowded and broadly attached, variable in form from broad to scales SMMC SAND ISCOTIATIC 3 6 as ¢ ccc acne adie is bio «sors Shui oie Sue sada nape Nae asada ate lie 1. Araucaria 1. Seed coat independent and extended into a wing, scale completely fused with bract which is thin but not wing-like. Leaves opposite decussate, dispersed, and narrowed to a basal petiole, broad and more or less SATEEN ELISL INC o's «nse wialel a0 atnicihte bie aa ad tenee Pasay dt n PISL, casts ct One Lae Sea 2. Agathis 1. ARAUCARIA Juss. Gen. PI. (1789) 413; RicHarp, Comm. Bot. Conif. & Cycad. (1826) 153; D.Don, Trans. Linn. Soc. 18 (1841) 163; Link, Linnaea 15 (1841) 541; ENDL. Gen. Pl. Suppl. 2 (1842) 26; Syn. Conif. (1847) 184; Carr. Traité Gén. Conif. (1855) 413; Man. PI. 4 (1857) 360; Gorpon, Pinetum ed. 1 (1858) 21; HENKEL & HocusteTtTer, Synop. Nadelholz. (1865) 2; Parv. in DC. Prod. 16, 2 (1868) 369; BENTH. & Hook. Gen. PI. 3 (1880) 423; EICHLER in E. & P. Nat. Pfl. Fam. 2, 1 (1889) 67; Sewarp & Forp, Trans. R. Soc. Lond. 198 (1906) 317; BARSALI, Atti Soc. Tosc. Sci. Nat., Mem. 25 (1909) 145; DALLIMORE & JACKSON, Handb. Conif. (1923) 150; Pitcer in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 263; Gaussen, Gymn. Act. & Foss. fasc. 11, ch. 14 (1970) 7; pe Laus. Fl. Nouv. Ca- 422 FLORA MALESIANA [ser. I, vol. 103 léd. et Dép. 4 (1972) 80; SmBa, Phytologia Mem. 8 (1986) 38. — Dombeya LAMK. Enc. Méth., Bot. 2 (1786) 301, t. 828, non L’HERIT. nec Cav. — Co- lumbea SAuIsB. Trans. Linn. Soc. 8 (1807) 317. — Colymbea SPRENG. Syst. Veg. 4, 2 (1827) 888 (refers to SALISBURY, but ‘corrects’ the spelling); SteuD. Nom. Bot. ed. 2, 1 (1840) 399. — Fig. 67, 68. Monoecious or sometimes (temporarily?) dioecious small to immense trees mostly with limited and very formal branching elements consisting of long sweeping primary branches in false whorls along the main trunk often turned apically upward candelabra-like, then in most species only one additional rank of branches. The first branches sooner or later deciduous and in open situations replaced by adventitious branches thus producing a variety of double-crown forms. Apex of a resting shoot a cluster of incompletely formed leaves. Leaves spirally placed, broadly attached, crowded, multi-veined when broad and even sometimes in the needle-shaped examples, becoming uniform in size along a branch, but sometimes quite variable in the juvenile forms, amphistomatic. Pollen cones subtended by a cluster of reduced, leaf-like, sterile bracts, often broadened at their bases and where the mature leaves are needle-like these bracts are at least somewhat broader and flatter. Fertile bract of the seed cone broad and often extended laterally into membranous wings, the apex provided with a prominent narrow spur above the thickened apical margin. Seed-bearing scale only partly fused with the associated bract, its apex a free acute scale-like ‘ligule’ + reaching the base of the spur on the fertile bract. Seed coat fused with its scale. Distr. Across New Guinea, coastal Queensland, New Caledonia, Norfolk Island, S. & Central Chile, and southern Brazil 19 spp. in two sections. Fig. 65. Fossils. In Jurassic times there was evidently an important centre of development and distribution in the Inda—Australia—Antarctic region, from whence it subsequently spread to the Kerguelen and southern Cape Colony on one hand and to Patagonia on the other. The close of the Mesozoic era seems to have witnessed its disappearance from Peninsular India, South Africa and New Zealand. Both sections had an Eogene centre in Antarctica and southern South America, one of which survives still on both sides. The oldest find of Arau- caria was from probably Late Triassic in N. Central India (then situated in the southern hemisphere) where it remained until the Early Cretaceous whereupon it disappeared (FLorIN, K. Svensk. Vet. Ak. Hand. III, 19, 1940, 81, map 5). KEY TO THE SECTIONS 1. Juvenile leaves bifacially flattened, cotyledons 2, pollen cones lateral............. 1. Sect. Araucaria 1. Juvenile leaves acicular, cotyledons 4, pollen cones terminal.....................-. 2. Sect. Eutacta 1. Section Araucaria Sect. Colymbea ENDL. Gen. Pl. Suppl. 2 (1842) 26; Syn. Conif. (1847) 185; Carr. Traité Gén. Conif. (1855) 414; Man. Pl. 4 (1857) 360; Gorpon, Pinetum ed. 1 (1858) 21; HENKEL & HocusTETTER, Synop. Nadelh6lz. (1865) 2; PARL. in DC. Prod. 16, 2 (1868) 370 (‘Columbea’); EICHLER in E. & P. Nat. Pfl. Fam. 2, 1 (1889) 69; SEwarD & Forp, Trans. R. Soc. Lond. 198 (1906) 317; P1LGER 1988] CONIFERALES (de Laubenfels) 423 “ — n - ak = ae ae J Fig. 65. Range of the genus Araucaria Juss. with the number of species. in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 263; WILDE & Eamgs, Ann. Bot. n.s. 16 (1952) 44 (‘Columbea’); GAUSSEN, Gymn. Act. & Foss. fase. 11, ch. 14 (1970) 7. — Sect. Intermedia WuirtE, J. Arn. Arb. 28 (1947) 260; WILDE & Eames, Ann. Bot. n.s. 16 (1952) 44; GAUSSEN, Gymn. Act. & Foss. fasc. 11, ch. 14 (1970) 7. — Sect. Bunya WILDE & Eames, Ann. Bot. n.s. 16 (1952) 44. — Subg. Colymbea ANTOINE, Conif. nach Lambert, Loudon & Anderen (1846) 99; Carr. Traité Gén. Conif. 2 (1867) 596. Cotyledons 2, either hypogeal or epigeal, each cotyledon apparently formed of two fused units. Juvenile leaves narrow but distinctly flattened, spreading, often twisted into a horizontal plane; mature leaves broad and often with an acuminate tip. Pollen cones lateral. Ligule elongated and more or less constricted above the seed; cross section of the cone scale complex narrowed to a distinct and generally elongated neck above the seed, margins of the bract with or without broad membranous lateral wings. Distr. In E. New Guinea, coastal Queensland, S. Central Chile and S. Brazil 4 non-overlapping spp., of which | in Malesia. Ecol. Trees of moist forests mostly rising above the associated trees and often colonizing disturbed areas and protecting the regrowth into that area of other trees. Notes. Sect. Intermedia was established for Araucaria klinkii (=A. hunsteinii) because it differs from the rest of sect. Araucaria while resembling sect. Eutacta in epigeal cotyledons and broad membranous wings on the cone scales, but it does have two cotyledons and the spreading flat juvenile leaves of sect. Araucaria. One could also add that mature leaves have the hooked tip seen elsewhere only on some species of sect. Eutacta but the apex of the cone scale complex and the laterally placed pollen cones conform to sect. Araucaria, Young plants of A. Aunsteinii are indistinguishable from those of A. bidwillii. Sect. Bunya was established for A. bidwillii because the cone scale complex has thick woody wings as opposed to no wings in the two American species and because of the double vascular supply to the bract and scale, The mature seed also separates from the scale complex. Srockey (Canad. J. Bot. 59, 1981, 1932) argues for the usefulness of these monotypic sec- 424 FLORA MALESIANA [ser. I, vol. 10° tions based on a variety of fossil material, but the differences with the residual sect. Araucaria hardly seems sufficiently important. 1. Araucaria hunsteinii K.Scu. Fl. Kaiser Wilhelms Land (1889) 11, t. 4, f. 8; Wars. Monsunia | (1900) 187, t. 10, f. B; SEewARD & Forp, Trans. R. Soc. Lond. 198 (1906) 324, f. 9; BARSALI, Atti Soc. Tosc. Sci. Nat., Mem. 25 (1909) 158; WitpE & EAMES, Ann. Bot. n.s. 16 (1952) t. 2, f. 10; GAUSSEN, Gymn. Act. & Foss. fasc. 11, ch. 14 (1970) 16, f. 536; How- CROFT, For. Genet. Res. Inf. n. 8, FAO For. Occ. Pap. 1987/2 (1987) 31; StrBa, Phytologia Mem. 8 (1986) 41. — A. schumanniana Wars. Monsunia | (1900) 187, t. 10, f. A; Wipe & Eames, Ann. Bot. n.s. 16 (1952) t. 2, f. 11; GAUSsEN, Gymn. Act. & Foss. fasc. 11, ch. 14 (1970) 15, f. 535. — A. klinkii Laut. Bot. Jahrb. 50 (1914) 48, f. 1; LANE-POOLE, For. Res. Terr. Papua New Guinea (1925) 72; WILDE & EAMES, Ann. Bot. n-s. 16 (1952) t. 2, f. 7: t 3% GAUuSSEN, Gymn. Act. & Foss. fasc. 11, ch. 14 (1970) 15; f. 534 left: Forest emergent 50—89 m tall with a clear bole of 35—60 m and up to 2 m diam. Branches in loose false whorls of 5 or 6 and sometimes rising towards their apex where the leaf-bearing shoots are clustered, forming a rounded crown on the upper part of the tree. In open sites after the first branches are shed a second set of smaller adventitious branches develop on the middle part of the bole. Outer bark reddish brown, rough, peeling in horizontal strips leaving a thick dark red corky flaky underbark which weathers to shades of brown. Much colourless resin is pro- duced. Two cotyledons c. 35 by 1.5—2.5 mm at the base, tapering gradually to the narrow blunt apex, flat, their surface similar to the shorter broader acute first leaves which contain half a dozen parallel vascu- lar strands evenly spaced, while the vascular strands of the cotyledons are separated into two groups by a slight gap along the centre. Juvenile leaves linear- ovate, narrowed to a decurrent base and tapering to a slightly acuminate pungent apex, very variable in size, less than 2 by 1 mm during resting phases to at least 2.5 by 0.5 cm in the first flush of growth and be- coming larger and more lanceolate as the plant ma- tures, twisting sharply at the base to attain a hori- zontal position. Adult leaves produced in full sun- light, often in five distinct rows, narrowed slightly at the base to a broad decurrent portion 10-15 mm long, ovate-lanceolate, nearly uniform along all but the ends of the branch, 7—15 by 1.2—2 cm, an asym- metrical dorsal ridge prolonged from the junction of the two subtending leaves, ventrally concave, in- flexed at the narrow acute apex. Pollen cones cluster- ing near the ends of foliage branches each in the axil of a leaf, subtended by a cluster of reduced leaves the first few more or less decussate and up to 25 mm long but not always remaining attached when the cone falls, the mature cone linear, 16—22 by 1.8—2.5 cm. Microsporophyll on a c. 4 mm peduncle, the apical part 5—10 by 2—2.5 mm and more or less linear but narrowing to an acute apex, slightly keeled on the dorsal side, margins membranous and somewhat ser- rate, expanded at the base on the other side of the peduncle to accommodate about 10 pollen sacs. Seed cones terminal on robust short branches, subtended by numerous reduced leaves, the immature cone ovoid with only the numerous lanceolate spurs vis- ible, mature seed cones obovoid to cylindrical with the apex conical to slightly depressed, 18—25 by 12.5—16 cm, the exposed slightly expanded outer edge of each seed scale complex more or less rhom- boidal in outline, the included seed 3—4 by 0.8—1 cm imbedded in the complex with the ligule extending another 2 cm but no wider than the seed and tapering at first only slightly and then more so near the acute free apex, the thick part of the fertile bract sharply expanded above the seed to its widest and thickest part at the level of the free tip of the ligule and then forming a blunt rhomboidal end or apophysis that is visible on the surface of the mature cone and includes a narrow lateral ridge on each side and terminates in a spur 9—15 mm long which is often broken off be- fore the cone reaches maturity, the two edges of the cone bract expanded into broad blunt membranous wings each as much as 4 cm wide. Distr. Malesia: E. New Guinea, in several large stands in large valleys and numerous small stands which are often clustered and with higher mountain areas intervening between the regions of occurrence (B.GrAyY, Papua New Guinea Dept. For. Res. Bull. 1, 1973, 1—56). Fig. 66. vu ra Kt GUY f° Re Ry f ¢ i Y x ae < ‘eal a owe s y. J_d4— ny pee F aca “A 8 Nee oO | ca es ry i i am: | - } 4 “ é t | J Lai yey — \ ‘as Fig. 66. Range of Araucaria hunsteinii K.Scu. Ecol. Emergent in the submontane oak forest on well drained sites over a variety of soils from 520 to 2100 m. This is the tallest tree of Malesia. There has been much speculation concerning the origin of the disjunct gregarious stands. Natural regeneration 1988] CONIFERALES (de Laubenfels) 425 does occur under forest conditions, but most vigor- ous germination is associated with open disturbed conditions and it is generally believed that prehistoric disturbance played a major role in producing the cur- rent distribution. Heavy commercial exploitation, pressure on regeneration by feral pigs, and anthropo- genic fires have all caused serious reductions in many stands. Uses. The basis of a major plywood industry at Bulolo which results from the fine quality of the wood and the impressive log sizes. The firm, light wood is easily worked and is yellowish brown in col- our with attractive purplish streaks in the heartwood. Vern. Pa’a, Watut-Bulolo, pai, Waria-Kaisinik, gerau, Waria, bimu, Toma, yanguman, Agaun, yomejo, Kotte-Pindui, karina, Bembi-Madang, rassu, Ongoruna, nd’uk, Wareng, kembaga, saa’vara, Taiora, sowes, Erave, Mt Matmuri. Note. Howcrort (/.c.: 5, 31) has distinguished a glaucous variety that corresponds to A. klinkii. Not only are fresh leaves glaucous, but the cones are also gray-blue due to a white exudate on their surface. 2. Section Eutacta ENDL. Gen. PI. Suppl. 2 (1842) 26; Syn. Conif. (1847) 186; CARR. Traité Gén. Conif. (1855) 418; Man. Pl. 4 (1857) 361; Gorpon, Pinetum ed. | (1858) 26; HENKEL & HocusTETTER, Synop. Nadelhdlz. (1865) 9; PaRL. in DC. Prod. 16, 2 (1868) 372; EIcHLeR, in E. & P. Nat. Pfl. Fam. 2, 1 (1889) 69; SEwarp & Forp, Trans. R. Soc. Lond. 198 (1906) 318; BARSALI, Atti Soc. Tosc. Sci. Nat., Mem. 25 (1909) 157; Pmrcer in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 265; FRANCO, Port. Acta Biol. Sist. ser. B, Julio Henriques (1949) 24; WILDE & Eames, Ann. Bot. n.s. 16 (1952) 43; GaussEN, Gymn. Act. & Foss. fasc. 11, ch. 14 (1970) 7; pE Laus. Fl. Nouv. Caléd. et Dép. 4 (1972) 81. — Sect. Eutassa (SALISB.) BENTH. & Hook. Gen. Pl. 3 (1800) 437. — Eutassa SAuissB. Trans. Linn. Soc. 8 (1807) 316. — Eutacta Linx, Linnaea 15 (1841) 543 (refers to SALIs- BuRY, but ‘corrects’ the usage). — Subg. Eutacta (LINK) ANTOINE, Conif. nach Lambert, Loudon & Anderen (1846) 99; Carr. Traité Gén. Conif. ed. 2 (1867) 604. Cotyledons 4, epigeal. The first leaves following the cotyledons small elongated triangular scales with juvenile leaves appearing on lateral shoots or much later on the leader. Juvenile leaves acicular, four angled in cross section, straight or falcate and never twisted into a horizontal plane. Adult leaves acicular (to broad and concave towards the ventral side and with an asymmetrical dorsal ridge prolonged from the junction of the two subtending leaves), never with an acuminate tip. Pollen cones terminal. Ligule narrowing abruptly above the seed generally without any con- striction; the thickened apex of the bract (apophysis) directly above the apex of the seed; margins of the bract with broad membranous lateral wings. Distr. NE. coastal Australia (1 sp.), New Guinea (1 var.), Norfolk Island (1 sp.), New Caledonia (incl. Loyalty Is.) (13 spp.); in Malesia only the one variety in New Guinea. 2. Araucaria cunninghamii Air. ex D.Don in Lamb. Pinus ed. 2, 3 (1837) t. 79; Sweet, Hort. Brit. 2 (1830) 475, nomen; Lams. Pinus ed. 3 (1832) no pages, nomen; Loup. Arb. & Fruct. Brit. 4 (1838) 2443, t. 2303-2305 et suppl. 2603, f. 2545; Forses, Pin. Wob. (1839) 157, t. 52; ANrorne, Conif. nach Lambert, Loudon & Anderen (1846) 102, t. 43 & 44; Enpi. Syn. Conif. (1847) 187; Carr. Traité Gén. Conif. (1855) 419; Man. PI. 4 (1857) 361; Gorpon, Pinetum ed. | (1858) 27; HenKeL & HOCHSTETTER, Synop. Nadelhdlz. (1865) 9; Part. in DC. Prod. 16, 2 (1868) 372; SieBoLp, Flor. Jap. 2 (1870) t. 139; Sewarp & Forp, Trans. R. Soc. Lond. 198 (1906) 325, f. 8c; BARSALI, Atti Soc. Tosc. Sci. Nat., Mem. 25 (1909) 167; Wuire, J. Arn. Arb. 10 (1929) 200; ibid. 28 (1947) 259; Franco, Bot. Soc. Brot. 2, 23 (1949) 162; Gaussen, Gymn. Act. & Foss. fase. 11, ch. 14 (1970) 32; Remty, Dept. For. Qld. Res. Pap. FLORA MALESIANA [ser. I, vol. 102 Fig. 67. Araucaria cunninghamii Ait. ex D.Don var. papuana LAUT. in West New Guinea, Kebar Valley, Vogelkop Peninsula, 600 m alt. (Photogr. J.F.U.ZrEck, 1954). n. 4 (1974); StrBA, Phytologia Mem. 8 (1986) 40. — Eutacta cunninghamii (Ait.) Link, Linnaea 15 (1841) 543; Carr. Traité Gén. Conif. ed. 2 (1867) 608. — Eutassa cunninghamii Spacw, Hist. Nat. Veg. Phan. 11 (1842) 362. — Fig. 67, 69. Forest emergent, 30—60 m tall, with a clear bole of 20—40 m and up to 2 m diam. Major branches tend to be in false whorls and tend to be rather straight, growing upwards at a slight angle but gradually de- clining with weight, persisting in open growth situa- tions. Subsequent ramification more complex and denser than in any other Araucaria giving the tree the appearance of a cypress when young and a spruce when older. Outer bark at first in nearly smooth hori- zontal peeling strips or hoops which become smaller and rough with maturity, red in the interior but weathering to dark brown or black. There is a thick white resinous exudate. Cotyledons linear, c. 2 cm by 1.5—1.8 mm, narrowing to an acute apex, with sever- al evenly spaced vascular strands. Juvenile leaves straight, linear-lanceolate, pungent, bilaterally flat- tened but laterally keeled, briefly decurrent forming a sharp rib on the stem, quite variable in size being tiny at the base of a shoot, most often c. 1 by 0.1 cm, on vigorous shoots up to 2.5 by 25 cm; leaves on the leader, particularly at the seedling stage, reduced to triangular spreading bifacially flattened scales c. 2 mm long. Leaves on older plants gradually becoming 1988] CONIFERALES (de Laubenfels) 427 falcately curved forward and acicular. Adult leaves crowded and curved so that their sharply pointed api- ces are directed slightly inward, four-angled but about twice as wide as thick, the ultimate leafy bran- ches c. 5 mm in diameter with lanceolate leaves c. 5 by 1.5 mm; vigorous branches at least 1 cm in diameter with leaves 7—9 by 2 mm. Pollen cones ter- minal on foliage shoots, sometimes rather short shoots, subtended by a cluster of numerous leaf-like bracts about the same size as the leaves but distinctly thinner and more crowded, the cone 4—8 cm long and 8—10 mm in diameter, linear but tapering slight- ly to a more or less blunt apex, formed of numerous microsporophylls. Each microsporophyill on a stalk 2-3 mm long, the apex extended into a triangular flat apical part 1.2—1.8 mm long, slightly keeled on the dorsal side, margins narrow and slightly serrate, with five or more pendant pollen sacs along the base. Seed cones terminal on robust shoots with a more or less abrupt transition to the fertile scales whose apical spines are like the leaves but bent backward, the mature spine-covered cones ovoid shaped, 6—10 by 5—7 cm. The cone scales complex less the spine 23-29 mm long and including the membranous wings c. 34 mm wide, the thickened end or apophysis up to 24 mm wide with a tetragonal central part c. 5 mm thick and bearing a strong central ridge, from the upper part of the seed to the apophysis the thick- ened scale extended laterally by firm lobes; a ligule covering the seed, 7—9 mm wide, narrowing sharply above the seed apex and then elongated into a tri- angular free apex c. 2 mm long and touching the edge of the apophysis; membranous wings bluntly round- ed and c. 12 mm wide; seed completely imbedded be- tween the scale and the ligule but indicated by an almond-shaped bulge 2 by 0.7 cm and tapering to- wards the micropyle at the base of the cone scale. Distr. There are two varieties, the type in Austra- lia and the other in New Guinea. var. papuana Laut. Bot. Jahrb. 50 (1913) 51; SmBa, Phytologia Mem. 8 (1986) 40. — A. beccarii WaARB. Monsunia | (1900) 187; Gress, Arfak (1917) 83, f. 5. — A. cunninghamii auct. non Ait.: Becc. Malesia 1 (1877) 180; F.v.M. Vict. Nat. 4 (1887) 121; Descr. Not. 9 (2) (1890) 65; LANE-POoLE, For. Res. (1925) 73; Howcrort, For. Genet. Res. Inf. n. 8, FAO For. Occ. Pap. 1979/1 (1979) 9. — Fig. 67, 69. Bark, particularly of younger plants, dark plum to red-brown, weathering on older plants to gray or blackish. Juvenile leaves up to 23-27 mm long, even on ultimate branches, contrasting sharply with Aus- tralian material: where planted side by side the Aus- tralian seedlings have leaves no more than half as long. Generally reported to be slightly bigger and more vigorous (trees 50~70 m, pollen cone 9-10 cm long, seed cone 7—12 by 6—8 cm). Pollen cones pro- duced in the middle part of the mature tree and seed cones in the upper part. Distr. Malesia: New Guinea, scattered in isolated to extensive stands from one end of the island to the other, both in the central range and along the north coast, including Japen and Ferguson Is. (B.GRAY, Papua New Guinea Dept. For. Res. Bull. 1, 1973, 1—56). Fig. 68. Fig. 68. Range of Araucaria cunninghamii Att. ex D.Don var. papuana LAuT. Ecol. Emergent in rain-forests from 60—2745 m in a variety of rain-forest soils usually along ridges, but occasionally in swampy conditions, most often in the submontane oak forest. The higher elevation oc- currences are in the more southerly part of New Guinea. Vigorous regeneration has been noticed in abandoned gardens and on old burn sites confirming that A. cunninghamiiis a pioneer tree and a nurse for the invasion of rain-forest. On the other hand lum- bering, fire, pigs, and agriculture all contribute to the destruction of natural stands. Uses. The same as for A. Aunsteinii, with which it often grows. The heartwood is difficult to distin- guish from the sapwood. Vern. Pien, Pidgin, ungwa, Kapauku, sumgwa, Manikiong, a//oa, Marconi R., kiriwi, Wandammen, ningwik, Tambuni Valley, makut, Pikpik, domooi- merr, tororomooi, Dajo, jarujosuwa, Tanahmerah, flabbito, Wapi, d’li, Telefomin, escera, Foie, sari, Bembi, bontuan, Kaigorin, wariri, Gurumbu, nimola, Esa’ala. Note. Howcrort (/.c.) has noted in some areas of New Guinea trees that have more gracile foliage, particularly in the juvenile stage. These differences could well correspond to the type variety which therefore might include New Guinea within its range. While noting that there are slight differences, recent authors have chosen not to use the variety in describ- ing New Guinea material perhaps because in its origi- nal description the distinctions actually given are not valid. 428 FLORA MALESIANA [ser. I, vol. 103 Fig. 69. Araucaria cunninghamii Arr. ex D.Don var. papuana Laut. Tree, 49 m high, on slopes of Mt Cycloop above Lake Sentani in West New Guinea, 700 m (Photogr. F.W.RAPPARD, 1956). 1988] CONIFERALES (de Laubenfels) 429 2. AGATHIS SALIsB. Trans. Linn. Soc. 8 (1807) 311, t. 15, nom. cons. (unnecessarily); RICHARD, Comm. Bot. Conif. Cycad. (1826) 83, t. 19; Wars. Monsunia 1 (1900) 182; SEwARD & ForD, Trans. R. Soc. Lond. 198 (1906) 310; DALLIMORE & Jackson, Handb. Conif. (1923) 138; Pircer in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 266; MEuER Drees, Bull. Jard. Bot. Btzg III, 16 (1940) 455; FRAN- co, An. Inst. Sup. Agron. 18 (1951) 101; GaussEN, Gymn. Act. & Foss. fasc. 11, ch. 14 (1970) 75; pe Laus. Fl. Nouv. Caléd. et Dép. 4 (1972) 126; WuHitTM. Trop. For. Pap. 11 (1977) 3; Pl. Syst. Evol. 135 (1980) 46, f. 1—5; SmrBa, Phyto- logia Mem. 8 (1986) 31. — Dammara Link, Enum. Hort. Berol. Alt. 2 (1822) 411, given in synonymy with Agathis [non GAERTN. Fruct. Sem. Pl. 2 (1790) 100, t. 103, f. 1, Burseraceae]; ENDL. Syn. Conif. (1847) 188; BLuME, Rumphia 3 (1847) 211; Carr. Traité Gén. Conif. (1855) 424; Man. PI. 4 (1857) 363; Gor- DON, Pinetum ed. 1 (1858) 77; HENKEL & HOCHSTETTER, Synop. Nadelholz. (1865) 209; Part. in DC. Prod. 16, 2 (1868) 374. — Fig. 70—85. Monoecious trees often of immense size with clear straight boles below the globular crown, the large branches often turning irregularly upward; young trees with a conical shape. Bark at first quite smooth and light gray to reddish brown, peeling with large thin irregular flakes that gradually become thicker leaving a pitted somewhat rough reddish brown surface on larger trees. The two cotyledons are broad and lanceolate with an acute apex, the several vascular strands at least at first divided into two groups. Following the cotyledons the leaves are little more than triangular scales with a distinct central vein and several lateral veins. The first full leaves appear in pairs on lateral shoots. Juvenile leaves distinctly larger than adult leaves, particularly those adult leaves exposed to the sun, more or less acute, varying among the species from oval and acuminate to lanceolate. Adult leaves bluntly acute to rounded at the apex, rare- ly acuminate or lanceolate, oval to linear, sometimes lens-shaped, with con- siderable variation even along a single shoot where for example the first leaves may be substantially narrower than the later ones, generally somewhat reduced on seed cone bearing shoots, narrowed at the base to a brief broad petiole which is often twisted to place the leaves in a horizontal position, opposite decussate, decurrent, dispersed along the branch so that individual leaves do not overlap, with many parallel veins that converge no more than slightly towards the apex, resin canals alternating with the veins, more or less hypostomatic. Foliage buds globular, tightly covered with several pairs of overlapping scales. Pollen cones appearing mostly on larger trees well after the seed cones first appear, lateral and often in the axils of both of an opposite pair of leaves or occasionally ter- minal, subtended by several pairs of scales which form the sessile to briefly pedunculate pollen cone buds with the lowest pair sometimes expanded into reduced spreading leaves, more or less cylindrical with numerous small spirally placed microsporophylls. Seed cone bracts also spirally placed, their thickened 430 FLORA MALESIANA [ser. I, vol. 103 a lan Ba f ) me / )} f Hi | a iE spathulata + Cia ae i - | If il |/ { Ze wa 9 JU b j | WW, kinabaluensis lenti ZS D> = LOO ea ‘ \ fon 7) if — NII | (/( | een ee I | | | | | PAE lhs h | | ec Ul rumpfii oe g T / i owN i f By : ) | \ / \ Y oe flavescens | | Saeed ie | dii/! | | 7 gf borneensis robusta celebica k ir i labillardieri e Fig. 70. Agathis cone elements. From left to right: lateral profile of microsporophyll, facial view of micro- sporophyll (from the angle indicated by arrow in the first figure), lateral profile of seed scale, facial view of seed scale (upper seed-bearing face), and seed. Microsporophylls in mm, scales and seeds in cm. — Lateral scallops of the seed scale and shape of seed wing shown are representative; these delicate structures show con- siderable variation. Orientation of the upper edge of the seed scale varies to which part of the cone it is in. Fully developed seed scales are formed in the middle part of the cone; numerous imperfectly formed scales occur towards the cone base and apex. Seed cone scales and seeds are laterally asymmetrical and both left- handed and right-handed cones are produced. — N.B.: under h, read philippinensis instead of rumpfii. apical margin blunt or in some species with a projecting flattened ‘beak’, the lateral margins thin and broadly expanded but not membranous, normally in- dented near the base to form a ‘scallop’ which is usually much larger on one side than the other or more often one side has only a kink, quite variable especially near either end of the cone but more regular in the central fertile part, deciduous when mature. Seed scale complex fused with the bract. Inverted seed attached along its base, more or less flattened and oval-shaped, the margin on one side greatly expanded from the basal part into an oval membranous wing, the other margin blunt or more often with a rudimentary wing or sporadically the seed with two wings (cones and their elements come in both left and right handed ver- sions). Seed cone oval to spherical. 1988] CONIFERALES (de Laubenfels) 431 Fig. 71. Range of the genus Agathis SauisB. Figures above the hyphen indicate the number of endemic species, that below the hyphen the total number of species. Distr. Three sections with 21 spp. from Malaya and the Philippines across New Guinea and the coast of Queensland to Fiji and northern New Zealand; in Malesia 11 spp. There is a gap in the Solomons. Fig. 71. Fossils: Fossil wood attributed to Agathis has been found in the Upper Cretaceous and Tertiary of New Zealand and from the Jurassic to the Tertiary in Australia, as well as in the Tertiary of Western Australia. Cone scales have been found in the northern hemisphere but the identification is uncertain (FLORIN, K. Sven- ska Vet. Ak. Handl. III, 19, 1940, n. 2, 82; Acta Horti Berg. 20 (4), 1963, 180, f. 15 map). Ecol. The majority belong to lowland rain-forests. KEY TO THE SECTIONS 1. Dorsal part of the microsporophyll not at all angled. 2. Seed scale only slightly angled to completely blunt. Spp. 1-8... ....... cece eee eens 1. Sect. Agathis 2. Seed scale with a distinct beak (seed cone small, 5—6 cm diam. by 6—7 cm long). Sp. 9 2. Sect. Rostrata 1. Dorsal part of the microsporophyll sharply angled. Spp. 10-11 ............ 3. Sect. Prismobracteata 1. Section Agathis Sect. Macrobracteatae MEUER Drees, Bull. Jard. Bot. Btzg III, 16 (1940) 457. — Sect. Microbracteatae MeveER DREES, /.c. 461. FLORA MALESIANA [ser. I, vol. 103 er Lene rer i : Can aN . Ta x Fig. 72. Flaky bole of Agathis borneensis WarB. in heath forest on podsolized white sandy terrace, c. 20 m alt., Brunei (Photogr. P.S.AsHToN, May 1959). 1988] CONIFERALES (de Laubenfels) 433 Large trees. Pollen cones with spoon-shaped microsporophylls without angled creases, rarely sessile. Seed cones in most cases at least 7 cm long and the seed bracts always blunt along their apical margins. Distr. In the same territory as the genus less the more southerly areas 13 spp., of which 8 in Malesia. KEY TO THE SPECIES 1. Juvenile (and more accessible) leaves not at all acuminate, adult leaves at least 6 cm long or else glaucous on the underside. 2. Leaves not glaucous on the underside, at least 6 cm long. Pollen cones at least 4 cm long. Microsporophylls slightly acute and nearly as long as wide or large (over 5 mm long). 3. Pollen cone at least 2 cm in diameter. Microsporophyll c. 6 mm wide, over 5 mm long, and blunt, resin MRED AMOS ED AIL OO 61h MR oe MEF 5 os atts oxidised ee nd abe renee, oe ie 1. A. borneensis 3. Pollen cone more than 14 mm in diameter. Microsporophyll no more than 2.5 mm wide, 2 mm long, and slightly acute, resin canals in the leaves solitary. 4. Adult leaves never broadly rounded at the apex. Pollen cones 12—14 mm in diameter. Apex of micro- sporophyll at least 2 mm long and wide. Seed bract at least 42 mm wide and 32 mm high with more or less straight margins and the apex bluntly ridged ....................0.-.0000- 2. A. celebica 4. Adult leaves acute to broadly rounded at the apex. Pollen cones 9—12 mm in diameter. Apex of micro- sporophyll less than 2 mm wide and 1.5 mm long. Seed bract less than 42 mm wide and 32 mm high with more or less rounded margins and the apex sharply rigid.................. 3. A. spathulata 2. Leaves glaucous on the underside, no more than 6 cm long. Pollen cones less than 4 cm long. Microsporo- phylls blunt (much wider than long), less than 2 mm long. 5. Adult leaves lens-shaped, S—7 cm long. Pollen cones 3—4 cm by 9-10 mm. Microsporophyll 2—2.5 mm ides Seed cone'c. 6m: in'diameter, spherical ........ Ma. gos. cede cece ec ee ees 4. A. lenticula 5. Adult leaves orbicular, blunt, 2.4—4 cm long. Pollen cones 8—14 by 4—6 mm. Microsporophyll 1.2—1.5 poe seed cone ci 4:5, cmin diameter) elongated 2. |... sawetrerncu oe ees 5. A. orbicula 1. Juvenile leaves distinctly acuminate. Adult leaves less than 6 cm long and not glaucous on the underside. 6. Adult leaves not acuminate. Microsporophyll helmet-shaped, the stalk attached near the centre of the apical part, at least 2 mm wide. Apex of the seed cone bract bluntly ridged. 7. Seed sharply angled opposite the wing. Mature pollen cone mostly at least 3.5 cm long by 10 mm in dia- meter. Seed cone bract 2.8—3.2 cm high. Adult leaves at least 4.5cm ......... 6. A. philippinensis 7. Seed bluntly rounded opposite the wing. Mature pollen cone 2—3.5 cm long by 7-10 mm in diameter. Seed cone bract 2.6—2.9 cm high. Adult leaves 3—4 cm long.................005: 7. A. flavescens 6. Adult leaves acuminate or small and blunt. Microsporophyll shingle-shaped, the stalk attached below the centre of the apical part, 1.6—1.8 mm wide. Apex of the seed cone bract sharply ridged 1. Agathis borneensis Wars. Monsunia | (1900) 184, t. 80; DaLttimore & JAcKsoNn, Handb. Conif. (1923) 143; Mever Drees, Bull. Jard. Bot. Btzg III, 16 (1940) 459; Gaussen, Gymn. Act. & Foss. fasc. 11, ch. 14 (1970) 96, t. 573; pe Laus. Blumea 25 (1979) $32, t. 1; Wuitm. Pl. Syst. Evol. 135 (1980) 54, t. 1 f.3,t.2.f.3,1.4; VeLDKAmp & DE Laus. Taxon 33 (1984) 345; Sirsa, Phytologia Mem. 8 (1986) 32. — Pinus dammara Lams. Descr. Pinus | (1803) 61, t. 38 (& 38a), nom. rej.; VELDKAMP & DE Laus. Taxon 33 (1984) 337. — A. loranthifolia SAuiss. Trans. Linn. Soc. Lond. 8 (1807) 312, t. 15, nom. superfl. — Dammara loranthifolia (SAuiss.) Link, Enum. Hort. Berol. Alt. 2 (1822) 411. — Dammara orientalis Lamp. Descr. Pinus 2 (1824) 15, nom. superfl.; Gor- pon, Pinetum ed. | (1858) 79. — A. dammara (Lams.) RicHarp, Comm. Bot. Conif. & Cycad. (1826) 83, t. 19. — Abies dammara (LAmB.) Desr. 8. A. kinabaluensis Tabl. Ecol. Bot. ed. 3 (1829) 356. — Dammara orien- talis var. orientalis CARR. Traité Gén. Conif. (1855) 426; HENKEL & HOCHSTETTER, Synop. Nadelhdlz. (1865) 210. — A. beccarii Wars. Monsunia | (1900) 184, t. 8F; DALLIMORE & JACKSON, Handb. Conif. (1923) 142; Mever Drees, Bull. Jard. Bot. Btzg III, 16 (1940) 458, f. 1. — A. macrostachys Wars. Mon- sunia | (1900) 183, t. 8A. — A. rhomboidales Wars. Lc. 184, t. 8C; Mever Drees, Bull. Jard. Bot. Btzg Ill, 16 (1940) 460; Harrison in Dallimore & Jack- son, Handb. Conif. ed. 4 (1966) 103. — A. alba Foxw. Philip. J. Sc. 4 (1909) Bot. 442. — A. latifolia Meuver Drees, Bull. Jard. Bot. Btzg III, 16 (1940) 459. — A. dammara ssp. dammara Wut. P1. Syst. Evol. 135 (1980) 56 (WuHiTMoreE described A. celebica and A. philippinensis). — Fig. 72, 73. Huge tree to 55 m tall. Juvenile leaves ovate- lanceolate, up to 14 by 4cm. Adult leaves ovate with 434 FLORA MALESIANA [ser. I, vol. 103 Fig. 73. Agathis borneensis WaARB. a. Mature foliage shoot; b. leaf variation of shaded branches or alter- nating with a; c. juvenile leaf; d. young seed cone; e. mature pollen cone, all x0.4; f. profile view of microsporophyll, «1.25; g. seed; h. end view and facial view of the seed cone scale; i. profile view of the upper edge of the same. a more or less acute apex, 6—12 cm by 20—35 mm, tapering at the base to ac. 5 mm petiole. The most common foliage leaf c. 7 by 3 cm but branches with relatively long and narrow leaves often interspersed with the more usual type and more general on younger trees. Resin ducts in pairs one above the other between most vascular strands rather than the prevailing solitary duct elsewhere in the genus. Mature pollen cones oblong, 4—7 cm by 20—25 mm, rounded at the apex, subtended by a 2—10 mm ped- uncle, the apex of the microsporophyll spoon- shaped, 5.5—6.5 by 4—5 mm, the apex a broad semi- circle. Mature seed cones oval, 6—8.5 by 5.5—6.5 cm. Seed bract roughly triangular but well rounded at the upper corners, a low thick ridge along the apical mar- gin, a strongly hooked 6 mm scallop on one side of the base, the other side with no more than a kink, 26—28 by 36—40 mm. Seed c. 12 by 9 mm, blunt at one upper corner and a broadly rounded wing c. 20 by 16 mm at the other corner. Distr. Malesia: throughout Borneo and more re- stricted areas in Malaya and N. Sumatra. Fig. 74. Fig. 74. Range of Agathis borneensis WARB. Ecol. Scattered in upland rain-forest from low elevations to 1200 m throughout its range and in dense nearly pure stands on low-lying sandy peat soil in many parts of Borneo and in one area in Malaya. It is of interest to note that Dacrydium pectinatum has a similar ecology. Vern. Malaya: dammar, dammar daging, dam- mar minyak, tulong, M; Sumatra: hedje, Tapanuli; Borneo: bindang, Sarawak, bulu, Iban, salang, Ke- dayah, tambunan, Sabah, manggilam, Dusun, bangalan, bengalan, Sampit, Pilau, toga, W. Kutai, bembuéng, SE. Borneo, nuju, Dajak, enghatan, Sanggan, pisau, putut, Sintang. Note. If the proposal to reject Pinus dammara Lame. in favour of Agathis borneensis WARB. is not accepted, the proper name of this species would be Agathis dammara (LAMB.) RICHARD, a name hereto- fore (incorrectly) associated with different species of Celebes, the Moluccas and the Philippines (see under A. celebica and A. philippinensis). If various closely related species were combined as varieties or sub- species under this species, there would be no reason to reject the name Agathis dammara. 2. Agathis celebica (KooRD.) Wars. Monsunia 1 (1900) 185; Dattmore & Jackson, Handb. Conif. 1988] (1923) 143; MEER DrREEs, Bull. Jard. Bot. Btzg III, 16 (1940) 461; pE Laus. Kalikasan 7 (1978) 146; Blu- mea 24 (1978) 504, f. 2; SmBa, Phytologia Mem. 8 (1986) 32. — Dammara alba RumPH. ex Hassk. Tijd. Nat. Gesch. Phys. 9 (1842) 179; Part. in DC. Prod. 16, 2 (1868) 374. — Dammara rumphii PREsL, Epim. Bot. (1851) 236, nom. superfl. — Dammara orien- talis var. pallens CARR. Traité Gén. Conif. (1855) 426. — Dammara orientalis var. alba KNIGHT ex HENKEL & HOCHSTETTER, Synop. Nadelhdlz. (1865) 211. — Dammara alba var. alba Hassk. Abh. Naturf. Ges. Halle 9 (1866) 180. — Dammara alba var. celebica Hassk. /.c. — Dammara celebica Koorp. Meded. Lands PI. Tuin 19 (1898) 263. — A. dammara auct. non RICHARD: WARB. Monsunia 1 (1900) 182, t. 9, f. 1; HARRISON in Dallimore & Jack- son, Handb. Conif. ed. 4 (1966) 98; DE Laus. Blumea 24 (1978) 503, f. 1. — A. alba auct. non Foxw.: MERR. Rumph. Herb. Amb. (1917) 76; MEu- ER Drees, Bull. Jard. Bot. Btzg III, 16 (1940) 466; DALLIMORE & JACKSON, Handb. Conif. ed. 3 (1948) 178. — A. beckingii MEIER DREES, Bull. Jard. Bot. Btzg III, 16 (1940) 463. — A. hamii MEIER DREEs, lc. 462, f. 1. — A. loranthifolia auct. non SALIsB.: MEueER Drees, /.c. 464. — A. celebica ssp. celebica VELDKAMP & WuHiTM. in Veldkamp & De Laub. Taxon 33 (1984) 345. — A. dammara ssp. dammara auct. non WuHiTM.: WuHITM. PI. Syst. Evol. 135 (1980) 57, p.p. Huge tree to 65 m tall. Juvenile leaves ovate- lanceolate, up to 15 by 4.6 cm. Shade leaves on ma- ture trees acute and roughly 9 by 3 cm. Leaves from fully exposed branches well rounded at the apex but still tapering, not blunt, 6—8 by 2—3 cm, tapering at the base to a S—10 mm petiole. Pollen cones after shedding pollen 4—6 by 1.2—1.4 cm or possibly even larger, subtended by a short peduncle c. 3 mm long, normally axillary but sometimes terminal (A. beck- ingii). The apical part of the microsporophyll spoon- shaped, spreading, with the stalk attached well behind its centre, c. 2.5 mm by 2 mm and slightly angled at the apex. Seed cone oval, 9—10.5 by 7.5—9.5 cm. Seed bract with a low thick ridge along the apical margin exposed in the unopened cone to within a few mm of its lateral margins, roughly tri- angular in shape, the lateral margins nearly straight with a small scallop 4—8 mm above the base on one side, the upper corners more or less angular and rigid, 32—36 by 42—45 mm. Seed c. 14 by 9 mm with a short acute projection on one upper corner and a broad rounded wing c. 24 by 16 mm at the other. Distr. Malesia: Celebes and Moluccas to Pala- wan in the Philippines and probably other parts of southern Philippines. Fig. 75. Ecol. A forest emergent scattered and locally common in lowland rain-forest from near sea-level to 1200 m. CONIFERALES (de Laubenfels) 435 =] Fig. 75. Range of Agathis celebica (KOORD.) WARB. Vern. Dammar, dammar radja, M, hulontuu, Malili, Kawo, Maliki, kisi, Buru, sa/o, Ternate, dayungon, Samar, Philippines. Note. WHITMORE (1980) insists that he can find no difference between this and A. philippinensis among others and certainly the vast majority of the collected materials (shade leaves and immature re- productive organs) is enigmatic. WHITMORE’s de- scriptions show he is dealing with immature material and the notion that pollen cones continue growth after shedding their pollen is inadmissible. Groups of both species grow side by side at Bogor, where prop- erly mature material can be compared and sharp dif- ferences easily seen. Because this is the only species known on Ambon, it is the species which RUMPHIUS (Herb. Amb, 2, 1741, 174, t. 57) has in mind and, like the closely related A. borneensis, which occupies similar habitats west of the Makassar Strait, is an im- portant producer of the resin known as dammar. 3. Agathis spathulata DE Laus., sp. nov.; SILBA, Phytologia Mem. 8 (1986) 37, nom. inval. — A. 436 FLORA MALESIANA robusta ssp. nesophila Wuitm. Pl. Syst. Evol. 135 (1980) 64. Arbor magna ad 60 m alta. Folia adulta acuta 9—10 cmx20—30 mm ad spathulata 7-9 cmx 18—20 mm. Strobili masculini 9-13 mm diametri, 4—7 cm longi, apicis microsporophyllis 1,5—2 mm latis, 1,3—1,5 mm longis. Squamae femineae 35—42 mm latae, 27—32 mm altae, marginibus rotundibus apicis costis acutis. Holotypus: DE LAUBENFELS P74] (12): Huge tree to 60 m tall. Juvenile leaves ovate- lanceolate, not at all acuminate, up to 13 by 4.5 cm. Adult leaves bluntly acute, 9-10 cm by 20—30 mm to broadly rounded and 7—9 cm by 18—20 mm, tapering at the base to a 5-10 mm petiole. Mature pollen cones 4—7 cm by 9—13 mm, subtended by a 2—9 mm peduncle, the apex of the microsporophyll spoon- shaped without ridges, 1.5—2 by 1.3—1.5 mm, broad- ly acute. Seed cone oval, 8.5—10 by 6.5—7.5 cm. Seed bract with a thin sharp ridge along the apical margin, broadly rounded at the upper corners, a small 6 mm scallop on one side of the base and a larger 10 mm scallop on the other, 27—32 mm high by 35—42 mm broad. Seed c. 10 by 6 mm, with a sharp projection on one upper corner and an elongated bent wing c. 24 by 10 mm at its widest at the other corner. Distr. Malesia: SE. New Guinea, E. Highland, near Obura. Fig. 76. Fig. 76. Range of Agathis spathulata DE LAvB. (squares) and A. /abillardieri WarRB. (dots). Ecol. Scattered as a rain-forest emergent or sur- viving in small exposed groves between 900 and 1980 m. Vern. Asong, muwaka, ogapa. Note. The seed cone scale and seed, though smaller, resemble those of A. robusta, a lowland spe- cies, but the pollen cone in particular is quite differ- ent. 4. Agathis lenticula pe Laus. Blumea 25 (1979) 537, f. 4; SirBa, Phytologia Mem. 8 (1986) 34. — Fig. 77. Large tree to 45 m tall. Juvenile leaves to 11 cm by 47 mm, ovate, tapering to an acute, scarcely acumi- [ser. I, vol. 10° Fig. 77. Agathis lenticula DE LAuB. a. Mature foliage shoot; b. extreme leaf shape; c. juvenile leaf; d. mature pollen cone, all 0.4; e. profile view of microsporophyll, x 1.25; f. seed; g. end view and fa- cial view of the seed cone scale; h. profile view of the upper edge of the same. nate apex and more sharply at the base. Adult leaves lens-shaped, more or less acute, 5—7 cm by 18—24 mm, tapering to a 3—7 mm petiole, glaucous on the underside. Mature pollen cones 3—4 cm by 9-10 mm, subtended by a 2—6 mm peduncle, the apex of the microsporophyll spoon-shaped and spreading, 2—2.5 by 1.5—2 mm, apex blunt. Seed cone nearly spherical, c. 7 by 6 cm. Seed bract with a thin sharp ridge along the apical margin, lateral margins broad- ly rounded with a strongly hooked scallop 7 mm above the base on one side, c. 27 mm high by 38—45 mm broad. Seed c. 11 by 7 mm with a slight blunt to no projection on one upper corner and a broadly rounded wing c. 8 by 14 mm at the other corner. Distr. Malesia: Sabah (Mt Kinabalu and the Crocker Range). Fig. 78. Ecol. Emergent in 1140—1680 m. Vern. Tanggilan, tengilan, tungilan, Dusun. Note. This species occurs just below A. kinaba- /uensis on Mt Kinabalu where the two can easily be compared in the field. The distinctive leaf shapes are readily apparent but in herbarium specimens the mountain rain-forest, 1988] ( Fig. 78. Range of Agathis lenticula DE LAvB. (circles) and A. orbicula pe Laus. (dots). glaucous leaf undersurface is generally not detect- able. Other differences are the larger pollen cones with blunt, not angled, microsporophylls and the lack of a sharp projection at the upper corner of the seed. Besides A. orbicula and A. endertii, other Agathis species with glaucous leaf undersides occur far to the east beyond Malesia. 5. Agathis orbicula DE Laus. Blumea 25 (1979) 540, f. 5; Smpa, Phytologia Mem. 8 (1986) 36. — Fig. 79. Tree to 40 m tall. A light yellow resin is produced in some abundance. Juvenile leaves ovate and blunt- ly acute, to 6.5 cm by 28 mm. Adult leaves ovate to orbicular, broadly rounded to slightly angled at the apex, 24—40 by 12—24 mm, tapering sharply at the base to a 3—7 mm petiole, glaucous on the underside. Mature pollen cones 8—14 by 4—6 mm, subtended by a 2—6 mm peduncle, the apex of the microsporophyll helmet-shaped, 1.2—1.5 by 1—1.2 mm, apex blunt. Seed cone oval, c. 7 by 4.5 cm. Seed bract with an acute ridge along the apical margin, ovate with a 4 mm scallop spreading nearly perpendicularly to each side of the base, c. 20 mm high and 33 mm broad. Seed unknown but leaving a blunt impression sug- gesting a shape like that of A. /enticula. Distr. Malesia: S. Sabah to Central Sarawak. Fig. 78. Ecol. Scattered in rain-forests and kerangas on low mountains and plateaus between 450 and 1050 m. Vern. Iban. Tumuh, Murut, tubu, Kenyah, bulok, 6. Agathis philippinensis Ware. Monsunia | (1900) 185, t. 8E; Dattmmore & Jackson, Handb. Conif. (1923) 147; Mever Drees, Bull. Jard. Bot. Btzg III, 16 (1940) 468; SirBa, Phytologia Mem. 8 (1986) 36. — Dammara rumpfii auct. non Presi: Presi, Epim. Bot. (1841) 236. — A. regia Wars. Monsunia | (1900) 183, t. 8B; Dattimore & Jackson, Handb. CONIFERALES (de Laubenfels) 437 Fig. 79. Agathis orbicula DE LauB. a. Mature foliage shoots; b. juvenile leaf; c. young seed cone; d. ma- ture pollen cone, all x 0.4; e. profile view of micro- sporophyll, x 1.4; f. end view and facial view of the seed cone scale; g. profile view of the upper edge of the same. Conif. (1923) 147. — A. alba auct. non Foxw.: Foxw. Philip. J. Sc. 4 (1909) Bot. 442. — A. dam- mara auct. non RICHARD: DE LAuB. Kalikasan 7 (1978) 144; Blumea 24 (1979) 499, f. 1. — A. dam- mara ssp. dammara auct. non WHITM.: WuITM. PI. Syst. Ecol. 135 (1980) 56, p.p., t. 5, f. 1 & 2. Huge tree to 60 m tall. Abundant white resin pro- duced. Juvenile leaves ovate and distinctly acumi- nate, to 7 by 3 cm. Adult leaves ovate, very slightly to distinctly acute, 4—6 by 1.5—2 cm, the smaller leaves, which probably derive from more exposed position, being the least acute, tapering at the base to a 5—8 mm petiole. Mature pollen cones 2.5—4.5 cm by 10-11 mm, subtended by a short peduncle, the apex of the microsporophyll helmet-shaped with the stalk attached close to its centre, 2—2.5 by 1.5—2 mm, the apex very slightly angled. Seed cone oval, 7-9 by 12 cm. Seed bract with a low thick ridge along apical margin, broadly rounded at the upper corners, a small 3—6 mm scallop on one side of the base, 28— 32 mm high by 35—45 mm broad. Seed c. 11 by 6 mm, broadly acute at one upper corner and with a wing c. 20 by 11 mm at its widest at the other corner. FLORA MALESIANA pests : a be 0 Fig. 80. Range of Agathis philippinensis WaRB. Distr. Malesia: Philippines to Celebes and Hal- maheira. Fig. 80. Ecol. Scattered and often emergent in upland rain-forest, mostly 1200-2200 m, occasionally as low as 450 m in Halmaheira, 900 m in Celebes and far to the north in Luzon to 250 m. Vern. Dammar, M, goga, solo, tjoga, Manado, molewaun, Teliwang, almaciga, Philippines, dayungon, Samar, dingan, Misamis. Note. Reports by foresters of ‘different’ trees in the southern parts of the Philippines may well apply to A. celebica which has been collected there. Most Philippine collections are of nondescript leaves and (rarely) immature cones. 7. Agathis flavescens RipLey, Kew Bull. (1914) 332; J. Fed. Mal. St. Mus. 6 (1915) 3; DatimmorE & Jackson, Handb. Conif. (1923) 143; MEVER DREEs, Bull. Jard. Bot. Btzg III, 16 (1940) 464; Sipa, Phytologia Mem. 8 (1986) 33. — A. dammara ssp. flavescens (RIDLEY) WuitM. Pl. Syst. Evol. 135 [ser. I, vol. 103 (1980) 59. — A. celebica ssp. flavescens (RIDLEY) VELDKAMP & WHITM. ex VELDKAMP & DE LAUB. Taxon 33 (1984) 346. Small to medium-sized tree to 12 m tall. Juvenile leaves ovate-lanceolate and slightly acuminate, up to 8 cm by 32 mm. Adult leaves ovate and, unlike most adult Agathis leaves, often wider before the middle, 3—4cm by 10—20 mm, rounded and blunt at the apex or very slightly angled, tapering at the base to a 3—5 mm petiole. Mature pollen cones 20—35 by 8—9 mm, subtended by a 2—8 mm peduncle, the apex of the microsporophyll helmet-shaped, 2 by 1.8 mm, the apex slightly angled. Seed cone oval, 6—7 by 7—8 cm. Seed bract with a thick ridge along the apical margin, broadly rounded at the upper corners, a large scallop more than one cm from the base on one side and a smaller c. 6 mm scallop on the other, 26—29 by 35—37 mm. Seed oval, c. 11 by 9 mm, blunt at one upper corner and with a broad wing 17 by 14 mm at its widest at the other corner. Distr. Malesia: the two or three highest peaks in Peninsular Malaya. Ecol. Scattered trees above a low summit scrub, 1200—1800 m, the leaves often yellow, suggesting adaptation to difficult nutrient conditions. AC f Fig. 81. Agathis kinabaluensis DE LAvuB. a. Mature foliage shoot; b. extreme leaf shape; c. juvenile leaf; d. mature pollen cone; all «0.4; e. profile view of microsporophyll, x 1.25; f. seed; g. end view and fa- cial view of the seed cone scale; h. profile of the up- per edge of the same. 1988] Note. This species corresponds in Malaya to A. philippinensis which it strongly resembles, differing in smaller cones, somewhat differently shaped leaves and seeds and especially by the blunt corner of the seed opposite the wing. 8. Agathis kinabaluensis DE LAusB. Blumea 25 (1979) 535; Sma, Phytologia Mem. 8 (1986) 34. — Fig. 81. Small to large tree to 36 m tall. Juvenile leaves ovate and strongly acuminate, to 9 cm by 44 mm. Adult leaves ovate, slightly acuminate or occa- sionally on the smaller leaves round and blunt, 3.5—7 cm by 18—32 mm, tapering at the base to a 4—7 mm petiole. Mature pollen cones 18—30 by 8—10 mm, nearly sessile to a4 mm peduncle at the base, the apex of the microsporophyll spoon-shaped, spreading slightly, 1.6—1.8 by 1.2—1.6 mm, the apex very CONIFERALES (de Laubenfels) 439 slightly angled. Seed cone oval, 7.5—8.5 by 11 cm. Seed bract with a prominent narrow ridge along the apical margin, broadly rounded at the upper corners, a smaller 4 mm and a larger 6 mm scallop on either side of the base, 28—32 by 40—45 mm. Seed c. 11 by 7 mm, prominently acute at one upper corner and with a broad wing 20 by at least 12 mm at the other corner. Distr. Malesia: Mt Kinabalu in Sabah. Ecol. Inor rising slightly above the mossy forest and rather small where the forest is stunted, 1500—2400 m. Vern. Tumu, Murut. Note. This species in Borneo corresponds in many ways with A. philippinensis but is nevertheless substantially different. 2. Section Rostrata DE LAUB., sect. nov. Arboris. Squama feminea projectioni apici instructa. Strobilus femineus 5—6 X 6—7 cm. Strobilis masculinus sessilus. Typus: Agathis australis. Small to large trees. Leaves ovate and slightly angled at the apex to round and blunt, the leaf undersides distinctly glaucous, solitary resin canals between the vascular bundles. Juvenile leaves more acute. Pollen cones more or less sessile. Microsporophylls spoon-shaped, more or less acute and spreading. Seed cones spherical to oval and rather small, S—6 by 6—7 cm. Seed bract with a distinct projection or beak near the centre of the ridged apical margin. Seeds broadly oval with a blunt projection at one upper corner and a broad wing at the other corner. Distr. There are 3 spp., widely separated, in New Zealand (1 sp.), in New Caledonia (1), and in Malesia: Borneo (1). Note. The projection on the seed bract corresponds to the ligule in Araucaria and other conifers and for Agathis is a primitive trait. 9. Agathis endertii Mever Drees, Bull. Jard. Bot. Btzg III, 16 (1940) 470; Harrison in Dallimore & Jackson, Handb. Conif. ed. 4 (1966) 99; pe LaAus. Blumea 25 (1979) 534, f. 2; Sirtpa, Phytologia Mem. 8 (1986) 33. — Fig. 82. Tree to at least 48 m. Juvenile leaves apparently lanceolate. Adult leaves ovate and broadly acute to semicircular and blunt at the apex, 5—8 cm by 17—36 mm with considerable variation on individual speci- mens, tapering to a 3—6 mm petiole, glaucous on the underside. Mature pollen cones 26-38 by 7 mm, sessile, the apex of the microsporophyll spoon- shaped, spreading, 2.5 by 2 mm, the apex slightly angled. Seed cone oval, 4.5 by 7 cm. Seed bract with a prominent narrow ridge along the apical margin surmounted by a spreading blunt more or less trian- gular projection c. 8 by 3 mm, somewhat rounded at the upper corners, a small 4 mm scallop on one side at the base, at least 25 mm by 30 mm. Seed c. 11 by 8 mm, a short acute projection on One upper corner, a broad wing c. 18 by 14 mm at its widest at the other corner. Distr. Malesia: Borneo. Fig. 83. Ecol. In more or less isolated populations from near sea-level to 1440 m, often associated with sand- stone kerangas. Vern. Bulok, Iban. Note. The leaves more or less resemble in par- ticular A. borneensis (but lack the paired resin canals) so that sterile specimens cannot be readily identified in this otherwise quite distinct species. 440 FLORA MALESIANA [ser. I, vol. 10° Fig. 83. Range of Agathis endertii MEWER DREES. <€—- Fig. 82. Agathis endertii MEIJER DREES. a. Mature foliage shoot; b. leaf variation; c. juvenile leaf; d. young seed cone; e. mature pollen cone, all x 0.4; f. profile view of microsporophyll, x 1.25; g. seed; h. end view and facial view of the seed cone scale; i. pro- file view of the upper edge of the same. 3. Section Prismatobracteatae MEER Drees, Bull. Jard. Bot. Btzg III, 16 (1940) 470. Large trees. Microsporophylls with sharp creases dividing the apical part into three or more fa- cies. Pollen cones with a short peduncle. Seed bracts always blunt along their apical margins. Leaves not glaucous underneath. Distr. From New Guinea and New Britain to Queensland, New Caledonia and Vanuatu (New Hebrides) 5 spp.; in Malesia: 2 spp. KEY TO THE SPECIES 1. Apical part of pollen cone prismatic with a series of lateral facies surrounding a flat hexagonal upper face ; 10. A. labillardieri 1. Apical part of pollen cone divided into three lateral facies...............00eeeeeeeee 11. A. robusta 1988] CONIFERALES (de Laubenfels) 44] Y Fig. 84. Agathis labillardieri Warn. a. Iwig with pollen cones; b. seed cone, both «0.4, 10. Agathis labillardieri Wars. Monsunia | (1900) 183; MEWER Drees, Bull. Jard. Bot. Btzg III, 16 (1940) 471; Harrison in Dallimore & Jackson, Handb. Conif. ed. 4 (1966) 99; Wuirm. Pl. Syst. Evol. 235 (1980) 60; SitBa, Phytologia Mem. 8 (1986) 34. — Fig. 84, 85. Huge tree to 60 m. Juvenile leaves ovate and acu- minate, to 10 by 6 cm. Adult leaves ovate to oval lan- ceolate, acute, 6-9 cm by 20—24 mn, narrowing to a 5—7 mm petiole. Mature pollen cones 25—35 by 10-15 mm, subtended by a 2—6 mm peduncle, the apical part of the microsporophyll prismatic with a series of lateral facies surrounding a flat hexagonal upper face 1—1.5 mm wide and long and crowded so that only the upper face is exposed. Seed cone oval, 8.5—10 by 7.5—9 cm. Seed bract with a low thick Fig. 85. Cones of Agathis labillardieri Wars. of a specimen growing in Sarmi on the north coast of West New Guinea (Photogr. H.R.KARSTEL, 1957), ridge along the apical margin exposed in the un- opened cone to within a few mm of its margins, roughly triangular in shape with nearly straight later- al margins, with a distinct scallop c. 8 mm above the base on one side, slightly rounded at the upper cor- ners, 30-32 by 38—42 mm. Seed c. 12 by 7 mm with a short broadly acute projection on one upper corner and a broad rounded wing c. 20 by 15 mm at the other. 442 FLORA MALESIANA [ser. I, vol. 103 Distr. Malesia: throughout the western part of New Guinea and eastward to the margins of the Sepik Valley. Fig. 76. Ecol. From near sea-level to 1350 m or occasion- ally to 1800 m, often on ultrabasic soil and locally very common. Vern. Dammar, M, pen, Pidgin, /egatulus, Mooi lang., aisjier, Arguni, idjir, Irahutu, ¢ar, Kebar, waiui, Pousami, kessi, Roberai, Kuri, fuko, Manikiong, ufo, Kapauku, warkai, Kamora, Kokonao, osier, Itik, sao, Kwerba, wel, Iwer, nu, Wagu, aglo, Orne, koba, E. Sepik. 11. Agathis robusta (Moore) Batley, Synops. Queensl. Fl. (1883) 498; Wars. Monsunia | (1900) 185; BAKER & SmitH, Pines of Austr. (1910) 376; DALLIMORE & JACKSON, Handb. Conif. (1923) 148; Franco, An. Inst. Sup. Agron. 18 (1951) 105, f. 2—10; HyLaNnb, Brunonia 1| (1978) 105, f. 1; SiBa, Phytologia Mem. 8 (1986) 37. — Dammara robusta Moore ex F.v.M. Quart. J. Trans. Pharm. Soc. Vict. 2 (1806) 173. — Dammara palmerstonii F.v.M. Vict. Natur. 8 (1891) 45. — A. palmerstonii (F.v.M.) BAILEY, Queensl. Dept. Agric. Bot. Bull. 3 (1891) 17; DALLIMORE & JACKSON, Handb. Conif. (1923) 147. — A. robusta var. robusta Wuit. PI. Syst. Evol. 135 (1980) 63. — A. robusta var. nesophila auct. non WuHITM.: WHIT. /.c. f. 6. Tree to 48 m. Juvenile leaves ovate and acuminate, up to 13 by 3.8 cm. Adult leaves ovate and acute to rounded at the apex, 5—9.5 cm by 10—26 mm, taper- ing toa 3—10 mm petiole. Mature pollen cones 4—8.5 cm by 7—9 mm, subtended by a 2—9 mm peduncle, apex of microsporophyll sharply angled to form three lateral facies, c. 1 mm wide and long, the slight- ly angled along its margin front (apical) face directed inward to the bases of the next microsporophylls ahead on either side. Seed cone oval to elongated, 9-15 by 8—10 cm. Seed bract with a thin sharp ridge along the apical margin, rounded at the upper cor- ners, the curving lateral margins with a larger scallop c. 10-12 mm above the base on one side and a smaller one 5—8 mm above the base on the other side, 30—45 by 39—46 mm. Seed c. 10 by 6 mm with a long acute projection on one upper corner and a long bent wing c. 30 by 12 mm at the other. Distr. In four localized clusters, one in the cen- tral part of New Britain, another east of Port Mores- by in Papua New Guinea, the largest in the rain- forests of northern Queensland, and the last in southern Queensland on Fraser I. and the nearby mainland. Fig. 86. Fig. 86. Range of Agathis robusta (MOORE) BAILEY. Ecol. Locally common on a variety of soils in ex- posed positions following disturbance from near sea- level in Australia and 400 m further north to 900 m throughout its range. Vern. Naveil, New Britain. CUPRESSACEAE Both Cupressaceae and the closely related Taxodiaceae are important Holarctic families which also have representatives in the southern hemisphere. A few species of both extend into habitats on the margins of the tropics or into tropical highlands. Of 18 genera of Cupressaceae only Libocedrus reaches into Malesia. Occasional reports of Callitris in New Guinea have been based on similar ap- pearing specimens of Casuarina. 1988] CONIFERALES (de Laubenfels) 443 1. LIBOCEDRUS ENDL. Syn. Conif. (1847) 42; Carr. Traité Gén. Conif. (1855) 84; Masters, J. Linn. Soc. Bot. 30 (1895) 20; Wars. Monsunia | (1900) 189; DALLIMORE & JACKSON, Handb. Conif. (1923) 300; Li, J. Arn. Arb. 34 (1953) 17; Florin & BouTELsE, Acta Horti Berg. 17 (1954) 31; DE Laus. Fl. Nouv. Caléd. et Dép. 4 (1972) 145; Sma, Phytologia Mem. 8 (1986) 108. — Libocedrus subg. Eulibocedrus PILceR in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 389. — Papuacedrus Li, J. Arn. Arb. 34 (1953) 25; FLorin & BouTELJE, Acta Horti Berg. 17 (1954) 31; BOUTELJE, /.c. 198, t. 4, pl. 7 & 8; VAN ROYEN, Alp. Fl. New Guinea 2 (1979) 1. — Austrocedrus FLoRIN & BOUTELJE, Acta Horti Berg. 17 (1954) 28. — Fig. 88, 89. Monoecious evergreen trees or shrubs. Bark smooth but fissured, peeling in strips or flakes, fibrous, rich brown but weathering to blackish or gray. Leaves in alternating whorls of 3 or 4 soon reduced to opposite decussate, those of the seedling single veined and linear, c. 1 cm long, changing abruptly on lateral branches and throughout mature trees to specialized scale forms. Scale-bearing branches with small dorsally keeled facial scales alternating decussately with larger marginal leaves that are strongly bilaterally flattened and sometimes ex- tend outward wing-like, the basal margins of each pair meeting between the suc- ceeding facial leaves, the whole branch system further differentiated dorsiven- trally with a convex upper surface bearing few or even no stomata and a concave lower surface where the stomata are concentrated, the leaf differentiation dim- inishing (or more or less disappearing) on older more exposed plants. Foliage buds nothing more than the last pair of developing leaves. Fertile structures solitary, terminal, often on short lateral branches. Pollen cones cylindric, com- posed of decussate or more or less crowded scales each with 2—6 inverted pollen sacs. Seed cones woody, composed of two oval opposed fertile scales each bear- ing two erect ovules at their bases and two small triangular lateral sterile scales. The bracts are mostly fused to the outer surface of the scales, the apex of the bract a short to elongated acute spreading projection. Seed an elongated cone with two very unequal wings, a narrow wing along one side and an elongated more or less expanded wing at least twice as long as the seed on the other side and extending beyond and more or less outward from the seed apex (micropyle). Distr. There are 7 spp. of Antarctic forests and tropical highlands including New Guinea, New Caledonia, New Zealand, and S. Central Chile with adjacent parts of Argentina. In Malesia: 1 sp. with 2 varieties endemic to New Guinea and nearby islands. Fig. 87. Fossils. Foliage shoots and wood from the Eocene of Patagonia and Chile (Florin, K. Svenska Vet. Ak. Handl. II, 19, nr. 2, 1940, 82). Ecol. A wide range of forest and rain-forest habitats from lowland tropics to the tropical tree line and throughout the Antarctic forests (in Chile prospering on the dry margins of the forest). Pollination and seed dispersal strictly by wind. Seedlings germinating in abundance in rich humous soils. Uses. The aromatic wood is similar to that of Juniperus in appearance and uses, with light coloured sap- wood and reddish brown to purplish heartwood. Where large enough it is much appreciated for construction and furniture while the bark is sometimes used for roofing. 444 FLORA MALESIANA [ser. I, vol. 10° > aA Fig. 87. Range of the genus Libocedrus ENDL. with the number of species, all endemic. Notes. The relationships of Libocedrus are emphatically with the Holarctic Thuja group of genera (Tyu- joideae) within Cupressaceae making it phytogeographically much like the everywhere associated Nothofagus whose relatives are also in the north. Like the other members of Thujoideae, the leaves are strongly differen- tiated into lateral and facial types and even further like many of these genera the branches are also differ- entiated dorsiventrally (the Holarctic genus Calocedrus was for a long time included within Libocedrus). At- tempts as in Li (1953) to attach Libocedrus to southern hemisphere cypresses (Callitroideae) by ignoring the highly specialized foliage forms and describing the seed cones as ‘valvate’ are inadmissible. The so-called valvate appearance is due to the few cone scales, the lowermost scales of any Cupressaceous seed cone being the same so that this appearance occurs wherever the number of scales is reduced, as in Chamaecyparis noot- katensis. The species of New Guinea and of Chile have been placed into separate genera based on slight differences. The upper surface of the leaves of the Chilean species are so constricted that little or no space is left for stomata, making them more or less hypostomatic while other species are clearly amphistomatic but with rather few upper stomata. The New Guinea species was separated on the basis of spirally placed microsporophylls. In fact, simple opposite decussate pollen cones occur alongside crowded cones whose microsporophylls ap- pear to be whorled or perhaps spirally placed. FLoRIN & BOUTELJE (1954) carefully examined these cones and found each two decussate pairs of microsporophylls brought to the same level but certainly not spirally placed. On the other hand, they adduced some other minor distinctions for the New Guinea material, in particular that the stomate bands are more or less separated by narrow irregular stomate-free zones, a character not seen elsewhere in the genus. In my opinion these otherwise very similar species should not be separated generically by such unimportant distinctions. 1. Libocedrus papuana F.v.M. Trans. R. Soc. Vict. n.s. 1 (1889) 32; Wars. Monsunia 1 (1900) 189; K.Scu. & Laut. Fl. Schutzgeb. Siidsee (1901) 156; Nachtr. (1905) 51; Koorp. Nova Guinea 8 (1911) 613; Laut. Bot. Jahrb. 50 (1913) 52, f. 2A—G; LaNE- POOLE, For. Res. Terr. Papua New Guinea (1925) 74; SILBA, Phytologia Mem. 8 (1986) 109. — Thuja pa- puana (F.v.M.) Voss, Mitt. Deut. Dendr. Ges. 1907 (1908) 88. — L. torricellensis SCHLTR ex LAUT. Bot. Jahrb. 50 (1913) 52, f. 2H—-N. — Papuacedrus pa- puana (F.v.M.) L1, J. Arn. Arb. (1953) 25; FLORIN & BouTELsE, Acta Horti Berg. 17 (1954) 32, pl. 2, t. 1—3; Harrison in Dallimore & Jackson, Handb. Conif. ed. 4 (1966) 323; vAN RoyEN, Alp. Fl. New 1988] CONIFERALES (de Laubenfels) 445 Fig. 88. View south over the Archbold Expedition camp and Lake Habbema to snow-capped Mt Wilhelmina, from an altitude of 3265 m. Open forest of Libocedrus papuana F.v.M. in foreground centre and big, moss- cushioned Podocarpus compacta; alt. of Lake Habbema 3225 m, Mt Wilhelmina 4750 m (Photogr. L.J.Brass, August 1938). Guinea 2 (1979) 2, f. 33. — Papuacedrus torricel- lensis (SCHLTR) Li, J. Arn. Arb. 34 (1953) 25; FLORIN & Boutese, Acta Horti Berg. 17 (1954) 31, pl. 2, t. 4—6:; HARRISON in Dallimore & Jackson, Handb. Conif. ed. 4 (1966) 323. — Fig. 88, 89. Trees 2—50 m tall, rarely shrubby, pyramidal or spreading or even slightly pendulous with age. Leaves on older and more exposed trees becoming quite small, facial scales rhomboidal, acute, up to 1 mm long, the lateral scales nearly straight, their base between the half as long as the lateral scales, c. outermost edges of the next lower facial scales, spreading slightly and then straight or weakly con- vex, tips extending slightly beyond the outermost edges of the next higher facial scales and either straight or slightly spreading or (when convex) slight- ly incurved and + touching the base of the next later- al leaves but below the tips of the next facial leaves, down to c. 2 mm long. Branches darker above and often glaucous below. Pollen cones 4—25 mm long by 2—3 mm in diameter, the upper part of the micro- sporophyll roughly triangular, c. 1.5 mm long, with 2—4 pollen sacs. Seed-bearing scale broadly lanceo- late to almost elliptic, at least slightly rounded at the tip, 8-12 mm long and 4—6 mm wide, narrowing at the base and sometimes slightly constricted along the side where touched by the smaller lateral scales which are about halfaslongand + lanceolate. Bracts broad- ly fused to the dorsal side of their corresponding scale on its lower half with a short spreading acute to obtuse tip. The woody mature cone brown or black- ish, often with ridges radiating from the bracts to the margins of the scale. Seed 2—3 mm long, the wing twice as long and spreading upward witha bend part- way along the outer edge towards the straight inner edge forming an acutish rounded apex. var. papuana Apex of lateral leaves becoming widely spreading on young plants up to 6 mm from the stem and often 446 FLORA MALESIANA [ser. I, vol. 103 Fig. 89. Libocedrus papuana F.v.M. var. papuana. A. sterile twig; B. fertile twig; C. pollen cone; D. seed cone; E. scales with pollen sacs (from Laut. Bot. Jahrb. 50, 1913, 52, f. 2). falcately bent outwards but the tip always turned up- wards, up to 13 mm from base to tip, gradually be- coming smaller and less spreading as the tree ma- tures. Distr. Malesia: most of the length of New Guinea along the central range but also on the Huon Peninsula, the Torricelli Mts, and the Cycloop Mts, grading into the other variety towards the western end of New Guinea, particularly at higher elevations. Fig. 90. Ecol. A tree of mountain rain-forests and especially the mossy forest from 1500 m to the tree line in the central range, sometimes emerging from the alpine scrub to as high as 3800 m, as low as 620 m along the north coast. Above 3300 ma stunted tree less than 10 m tall. Vern. Tera, Garaina; gamuga, kamgenkuna, Togoba, Hagen, ogeleh, oleh, Chimbu, Masul, dza- gosa, dzasihanini, hanini, Asaro, Kefamo, guta, juta, Mairi, Watabung, kaibel-kombam, kaipil, ongol, Wahgi, Minj, mandilasop, mondalasap, mondolasop, wrenak, Mendi, ab, aiap, aip, hap, kap, Enga, Kepilam, aioba, hohoba, Lei area, bit, Yogom, iwunturra, Manki, ye-enka, Nauti, sukou, Wapi, Migote. var. arfakensis (GIBBS) DE LAUB., stat. nov. — L. ar- JSakensis Gipps, Arfak (1917) 84, f. 6a, b; DALLIMORE & Jackson, Handb. Conif. (1923) 301; Sma, Phytologia Mem. 8 (1986) 108. — Papuacedrus ar- fakensis (Gress) Li, J. Arn. Arb. 34 (1953) 25; FLORIN & BouTELse, Acta Horti Berg. 17 (1954) 32, 1988] CONIFERALES (de Laubenfels) 447 = Fig. 90. Range of Libocedrus papuana F.v.M. var. papuana (dots) and var. arfakensis (GIBBS) DE LAUB. (triangles). constricted to the position on older trees close to the base of the following facial leaves but between their base and tip the lateral leaves expanded outward as much as 3 mm and overall as much as 10 mm long, thus producing a strongly convex shape, gradually becoming smaller until they become indistinguish- able from the type variety on older and exposed trees. Distr. Malesia: from the Wissel Lakes through the Vogelkop Peninsula and probably including the material from Batjan and Ternate. Fig. 90. Ecol. Like the type variety from 1600 to 2500 m, but as low as 840 m along the north coast of the Vo- gelkop Peninsula. Vern. Autibo, dautibo, dautie, matu, matu- dautie, Kepauko, wonga, Arfak area, bootsjeka, butsga, Manikiong, pomoan, Manikiong, Hattam, duwak, nipau, tuwa, Kebar, sowa, swa, Kebar, And- jai, araum, eis, Karoon. Note. Despite the strikingly different juvenile leaves, intermediate specimens exist, especially at higher elevations and it has not been possible to separate specimens taken from older plants. t. 3; HARRISON in Dallimore & Jackson, Handb. Conif. ed. 4 (1966) 322. Apex of lateral leaves at first spreading but soon PINACEAE The pine family is one of the most characteristic families of the holarctic realm of which family a few genera reach the margins of the tropics in highlands, but only Pinus extends into tropical lowlands including Malesia. 1. PINUS Linn. Gen. PI. ed. 1 (1737) 731; Sp. Pl. (1753) 1000; LAMBERT, Pinus (1803); Par-. in DC. Prod. 16, 2 (1868) 378; ENGELMANN, Trans. St.Louis Acad. Sc. 4 (1880) 161; Masters, J. Linn. Soc. 35 (1904) 560; SHaw, Publ. Arn. Arb. 5 (1914) 24, t. 1—39; Pircer in E. & P. Nat. Pfl. Fam. ed. 2, 13 (1926) 331, t. 177, 178; GaussEN, Gymn. Act. & Foss. fasc. 6 (1960) 11, t. 324—369, pl. 24—29; Mirov, Genus Pinus (1967); Smpa, Phytologia Mem. 8 (1986) 129. — Fig. 91-93. Evergreen monoecious trees or rarely shrubs. Bark smooth or rough, par- ticularly on older trees, peeling in flakes often of very irregular shape. Leaves linear or lanceolate, often with minute serrations, spirally placed, soon replaced by scales in the axils of which appear reduced shoots in the form of bundles en- closed around their base by a sheath of scale leaves, adult leaves in the bundles linear, pungent, needle-like, the cross section of each leaf forming a sector of the circle formed by the entire bundle. Pollen cones numerous, axillary, cylin- drical, subtended by a cluster of overlapping scales similar to the foliage bud, microsporophylls scale-like with two inverted pollen sacs. Seed cones terminal FLORA MALESIANA [ser. I, vol. 102 Fig. 91. Pinus merkusii JuNGH. & DE VRIESE in primary forest habitat on slope of Mt Bandahara, Leuser Nature Park, Sumatra, 2000 m alt. (Photogr. W.J.J.O. DE WILDE, 1972). 1988] CONIFERALES (de Laubenfels) 449 on short scaly shoots, more or less cylindrical, consisting of numerous fertile scales which become woody, ripening in the second or third year, the apiculate bract fused with the scale, two inverted ovules on each scale. Seed egg-shaped, with a firm outer shell, usually with an expanded wing which is attached to the broad base of the seed. Distr. Over one hundred species across the middle and higher latitudes of the northern hemisphere and southward into Central America, Cuba, and Hispaniola as well as into Malesia, where two important pines of southeastern Asia (belonging to subg. Diploxylon) extend their range. Absent from the Indian subconti- nent, but occurring throughout the Himalayan range. Fig. 92. Pinus merkusii JUNGH. & DE VRIESE in fire-prone grassland, blang area along the Gajo-road, Sumatra (Photogr. J.C. VAN DER MEER Monr, 1925). Ecol. A wide range of forest and savanna habitats most characteristically following disturbance and thus even as large trees surrounded by rain-forest in some cases. Frequently occurring as a fire climax woodland or scattered in fire-prone grasslands. Pollination and seed dispersal by wind or in many cases the seed are gathered by birds or rodents or even collected for human food. Note. Pines are leading sources of lumber and pulpwood and some species such as the two described here yield large amounts of pitch and are tapped to make turpentine. Use as ornamentals is widespread and pines are favourite subjects of afforestation. The widespread use of pines in tropical tree plantations derives from the detailed information available for their silviculture and not because they are of particularly good quality wood. The fact is that much tropical pine wood production is of very poor quality and it is to be hoped that knowledge of the production of better quality woods will increase in the future. 450 FLORA MALESIANA [ser. I, vol. 103 x ; Fig. 93. Pinus merkusii JUNGH. & DE VRIESE. a. Twig with seed cone; b. twig with pollen cones; c. growing shoot; d. pair of needles; e. tips of needles, enlarged; f, g. pollen cones (from DE VRIESE, PI. Nov. Ind. Bat. Ori, 1845 5h 2): 1988] CONIFERALES (de Laubenfels) 451 KEY TG °THE SPECIES 1. Leaves two per bundle. Mature seed cone ovate .. 1. Leaves three per bundle. Mature seed cone globular 1. Pinus merkusii JUNGH. & DE VRIESE in De Vriese, Pl. Nov. Ind. Bat. Or. (1845) 5, t. 2; Bot. Zeit. 4 (1846) 13; ENpi. Syn. Conif. (1847) 176; Carr. Traité Gén. Conif. (1855) 380; Gorpon, Pinetum ed. 1 (1858) 169; Mig. Fl. Ind. Bat. 2 (1859) 1069; HENKEL & HOCHSTETTER, Synop. Nadelhdlz. (1865) 43; pE Boer, Conif. Archip. Ind. (1866) 5; PARL. in DC. Prod. 16, 2 (1868) 389; VipaL, Sin. Atlas (1883) 43, t. 98, f. B; Rev. Pl. Vasc. Filip. (1886) 296; Merr. For. Bur. Bull. Philip. 1 (1903) 15; Foxw. Philip. J. Sc. 6 (1911) Bot. 169; SHaw, Publ. Arn. Arb. 5 (1914) 23, t. 198—200; DALLIMoRE & JACKSON, Handb. Conif. (1923) 415; GaussEN, Gymn. Act. & Foss. fasc. 6 (1960) 146, t. 344, 2 & 3; CriTcHF. & Littte, U.S. Dept. Ag. Misc. Publ. 991 (1966) 15, map 39; Mrrov, Genus Pinus (1967) 284, f. 3-51-52, t. 3-68-69, 553; CooLinc, Fast Growing Timber Trees Low. Tropics 4 (1968) 126; STEEN. Fl. Males. Bull. m. 25 (1971) 1948; PHENGKLAI, Thai For. Bull. 7 (1973) 1, f. 1; SiBa, Phytologia Mem. 8 (1986) 149. — P. sylvestris auct. non L.: Lour. Fl. Coch. 2 (1790) 579. — P. sumatrana JuNGH. Bot. Zeit. 4 (1846) 698. — P. finlaysoniana BLUME, Rumphia 3 (1849) 210. — P. latteri Mason, J. Asiat. Soc. 1 (1849) 74. — P. merkusii var. tonkinensis CHEV. Rev. Bot. Appl. Ag. Trop. 24 (1944) 7. — P. merku- siana COOLING & GAUSSEN, Trav. Lab. For. Tou- louse (1970) 1. — Fig. 91-93. Large tree 40—5S0 m tall, occasionally to 70 m. Pyramidal with heavy horizontal branches and thick fissured bark in most areas but in some regions trees have more slender, somewhat ascending branches and much thinner, smoothish bark resembling that in the upper crown of thick-barked populations. These two types hold true in artificial plantings. The red- dish bark of young trees changes to dark brown weathering to gray on older trees. Foliage buds long and narrow with awl-shaped scales. Needles in pairs, 16—19 cm by | mm, abruptly pointed, stomata on all faces, falling in the second year. Basal sheath 12—18 mm long, reddish. Pollen cones 18—25 by 5 mm. Seed cone cylindrical before opening, 5—11 by 3 cm and twice as thick after opening, generally falling soon after shedding seeds. Apophysis broadly tetra- gonal in shape with a smooth, almost depressed um- bo. Seed 7.5 by 4.5 mm, with a deciduous wing 25 by 8 mm. Distr. In Malesia common throughout the mountains of Atjch in Sumatra and scattered further south in Tapanuli with an isolated outlier near Mt Kerintji at c. 2° south of the equator and the only natural occurrence of pine south of the equator. The 1. P. merkusii 2. P. kesiya Fig. 94. Range of Pinus merkusii JUNGH. & DE VRIESE. same or a closely related pine is scattered throughout SE. Asia from E. Burma to the South China Sea and two small areas in the Philippines, one on Mindoro and the other near the west coast of Luzon in Zamba- les Prov. Fig. 94. Ecol. From low elevations to 2000 m, generally on poor quality acid podzolic soils over sandstone or fresh volcanic ash, sometimes on deeply leached acid basalt, rarely successfully competing on richer forest soils. Most stands show a clear relationship to fire or other disturbance and the pine can be seen to be ex- panding in recently disturbed areas. In Sumatra the habitat experiences heavy year-round precipitation, but the pine areas themselves definitely favour the drier sites. The Tapanuli populations, which have thin bark, are more sensitive to fire and do not de- scend below 1000 m. Elsewhere, including the Philip- pine islands, this pine grows in strongly seasonal en- vironments. Vern. Sumatra: dammar batu, dammar bunga, hejam, hujam, ujam, ujem, Atjeh, higi, Kerintji, ka- ju tussam, tussam, Battok, Tapanuli; Philippines: tapulao, Sambal, Luzon, agoo, aguu, salit, Philip. islands. Note. The similar pines of the southeastern Asian mainland and the Philippines differ in certain ways from those of Sumatra. The mainland pines, which grow in areas with a distinct dry season much stronger than any seasonality in Sumatra, have a dis- tinct grass stage for the seedling, a character not seen 452 at all in Sumatra. The needles are 19—24(—27) cm long, the seed cones have a more typical taper of most pines, not the unusual cylindrical Sumatran shape, and their umbos tend to be distinctly raised on an apophysis distinctly wider than high. Seeds are nearly twice as heavy. Sumatran pines have a multi- nodal leader (several nodes in each year’s growth) while the mainland form is uninodal. The new spe- cies, P. merkusiana, proposed for the mainland population, was described without a type specimen, but the earlier name, P. /atteri, is already available. The varietal name tonkinensis is also available. Like Sumatra, mainland areas have an ‘upland’ thin- barked form to which the varietal name applies should further nomenclatural divisions be needed. 2. Pinus kesiya RoyLE ex GoRDON in Loudon, Gard. Mag. 16 (1840) 8; HARRISON in Dallimore & Jackson, Handb. Conif. ed. 4 (1966) 436; StyLes & BURLEY, Comm. For. Rev. 51 (1972) 241; BuRLEy, Proc. IUFRO Breeding Symp. Gainesville (1972) 38; PHENGKLAI, Thai For. Bull. 7 (1973) 3; QuimsBo, Canopy 4 (9) (1978) 9, 14; PADoLina, Canopy 4 (10) (1978) 5; SryLes, Canopy (Aug. 1979) 5; in Armitage & Burley, Trop. For. Pap. 9 (1980) 199. — P. taeda auct. non L.: BLANCO, FI. Filip. (1837) 767; MERR. Bur. Govt. Lab. Publ. Philip. 27 (1905) 82. — P. in- sularis ENDL. Syn. Conif. (1847) 157; PREsL, Epim. Bot. (1851) 37; PaRL. in DC. Prod. 16, 2 (1868) 390; VIDAL, Sin. Atlas (1883) 43, t. 98, f. C; Rev. Pl. Vasc. Filip. 1 (1903) 15; Philip. J. Sc. 5 (1910) Bot. 325; Whiter. For. Bur. Bull. 10 (2) (1911) 26, t. 2, 3; Foxw. Philip. J. Sc. 6 (1911) Bot. 170; Saw, Genus Pinus (1914) 60, t. 23, f. 208—210; Wu, Acta Phyto- tax. Sinica 5 (1956) 145; RAIZADA & SAHNI, Ind. For. Rec. 5 (1960) 114, t. 5, f. 3; Back. & Baku. f. FI. Java 1 (1963) 91; CrircHr. & LiTTLE, U.S. Dept. Ag. Misc. Publ. 991 (1966) 14, map 38; Mrrov, Genus Pinus (1967) 297; Roso, Sylvatrop 3 (1978) 31; DE Laus. Kalikasan 7 (1978) 148; Roso, Canopy (March 1979) 10; SttBa, Phytologia Mem. 8 (1986) 145. — P. khasyana GrirF. Notul. Pl. Asiat. 4 (1854) 18, t. 367, 368, with ref. to descr.: 2 (1848) 58. — P. kasya Part. in DC. Prod. 16, 2 (1868) 390; BRANDIS, For. Fl. NW & Central India (1874) 508; Kurz, Fl. Burma 2 (1877) 499; GamBLE, Man. Ind. Timber (1881) 397. — P. khasia ENGELMANN, Trans. St. Louis Acad. Sc. 4 (1880) 179. — P. khasya Hook. f. Fl. Br. India 5 (1888) 652; GAMBLE, Man. Ind. Timber 2 (1902) 708; Merk. For. Bur. Philip. 1 (1903) 15; BRANDIs, In- dian Trees (1906) 690; DALtimoRE & JACKSON, Handb. Conif. (1923) 400; Burc NGoc-sanu, Adan- sonia 2 (1962) 337; GaussEN, Gymn. Act. & Foss. fasc. 6 (1960) 154, f. 345, 5, 7, incl. var. insularis (ENDL.) GAUuSSEN, /.c. f. 345, 6; NGUYEN KuHa, Ann. FLORA MALESIANA [ser. I, vol. 10° Sci. For. 23 (1966) 261; Mrrov, Genus Pinus (1967) 295. Tree to 35—45 m, the crown expanding on older trees. Bark reddish brown, thick and deeply reticu- lately fissured, breaking off in small thick irregular plates and thus sometimes becoming smoother and plate-like. Branchlets smooth, bright brown. Buds oblong cylindric and non-resinous with brown lan- ceolate scales free at the tips. Need/es in threes, rarely pairs, 12—24 cm by 0.5 mm, acuminate, stomata on most surfaces, falling after two years. Basal sheath 5—18 mm long, greyish brown. Pollen cones 18—30 by 5 mm. Seed cone ovoid to conical before opening, 4.5—10 by 3—5 cm, very persistent. Apophysis wider than long, pyramidal, the umbo with a small expand- ed and usually deciduous mucro. Seed 5—8 by 3 mm with a deciduous wing 20 by 8 mm. Distr. Across SE. Asia to E. India (Khasia) and a short way into China; in Malesia: Philippines (common in the northern part of Luzon). Fig. 95. Fig. 95. Range of Pinus kesiya RoyLE ex GORDON. Ecol. Most often in open pure stands following fire, but also mixed with oaks and Ericaceous species at intermediate elevations from 300 to 2700 m, often on steep slopes. Generally in the same area as P. merkusii, but usually at higher elevations. A few specimens in the Philippines have been considered hybrids between these two species, but actual hybri- dization has not been verified. Vern. Al-al, parua, saleng; boo boo, bot bol, bulbul, lf., tapulao, Zambales. Note. Pinus timorensis (an earlier name) was thought by some to be equal to P. kesiya (P. in- sularis) although the description is inadequate to confirm this. There are no native pines in Timor; perhaps this was a cultivated tree. 1988] CONIFERALES (de Laubenfels) 453 Doubtful or Excluded Callitris sp. mentioned from New Guinea by H.J.Lam, Nat. Tijd. Ned. Ind. 89 (1929) 304, 354; Sargentia 5 (1945) 143, 168, is according to VAN STEENIS, Acta Bot. Neerl. 2 (1953) 299 = Gymnostoma sp. (Casuarina- ceae). Podocarpus elata R.BR. — ENGLER, Bot. Jahrb. 7 (1886) 445, mentioned a specimen from Timor (Kupang Bay) to belong to this Australian species. PmGeR did not mention this specimen in his monograph and WASSCHER did not see it either (cf. Blumea 4, 1941, 471). Podocarpus palembanica Mia. Fl. Ind. Bat. Suppl. (1860) 252, 289. According to DE BoER (Conif. Arch. Ind., 1866, 4) it is not a conifer because of its non-coniferous wood. KOSTERMANS (Reinwardtia 2, 1953, 362) observed its stipules and nerves and identified it as a juvenile specimen of Ganua sp., later referred tentatively to a distinct species by VAN DEN AssEM (Blumea 7, 1954, 482): Ganua palembanica (MIQ.) VAN DEN ASSEM & KosTerM. (Sapotaceae). Thuja javanica Bur. f. Fl. Ind. (1768) 202, t. 64, f. 3. — Podocarpus javanicus (BURM. f.) MERR. Philip. J. Sc. 19 (1921) 338, pro nomen. BURMAN’s description and figure are based on a specimen of Java. In L there is a specimen in herb. VAN ROYEN, with a label ‘Thuya javanica é Java. Monoic.’ which seems to be in Bur- MAN’s handwriting and which can provisionally be accepted as the type specimen. According to HALLIER /f. (Meded. Rijksherb. n. 37, 1918, 92) it belongs to Juniperus chinensis, a conifer already cultivated in Java in early days. op 0h 1 ee q ; he py Pads mee hy. @ith Sie @ 2% cue ne a. cy taney a z , a an on * ena, Cerin . _ fies 7 ra, @ yg =a o POLYGALACEAE (R. van der Meijden, Leiden)! Herbs (sometimes saprophytic), shrubs, lianas or trees. Stipules absent but stem sometimes provided with a pair of glands at the nodes. Leaves simple, entire, usually spirally arranged, sometimes alternate, (semi)decussate or verticillate, sometimes scale-like or absent. /nflorescence usually raceme-like and un- branched, (supra- or extra-)axillary and/or terminal, sometimes thyrsoid or fas- ciculate, rarely flowers solitary. Bracts present; bracteoles basal, rarely (Salo- monia, Epirixanthes) absent. Flowers bisexual, more or less zygomorphous, rarely actinomorphous. Sepals 5, free and quincuncial, or the lower (abaxial) 2 connate, sometimes all connate, subequal or the lateral ones larger and then often wing-like (alae) and petaloid. Petals 3 or 5, free or variously united, occa- sionally also with the calyx, usually adnate to the base of the staminal tube or the filaments, subequal or more often unequal with the lower petal often keel- like and frequently pouched, lobed, or crested. Stamens 2—10, usually 8, fila- ments usually more or less connate except between the upper stamens, often ad- nate to the petals; anthers basifixed, tetra- or bi-, rarely trisporangiate, 1- or 2-locular, opening by a single and often oblique pore or by a longitudinal in- trorse slit. Ovary superior, usually 2-locular but occasionally 1-, 3-, 5-, 7- or 8-locular, sessile or sometimes stipitate; style simple but often variously dilated or lobed at apex, usually articulate with the ovary and nearly always deciduous in fruits. Ovules 1 per cell and subapical, or (in Xanthophyllum) 4—more in a 1-locular, bicarpellate ovary with 2 parietal placentas, anatropous, bitegmic and crassinucellate. Fruit various, a berry, capsule, samara or drupe. Distribution. About 15 genera and over 1000 species, widespread in temperate and tropical regions of the world, especially well-developed in South America and South Africa. In Malesia 6 genera, of which Polygala and Securidaca (not in Australia) are cosmopolitan, Xanthophyllum and Salomonia Indo-Australian, Epirixanthes Indo-Malayan. The sixth genus is Eriandra which belongs to the tropical American tribe Moutabeae, of which 3 genera are known in South America; Eriandra occurs in New Guinea and the Solomon Islands and represents a marked ex- ample of disjunct, tropical trans-Pacific affinities. Ecology. Very diverse, in wet or dry, open or closed, warm or rather cold habitats, in the lowlands or on mountains up to 3600 m; greatly varying in habit. Epirixanthes is a genus of small saprophytic (not parasitic) plants on humous soil in deep shade among litter of the rain-forest. Salomonia is an unseemly herb of open places. In Pol/ygal/a there are woody shrubs in the under- growth of the rain-forest, but most species are herbs of open country, several even preferring a seasonal climate; one species (§ Melchiora) is a climber of the rain-forest, like Securidaca. The species of Xanthophyllum and the monotypic Eriandra are trees of the rain-forest, sometimes of lofty size. In some species of Polygala the occurrence of endomycorrhiza (VA-mycorrhiza) has been dem- onstrated (HEUBL, 1984). Pollination. Most species seem to be adapted to cross-pollination, but thusfar actual observa- tions have only been reported for some Polygala species (see there). In Epirixanthes the structure of the flowers seems fit for cross-pollination (see there). Self-pollination, however, has been re- (1) For this revision the 1970 manuscripts have been used of three former students of Professor Van Steenis: F.C. Roest (Polygala sect. Chamaebuxus), L.P. Rijfkogel (Securidaca), and Mrs, H.M.Y.J. André de la Porte-Janss (Salomonia, Epirixanthes). (455) 456 FLORA MALESIANA [ser. I, vol. 103 ported for species in diverse genera, also for those species in which cross-pollination has been reported, or is suspected to be possible. It may be concluded tentatively that self-pollination is an effective second-chance possibility for reproduction in the Polygalaceae. Dispersal. Corresponding to the diversity in fruit and seed types there is a great variation in dis- persal types (VERKERKE, 1985). Especially in Polygala many dispersal types occur: myrmeco- chory, ornithochory, anemochory, diplochory, epizoochory. Ornithochory also occurs in Dicli- danthera, Carpolobia, Atroxima, and probably in some Xanthophyllum species. Moutabea fruits have endozoochorous dispersal by monkeys (VAN ROOSMALEN, 1985); this may also be true for some Xanthophyllum species. Myrmecochory (or perhaps also anemochory) probably occurs in Bredemeyera, Comesperma, and Epirixanthes; anemochory also in Monnina and Securidaca (VAN ROOSMALEN, /.c.), but in the latter also hydrochory is possible. Epizoochory is the possible means of dispersal for Sa/omonia. References: HEuBL, Mitt. Bot. Staatssamml. Munchen 20 (1984) 222; vAN ROOSMALEN, Fr. Guianan FI. (1985) 360; VERKERKE, J. Arn. Arb. 66 (1985) 385. Morphology. Recently the morphology and ontogeny of ovules, fruits and seeds have been described (and reviewed) by VERKERKE; Of Polygala: VERKERKE & BAUMAN (1980); of Xantho- phyllum: VERKERKE (1984); of the remaining genera: VERKERKE (1985). LEINFELLNER (1972) dem- onstrated that there is no principal difference in the ontogeny of the unilocular, multiovulate ovary of Xanthophyllum and the bi- to octoloculate, uniovulate ovary of the other Polygalaceae. The ontogeny of bi- and trisporangiate anthers of Polygala has been described by CHopat (1891) and VENKATESH (1956). Stipular outgrowths or nodal glands are pseudostipules in the sense of WEBERLING (VAN DER MEIJDEN, 1982: 3). Contrary to the idealistic opinion of CHopaAT (/.c.) (also adopted by HUTCHINSON, 1967) and CRONQUIST (1981), the primitive number of stamens is 8; the presence of 10 stamens (Diclidanthera; and a rare abnormality in Xanthophyllum) is a derived character. References: CHODAT, Monogr. I (1891); CRoNqQuisT, Integr. Syst. (1981) 763; HUTCHINSON, Gen. FI. Pl. 2 (1967) 338; LEINFELLNER, Oest. Bot. Z. 120 (1972) 51; VAN DER MEIJDEN, Leiden Bot. Ser. 7 (1982) 3; VENTAKESH, Bull. Torrey Bot. Club 83 (1956) 19—26; VERKERKE, Blumea 29 (1984) 409—421; J. Arn. Arb. 66 (1985) 353—394; VERKERKE & BOUMAN, Bot. Gaz. 141 (1980) 277-282. Vegetative anatomy. The Polygalaceae exhibit an interesting diversity in their leaf and wood anatomy, which has only fragmentarily been explored, especially for the Malesian genera. Hairs if present are unicellular, or more rarely uniseriate. The lower epidermis is papillate in a number of species. Stomata may be of the anomocytic, paracytic or anisocytic type (all three types occur within the genus Xanthophyllum). An adaxial hypodermis is frequently present in the woody species with coriaceous leaves. The vascular pattern in petiole and midrib ranges from a single collateral bundle to a closed cylinder with accessory bundles (again the whole range of the family is represented in Xanthophyllum). The nodes are unilacunar. Unusual tracheoidal idio- blasts in the leaf mesophyll are characteristic for Xanthophyllum. The secondary xylem of the trees and climbers is characterised by largely solitary vessels with simple perforations, fibres with distinctly bordered pits and heterocellular rays which are usually narrow (1—2-seriate), but may be much wider in the Moutabeae (e.g. Securidaca). Axial paren- chyma is mainly paratracheal in Po/lygala, apotracheally diffuse and diffuse-in-aggregates plus vasicentric to loosely aliform in Securidaca, and apotracheally banded plus vasicentric in Xan- thophyllum. Included phloem occurs in the wood of Securidaca (and other Moutabeae). Despite the anatomical distinctness of Xanthophyllum (mainly through its tracheoidal idioblasts), vegetative anatomy clearly witnesses affinity with other Polygalaceae (epidermal characters, overall leaf histology, solitary vessels and fibre type in the wood; the parenchyma distribution in some Xanthophyllum species is reminiscent of that of Securidaca). In its wood anatomy Xanthophyllum also recalls Trigoniaceae, especially Trigoniastrum. References: BRIDGWATER & Baas, [AWA Bull. n.s. 3 (1982) 115—125; Dicx1son, Bot. J. Linn. 1988] POLYGALACEAE (van der Meijden) 457 Soc. 67 (1973) 103—115; METCALFE & CHALK, Anatomy of the Dicotyledons 1 (1950) 133-138; Styer, J. Arn. Arb. 58 (1977) 109—145. Palynology. Pollen grains in Polygalaceae are mostly suboblate to prolate, sometimes equa- torially constricted (Epirixanthes cylindrica), and measure from 25 to 62 um. The apertural sys- tem is zonocolporate (‘stephanocolporate’) in all genera, the number of apertures ranging from 5 (Xanthophyllum papuanum) to 17, or up to 42 in Polygala. The endoapertures may be fused (‘synorate’) to form one broad, equatorial endoaperture, bordered by endexinous costae. In this way, the circle-shaped endoaperture divides a grain into two rigid halves, in which the colpi are no longer active in harmomegathic functioning. Volume accommodation in such a grain, and pos- sibly also in grains with poorly separated pores, is achieved by folding of the flexible parts of the mesocolpia which cross the endoapertural zone (MULLER, 1979). Similar apertural systems and harmomegathic mechanisms occur in the genus Ufrricularia of the unrelated family Len- tibulariaceae (THANIKAIMONI, 1966; HUYNH, 1968). Exine stratification is mostly obscure using light microscopy. LARSON & SKVARLA (1961) dem- onstrated the presence of ectexine, endexine, and a columellate infratectal layer in Polygala with electron microscopy. The tectum is mostly smooth, pitted, or perforate, sometimes somewhat fossulate or scabrate. Large perforations (‘aperturoid depressions’, ‘lacunae’) may occupy the poles. Pollen of the family Polygalaceae is very distinct. Supposed relationships with other families within the Po/ygalales are not supported by pollen morphological evidence. Pollen of the family Krameriaceae is certainly dissimilar to that of Polygalaceae (ERDTMAN, 1944, 1952; Simpson & SKVARLA, 1981). On the other hand, the distinct pollen type represents a strong argument for in- clusion of Xanthophyllum as a genus within Polygalaceae. Also in ascertaining the systematic position of Diclidanthera (ERDTMAN, 1944) and Eriandra (VAN ROYEN & VAN STEENIS, 1952) pollen morphology turned out to provide circumstantial or even deciding evidence. It is not yet possible to key out all Malesian genera of Polygalaceae on the strength of pollen characters. Only one rough separation can be made: pollen of Epirixanthes, Eriandra, Securidaca, and Xanthophyllum mostly has 12 or less apertures, while grains in Polygala and Salomonia rarely have less than 12. Pollen of Epirixanthes is characterized by its heavy equatorial belt and involute colpus margins. These features probably represent derived states in Polygalaceae. Pollen of Securidaca has clearly separated endoapertures. This state may be con- sidered as primitive, comparing with the synorate type in Polygalaceae. References: ERDTMAN, Bot. Notis. (1944) 80-84; Pollen morphology and plant taxonomy, Angiosperms (1952) 332—333; HuyNu, Pollen et Spores 10 (1968) 11—55; LARSON & SKVARLA, Pollen et Spores 3 (1961) 21-32; MuLLER, Ann. Missouri Bot. Gard. 66 (1979) 593-632; VAN RoyeN & VAN STEENIS, J. Arn. Arb. 33 (1952) 91—95; Stimpson & SKVARLA, Amer. J. Bot. 68 (1981); THANIKAIMONI, Pollen et Spores 8 (1966) 265—284. — R.W.J.M. VAN DER HAM. Phytochemistry & Chemotaxonomy. Chemical characters of the family were summar- ized and discussed by HEGNAUER (Chemotaxonomie der Pflanzen 5, 1969, 352—361, 459). A sup- plement will be included in volume 8. Glycosides releasing methyl salicylate on hydrolysis and saponins with triterpenic acids as their sapogenins are rather common in the family, especially in roots. The most characteristic genuine sapogenin of the family seems to be presenegenin, C3oH4¢O>, an acid-labile derivative of oleanolic acid; it was shown to be mainly sapogenin of roots of several members of Bredemeyera, Carpolobia, Polygala and Securidaca. Closely related sapogenins are polygalacic acid from Polygala paenea and bredemolic acid from Bredemeyera floribunda. Mono-, di- and trimethoxycinnamic acids as well as ferulic and sinapic acid occur widely; usually they are esterified with free sugars such as saccharose or with the sugar-part of the bidesmosidic saponins. Leaf flavonoids seem mostly to be glycosides of quercetin and kaemp- ferol. Polygalitol (= aceritol = 1,5-anhydroglucitol) is a characteristic hexitol derivative of roots of many polygalaceous plants; it occurs free and combined with sugars. Seeds usually store mainly proteins and fatty oils, but no starch. There is still littke known about the chemistry of 458 FLORA MALESIANA [ser. I, vol. 103 polygalaceous seed oils, but very unusual oils were shown to be produced by Monnina emarginata and Polygala virgata. Three groups of secondary metabolites have still to be mentioned notwith- standing the fact that they are known only from a few species of Polygala at present. These are polyhydroxylated xanthones, and naphthalin- and bibenzyl-y-butyrolactone-type lignans such as podophyllotoxin (e.g. Polygala polygama) and suchilactone (e.g. Polygala chinensis). Moreover, the Central American Polygala paniculata yielded a number of rutaceous coumarins and a diester of khellactone. The taxonomic relevance of chemical characters was discussed by HEGNAUER in 1969. It may be added now that the occurrence of isoprenylated coumarins and an obvious total lack of iridoids agree well with the assumption of sapindalean (sensu /ato) affinities. Tannins too seem to be total- ly lacking in Polygalaceae ; this, however, does not contradict the just mentioned assumption, be- cause within Sapindales s./. a strong tendency to replace tannins by other types of secondary meta- bolites is apparent. Finally it should not be forgotten that several chemical characters may prove valuable in future for infrafamiliar classification. — R. HEGNAUER. Taxonomy. CRONQUIST’s circumscription of the order Polygalales (1981: 763) reflects the general opinion of systematists; next to the Polygalaceae (with Xanthophyllum as a separate fam- ily) it includes the Trigoniaceae, Vochysiaceae, Malpighiaceae as well as the Tremandraceae and the Krameriaceae. Wood anatomical evidence (BRIDGWATER & BAAS, 1982) supports a close af- finity of Polygalaceae, Xanthophyllum and Trigoniaceae, but not with the other families. VAN DER MEIDEN (1982) found no arguments to include Tremandraceae and Krameriaceae. Although there seemed to be little evidence for the inclusion of Krameriaceae in the order (SIMPSON & SKVARLA, 1981; Smmpson, 1982), the first author recently supported the classification of Krameria- ceae next to Polygalaceae, mainly based on serological evidence (BUSE-JUNG, 1979). Splitting off Xanthophyllum as a separate family has been based on incomplete or erroneous knowledge of the genus. Of the three characters mentioned by CRONQUIST (/.c.), two do not hold: filaments are often partly and sometimes halfway connate in Xanthophyllum, and seeds with copious endosperm occur in four of the seven subgenera. Thus Xanthophyllum differs in a single though compound character from other Polygalaceae, viz. in the structure of the ovary (which ontogenetically is largely similar to other Polygalaceae, cf. LEINFELLNER, /.c.); this differs in the reduced septs, in the doubling of the number of ovules per carpel, and in the height of insertion of the ovules. On the other hand Xanthophyllum has a number of striking similarities with other Polygalaceae in the structure of the flowers, as well as in some vegetative characters (the presence of laminar and nodal glands). Also studies on the morphology of ovules, fruits and seeds (VERKERKE, 1984, 1985), of foliar anatomy (Dickison, 1973) and wood anatomy (BRIDGWATER & BAAs, 1982) do not present arguments to split Xanthophyllum from the Polygalaceae. Mainly based on differences in floral structure, CHODAT (1891) distinguished three tribes in the family, Polygaleae, Moutabeae and Xanthophylleae. Both the studies of StyER (1977) and VERKERKE (1984, 1985) revealed that the differences between Moutabeae and Polygaleae are unclear; the results of vegetative and of seed anatomy do not correspond with differences in the flowers. Therefore a formal subdivision of the family is not presented. References: BRIDGWATER & Baas, IAWA Bull. n.s. 3 (1982) 115—125; BusE-JuNG, Thesis, Kiel (1979); CHopaT, Monogr. I (1891); CRonquist, Integr. Syst. (1981) 763; Dickison, J. Linn. Soc. Bot. 67 (1973) 103—115; LEINFELLNER, Oest. Bot. Z. 120 (1972) 51; VAN DER MEUDEN, Leiden Bot. Ser. 7 (1982); Simpson, Taxon 31 (1982) 517—528; Simpson & SKVARLA, Amer. J. Bot. 68 (1981) 277—294; Styer, J. Arn. Arb. 58 (1977) 100—145; VERKERKE, Blumea 29 (1984) 409—421; J. Arn. Arb. 66 (1985) 353—394. Bibliographical note. R. Cuopat published a monograph of the family in two parts in Mem. Soc. Phys. Hist. Nat. Genéve, Suppl. 1890 (1891) 1—143, t. 1-12 and ibid. 31 (1893) 1—500, t. 13-35. Because of frequent mention of this basic work and the complicated reference, I refer to this work in simpler form, as follows: CHopat, Monogr. I (1891) and CHopaT, Monogr. II (1893). 1988] POLYGALACEAE (van der Meijden) 459 KEY TO THE GENERA 1. Herbs, sometimes woody at base. 2. Lateral sepals larger than the 3 outer ones and petaloid, about as long as the petals. Lower petal (keel) appendiculate apically (except P. tatarinowii). Stamens 8. Fruit laterally dehiscent, the margins entire. Seeamoitha lobed arilat, micropylar SIGG)..05 aetna Geese cies chee oe loons 1. Polygala 2. All sepals subequal, not petaloid, much shorter than the petals. Keel apically inappendiculate. Stamens 2—6. Fruit either indehiscent, or laterally dehiscent and then the margins dentate or spinose. Seed ex- arillate. 3. Autotrophic plant with (small) green leaves. Fruit laterally dehiscent, dentate or spinose along the margins, far exceeding the sepals. Pericarp pergamentaceous. Style S-curved at base, long. Disk absent. Bmences + or 6, rarely 5. Rachis of inflorescence winged..--- 2-4-2 -2 sence ee ere 3. Salomonia 3. Saprophytic, echlorophyllous plant with scale-like leaves. Fruit indehiscent, enclosed by the sepals. Peri- carp fleshy. Style straight or very short. Disk present (but indistinct), adnate to the base of the ovary, semi-annular or as a lobe. Anthers 3 or 5, rarely 2 or 4. Rachis terete............. 4. Epirixanthes 1. Shrubs, trees or lianas. 4. Sepals united at base, upper part of calyx caducous by a circumcision. Petals basally adnate to the calyx. Ovary 7- or 8-locular. Fruit (by abortion) often 4- or 5-locular. Seeds completely enclosed in an aril 5. Eriandra 4. Sepals free, caducous or 3 or 5 persistent. Petals free from the sepals. Ovary 1- or 2-locular. Seeds without an aril, or aril lobed. 5. Petals 5. Lateral sepals not petaloid and less than twice as large as the other sepals. Ovary and fruit PeIcemiar with'4 OF MOTE OVUIES 25.2.2 | ML ER WO Pe ns eters 6. Xanthophyllum 5. Petals 3 or with an additional pair of much-reduced ones. Lateral sepals (alae) petaloid and at least twice as large as the other sepals. Ovary 1- or 2-locular, each locule containing a single ovule. 6. Ovary and fruit 1-locular. Fruit a distinct samara, indehiscent. Twigs with a pair of glands at the nodes. Bees WiLnOlt appendages; clabrous!s cs soe RN Oe eee 2. Securidaca 6. Ovary and fruit 2-locular. Capsule dehiscent, without a large wing. Nodal glands absent, rarely (6. P. sumatrana) pseudostipules present. Seeds either with a lobed aril, or very long-hairy... 1. Polygala 1. POLYGALA LINNE, Sp. Pl. 2 (1753) 701; Gen. Pl. ed. 5 (1754) 315; DC. Prod. 1 (1824) 321; BENTH. & Hook. Gen. PI. 1 (1862) 136; HAssk. in Miq. Ann. Mus. Bot. Lugd.- Bat. 1 (1863) 151; CHopaT, Monogr. I (1891); ibid. II (1893); E. & P. Nat. Pfl. Fam.3, 4 (1896) 330; BLAKE, N. Amer. FI. 25 (1924) 305; MUKHERJEE, Bull. Bot. Soc. Beng. 12 (1961, ‘1958’) 29; Hutcu. Gen. FI. Pl. 2 (1967) 340; ADEMa, Blumea 4 (1966) 256. — Chamaebuxus (DC.) Spacu, Hist. Nat. Vég. Phan. 7 (1838, ‘1839’) 125; Opiz, Oekon. Neuigk. Verh. (1839) 526. — Badiera (non DC.) HAssk. Cat. Hort. Bog. (1845) 227, p.p. — Semeiocardium (non ZOLL.) Hassk. in Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 150. — Fig. 1—16. Annual or perennial herbs, shrubs, small trees or rarely lianas. Stems or twigs rarely with nodal glands, these sometimes transformed into spines. Leaves alter- nate or verticillate, sometimes (sub)opposite, rarely caducous, sometimes scale- like or apparently absent. /nflorescences raceme-like or rarely paniculate, (supra-)axillary, terminal (and then often overtopped by side-branches) or lateral, sometimes in dense heads. Sepals 5 or the lower pair connate, unequal, caducous or persistent in fruit, the two lateral ones (alae) at least twice as large as the other ones and often petaloid. Petals 3, unequal, halfway adnate to the staminal tube and sometimes mutually connate into a single 3-lobed petal; some- 460 FLORA MALESIANA [ser. I, vol. 103 times with an additional pair of reduced lateral petals; lower petal (keel) boat- shaped, clawed, its blade entire or 3-lobed or basally auriculate, at apex with or less often without 2 entire or variously incised appendages (crest). Stamens 8, rarely 6, monadelphous or partly dia- or triadelphous; anthers usually bisporan- giate by abortion of the outer microsporangiae, or sometimes tri- or tetraspo- rangiate, sessile or on a free filamentous stalk, opening by an apical pore or a V-shaped introrse slit common to both cells. Disk annular or variously reduced or often apparently absent. Ovary 2-celled, each locule with a single subapical ovule; style tip various, often 2-lobed, the apical lobe often sterile and variously dilated, the stigmatic lobe lateral or subapical. Capsule 2-celled or very rarely 1-celled by abortion, compressed contrary to the sept, often more or less winged, sometimes with a double wing, dehiscing by a marginal split, reniform to oblanceolate. Seeds various, usually at micropylar side with a lobed or un- lobed appendage (aril) and/or with an elongate appendage, at the opposite cha- lazal side sometimes with a variously shaped appendage, rarely appendages at both sides lacking; glabrous to hairy, or rarely with a coma of hairs. Distr. A polytypic genus comprising at least 500 spp. in tropical, subtropical, temperate and montane regions. The majority of species grow in tropical South and Central America, where Polygala also has its greatest infrageneric diversity. Both in North America and in South Africa secondary centres of speciation occur. Eurasia, North Africa, Malesia and Australia are rather poor in species. In Malesia the species can be referred to four little-related sections. Sect. Melchiora is monotypic and endemic to New Guinea, sect. Pseudosemeiocardium is common in Southeast Asia, sect. Chamaebuxus occurs over all continents except tropical South and Central America, and the largest (probably unnatural) sect. Polygala covers the same area as the genus. The majority of species in Malesia have rather restricted areas, with some notable exceptions, e.g., P. persicariaefolia, which also occurs in tropical Africa, and P. paniculata, an American species which is now a pantropical weed. Two species are endemic in Malesia. Ecol. Species of sect. Chamaebuxus belong to the undergrowth of rain-forests, in Malesia occurring in submontane and montane forests, between 400 and 3000 m altitude. The single species of sect. Melchiora is a true liana of the tropical rain-forest. The remaining herbaceous species are heliophilous species growing in open terrains, in open woodland, with a preference for grasslands in seasonal areas; this is especially true for 9. P. javana, 12. P. longifolia, 14. P. exsquarrosa, 17. P. wightiana, and 18. P. rhinanthoides, which are all restricted to areas subject to a distinct or pronounced dry season. Pollination. Self-pollination probably occurs in all species, although the flowers of the majority are attrac- tive to insects. Many large-flowered species show structures which seem to be adaptations to pollinating insects (FAEGRI & VAN DER PuL, 1979, for P. chamaebuxus), but in those species, too, self-pollination seems to occur frequently. In a number of species pollen grains are deposited directly on the stigma, often already in the unopened flower (VENKATESH, 1956). In many species the style tip bears special pockets in various ways in which the pollen grains are deposited from the anthers. At that time the stigma can theoretically still be pollinated with foreign pollen grains, as for instance in P. /utea (MILLER, 1971) and P. vulgaris (HEUBL, 1984). According to HEUBL’s observations, however, a visiting insect will effect self-pollination. Yet there are rather reliable records of hybrids of the latter species in nature (HEUBL, /.c.), and F-plants of artificially produced hybrids are often fertile. This must mean that cross-pollination (rarely) occurs in P. vulgaris. Only BRANTJES (1982) observed and described cross-pollination in the two Brazilian species P. monticola H.B.K. and P. vauthieri Cuopat. The pollination mechanism of those species is a very precise one, and if cross-pollination fails, the flowers are effectively self-pollinated. Perhaps such examples of precision cross-pollination can be discovered in many other Polygala species, be- cause it is rather unlikely that the great variation in style tips and stigma forms could have evolved in complete absence of gene-exchange. It is also possible that many species in which formerly cross-pollination was rare, have lost this possibility. 1988] POLYGALACEAE (van der Meijden) 461 References: BRANTIES, Pl. Syst. Evol. 141 (1982) 41-52; FAEGRI & VAN DER Pu, Princ. Poll. Ecol. 3rd ed. (1979) 165, f. 6; HEuBL, Bot. Mitt. Miinchen 20 (1984) 234; MILLER, J. Arn. Arb. 52 (1971) 267; VENKa- TESH, Bull. Torrey Bot. Club 83 (1956) 19. Taxon. In view of the surprisingly great variation in species-constant characters of diverse flowering and fruiting parts, it has been tried to split this large genus into a number of smaller genera. The large genus con- cept as adopted by R. CHopatT (Monogr. I, 1891, 93), the only author who revised all species, is still adopted, some minor questions regarding the status of some American groups set aside. CHoDAT’s infrageneric division of the genus, however, was unsuccessful, as had been pointed out by S.F.BLAKE in his revision of the North American species (1924), and more recently by ADEMA (1966) for the Malesian ones. This is probably due to the fact that two of CHopAT’s major characters to define his sections, viz. the presence or absence of sepals at fruiting state, and of carinal appendages, are not constant in the different lineages: parallel developments (reductions c.g. reversions of character states) will have been developed in many of these lineages. Until a new revision of the tropical American species has been made, it will be impossible to make a balanced reconsidera- tion of the status of the diverse groups of species within and outside the borderlines of this genus. In the Indo-Australian area a great diversity of species is present, probably caused by diverse migration lines. Perhaps the Australian genus Comesperma should also be included in Po/ygala; see note under 4. P. papuana. KEY TO THE SECTIONS AND SPECIES 1. Lateral sepals (alae) caducous at the beginning of fruit-setting. Disk present. Leaves usually with distinct petioles. 2. Annual herbs up to 30 cm high. Flowers, fruits and seeds up to 3 mm long. Spp. 1—3. — 1. Sect. Pseudo- semeiocardium 3. Keel without appendages at apex. Seed without tubercles, without black appendage at chalazal side 1. P. tatarinowii 3. Keel with a pair of appendages near apex. Seed tuberculate, with a small or large black appendage at chalazal side. 4. Capsule narrowly winged, longer than wide. Seed at chalazal side with a cylindrical appendage one third aallonoianithe seed itself js oite 1 ees ae eee te nee oo mes 5. E. pallida 1. Sepals connate for 1/4—3/4. Bracts persistent till or after the fruits have fallen. 3. Flowers subtended by a bract and a pair of subulate bracteoles. Fruits fully enclosed by the sepals 2. E. papuana 3. Bracteoles absent. Apex of fruits not enclosed by the sepals. 4. Bracts lanceolate, 2—2.5 by 0.5—0.6 mm...... 4. Bracts elliptic to obovate, 1.5—2 by 0.8—1.2 mm 1. Epirixanthes elongata BLUME, Cat. (1823) 82; NEES, Flora 8 (1825) 133; REUTER in DC. Prod. 11 (1847) 44; ZoxL. Syst. Verz. 3 (1855) 58; Mia. Fl. Ind. Bat. 1, 2 (1858) 128; Hassk. in Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 143; PENzic, Ann. Jard. Bot. Btzg 17 (1901) 142, t. 24—26; Pitcer in E. & P. Nat. Pfl. Fam., Nachtr. 3 (1908) 190; Wirz, Flora 1 (1910) 395; BAcKER, Schoolfl. Java (1911) 75; MeErRR. En. Born. (1921) 325; STEEN. Trop. Natuur 23 (1934) 51, f. 10; Put, Rec. Trav. Bot. Néerl. 31 (1934) 761; SMITINAND, Thai For. Bull., Bot. 2 (1955) 2; BACKER & Baku./f. Fl. Java 1 (1963) 200; HANSEN c.s. Dansk Bot. Ark. 25, 2 (1967) 83. — E. linearis BLUME, Cat. (1823) 82; REUTER in DC. Prod. 11 (1847) 44; Mia. eAieeL Tee UCTY, PEEL oe ee 4. E. kinabaluensis SO. Sec Bead, cosgeve cc ete et 3. E. cylindrica Fl. Ind. Bat. 1, 2 (1858) 128. — Salomonia aphylla GriFF. Proc. Linn. Soc. 1 (1844) 221; Trans. Linn. Soc. 14 (1845) 112; BENN. Fl. Br. India 1 (1872) 207; KinG, J. As. Soc. Beng. 59, ii (1890) 132; CHoparT in E. & P. Nat. Pfl. Fam. 3, 4 (1896) 342; Josepu, Bull. Bot. Surv. India 12 (1970) 73, f. 1—5. — Salomonia parasitica GriFF. Not. 4 (1854) 538, t. 1598, f. 5. — E. tenella Hook.f. Trans. Linn. Soc. 23 (1862) 158; C. MULL. in Walp. Ann. 7 (1868) 243. — Salomonia elongata (BLUME) KuRz ex Koorp. Exk. Fl. Java 2 (1912) 453; RipLey, Fl. Mal. Pen. 1 (1922) 140. — E. aphylla (GriFF.) MERR. Philip. J. Sc. 13 (1918) Bot. 142. — Fig. 18. Herb, up to 25 cm. Stem simple or branched in up- Fig. 18. Epirixanthes elongata BLuME. (Photogr. A. ELSENER, Borneo, 28 May 1964). 1988] POLYGALACEAE (van der Meijden) 491 per half, 0.5—2 mm wide, violet or reddish, set with minute clavate hairs or papillae. Leaves 2—3.5 mm long, ciliate-pilose, usually violet. Spike 0.5—6 cm by 2—3 mm, rather dense, acutish at apex. Bracts ovate, c. 1.2 by 0.5 mm, cuspidate, ciliolate-papillose, erect, caducous before the flowers are full-grown; bracteoles absent. Flowers 2—2.5 mm long, yellowish white. Sepals free, subequal, c. 1 mm long, ciliolate- papillose. Anthers 5, rarely 4, sessile or stipitate. Disk lobe present adaxially at the base of the ovary, difficultly discernible, not accrescent after flower- ing. Ovary + orbicular; style longer than the ovary, c. 0.6 mm long, straight or apically slightly curved upwards, apically widened into the slightly 2-lobed stigma; gynoecium at first longer than the androe- cium, later equally long. Fruit reniform to broadly elliptic, c. 0.6 by 0.9 mm, enclosed by the slightly longer sepals. Distr. E. India, S. Burma (Tavoy), N. Vietnam, S. China; in Malesia: Malay Peninsula, Sumatra, W. Java, Borneo, Moluccas (Ambon, Ceram). 2. Epirixanthes papuana J.J.SmiTH in Fedde, Rep. 10 (1912) 286; Nova Guinea Bot. 8 (1914) 897; STEEN. Trop. Natuur 23 (1934) 51; BACKER & Baku. FI. Java 1 (1963) 200. — Salomonia cylindrica (BLUME) Kurz, p.p.: K.Scu. & Laut. Nachtr. Fl. Schutzgeb. Siidsee (1905) 285. Herb to 25 cm, generally purplish red in most of its parts except the corolla. Stem often repeatedly bran- ched in apical part, 1-3 mm wide, glabrous. Leaves 1.5—2.5 mm long, glabrous. Spike 0.5—6(—12) cm by 2.5—4.5 mm, very dense, rounded at apex. Bracts el- liptic to obovate, 1—2 by c. 1 mm, rounded apically, glabrous, patent with slightly upturned apex, per- sistent till or after the fruits have fallen, at last re- flexed; bracteoles always present, subulate, c. 1 mm long. Flowers 1.5—2.7 mm long, white but purple in bud. Sepals connate for 1/2—3/4, subequal, gla- brous. Stamens usually 3 and then anthers sessile, rarely 2 and then filaments free to halfway. Disk very indistinct, semi-annular, enclosing and fully adnate to the basal part of the lateral and lower side of the ovary, hardly accrescent in fruit. Ovary broadly el- liptic; style up to 0.1 mm long; stigma very asymmet- rical, 0.2 mm long, the upper stigmatic lobe oblique, rounded, the lower lobe narrower and longer, direct- ed a little downwards, hollow at apex; gynoecium shorter than androecium. Fruit broadly obovate, 0.7—0.9 by 0.7—0.9 mm, at base tightly enclosed by the fully adnate and very indistinct disk, the whole enclosed by the slightly longer sepals. Distr. Malesia: N. Sumatra (incl. Enggano 1.), W. Java, Borneo, Philippines (Luzon), Moluccas (Talaud, Ceram), New Guinea. Also in the Solomon Is. (San Cristobal). 3. Epirixanthes cylindrica BLUME, Cat. (1823) 82; NEEs, Flora 8 (1825) 133; REUTER in DC. Prod. 11 (1847) 44; ZoLL. Syst. Verz. 3 (1855) 58; Mia. Fl. Ind. Bat. 1, 2 (1858) 128, t. 15; HAsskK. in Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 143; Becc. Malesia 3 (1890) 325; Kina, J. As. Soc. Beng. 59, ii (1890) 132; PEN- zic, Ann. Jard. Bot. Btzg 17 (1901) 142, t. 20—23; Merr. Philip. J. Sc. 1 (1906) Suppl. 203; PiLGer in E. & P. Nat. Pfl. Fam., Nachtr. 3 (1908) 190; BacKER, Schoolfl. Java (1911) 76; WENT, Nova Guinea Bot. 8 (1909) 169; Koorp. Exk. Fl. Java 2 (1912) 453; Merr. En. Born. (1921) 325; En. Philip. 2 (1923) 386; STEEN. Trop. Natuur 23 (1934) 52; HEND. Mal. Nat. J. 4 (1949) 27; BACKER & BAKH./. Fl. Java 1 (1963) 200. — Salomonia cylindrica (BLUME) Kurz, J. As. Soc. Beng. 43, ii (1874) 79; Cuopat in E. & P. Nat. Pfl. Fam. 3, 4 (1896) 342; K.Scu. & Laut. Nachtr. Fl. Schutzgeb. Stidsee (1905) 285, p.p.; RmpLey, Trans. Linn. Soc. II, Bot. 9 (1916) 19. Herb, up to 25 cm, (sub)glabrous. Stem simple or branched, 1.5—2 mm wide. Leaves 2—4 mm long, slightly thickened at base, minutely sparsely papil- lose. Spike 0.5—6 cm by 4.5—5.5 mm, very dense, rounded at apex. Bracts elliptic to obovate, 1.5—2 by 0.8—1.2 mm, not thickened at base, at apex acutish and finely mucronate, imbricately covering the flower buds, patent with slightly upturned apex, per- sistent till or after the fruits have fallen, at last re- flexed; bracteoles absent. Flowers c. 2.5 mm long. Sepals connate + halfway, unequal, slightly cucul- late, c. 1.5 mm long. Anthers 3 or 5, rarely 4, sessile. Disk rather distinct, enclosing the lower and lateral side of the ovary and adnate to it, expanding gradual- ly in fruit and then visible as a rim near the base of it. Ovary broadly elliptic; style shorter than the ovary, subcylindrical, c. 0.1 mm long, unequally 2-lobed, the larger upper lobe stigmatic, the lower one tooth-like and hollow at apex; gynoecium some- what shorter than the androecium. Fruit broadly el- liptic, c. 1 by 1 mm, at base tightly enclosed by the disk, the whole enclosed by the somewhat shorter sepals. Distr. Burma; in Malesia: Sumatra, W. Java, Borneo, New Guinea. Note. Closely allied to the next species; see there. 4. Epirixanthes kinabaluensis WENDT, sp. nov. A E. cylindrica bracteis (ovatus-jlanceolatis, 2— 2,5 0,5—0,6 mm, parum crassinervatis ad basin, gradatim attenuatis et mucronatis, gemmis florium partiter expositis, persistentibus tempore fructifican- di differt. — Typus: CLEMENS 30039 (A holo; BO, G, L, NY, UC). See note | under the genus. Herb up to 30 cm, glabrous. Stem simple or branched, 1—2.5 mm wide, tinged reddish. Leaves 2~3.5 mm long, slightly swollen at base, brown. 492 FLORA MALESIANA [ser. I, vol. 103 Spike 1.5—6 cm by 4—5 mm, very dense, rounded at apex. Bracts (ovate-)lanceolate, 2—2.5 by 0.5—0.6 mm, at base slightly thickened along midrib, very gradually narrowed into the acute and finely mucro- nate apex, not fully covering the flower buds, patent with upturned apex at flowering time, persistent until the fruit has fallen, then reflexed and eventually ca- ducous; bracteoles absent. Flowers c. 2 mm long, milk-white with yellow-brown calyx. Sepals halfway connate, unequal, slightly cucullate, 1.5 mm long. Anthers 5, sessile. Disk rather distinct, enclosing the basal part of the lower and lateral side of the ovary and adnate to it, expanding gradually in fruit and then visible as a rim near the base of it. Ovary broad- ly elliptic; style shorter than the ovary, subcylindri- cal, c. 0.1 mm long, unequally 2-lobed, the larger up- per lobe stigmatic, the lower one tooth-like and hollow at apex; gynoecium somewhat shorter than the androecium. Fruit broadly elliptic, c. 1 by 1 mm, at base tightly enclosed by the disk, the whole en- closed by the somewhat shorter sepals. Distr. Malesia: Sumatra, Borneo. Note. Very closely allied to E. cylindrica, from which it differs only by the narrower and longer bracts. 5. Epirixanthes pallida WENDT, sp. nov. Sepala libera. Bracteae ellipticae, 2—3 x 1—1,5 mm, compressae ad basin, obtusae, marginibus hya- linis, pallidae, imbricatae apicem spicae tectae, ante tempus florendi caducae. Stylus brevis. — Typus: BROOKE 10498 (L). See note | under the genus. Herb up to 15 cm, glabrous, straw-coloured. Stem simple or branched in upper half, 2—2.5 mm thick. Leaves 2—4 mm long, swollen at base. Spike 1—5 cm long, 2.5—3 mm thick, very dense, rounded at apex. Bracts elliptic, 2-3 by 1—1.5 mm, at base com- pressed along the midrib, at apex obtuse, the margin nearly hyaline, pallid in dry state, imbricately cover- ing the younger part of the spike and completely hid- ing the flower buds, caducous at the beginning of flowering, thus leaving the fruiting calyx exposed; bracteoles absent. Flowers c. 1.8 mm long, whitish. Sepals free, unequal, the larger ones slightly cucul- late, 0.8—1 mm long. Anthers 5, sessile. Disk rather distinct, enclosing the basal part of the lower and lat- eral side of the ovary and adnate to it, expanding gradually in fruit and then visible as a rim near the base of it. Ovary elliptic, style shorter than the ovary, subcylindrical, c. 0.2 mm long, unequally 2-lobed, the larger upper lobe stigmatic, the lower one tooth- like and hollow at apex; gynoecium somewhat shor- ter than androecium. Fruit broadly elliptic or nearly didymous, c. 0.6 by 0.8 mm, at base tightly enclosed by the disk, the whole enclosed by the siightly longer sepals. Distr. Malesia: Borneo, Celebes. Note. A distinct species, easily recognizable by its relatively large bracts which imbricately cover the upper part of the spike but are very early caducous. 5. ERIANDRA ROYEN & STEEN. J. Arn. Arb. 33 (1952) 91, t. 1; STEEN. Blumea 12 (1964) 319; Buren. Gen. Fi Pl. 2:(1967)).343:. Tree. Twigs apparently without nodal glands. Leaves alternate, petioled. In- florescences fasciculate, axillary, short; bracts and bracteoles + persistent; bracts at very base with a pair of inconspicuous glands. Sepals 5 (or 4), equal, connate in basal half and adnate to the base of the petals, caducous as a whole together with the petals at the beginning of fruit-setting by a circumcision, the connate basal part persistent. Petals 5 or 4, connate for 3/4, basally adnate to the sepals and halfway to the staminal tube, equal, the free parts half-orbicular. Stamens (8—)10, monadelphous; anthers hairy all round, tetrasporangiate, the inner microsporangiae fused in upper half, shorter than the outer ones; de- hiscing at inner side by a sickle-shaped slit common to both loculi. Disk annular, inconspicuous, hairy. Ovary globose, 7—8-locular, each locule with 1 ovule; style terminal, straight; stigma capitate, slightly lobed. Fruit a fleshy globular berry, apically with a remnant of the style, basally with a remnant of the calyx tube. Seeds c. 4, flattened, radially, completely enveloped by an aril; testa very densely hairy; albumen fleshy. 1988] POLYGALACEAE (van der Meijden) 493 Distr. Solomon Islands; in Malesia: New Guinea. Monotypic. Ecol. Primary and secondary rain-forests, 0—150(—660) m. Notes. In flowering and fruiting characters resembling the tropical American genus Diclidanthera MartT., a genus of small trees and lianas differing in the 5-locular ovary (and fruit), the glabrous anthers, the constrict- ed corolla tube and the exarillate seed. As the nearly regular flowers and the many-celled ovary are probably derived in the family, it is probable that both genera are closely allied. STYER (J. Arn. Arb. 58, 1977, 124) studied the anatomy of Eriandra, and compared his results with those of the South American genera Barnhartia GLEASON, Moutabea AuBL. and Diclidanthera. He concluded (/.c. 139) that Eriandra, although a tree, has a ‘lianous ancestry’. 1. Eriandra fragrans ROYEN & STEEN. J. Arn. Arb. 33 (1952) 91, t. 1; STEEN. Blumea 12 (1964) 319. Tree up to 32 m, 70 m dbh. Twigs glabrous. Leaves oblong to (ob)lanceolate, 7—40 by 3—10 cm, nar- rowly cuneate at base, acutish or shortly acuminate at apex, coriaceous, glabrous above, slightly hairy on midrib beneath, in basal part beneath with 1—4 in- distinct glands 0.2 mm diam.; midrib sunken above in basal half, nerves S—16 pairs, finely prominent at both sides; petiole 1—2.5 cm, flattened above, wrin- kled below, apically with a pair of indistinct glands 0.2 mm diam. Axis of raceme-like fascicle 2-20 mm long, shortly hairy to glabrous, at the nodes near the bracts with a pair of inconspicuous, tiny, slightly prominent glands. Flowers S—6 mm long, white, fra- grant, on 1—5 mm long pedicels. Sepals obovate- spathulate, 4—5.5 mm long, glabrous except for the margin, the outer ones ciliate to base, the inner ones ciliate only apically. Petals free over c. 3 mm, the free parts reflexed during flowering; glabrous except for the ciliate incurved margins. Staminal tube glabrous outside, hairy inside; anthers elliptic, 3-locular in up- per half, 4-locular at base, c. 1 mm long, hairy all round. Ovary glabrous; style 3—4 mm long, densely hairy. Fruit 2—4 cm diam., apically with a hairy rem- nant of the style. Seeds elliptic, c. 1.5 by 1 cm; aril c. 0.2 mm thick; indumentum of testa very dense, 1—2 mm thick. Distr. Solomon Islands (New Georgia, Kolom- bangara); in Malesia: New Guinea. Ecol. Primary and secondary rain-forests, usual- ly in the valley bottoms or on plains near the coast, usually on well-drained sandy clay, sometimes on limestone, 0—150(—660) m. 6. XANTHOPHYLLUM Roxs. Pl. Corom. 3 (1820, ‘1819’) 81, nom. cons.; BENN. FI. Br. India 1 (1872) 208; CHopaT, Bull. Herb. Boiss. 4 (1896) 254; in E. & P. Nat. Pfl. Fam. 3, 4 (1896) 343; GAGNEP. in Desv. J. Bot. 21 (1908) 241; GAGnepP. FI. Gén. I.-C. 1 (1909) 242; CHopaT in Merr. Pl. Elm. Born. (1929) 133; Hutcu. Gen. FI. PI. 2 (1967) 339; No, Tree Fl. Mal. 1 (1972) 352; MeEDEN, Leiden Bot. Ser. 7 (1982). — Pelae [HERMANNUS, FI. Zeyl. (1717) 24 (‘Paelae’)| ADANSON, Fam. 2 (1763) 448, nom. rejic. — Eystathes Lour. Fl. Coch. 1 (1790) 235, nom. rejic. — Jakkia BLuME, Cat. (1823) 17; Bijdr. (1825) 60 (‘Jackia’), non Jackia WALL., nec Jackia SPRENGEL. — Skaphium Mia. FI. Ind. Bat., Suppl. (1861) 357. — Ba- nisterodes [L., Fl. Zeyl. (1747) 192 (‘Bannisterioides’)| O. K. Rev. Gen. Pl. 1 (1891) 45, nom. illeg. — Fig. 19-26. Shrubs or trees. Twigs terete, sometimes with annular or cap-like nodal glands. Axillary buds 2—8, serial, each with 2 budscales. Leaves alternate (or spiral), sometimes shifted-decussate, stalked, nearly always with glands beneath, mostly near base of blade. /nflorescence axillary, branched or sometimes unbranched, panicle- or raceme-like. F/owers solitary or in lower half with 3 (rarely more) together. Sepals 5, free, usually slightly unequal, usual- ly caducous after flowering. Petals 5, usually unequal, free; lower petal (carina) usually boat-shaped, clawed, apically inappendiculate. Stamens nearly always 494 FLORA MALESIANA [ser. I, vol. 10° 8, rarely 7, 9, or 8—10; 4 epipetalous, 2 placed at the base of the carina and ad- nate with it or not, 2 alternipetalous and opposite the lateral sepals; filaments free or connate basally, rarely halfway connate, very rarely triadelphous; an- thers tetrasporangiate, opening introrsely with slits. Disk annular. Gynoecium usually as long as androecium; ovary usually shortly stipitate, syncarpous, com- posed of 2 median carpels, 1-locular or sometimes semi-2-locular; style ter- minal; stigma slightly bilobed or sometimes peltate; ovules 2-seriate, 4 or (6—)8—c. 20, very rarely c. 40. Fruit indehiscent (or very rarely irregularly 2-valved, not in Mal.), globular or rarely ellipsoid, c. 1—15 cm, usually with a hard pericarp. Seed(s) 1 or 4—20, inappendiculate, glabrous; embryo large, flat or thick; albumen copious to nearly absent. Distr. About 94 species in tropical Southeast Asia, northwards to S. China and Hainan, throughout Malesia to Australia (N. Queensland); 76 spp. do not occur outside Malesia; 2 are endemic to Queensland. Not a single species crosses WALLACE’s plant-geographical demarcation line. Although only 7 spp. occur east of this line, at subgeneric level the greatest variation is found there. For that reason it is assumed that the genus originated in Austro-Malesia. Ecol. By far the majority of the species grow in the tropical lowland rain-forest, mostly below 500 m. About 10 spp. (of which 3 extra-Malesian ones) are usually found between 500—1200(—1500) m, but some- times also lower. About a dozen species are sometimes found in freshwater swamp forests, but they are not confined to that habitat, except 63. X. ramiflorum. Also nearly all of the continental Southeast Asian species seem to be confined to the rain-forest; 60. X. /anceatum is restricted to stream banks and swamps; 62. X. virens occurs in monsoon forests, tolerating rather long dry periods. Otherwise the ecology is for all species ‘rain-forest’, not specifically repeated. Note. From the subdivision of the genus it appears that both the fruit and seed, and the flower structure are important for specific distinction, in addition to vegetative characters. Unfortunately the mature fruit is yet unknown in several species (spp. 4, 16, 25, 34, 35, 36, 37, 38, 41, 51, 52, 58) and flowers are unknown from spp. 40, 49, 61, 67, 68, and 76. Besides, specimens in fruit seldom carry flowers too, and the reverse. These factors have complicated the key and necessitated to enter many species twice or even thrice. KEY ITO THE SUBGENERA 1. Fruit 4—more-seeded. 2 EmMbGEYOutnIck, covered by little endosperm. Spp).72—76 22. 6-6-0. -- ose V. Subg. Brunophyllum 2. Embryo flat, covered by copious endosperm. Spp. 69-71 ...........00cceeeee IV. Subg. Exsertum 1. Fruit 1- (exceptionally 2-)seeded. 3. Stamens triadelphous. Embryo flat, covered by copious endosperm. Spp. 64—68 Wl. Subg. Triadelphum 3. Stamens monadelphous or free. Embryo thick, without or with little endosperm. 4. Stigma peltate. Twigs at the nodes with a pair of annular glands. Sp. 63...... Il. Subg. Coriaceum 4. Stigma small, slightly bilobed. Nodal glands absent on the twigs. Spp. 1—62 I. Subg. Xanthophyllum KEY TO THE SSPECHLES 1. Petiole and lower side of leaves densely patently hairy, at least on the midrib. 2. Venation of 3rd order scalariform. 3. Inflorescence axis at the nodes with 2 crateriform or cup-like glands. 4. Leaves glaucous-papillose beneath, with 6—8 pairs of secondary nerves; laminar glands indistinct or on- PeaEMCTUAS A OUeS CISHNICE ... Bee eee ore en’ cee ota Peo ee Tee ee 3. X. sulphureum 4. Leaves smooth beneath, with 12 or 13 pairs of secondary nerves; laminar glands numerous, distinct 4. X. cockburnii 3. No glands present at the nodes of the inflorescence axis. 5. Leaves glaucous-papillose beneath. Ovary and fruit hairy all round or in 4 rows in upper half 12. X. rufum << 1988] POLYGALACEAE (van der Meijden) 495 5. Leaves smooth beneath. Ovary and fruit glabrous or hairy in 2 rows in upper half. 6. Inflorescence brownish hairy. Sepals persistent in fruit, the outer c. 2—3 mm long, the inner c. 3.5—5 EMITS 2 Se Odes leds UR PE Oe Se ae eee eee eee 11. X. velutinum 6. Inflorescence axis greyish white-hairy. Sepals caducous in fruit, the outer c. 4 mm long, the inner 5—6 errmmmripes (8 oo. JEG. bin tg blade kee ee eee Be eee 8. X. pubescens 2. Venation of 3rd order finely reticulate. 7. Ovules 4. Carina appressedly hairy outside, other petals appressedly hairy apically 57. X. malayanum 7. Ovules 8—16. Petals (sub)glabrous outside. 8. Sepals and inflorescence axis with hairs c. 1 mm long...................... 50. X. trichocladum 8. Sepals and inflorescence axis with hairs 0.1—0.2(—0.4) mm long. BoP eat-pide-bullate between: the tertiary Merves 2. a.) 2c). tre ek ee eee 49. X. reticulatum 9. Leaf-blade flat or only the midrib raised in a shallow depression. 10. Pedicel 2.5—4.5(—5) mm long. Anthers 0.9—1.4 mm long. Flowers 11—12 mm long 48. X. purpureum 10. Pedicel 7.5—14 mm long. Anthers 1.5—2.2 mm long. Flowers 12.5—16.5 mm long. 11. Petiole S—6 mm long. Flowers 16—16.5 mm long. Anthers c. 2.2 mm long 46. X. beccarianum 11. Petiole 1.5—2.5(—3) mm long. Flowers c. 12.5 mm long. Anthers 1.5—1.6 mm long 47. X. pedicellatum 1. Petiole and lower side of leaves shortly appressedly hairy or glabrous. 12. Leaves glaucous-papillose beneath. 13. Venation of 3rd order scalariform. Ovary hairy on median ribs. Fruits irregularly 4-sulcate 10. X. schizocarpon 13. Venation of 3rd order reticulate. Ovary glabrous or, if hairy, not especially hairy on the ribs. Fruit not 4-sulcate. 14. Ovary glabrous. 15. Ovules 4. Anthers 0.3—0.4 mm long. 16. Petiole (6—)7—11(—14) mm. Axillary buds very densely shortly hairy, not soon glabrescent. Inner sepais @°2-5 mm long. Anthers hairy at. base t joc. iis . tetas Shien 14. X. nigricans 16. Petiole 5—6.5 mm. Axillary buds soon glabrescent. Inner sepals c. 4 mm long. Anthers glabrous or wetinvery few: hairs-at base -.0sdie wa abot wr ath liws seas renee eo Sree aie 15. X. borneense 15. Ovules 8—18. Anthers at least 0.7 mm long. Risen EGS 3—7 im Ong f..e. x-oueys ie os Ewe Wea dee eee 43. X. penibukanense 17. Axillary buds up to 1 mm long. 18. Petiole (S—)8—12 mm long. Flowers up to 8.5 mm long. Fruit l-seeded, up to 1.2 cm long 63. X. ramiflorum 18. Petiole 1.5—3 mm long. Flowers 15—16 mm long. Fruit 10- or more-seeded, at least 5 cm long 72. X. brevipes 14. Ovary hairy. 19. Ovules 8—16. 20. Petiole distinctly transversely wrinkled. 21. Petiole 8-15 mm long, gradually widened into the attenuate leaf base. Lower pair of nerves reach- ing beyond the middle of the leaf-blade. ......6...cseccseceusiscnecse 43. X. penibukanense 21. Petiole 3—6(—7) mm long, abruptly ending into the cuneate or cordate leaf base. Nerves in the lower part of the leaf blade not reaching the middle of the blade ............ 42. X. discolor 20. Petiole not transversally wrinkled. 22. Ovary and style appressedly hairy. Petiole (8—)10—14 mm long. Nerves 11~—14 pairs 44. X. pseudoadenotus 22. Ovary and style patently hairy. Petiole 4—9 mm long. Nerves 6—13 pairs .... 45. X. pulchrum 19, Ovules 4. 23. Axillary buds at least 4 mm long. 496 FLORA MALESIANA [ser. I, vol. 103 24. (Upper) axillary bud situated (1.5—)3—15 mm above the leaf axil and placed on a 0.5—1.5 mm long Stale, (ya, ew, ae A EE ee: Reece eee 41. X. korthalsianum Note: Compare also 6/. X. lateriflorum. 24. Axillary bud situated in the leaf axil, sessile. 25. Scales of axillary bud at apex with 2 distinct thickenings..................... 36. X. clovis 25. Scales of axillary bud not thickened at apex. 26: Petrole 263 Uri. As Pee IP ERE CUE a eae nda She eet teat 35. X. petiolatum 26. Petiole (excl. attenuate leaf base) up to 17 mm. 27. Axillary buds half-patent to patent, with their flat side perpendicular to the twig 39. X. bracteatum 27. Axillary buds erect, usually with their flat side pressed against the twig. 28. Petals 6-8 mm long. Outer sepals 1.5—2.5 mm long, inner side 2.5—3.5 mm long. Anthers OF4=05 mmrlon gees «te Se Sas SIE, RN A TE Re 28. X. griffithii 28. Petals c. 13 mm long. Outer sepals c. 3—4 mm long, inner sepals 3.5—5.5 mm long. Anthers GROMPMM-LOMNE ee pe et ects SO, ee eee ee ae is ee oe ee 38. X. angustigemma 23. Axillary buds at most 1.5 mm long. 29. Leaf-blade 1.5—5(—6) by 0.5—1.5(—2) cm. Flowers solitary or in an up to 3-flowered inflorescence, 10S hemmulong Petalsve: 7 mmilong: Pedicel 1 -5—2i 5mm long: F222. eee see eee 58. X. novoguinense S4eseetals 10—12 mm long. Pedicelis—4-5) min ONG) = a2e6 een ec ae oles eee 59. X. ngii 12. Leaves smooth beneath. 35. Ovary hairy all round, not glabrous at lateral side. 36. Most axillary buds, at least in upper half of the twigs, 3 mm long or longer. 37. Axillary buds (8—)11—20(—30) by 6—12(—14) mm, + 1—3 times as long as wide, flat, at base attenuate and not thickened, at apex rounded to obtuse. (Flowers unknown) ....... 40. X. heterophyllum 37. Axillary buds up to 11 mm long and basally thickened, or more than 3 times as long as wide and + acute, or smaller than 8 mm. 38. Axillary buds erect, flattened against the twig, very densely shortly hairy...... 37. X. reflexum 38. Axillary buds half-patent to patent, glabrous or sparsely shortly hairy. 39. Axillary buds at base narrow, not thickened. 40. Leaf base cordate with upturned margins. Leaves with 14—20 pairs of nerves. Petals 16 mm long 25. X. brigittae 40. Leaf base attenuate and flat. Leaves with 5—7 pairs of nerves. Petals 10—12 mm long 31. X. incertum 39. Axillary buds at base wide and strongly thickened. 1988] POLYGALACEAE (van der Meijden) 497 41. Inflorescence unbranched. Petiole 4—6.5 mm. Leaves with 3—5 pairs of nerves 19. X. neglectum 41. Inflorescence branched. Petiole 8—14 mm. Leaves with 7 or more pairs of nerves. 42. Petiole (18—)25—30 mm. Leaf-blade 22—42 cm long, finer nervation beneath indistinct 34. X. ceraceifolium 42. Petiole 8—18 mm, exceptionally longer. Nervation at lower side of leaf-blade distinctly prom- inent, or leaf-blade much shorter than 20 cm. 43. Leaf-blade up to 3 times as long as wide. Petiole 8—14(—16) mm. Nerves (6 or) 7—9(—11) pairs. Leaf base never cordate. Anthers 0.4—0.6(—0.7) mm long.............. 30. X. vitellinum 43. Leaf-blade c. 4—6 times as long as wide. Petiole (8—)15—18(—24) mm, if shorter than 15 mm, then leaf base cordate. Nerves (9—)13—20 pairs. Anthers (0.6—)0.7—1.2 mm long. 44. Leaf base cuneate to rounded, or cordate with the margins curved upwards and connate above the apex of the petiole. Petals (8.5—)9.5—12.5(—14.5) mm long. Anthers (0.6—)0.7—0.9(—1) mimi Ong. 2c. sh bets ees Sos ee A a ee eee 32. X. adenotus 44. Leaf base cordate, flat, or only with little upturning margins. Petals 15—18.5 mm long. An- thers) lol? mmuloniore. sin es ccc ight at aeeceeee oom 33. X. palawanense 36. Most axillary buds up to c. 2 mm long. 45. Midrib sunken above over most of its length. 46. Petals + equal in size and form. Fruit black, 2—6 cm diam., seeds 4—12. (Compare also 70. X. IPITITETICATION:. (-daes en Sex cone hs DAES ees: ees =. Se ee ee eas 71. X. stipitatum 46. Petals unequal. Fruit not black, up to 2 cm diam., 1-seeded. 47. Flowering. ae. Upper. petals hairy. inside;to. near apex = sts-sieaw We Di sien Pees Ske Saeed eae ce 55. X. wrayi 48. Upper petals hairy inside only at base. 49. Nerves 12—20 pairs. Filaments 1—3 mm connate ..........6.2.2csceeess 56. X. venosum 49. Nerves 3—7 pairs. Filaments free at base. 50. Petiole glabrous or hairy only in the upper groove. Finer venation beneath not very distinct 54. X. eurhynchum 50. Petiole densely hairy all round. Finer venation at lower side very distinctly prominent 53. X. retinerve 47. Fruiting. 51. Fruit neither densely set with thick warts, nor ribbed. 52. Petiole glabrous or hairy only in the upper groove. Fruit densely hairy, smooth to slightly wrinkl- ed or 2—4-sulcate, sometimes finely rugose, up to 1.5 cm diam......... 54. X. eurhynchum 52. Petiole densely hairy all round. Fruit rather sparsely hairy, smooth, up to 2.2 cm diam. 53. X. retinerve 51. Fruit densely set with thick warts, or verrucose and ribbed. 53. Fruit apically shortly but distinctly beaked, strongly verrucosely ribbed .... 56. X. venosum 53. Fruit apically rounded, densely set with thick warts...... 54. X. eurhynchum, 55. X. wrayi 45. Midrib flat above or prominent, then sometimes with a groove in basal half. 54. Region of the twig above the leaf axil as well as the 2—4 very small and inconspicuous axillary buds densely shortly hairy. 55. Petiole glabrous or hairy only in the upper groove. Finer venation beneath not very distinct. Fruit either densely set with thick warts or densely hairy and smooth to slightly wrinkled or 2—4-sulcate 54. X. eurhynchum 55. Petiole densely hairy all round. Finer venation beneath very distinctly prominent. Fruit rather sparsely hairy, SMOOES. 02:34 asin: CIe Ree lec 8 00 De ee 53. X. retinerve 54. Twigs glabrous above the leaf axil. Axillary buds glabrous or sparsely hairy and soon glabrescent. 56. Axillary buds (2 or) 3—7, the upper one situated at least 1-2 mm above the leaf axil. 57. Upper axillary buds placed on a 1—2.5 mm long stalk. (Flowers unknown) 61. X. lateriflorum SI; Uppers axillary buds.sessile: Oni tHe tWe. cas abark wee mnie air west udanse odes 62. X. virens 56. Axillary bud(s) seemingly 1, or 2, rarely 3, close together in the leaf axil. 58. Ovules 8—12. (Compare also 7. X. ferrugineum) ........6..00 0005: 51. X. erythrostachyum 498 FLORA MALESIANA [ser. I, vol. 10° 58. Ovules 4. 59. Inflorescence branched. 60. Leaf-blade c. 4—6 times as long as wide. Petiole (8—)15—18(—24) mm. Nerves (9—)13—20 pairs. Anthers)(0.6—)0.7—0:9(—D)smmylongiewiy ocacten- Gossett ac eee ee oat 32. X. adenotus 60. Leaf-blade up to c. 3 times as long as wide. Petiole 8—14(—16) mm. Nerves (6 or) 7—9(—11) pairs. Anthers 0:4—0.6(—0.7 muon eter snes satin ee en ce a oles tte 30. X. vitellinum 59. Inflorescence unbranched or sometimes with a side-branch at base. 61. Ovary (half-)patently hairy, fruit patently hairy. 62. Nerves c. 8—10 pairs. Pedicel of flowers 4.5—6 mm, of fruits 8—11 mm long. Petals 13—19 mint long: Anthers:0'8 lez immitllongecssn. ast: a hee ee tes nates 24. X. ancolanum 62. Nerves 3—5 pairs. Pedicel of flowers c. 2 mm, of fruits c. 4.5 mm long. Anthers 0.3—0.4 mm long eb Si. PRED Watton spatiale ha woe ane ae els ead eats cists ee a) eer 19. X. neglectum 61. Ovary and fruit appressedly hairy. 63. Inner sepals 2—2.5 mm long. 64. Leaf base truncate to cordate. Carina auriculate. Filaments for 0.7—1 mm connate 21. X. tardicrescens 64. Leaf base cuneate to rounded. Carina gradually enlarged, not auriculate. Filaments free 18. X. subcoriaceum 63. Inner sepals 2.8—4.5 mm long. 65. Anthers 0.4—0.7 mm long. Gynoecium somewhat shorter than the carina. Laminar glands NUMELOUS epee eee ly eee Ae thee ae ei bre een et aes 26. X. tenuipetalum 65. Anthers 1—1.2 mm long. Gynoecium 0.5—2 mm longer than the carina. Laminar glands 1—6. 66. Leaf base cordate with upturned margins. Nerves 14—20 pairs. Petiole 15—18 mm 25. X. brigittae 66. Leaf base attenuate and flat. Nerves 5—7 pairs. Petiole6—10 mm . 23. X. philippinense 35. Ovary glabrous, at least at the lateral sides. 67. Nerves of 3rd order scalariform. 68. Inflorescence axis at the nodes with 2 crateriform glands. 69. Leaf base cuneate to rounded. Leaf-blade 5—25 by 2—12 cm. Nerves 4—10 pairs 1. X. flavescens 69. Leaf base cordate. Leaf-blade 20—50 by 6.5—20 cm. Nerves 14—20 pairs....... 2. X. bullatum 68. No glands present on inflorescence axis. 70. Ovary and fruit hairy on 4 ribs in apical half, hairs brown ............. 13. X. macrophyllum 70. Ovary and fruit glabrous or hairy in 2 rows, hairs greyish. (Virdribstlatibeneathi(prominentrabove) estes. e-mate ere. toca see ee mes 9. X. resupinatum 71. Midrib prominent beneath (prominent to sunken above). 72. Inflorescence axis branched or unbranched, very densely set with flowers or scars of pedicels 5. X. hosei 72. Inflorescence axis branched, except in upper part not densely set with flowers or scars of pedicels. 73. Pedicels 1—1.5(—4) mm. Sepals thickened basally, not fleshy, when dry medium brown with light coloured margin; outer sepals very sparsely minutely hairy ............. 6. X. ferrugineum 73. Pedicels (2—)4—10 mm long. Sepals, if thickened, somewhat fleshy, without a lighter coloured Maren: Outer sepals more densely hdinyea nes ee ee elec ee eee ee 7. X. affine 67. Nerves of 3rd order reticulate. 74. Ovules 4. 75. Petiole distinctly transversally wrinkled, 3.5—4.5(—6.5) mm. Nerves 3 or 4 pairs. Inflorescence and pecreers (subjelabrous . «.«5d0.4.50%020. ae eigeahtaeeante Se Oe 16. X. ovatifolium 75. Petiole smooth, 6—11 mm. Nerves 4—6 pairs. Inflorescence and pedicels more or less densely shortly Dainy aeetatess ket 2B), to 08) We Bb ie te Oe Ten Sed . leet Le en ee 17. X. tenue 74. Ovules 8 or more. 76. Laminar glands rather numerous and distinct, 0.5—1 mm diam., nearly all situated in the ‘axil’ of secondary nerves and midrib. (Flowers and ripe fruits unknown) .......... 68. X. hildebrandii 76. Laminar glands scattered or situated at the margin, not in the ‘axil’ of secondary nerves and midrib. 1988] POLYGALACEAE (van der Meijden) 499 77. Flowering (flowers unknown in 67. X. contractum and 76. X. chartaceum). 78. Stamens triadelphous, i.e., 6 stamens connate in 2 groups, and the remaining 2 stamens connate. 79. Laminar glands sometimes situated near margin of leaf-blade but never present on the margin RSeie Antners completely: Pla brOUS 4) cae ee eee ce ee eee eee 66. X. montanum 79. Leaf-blade in upper half with at least 6 glands situated on the margin. Anthers shortly hairy along slits and at base. 80. Filaments and style densely and rather shortly patently hairy ........... 65. X. celebicum enaments;and style.clabrous/sry9-)..9!> eetts 15 7b. Es eee eee 64. X. ellipticum 78. Stamens either free or all connate except between upper petals. 81. Petals unequal, the lower middle one (carina) very distinct from the upper ones. 82. Petals black when dry. Twigs at the nodes with rather distinct annular glands. Filaments partly Beewwithr lanate Hairs Vell, estes eaters coe Cie Sete ee cee ee ORS ey ene 73. X. obscurum 82. Petals reddish orange when dry. Twigs without nodal glands. Filaments with short, straight marsimear Bases fr). ra G4- ast fa tes Lee ela che tate nets Rela a pee ee eee 52. X. laeve 81. Petals nearly equal in size and form. 83. Pedicel 1.5—4 mm. Stamens shorter than petals. 84. Filaments (nearly) free. Petals 7—8.5 mm long. Outer sepals 2.5—3.5 mm long, inner sepals Se NT NONE chr. Bates as stele oo aoe arcene Carre hens AC A Ee a ae 74. X. papuanum 84. Filaments connate to nearly halfway. Petals 9.5—12 mm long. Outer sepals 4—5.5 mm long, mmner sepals,.6= 7mm donets hy! 66 /sc7se.is See SR a ee eee 75. X. ecarinatum 83. Pedicel at least 8 mm. Stamens longer than petals. 85. Petals 14—16 mm long. Stamens 17—22 mm long. Anthers hairy from base to apex, 1—1.3 mm DE ee Siar Reenetnets ee ate. giae Rin Westie eae ot omits Clee Ace ene iE Salant a c 69. X. suberosum 85. Petals 7-12 mm long. Stamens 11—13(—16) mm long. Anthers glabrous or with a few hairs at base, 0.7—0.9 mm long. 86. Petals (8—)9—11(—12) mm long. Style hairy to apex. Laminar glands 6—10(—20) 70. X. amoenum 86. Petals 7—7.5(—8) mm long. Style glabrous in upper half. Laminar glands 0—2(—4) 71. X. stipitatum 77. Fruiting. 87. Fruit small, less than 2 cm diam., 1- (or 2-) seeded (mature fruit not known in 52. X. /aeve and 67. X. contractum). 88. At least 6 glands situated on the leaf margin itself, in upper part of the leaf-blade. aE ACL. 3—O(—8)) SIN C2 4:4: 2.6'5/ ace piem, orci aco oats ene erstet ares Aaa ee eee 64. X. ellipticum PE OAICOM T= 9 5. TIN ok ost eunlarmayepevegaed Sy ty Paks oie weg A wo tie Gaeta: Meee 65. X. celebicum 88. Laminar glands absent from margin of leaf-blade. SO SMSATIb (NrOminent aADOVE sis: - eis) 20 Mosere se 9 Sy elae cle ares Wie. Saree 67. X. contractum 90. Midrib sunken above. 91. Twigs at the nodes with distinct annular glands. Pedicel 4—5.5 mm.... 66. X. montanum 91. Twigs without nodal glands. Pedicel 8-15 mm. (Fruits unknown) .......... 52. X. laeve 87. Fruit large, many-seeded. 92. Pericarp strongly wrinkled outside when dry. 03. bruit globular to, pear-shaped, blacks... sasied os austin meat ne Oealecaets 76. X. chartaceum 93. Fruit ellipsoid, attenuate at apex and base when dry, orange to brown.. 75. X. ecarinatum 92. Pericarp remaining smooth when dry. 94. Fruit black. Seeds with copious albumen, with thin cotyledons ......... 69. X. suberosum 70. X. amoenum 71. X. stipitatum 94. Fruit not black. Seeds (nearly) without albumen, with very thick cotyledons. 95. Seeds sticking to the pericarp when dry .....:.cccecoscsecevnsesnens 74. X. papuanum 95. Seeds sticking together in drying, leaving shining reddish marks on inner side of the pericarp 73. X. obscurum 500 FLORA MALESIANA [ser. I, vol. 103 I. Subgenus Xanthophyllum Nodal glands absent. Axillary buds sometimes very long, usually thickened basally, subacute, sometimes hairy. Petiole sometimes with glands, sometimes hairy. Leaf-blade sometimes hairy, base rarely cordate; tertiary nerves finely reticulate, or scalariform. /nflorescence usually branched. Sepals usually dense- ly minutely greyish hairy out- and inside, sometimes with longer or differently coloured hairs, the inner ones often subglabrous in marginal area and often slightly keeled. Petals unequal, ciliate above base; carina unguiculate, boat- shaped, auriculate or not, usually densely hairy outside; lateral and upper petals narrow, usually oblanceolate to + linear, more or less flat, slightly constricted basally, usually glabrous outside, usually slightly hairy inside above insertion of filaments, upper petals often reflexed. Ovary sometimes semi-2-locular, glabrous to hairy; style usually hairy; stigma small, slightly 2-lobed, very rarely wider than apex of style; ovules 4 to c. 20. Fruit indehiscent, usually globular, usually c. 1.5—2 cm diam. Seed(s) 1 (or 2), rarely up to 4; testa 1- or 2-layered; albumen present or not; embryo globular, plumule not differentiated. KEY TO THE SECTIONS 1. Tertiary nerves scalariform. Testa sticking to the inner side of the fruitwall in drying. Albumen absent. Spp. Ha ESI I RES isons ere NR Oc HIE GE PEO CRETE SCENT ceed ANCE NRE TT 1. Sect. Xanthophyllum 1. Tertiary nerves finely reticulate. Testa sticking to the rest of the seed in drying. Albumen present, though SOMLEMMMESTSCATCO SS 0]2) UA — Ole crea reastentre rene fe Scr sreMeye aD eve aifavrayene) cysts siese 0s ine [a shen BveLOneae 2. Sect. Eystathes 1. Section Xanthophyllum Twigs usually densely patently hairy, sometimes glabrous. Petio/e usually more or less smooth, sometimes with glands. Leaf-blade: tertiary nerves scalariform, prominent. Petals: carina usually shortly unguiculate. Ovary sometimes semi-2-locular (the marginal areas of the carpels touching the other ones along their length), glabrous, sometimes hairy in 2—8 rows, rarely hairy all round; ovules 6—16, or (not in Mal.) 4—10, in 7. X. affine very rarely 5. Fruit globular or sometimes pointed or 4-sulcate, smooth or sometimes tuberculate. Seed(s) 1 or occasionally 2 (in /. X. fla- vescens often 2); testa reduced, without hard inner layer, sticking to the pericarp in drying, the raphe widely branched, hypostase not developed; albumen absent in ripe seed; radicle not exserted. 1. Xanthophyllum flavescens Roxs. Pl. Corom. 3 (1820) 82, t. 284, f. 2; WaLL. Cat. (1831) 4198; Roxs. Fl. Ind. ed. Carey 1 (1832) 221; WaLp. Rep. 1 (1842) 248; Drury, Handb. Ind. FI. 1 (1864) 56; Kurz, J. As. Soc. Beng. 42, ii (1873) 79, 80; Prelim. Rep. For. Pegu (1875) 26; For. Fl. Burma | (1877) 81; Prain, Bengal PI. (1903) 236; BRANDis, Indian Trees (1906) 44, excl. syn.; MEIDEN, Leiden Bot. Ser. 7 (1982) 64. — X. excelsum (BLUME) Mia. FI. Ind. Bat. 1, 2 (1858) 129; Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 274; Bin- NEND. Ann. Jard. Bot. Btzg 1 (1876) 172; Kinc, Mat. Fl. Mal. Pen. (1890) 142, p.p. (sub X. affine); K. & V. Icon. Bog. 1, 1 (1897) 11, p.p. (sub X. affine B ex- celsa); BOERL. Cat. Hort. Bog. (1899) 58; K. & V. Bijdr. Booms. 5 (1900) 298; Racrs. Ann. Jard. Bot. Btzg 17 (1900) 53; BoorsMa, Bull. Dép. Agr. Ind. Néerl. 16 (1908) 3; GAGNEP. in Desv. J. Bot. 21 (1908) 253; Fl. Gén. I.-C. 1 (1909) 246; BACKER, Schoolfl. Java (1911) 80; Koorp. Exk. Fl. Java 2 (1912) 453; Merr. En. Born. (1921) 326, p.p.; RIDLEY, Fl. Mal. Pen. 1 (1922) 143, p.p. (sub X. af- fine); MERR. En. Philip. 2 (1923) 386, p.p.; BuRK. Gard. Bull. S. S. 3 (1923) 35, p.p.; ENDERT, Tectona 18 (1925) 97; DocTERS VAN LEEUWEN, Zoocecidia ni Me 1988] POLYGALACEAE (van der Meijden) 501 (1926) 273; HEYNE, Nutt. Pl. (1927) 901; Cras, FI. Siam. En. 1 (1931) 105; Burk. Dict. (1935) 2268; GaGneP. FI. Gén. I.-C. Suppl. 1 (1939) 222; MASAMUNE, En. Phan. Born. (1942) 380; Wyatrt- SmiTH, Mal. For. Rec. 17 (1952) 80, 363, p.p.; ibid. 23? (1963) f. 10, p.p. (sub X. affine); BACKER & BakH./. Fl. Java 1 (1963) 201; No, Fed. Mus. J. n.s. 13 (1971) 137; Tree Fl. Mal. 1 (1972) 354, f. 1, p.p. — Jakkia excelsa BLUME, Bijdr. (1825) 62 (‘Jackia’); G.Don, Gen. Hist. 1 (1831) 368. — Monnina excelsa (BLUME) SPRENGEL, Syst. Veg. 3 (1827) 265; STEUD. Nom. ed. 2, 2 (1841) 157. — X. adenopodum Mia. Fl. Ind. Bat., Suppl. (1861) 393; Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 277. — Banisterodes excelsum (BLuME) O. K. Rev. Gen. PI. 1 (1891) 46, nom. illeg. — X. affine var. adenopodum (Miq.) K. & V. Icon. Bog. 1, 1 (1897) 11. — X. glandulosum MERR. Philip. Gov. Lab. Bur. Bull. 35 (1905) 34; GAGNEP. in Desv. J. Bot. 21 (1908) 252; MerR. En. Philip. 2 (1923) 485. — X. floriferum Emer, Leafl. Philip. Bot. 5 (1913) 1674. — X. multiramosum ELMER, l.c. 1676; MerRR. Philip. J. Sc. 27 (1925) 29. — X. loheri Merr. /.c. 29. — X. pallidum Ri.ey, Kew Bull. (1938) 113; MASAMUNE, En. Phan. Born. (1942) 380. Tree, 8—36 m high, up to 1 m dbh. Twigs glabrous. Petiole (5—)7—15 mm, often seemingly longer when leaf base is attenuate, mostly in apical half with two rather distinct glands. Leaf-blade (5.2—)7—18(—25) by (2.3—)3—7(—12) cm, base often long attenuate with the margins of the leaf-blade curved upwards and gradually grading into the ridges of the petiole; above yellowish green to yellowish brown; beneath: secondary nerves (4—)6—8(—10) pairs, usually form- ing a rather distinct intramarginal nerve, venation sometimes indistinct; glands 1-10, scattered, 0.3—0.6(—0.9) mm diam., basal glands usually ab- sent if petiole bears glands. /nflorescences often several together on one thickened node, rarely un- branched, as long as or longer than the leaves; axes slender, somewhat flattened, slightly grooved, thick- ened on the nodes and there with usually distinct, protruding glands, brownish, densely minutely pa- tently hairy. Flowers usually with 3 together. Pedicel (2.5—)3—4.5 mm, rarely longer, densely minutely half-patently greyish hairy. Sepals: outer sepals 2.3—3(—3.6) by 1.4—2.7 mm; inner sepals 3—4.1 by 2.3—3.7 mm. Petals bright yellow, or white and the upper petals with a yellow spot, or fully white, when dry yellowish, the longest one (6—)7—8(—8.5) mm long; carina basally more or less densely hairy, in up- per part outside densely patently hairy to glabrous, inside rather densely hairy to glabrous; lateral petals usually glabrous outside, mostly glabrous inside, up- per petals reflexed, ciliate basally or up to 2/3 their length, usually glabrous outside, inside patently hairy or at base only. Stamens: filaments very rarely ¢. 1 mm connate; anthers (0.4—)0.5—0.6(—0.65) mm long. Ovary glabrous to hairy in apical region on 4 ribs, very rarely (in one coll.) loosely hairy all over, often semi-2-locular; style sometimes glabrous in basal part; ovules 8—12, rarely 6. Fruit 1- to often 2-locular, the second chamber often reduced and seemingly enveloped by the pericarp; usually globular, up to c. 2 cm diam., smooth to irregularly low-tuberculate, rarely densely tuberculate, rather dull to somewhat shiny, yellowish to brownish; peri- carp usually hard; pedicel up to 6(—7) mm long. Seed(s) 1 or 2. Distr. Throughout tropical continental SE. Asia; in Malesia: Sumatra (incl. Simalur, Banka), Malay Peninsula, Borneo, Philippines (Luzon, Mindoro, Palawan, Masbate, Guimaras, Negros, Mindanao). Note. Closely resembling 7. X. affine, and nearly always combined with that species in one taxon. The only reliable character distinguishing it from the lat- ter species is the presence of nodal glands in the inflo- rescence. Due to the variability of X. affine it is diffi- cult to identify sterile collections of both species. Dif- ferences are found in the petiole; in _X. flavescens the petiole is glabrous and often bears a pair of glands, in X. affine the petiole is minutely hairy, and eglan- dular. 2. Xanthophyllum bullatum KiNG, J. As. Soc. Beng. 59, ii (1890) 142; BurK. & HEND. Gard. Bull. S. S. 4 (1928) 222; No, Tree Fl. Mal. 1 (1972) 356, f. 1; Corner, Gard. Bull. Sing. Suppl. 1 (1978) 146; Mev- DEN, Leiden Bot. Ser. 7 (1982) 66. Shrub or small tree, 3—4.5 m high, 6 cm dbh. Twigs appressedly hairy. Petiole c. 12—22 mm, rather densely minutely appressedly hairy. Leaf- blade 20—50 by 6.5—25 cm; base cordate; above yellowish green, bullate between midrib and second- ary nerves, midrib strongly prominent to flat, sec- ondary nerves rather strongly prominent, venation finely prominent; beneath creamish brown to light brownish green, minutely hairy on the nerves, sec- ondary nerves 14—20 pairs, protruding, mostly forming a nearly complete, rather distinct intramar- ginal nerve, venation sometimes rather obscure; glands rather few and mainly along the margin, or numerous and scattered, 0.3—0.6 mm diam., basal glands (1 or sometimes 2 pairs) at very base on mid- rib. Inflorescences shorter than the leaves, mostly elongate and often very densely set with flowers; axes appressedly to patently minutely hairy, light to very dark brownish, with large, protruding nodal glands. Pedicel 5—7 mm, very densely minutely appressedly reddish brown hairy. Sepals minutely appressedly reddish brown hairy; outer sepals c. 4.5 by 3 mm, with or without protruding glands; inner sepals c. 6.5 by 3.7 mm, keeled. Petals white, when dry yellowish, the longest one 12 mm long; carina in apical part rather shortly appressedly hairy outside, minutely so 502 FLORA MALESIANA [ser. I, vol. 10° inside, further glabrous; other petals glabrous out- side, glabrous or ciliate at base, shortly hairy or not inside in basal part, the lateral petals strongly bent, the upper petals strongly S-curved. Stamens: fila- ments glabrous or sparsely shortly hairy at base; an- thers 0.8—1.2 mm long, ciliate along slits. Ovary slightly ribbed; style glabrous in basal part, apically shortly appressedly hairy; ovules 8—10. Fruit globu- lar, up to 2 cm diam., dull, rather smooth, light brown; pericarp rather thin, not very hard. Distr. Malesia: Malay Peninsula (Perak, Treng- ganu, Selangor). 3. Xanthophyllum sulphureum KiNG, J. As. Soc. Beng. 59, ii (1890) 143; GAGNEP. in Desv. J. Bot. 21 (1908) 252 (‘sulfureum’); RipLey, Fl. Mal. Pen. 1 (1922) 145; Wyatt-SmitH, Mal. For. Rec. 17 (1952) 81, 363; Nc, Tree Fl. Mal. 1 (1972) 363, f. 4, excl. FRI 8041; MepEN, Leiden Bot. Ser. 7 (1982) 67, f. 10c. Small to large tree, 30-50 m high. Twigs finely reticulately wrinkled, brownish hairy, glabrescent. Axillary buds up to 2.8 mm long but often much smaller, rounded, densely hairy. Petiole 9-13 mm, very densely patently hairy. Leaf-blade 9—20 by 2.8—7.5 cm; above yellowish green, dull, midrib basally slightly prominent to slightly sunken and rather densely hairy, upwards sunken and glabrous, nerves and venation rather obscure; beneath glaucous-papillose, rather densely hairy especially on the nerves, secondary nerves 6—8 pairs, forming a rather distinct intramarginal nerve; basal glands per- haps always present but mostly hidden by the very prominent midrib and then scarcely visible, rarely not so and then distinct, c. 0.5 mm diam., other glands apparently few or possibly absent. Jnflores- cences shorter than the leaves; axes rather smooth, very densely brownish hairy, with distinct, large, prominent ‘cap-like’ nodal glands. Pedicel c. 4 mm, rather stout, very densely patently brownish pubes- cent. Sepals densely brownish patently shortly hairy at both sides; outer sepals 4 by 3.7 mm; inner sepals 5 by 4.2 mm, some with glandular spots, distinctly keeled. Petals yellow, when dry yellowish, the long- est one 11 mm; carina very densely (nearly velvety) patently pubescent outside, densely pubescent inside in apical part; other petals woolly-tufted outside. Stamens : filaments nearly glabrous; anthers 0.5 mm long, cohering around the style. Ovary containing 16 ovules. Fruit globular, c. 2cm diam., light brownish, dull, finely tuberculate; pericarp hard. Distr. Malesia: Malay Peninsula (Perak, Kelan- tan, Pahang). 4. Xanthophyllum cockburnii MEIDEN, Leiden Bot. Ser. 7 (1982) 67. Tree, 20 m high, 30 cm dbh. Axillary buds densely patently hairy. Petiole 8-12 mm, densely hairy. Leaf-blade 11—17 by 3.5—7 cm, base rounded to ob- tuse; above slightly bullate between midrib and sec- ondary nerves, greenish or brownish yellow, midrib rather obscure, finely prominent to flat, secondary nerves obscure; beneath creamish to greenish yellow, rather densely patently hairy on the nerves, second- ary nerves 12 or 13 pairs, forming a complete intra- marginal nerve, fine venation obscure; glands c. 1S—20, 0.2—0.4 mm diam., those at very base distinct, situated on midrib. /nflorescences shorter than to as long as the leaves; axes very densely red- dish brown pubescent, with rather large nodal glands. Pedicel 4.5 mm. Sepals densely patently red- dish brown hairy; outer sepals 3.8—4 by 3 mm; inner sepals 3.8—4 by 3—3.3 mm, keeled. Petals white, when dry yellowish, the longest one 11 mm; carina light yellowish velutinous outside, rather densely hairy inside in apical part; other petals glabrous or slightly hairy apically, not ciliate at base. Stamens: anthers 0.8—0.85 mm long, minutely hairy. Ovary containing 8 ovules. Fruit unknown. Distr. Malesia: Malay Peninsula (Pahang, near Johore border, Upper Anak Endau R.). Once col- lected. 5. Xanthophyllum hosei Ripley, Kew Bull. (1938) 113; MASAMUNE, En. Phan. Born. (1942) 380; MEu- DEN, Leiden Bot. Ser. 7 (1982) 69. Small tree, up to 3.5 m high, dbh 3 cm. Twigs minutely appressedly hairy, soon glabrescent, or gla- brous. Petiole 7-10 mm, minutely appressedly hairy. Leaf-blade 12—21 by 4.5—10 cm, base round- ed to more or less obtuse, apex rather suddenly cuspidate to acuminate; above slightly bullate be- tween midrib, secondary nerves, and intramarginal nerve, dark greyish green, midrib sunken, rather wide, other nervation finely prominent; beneath lighter coloured, sparsely minutely appressedly hairy in basal part especially on the nerves, secondary nerves 7—11 pairs, forming a distinct, (nearly) com- plete intramarginal nerve; glands 8 to very numer- ous, scattered, (0.2—)0.3—0.4 mm diam., basal glands larger. J/nflorescences branched or un- branched, up to 12 cm long; axes slightly thickened, curved, smooth, dull, minutely very densely appres- sedly hairy, internodes (except at very base) very short, c. 0.5—1 mm long, thus giving the inflores- cence a very dense-flowered appearance. Pedicel 2.5—4 mm, grooved, densely minutely appressedly hairy. Sepals glabrous to sparsely minutely appres- sedly hairy inside; outer sepals 2.1—2.5 by 2.5—2.8 mm, with 2—4 glandular spots, minutely densely ap- pressedly hairy outside; inner sepals 3.5—3.8 by 2.2—2.6 mm, rather distinctly keeled and there densely minutely appressedly hairy, further more or less glabrous outside. Petals yellowish when dry, the 1988] longest one c. 8.5 mm; carina sparsely appressedly hairy outside in middle part, glabrous inside; other petals ciliate to halfway, further glabrous. Stamens: filaments wide at base, sparsely appressedly hairy adaxially and abaxially to + halfway, further gla- brous; anthers 0.85—1 mm long, ciliate along slits. Ovary nearly sessile, ribbed; style glabrous in basal 1/3 part, upwards rather densely appressedly hairy till the apex; ovules 7—10. Fruit (very young) not ribbed, finely pustulate; style persistent, recurved; sepals persistent. Distr. Malesia: Borneo (Sarawak). 6. Xanthophyllum ferrugineum MEUDEN, Bot. J. Linn. Soc. 67 (1973) 118; Leiden Bot. Ser. 7 (1982) 69. Tree, up to 25 m high, up to 40(—60) cm dbh. Twigs glabrous. Petiole 9-14 mm. Leaf-blade 8—20 by 3—7 cm; above bright yellow to yellowish green, nerves finely prominent to rather obscure but mostly more distinct above than beneath; beneath midrib not very prominent, secondary nerves c. 5—7 pairs, finely prominent to rather obscure, intramarginal nerve indistinct, tertiary nerves finely prominent to obscure, fine venation mostly obscure; glands rather few, mainly situated near margin, 0.4—0.6 mm diam., basal glands similar. /nflorescences shorter than the leaves; axes more or less flattened, yellowish brown, densely to rather sparsely shortly patently hairy. Pedicel 1—1.5(—4) mm, densely minutely ap- pressedly greyish hairy. Sepals possibly persistent in fruit, medium-brown with light brown margin when dry; outer sepals 3.7—4.8 by 3.5—4 mm, (sub)gla- brous outside, often with (sometimes minute) glan- dular spots; inner sepals 4—5 by 4—4.2 mm, thick- ened in middle basal part, minutely appressedly hairy especially on the midrib, glabrous along the margin. Petals yellow, or white and the upper petals with a yellow spot, when dry yellowish, the longest one 8.5—10 mm long; carina very densely patently pubes- cent outside, densely pubescent in apical part inside; other petals glabrous or with few hairs in apical part outside. Stamens: anthers 0.5—0.7 mm long, minute- ly hairy. Ovary smooth or ribbed, glabrous, rarely sparsely appressedly hairy all round, apically often greyish hairy in 4 short rows; ovules 8—14. Fruit (im- mature) partly enclosed by the persistent sepals, ovoid, finely tuberculate, dull, yellowish brown. Distr. Malesia: Borneo (Sarawak, Brunei, Sabah, Tawau). 7. Xanthophyllum affine Kortu. ex Mig. Ann. Mus. Bot. Lugd.-Bat. | (1864) 271; Kurz, J. As. Soc. Beng. 42, ii (1873) 80; Benn. Fl. Br. India | (1874) 209; Kurz, For. Fl. Burma | (1877) 82; F.-ViLt. Nov. App. (1880) 14; Kina, J. As. Soc. Beng. 59, ii (1890) 142, p.p.; Cuopat in E. & P. Nat. Pfl. Fam. 3, 4 POLYGALACEAE (van der Meijden) 503 (1896) 345; K. & V. Icon. Bog. 1, 1 (1897) 11, p.p. (a genuina, t. 11, p.p., excl. f. 12-14; Rm ey, J. Str. Br. R. As. Soc. n. 33 (1900) 45; BRANDIS, Indian Trees (1906) 44; GAGNEP. in Desv. J. Bot. 21 (1908) 253; Rmp.ey, J. Fed. Mal. St. Mus. 4 (1909) 5; J. Str. Br. R. As. Soc. n. 59 (1911) 73; MERR. En. Born. (1921) 326, p.p. (sub X. excelsum); RIDLEY, Fl. Mal. Pen. 1 (1922) 143, p.p.; BuRK. Gard. Bull. S. S. 3 (1923) 35, p.p. (sub X. excelsum); MERR. En. Philip. 2 (1923) 386, p.p. (sub X. excelsum); BAKER, J. Bot. 62 (1924) Suppl. 7; Burk. & HEND. Gard. Bull. S.S. 3 (1925) 345; HEND. ibid. 4 (1928) 222; Watson, Mal. For. Rec. 5 (1928) 249; CHopaT in Merr. PI. Elm. Born. (1929) 133; Crarp, Fl. Siam. En. 1 (1931) 105; Burk. Dict. (1935) 2268, p.p.; KeirH, N. Born. For. Rec. 2 (1938) 225; DocTERS VAN LEEUWEN, Ned. Kruidk. Arch. 51 (1941) 166, 167; MASAMUNE, En. Phan. Born. (1942) 379; Wyatt-SmitTH, Mal. For. Rec. 17 (1952) 80, 363, p.p. (sub X. excelsum); BALAN MENON, ibid. 19 (1956) 34; WyatrT-SMITH, ibid. 23? (1963) f. 10, p.p.?; MEWER, Bot. News Bull. Sandakan 7 (1967) 88; Nc, Fed. Mus. J. n.s. 13 (1971) .1373 Tree. Fl. Malls la(1972)):3545: i. allay pes Corner, Gard. Bull. Sing. Suppl. 1 (1978) 146, 211 (excl. n. 29341); MEUDEN, Leiden Bot. Ser. 7 (1982) 70, f. 2, 3A g—i, 10C, 11. — Banisterodes affine (Mig.) O. K. Rev. Gen. Pl. 1 (1891) 46, nom. illeg. — X. sarawakense CHopat, Bull. Herb. Boiss. 4 (1896) 262 (‘sarawakensis’); GAGNEP. in Desv. J. Bot. 21 (1908) 252; MASAMUNE, En. Phan. Born. (1942) 381. — X. excelsum var. affine (M1Q.) BOERL. Cat. Hort. Bog. (1899) 59; HEYNE, Nutt. Pl. (1927) 901. Tree, rarely a shrub, up to 15(—30) m, up to 40(—95) cm dbh. Twigs mostly very densely, some- times rather sparsely, patently to appressedly mi- nutely greyish hairy, glabrescent. Axillary buds 2 (or 3), often seemingly single, the upper bud often stalked, rarely supra-axillary for more than 0.5 mm. Petiole (S—)6—11(—17) mm, more or less densely minutely greyish hairy all round or at the upper side only, eglandular. Leaf-blade rarely linear-lanceo- late, 6.5—18(—35) by (2—)3—8(—13) cm, base cuneate to rounded-attenuate, sometimes attenuate with the margins of the leaf-blade curved upwards and grad- ually grading into the ridges on the petiole, apex sometimes cuspidate; above sometimes slightly bul- late between secondary nerves, yellowish to rather dark greyish green, often with a lighter strip along midrib and base of secondary nerves, midrib promi- nent to flat, rarely slightly sunken, in basal part often with a groove and often minutely hairy; beneath con- colorous or more brownish, sometimes more or less densely minutely hairy in basal part, secondary nerves 5—16 pairs, usually forming a more or less dis- tinct intramarginal nerve in the apical half, intramar- ginal nerve rarely (nearly) complete to the base; glands very variable in number and size, usually dis- 504 FLORA MALESIANA [ser. I, vol. 10° tinct. /nflorescences shorter than the leaves; axes smooth, minutely greyish hairy; bracts sometimes with distinct glands. Pedice/ (2—)4—11 mm, rarely longer, densely minutely greyish hairy. Sepals persis- tent or not in fruit, sometimes with small glands; outer sepals 1.7—5.5 by 1.8—4.7 mm, inner ones 2.1—7.8 by 2.1—5.2 mm. Petals white (or rarely pink), the upper ones with an orange-yellow spot, when dry yellowish, the longest one 6—17 mm; carina appressedly to half-patently, sometimes patently hairy outside, inside appressedly hairy or sometimes glabrous; other petals sometimes quite glabrous, usually hairy outside at base, upper petals hairy in- side or only at base. Stamens: filaments glabrous to sparsely appressedly hairy in basal part; anthers 0.5—1.3 (—1.6) mm long. Ovary often semi-2- locular, smooth to distinctly ribbed, sometimes tu- berculate, glabrous or sometimes greyish hairy in 2(—4) rows in apical part; ovules usually 8—12, very rarely 6 or 5, rarely up to 16. Fruit globular or somewhat longer than wide, rounded or slightly but sometimes rather sharply beaked apically, up to 1.5 cm diam., smooth to pustulate or tuberculate, some- times ribbed, creamish to brown; pericarp not very thick; sepals (sub)persistent in fruit or not. Distr. SE. Asia (Khasya, Laos, Thailand); in Malesia: Sumatra (also Banka), Malay Peninsula, Borneo (Sarawak, Brunei, Sabah), Philippines (Pa- lawan, Culion, Busuanga). Notes. Very variable, and probably not homoge- neous; for a discussion, see VAN DER MEIJDEN, /.c. 71-73. Most authors have included /. X. flavescens in this species; see note under that species. 8. Xanthophyllum pubescens MEIDEN, Leiden Bot. Ser. 7 (1982) 73. — Xanthophyllum spec. B NG, Tree Fl. Mal. 1 (1972) 365, f. 5. Shrub or small tree, up to 12 m, 5 cm dbh. Twigs more or less brownish hairy. Axillary buds densely patently rather shortly hairy. Petiole 10-12 mm, hairy like twig. Leaf-blade 15—28 by 5.5—9.5 cm; above greenish, more or less densely hairy on the nerves, glabrescent, midrib flat to slightly promi- nent; beneath lighter coloured, rather densely patent- ly hairy, secondary nerves 8 or 9 pairs, forming a more or less distinct intramarginal nerve; glands (2—)7—14(—18), scattered, (0.5—)0.6—0.8(—1) mm diam. /nflorescences up to 10 cm long; axes densely patently greyish hairy; in basal part flowers with 3 to- gether; bracts with 2 large glands. Pedicel 5—8 mm, very densely minutely greyish patently hairy. Sepals densely to very densely patently greyish hairy out- side; outer sepals 3.8—4 by 2.7—4 mm; inner sepals 5.2—6.1 by 3.8—4.5 mm, slightly keeled. Petals white, the upper ones with a yellow spot, when dry yellowish, the longest one 12—13 mm; carina very densely shortly half-patently whitish hairy outside, in apical part rather densely hairy inside; other petals glabrous or basally ciliate, or the upper petals minutely hairy inside above the base. Stamens: an- thers 1.1—1.5 mm long, sometimes cohering around the stigma, ciliate along slits. Ovary containing 8—13 ovules. Fruit (immature) obovoid, dull, finely pustulate. Distr. Malesia: Malay Peninsula (Pahang, Jo- hore). 9. Xanthophyllum resupinatum MeEUDEN, Bot. J. Linn. Soc. 67 (1973) 120; Leiden Bot. Ser. 7 (1982) 74. Tree, up to 25 m, 40 cm dbh. Twigs glabrous. Axil- lary buds very inconspicuous, + enclosed between basal part of the petiole and the twig. Petiole 4.5—7 mm, often appearing longer because of the narrow leaf base, transversely wrinkled. Leaf-blade 6—12.5 by 2.7—5 cm, attenuate into a narrow petiole-like part; above (very) shiny, greenish to dark olive- greenish, midrib distinctly prominent; beneath yel- lowish brown to greenish brown, very shiny, midrib flat to faintly sunken, rarely faintly prominent or a narrow crest, secondary nerves 4—7 pairs, forming an incomplete, indistinct intramarginal nerve, tertiary nerves rather indistinct to obscure; glands very few, mostly situated above the middle, 0.3—0.7 mm diam. Inflorescences shorter than the leaves; axes grooved, densely minutely appressedly greyish hairy. Pedicel slightly grooved, 4—5 mm, very densely minutely ap- pressedly greyish hairy. Sepals dark brown to black- ish; outer ones c. 3 by 2.4—3 mm, with scattered, in- distinct, small glandular spots, sometimes with 2 protruding glands halfway; inner sepals c. 4—4.8 by 2.8—3.5 mm, slightly keeled, with few glandular spots. Petals white, when dry dark brownish, the longest one 7—9 mm; carina very densely half-patent- ly greyish pubescent outside, + hairy inside in apical part; other petals glabrous. Stamens: anthers 0.5—0.7 mm long, faintly hairy. Ovary dark brown- ish, ribbed, slightly appressedly hairy on the ribs in apical part, or glabrous; ovules 9-14. Fruit 1.2 cm diam., blackish, slightly shiny, finely pustulate; peri- carp rather thin, hard. Distr. Malesia: Borneo (Sarawak, E. Sabah, N. Kalimantan). 10. Xanthophyllum schizocarpon CHopatT in Merr. Pl. Elm. Born. (1929) 136; MAsAMUNE, En. Phan. Born. (1942) 381; MEIDEN, Leiden Bot. Ser. 7 (1982) 74. Tree, 10—25 m, 35 cm dbh. Twigs minutely appres- sedly hairy in younger parts to glabrous. Axillary buds 2 or 3, the upper one 0—3 mm above the axil. Petiole 5—7 mm, minutely appressedly hairy to gla- brous. Leaf-blade 4.5—12 by 1.5—4.5 cm; above 1988] greyish green, midrib sunken; beneath glaucous- papillose, minutely appressedly hairy on nerves, sec- ondary nerves 6 or 7 pairs, not forming an intramar- ginal nerve, finer venation indistinct; glands numer- ous, scattered, c. 0.2 mm diam., basal glands (if pres- ent) c. 0.5 mm diam. /nflorescences often shortly supra-axillary, often 2 per leaf axil, unbranched, shorter than the leaves; axes faintly grooved, brown- ish, minutely appressedly hairy; lower bracts oppo- site. Pedicel 6—8 mm, slightly grooved, minutely densely more or less appressedly greyish hairy. Sepals: outer ones 2.5—3.8 by 2.3 mm; inner ones 3.1—4.3 by 3.2 mm, + keeled. Petals yellow, when dry yellowish, the longest one 6—7 mm; carina rather densely appressedly hairy outside, shortly hairy in- side in apical part; lateral petals glabrous inside, fur- ther like the upper ones, upper petals glabrous to shortly hairy outside, shortly patently hairy inside up to 2/3. Stamens: 8 (or 9); anthers c. 0.3 mm long. Ovary slightly ribbed, brownish, appressedly hairy on median ribs; ovules 6—8. Fruit irregularly globular, c. 1.5 cm diam., mostly wider than high, ir- regularly 4-sulcate, dull, light greenish to yellowish brown; pericarp hard and irregular in thickness. Distr. Malesia: Borneo (Sarawak: Mersing Hill & Lambir; Sabah: Tawau). 11. Xanthophyllum velutinum CuHopart, Bull. Herb. Boiss. 4 (1896) 259; MEuDEN, Leiden Bot. Ser. 7 (1982) 75. Tree, 5—25 m, 60 cm dbh. Twigs densely set with short and long hairs, longer hairs 0.3—0.8 mm long. Axillary buds 1—3 mm long, hairy like the twig. Petiole (4—)8—14(—16) mm, hairy like the twig. Leaf- blade (6—)9—22 by 3-9 cm, apex cuspidate or sometimes acuminate; above greyish green, in basal part sometimes sparsely hairy especially on the nerves and midrib, midrib distinctly sunken, nerves and intramarginal nerve slightly sunken to flat and rather obscure, venation obscure; beneath brownish, hairy like the twig, rarely rather sparsely hairy, sec- ondary nerves 8—11 (or 12) pairs, strongly promi- nent, forming a strongly prominent intramarginal nerve; glands (7—)10 or more, scattered, (0.2—)0.3— 0.4(—0.6) mm diam., basal glands 0.9—1.2 mm long. Inflorescences about as long as the leaves; axes ribbed, very densely patently brownish hairy in the basal part; flowers with 3 together. Pedicel 1.5—6.5 mm, ribbed, very densely patently brown pubescent, rarely shortly hairy. Sepals persistent in fruit, greyish hairy; outer sepals 2.1—3.2 by 1.7—3 mm, with 2-4 glandular spots; inner sepals 3.5—5.3 by 2.4—3.4 mm, distinctly keeled. Petals white or yellow, when dry brownish orange, the longest one 9.5—10.5 mm long; carina more or less densely appressedly hairy outside, sparsely appressedly hairy inside in middle part; other petals ciliate in basal half, glabrous out- POLYGALACEAE (van der Meijden) 505 side, upper petals reflexed. Stamens: filaments rather sparsely appressedly hairy till apex; anthers (0.75—)0.8—1.2 mm long, usually cohering around the stigma, ciliate along slits. Ovary sessile, distinctly ribbed, glabrous or sometimes hairy on 2 ribs down to halfway; style reflexed at end of anthesis; ovules (6—)8—12. Fruit (immature) ovoid, very sharply beaked, with 8 or more distinct ribs, finely pustulate to tuberculate, dull, greyish brown; pericarp thick, hard; pedicel crowned by persistent sepals. Distr. Malesia: Borneo (Sarawak, Sabah). Brunei, 12. Xanthophyllum rufum Benn. FI. Br. India 1 (1874) 210; Kinc, Mat. Fl. Mal. Pen. (1890) 144; MarnGay, Kew Bull. (1890) 113; GAGNEP. in Desv. J. Bot. 21 (1908) 252; RrpLey, Fl. Mal. Pen. 1 (1922) 145; Watson, Mal. For. Rec. 5 (1928) 249; Burk. Dict. (1935) 2269; WyatTtT-SmiTH, Mal. For. Rec. 17 (1952) 81, 362; BALAN MENON, ibid. 19 (1956) 34; Wyatt-SitH, ibid. 237 (1963) f. 5, 9; MEWER, Bot. News Bull. Sandakan 7 (1967) 87; No, Tree Fl. Mal. 1 (1972) 361, f. 4; MEWDEN, Leiden Bot. Ser. 7 (1982) 76, f. 3A-1, 12. — Banisterodes rufum (A.W.BEN- NETT) O. K. Rev. Gen. PI. 1 (1891) 46, nom. illeg. — X. flavum Riwiey, Kew Bull. (1925) 77. — X. he- teropleurum CuHopat in Merr. Pl. Elm. Born. (1929) 134; MASAMUNE, En. Phan. Born. (1942) 380; No, Tree Fl. Mal. 1 (1972) 363. — Fig. 19. Tree, up to 32 m, 40 cm dbh. Twigs rufous-hairy. Axillary buds to 1.8 mm long, but often much smal- ler, densely pubescent. Petiole 7-21 mm, densely patently pubescent, more or less glabrescent, apically often with large glands. Leaf-blade 8—25 by 4-13 cm, base sometimes cordate; above bright (neon- like) yellow-green to light greyish green, midrib and nerves slightly sunken, venation indistinct; beneath light brownish to brownish green, papillose, rather densely hairy, secondary nerves 5—9 pairs, forming a rather distinct intramarginal nerve in apical or rarely also in basal part; glands very numerous, c. 0.1 mm diam., basal glands large, situated on the midrib at the very base of the leaf and then obscure, or at very apex of the petiole and then conspicuous. /n/flores- cences sometimes unbranched, shorter than to as long as the leaves; axes finely ribbed, very densely rufous-hairy; bracts conspicuous, persistent, often with distinct glands, bracteoles small, subpersistent. Pedicel 4.5—7(—10) mm, ribbed, very densely rufous-hairy. Sepals persistent or not, sometimes present under fully ripe fruits (then up to c, 12 mm long), very densely rufous-hairy outside, glands pres- ent or not, inconspicuous; outer sepals c. 4—6 by 4-6 mm, mostly slightly ribbed; inner sepals c. 5—7 by 4-6 mm, keeled. Petals white, the upper ones with a yellow spot, when dry yellowish, the longest one 12—15 mm; carina very densely yellowish brown pu- 506 FLORA MALESIANA [ser. I, vol. 103 \ \\ \ A \\ Nest eciatic! \\, \ \ \ NS AN NS KK. KK. WS SS Fig. 19. Xanthophyllum rufum BENN. a. Habit, x 0.5; b. base of leaf, x 2; c. flower; d. flower, longitudinal section, gynoecium removed; e. carina with two stamens enclosed; f. gynoecium; all x 3; g. ovary, longitu- dinal section, x6 (SAN 36554). 1988] POLYGALACEAE (van der Meijden) 507 bescent outside, inside hairy in apical part; other pet- als pubescent outside in apical part or only apically tufted. Stamens: filaments glabrous; anthers 0.4—0.6 mm long, shortly hairy at base or glabrous. Ovary very densely rufous-pubescent all round or pubescent in 4(—8) rows, the median rows mostly longer than the lateral ones; style very densely rufous-pubescent in apical part; ovules 12—14. Fruit globular, up to 1.8 cm diam., often with 2—4, some- times hairy ridges running down from the style-scar, yellowish green, dull, finely tuberculate; pericarp rather thick, hard; not rarely some sepals present. Distr. Malesia: Sumatra, Malay Peninsula, Borneo. Note. In most Bornean collections the ovary is hairy all round (or sometimes hairy in 8 rows), and the sepals are more or less persistent in fruit (‘X. heterophyllum’ CHopat). In Malaya (‘X. rufum’) and Sumatra (‘X. flavum’ RIDLEY) the ovary is usual- ly hairy in 4 rows, and the sepals are soon dropping off. These differences are not consistent in Malaya, however, as sometimes the ovary is hairy all round, or the sepals are persistent. Apart from those I did not find other differences; therefore it is not neces- sary to distinguish varieties based on these conspic- uous but minor and grading differences. 13. Xanthophyllum macrophyllum BAKER, Kew Bull. (1896) 21; Arry SHAw, Kew Bull. (1940) 252; MASAMUNE, En. Phan. Born. (1942) 380; MEER, Bot. News Bull. Sandakan 7 (1967) 88; MEIDEN, Leiden Bot. Ser. 7 (1982) 78. Shrub or tree, up to 25 m, up to 30 cm dbh. Twigs glabrous. Petiole 10-18 mm, with (O—)2(—4) very distinct glands. Leaf-blade (10—)14—28 by (3.8—)4.5—10 cm, base often curved upwards and de- current-attenuate; above sometimes slightly bullate between the secondary nerves, greenish, midrib slightly to distinctly prominent, rarely slightly sunken, nerves finely prominent; beneath mostly brownish green, secondary nerves 7—10 pairs, form- ing a nearly complete rather prominent intra- marginal nerve; glands mostly rather few, scattered, 0.4—0.5 mm diam. Jnflorescences sometimes un- branched, shorter than the leaves; axes grooved, ap- pressedly brown-velvety; bracts often opposite, with 2 small indistinct glands. Pedice/ 2-12 mm, appres- sedly brown-velvety. Sepals shortly appressedly brownish hairy outside, ribbed inside; outer sepals 5—6.5 by 3.8—6.2 mm; inner sepals 6—7.2 by 3.7—6.2 mm. Petals yellow, or white and the upper ones with a yellow spot, when dry yellowish to dark- brown, the longest one 13—16 mm long; carina ap- pressedly velvety outside, more or less densely hairy inside in apical part; other petals glabrous or hairy outside in apical part. Stamens: anthers 0.55— 0.75(—0.8) mm long, shortly hairy. Ovary sessile or shortly stipitate, creamish brown, often about 8-ribbed when dry, the median ribs most prominent and hairy over 1/3—2/3 their length, the other ribs hairy in apical part only; ovules 6—14. Fruit c. 2 cm diam., brownish to blackish, ribbed in apical part, thick-walled. Distr. Malesia: Borneo (Sarawak, Sabah). 2. Section Eystathes (Lour.) MEUDEN, Bot. J. Linn. Soc. 67 (1973) 117; Leiden Bot. Ser. 7 (1982) 81. — Eystathes Lour. Fl. Coch. 1 (1790) 235. Twigs sometimes hairy. Axillary buds sometimes 3(—7), sometimes erect. Leaves sometimes shifted-decussate. Leaf-blade: tertiary nerves finely reticulate, mostly distinctly protruding. /n- florescences: bracts sometimes opposite in basal part. Petals: carina usually distinctly ungui- culate. Stamens: filaments sometimes connate over up to 3 mm, rarely occasionally triadelphous, sometimes with a knob-like appendage at inner side. Ovary mostly densely hairy all round, rarely glabrous; stigma very rarely wider than the apex of the style; ovules 4 or 8— 16 (rarely more, excep- tionally 5 or 6). Fruit mostly globular, smooth or rarely tuberculate. Seed(s) 1 or occasionally 2, very rarely up to 4; testa with or without a hard inner layer; albumen thin but distinct; radicle exserted or not. KEY TO THE SUBSECTIONS 1. Testa with a hard inner layer. Albumen forming a distinct layer. Embryo laterally near the base with 2 flat- tened areas. Radicle exserted. Spp. 14—S7...0.ccscccvsenvcuteccncwavecsvegase 2a. Subsect. Jakkia 1. Testa without a hard inner layer. Albumen very thin. Embryo without flattened areas near the base. Radicle mot exserted;. SDP, 58x62 i! die deated hat vile ARs 0 « altnate oblate ied «jalan gDWie iT 2b. Subsect, Eystathes 508 FLORA MALESIANA [ser. I, vol. 10° 2a. Subsection Jakkia (BL.) MEVDEN, Leiden Bot. Ser. 7 (1982) 81. — Jakkia BLUME, Cat. (1823) 17. Twigs and inflorescence axes without nodal appendages. Axillary buds mostly 2 (seemingly single) and close together, rarely up to 4 or the upper one supra-axillary. Seed 1 or seeds occa- sionally 2; testa with a hard inner layer; albumen forming a rather thin, distinct layer, which is very thin at the lateral sides of the cotyledons near the base of the embryo; embryo laterally near the base with 2 flattened areas, radicle exserted. 14. Xanthophyllum nigricans MEVDEN, Bot. J. Linn. Soc. 67 (1973) 119; Leiden Bot. Ser. 7 (1982) 82. Small tree, up to 15 m, 20 cm dbh. Twigs finely longitudinally wrinkled. Axillary buds 2 or 3 (or 4), 1.5—2.2 mm long, very densely shortly patently light- brown hairy. Petiole (6—)7—11(—14) mm. Leaf- blade 3.5—12 by 1.2—5.5 cm; above greyish olive- green (to brownish); beneath concolorous, or bluish because of thin waxy layer, (glaucous-)papillose, sec- ondary nerves 5 or 6 (or 7) pairs, forming a more or less distinct intramarginal nerve; glands rather scarce or sometimes apparently absent, situated on or near the midrib, 0.1—0.2 mm diam., basal ones often slightly larger. Jnflorescences unbranched, shorter than the leaves; axes flattened, angular, black, very sparsely appressedly hairy; flowers with 1—3 to- gether. Pedicel 2—2.5 mm, black, nearly glabrous. Sepals glabrous outside, densely very shortly more or less patently hairy inside at base, further glabrous in- side; outer sepals 1.8 by 1.7 mm; inner sepals 2.5 by 2.1 mm. Petals white, when dry dark reddish, very sparsely appressedly hairy outside, faintly ciliolate apically, the longest one 8 mm long. Stamens: an- thers c. 0.3 mm long. Ovary black, glabrous; style glabrous or basally very sparsely appressedly hairy; ovules 4. Fruit globular, up to 1.3 cm diam., dull, slightly wrinkled, brownish; pedicel up to 3 mm long. Distr. Malesia: Borneo (Brunei, Sabah). 15. Xanthophyllum borneense Mig. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 277; Masamune, En. Phan. Born. (1942) 379; MEIsDEN, Leiden Bot. Ser. 7 (1982) 82. — X. glabrescens RIDLEY, Kew Bull. (1938) 113; MASAMUNE, En. Phan. Born. (1942) 380. Small tree, up to 5 (—8?) m high. Twigs more or less reticulately wrinkled. Axillary buds 2 (or 3), 1.2—1.8(—3) mm long. Petiole c. 5—6.5 mm. Leaf- blade 4—16 by 1.5—7 cm, margin undulate, apex acu- tish; above slightly bullate to flat between secondary nerves, greenish to brownish; beneath glaucous- papillose, secondary nerves 4—6 pairs, forming a rather distinct intramarginal nerve; glands numerous, scattered, 0.1—0.2 mm diam. J/nflores- cences unbranched, about as long as the leaves; axes flattened basally, angular, dull, light brownish, gla- brous. Pedicel 1.5—2.5 mm, dull, brownish, gla- brous. Sepals glabrous outside; outer sepals 3 by 2.1 mm; inner sepals 3.8 by 2.8 mm. Petals light brown- ish or orange when dry, minutely ciliate apically, the longest one 10—11 mm long; carina sparsely appres- sedly minutely hairy outside. Stamens: anthers c. 0.3 mm long, glabrous or with very few short hairs at base. Ovary completely glabrous; style very sparsely more or less appressedly hairy; ovules 4. Fruit globu- lar to broadly ovoid, c. 1.8 cm diam., smooth, brown; pedicel c. 5 mm. Distr. Malesia: Borneo (Sarawak, Sabah, SE. Borneo). 16. Xanthophyllum ovatifolium CHopDAT, Bull. Herb. Boiss. 4 (1896) 258; MEIDEN, Leiden Bot. Ser. 7 (1982) 83. Axillary buds 2 (or 3), 1—1.8 mm long. Petals 3.5—4.5(—6.5) mm; glands present or not. Leaf- blade 3.5—9.5 by 1.4—6 cm, apex sometimes cuspi- date; above rather dull, brownish to greenish, midrib little prominent at base; beneath dull, secondary nerves c. 3 or 4 pairs, not forming an intramarginal nerve; glands 8—20, usually situated halfway be- tween margin and midrib, 0.4—0.5 mm diam., basal glands usually present, 0.5—0.6 mm diam. /nflores- cences shorter than the leaves, unbranched; axes slightly flattened, brownish, glabrous to very sparse- ly appressedly hairy. Pedicel 7-8 mm, glabrous. Sepals glabrous outside; outer ones c. 2.8 by 1.7 mm; inner ones 3—3.5 by 1.8—2 mm. Petals white, when dry light brownish, ciliate at apex and base, further glabrous, the longest one 9-10 mm. Stamens: an- thers 0.3—0.4 mm long, glabrous. Ovary glabrous; style sparsely appressedly hairy basally; ovules 4. Fruit unknown. Distr. Malesia: Borneo (Sarawak). 17. Xanthophyllum tenue CHopar in Merr. Pl. Elm. Born. (1929) 135; MASAMUNE, En. Phan. Born. (1942) 382; MeverR, Bot. News Bull. Sandakan 7 (1967) 88; MEUDEN, Leiden Bot. Ser. 7 (1982) 83. Tree, up to 25 m, 40 cm dbh. Twigs glabrous to rather densely very shortly patently hairy. Axillary buds 2 (or 3), 1.5—2.5 mm long, shortly hairy, some- 1988] times irregular and larger because of cork-forming in apical region. Petiole 6—11 mm, not transversely wrinkled, shortly patently hairy to glabrous; glands absent or small. Leaf-blade (S—)7—16 by (2-) 2.5—6.5 cm; above slightly bullate to flat between secondary nerves, dull, greyish green to light reddish brown, midrib glabrous to patently minutely hairy in basal half; beneath mostly rather dull, yellowish green, glabrous to minutely patently hairy, midrib slightly prominent to flat, secondary nerves 4—6 pairs, mostly forming a rather indistinct intramar- ginal nerve, venation mostly not very prominent, sometimes rather indistinct; glands (0O—)4—25, often situated near midrib, 0.3—0.5(—0.7) mm diam.; basal glands often present, relatively large. Jn- florescences much shorter than the leaves, un- branched; axes slightly angular, sparsely to rather densely patently shortly hairy. Pedicel (2.S—) 4—6(—10) mm, rather sparsely to densely minutely hairy. Sepals nearly glabrous outside; outer sepals 2.1—2.8 by 1.3—2.5 mm, inner sepals 2.5—4(—4.5) by 1.5—3.2 mm. Petals yellowish or white, when dry orange, the longest one 8—9(—11.5) mm; carina sparsely to rather densely appressedly hairy outside, inside hairy to halfway or up to the apex; other petals glabrous to sparsely hairy outside, lateral petals hairy inside to halfway, upper petals hairy inside to apex. Stamens: filaments nearly free or connate over up to 1.5 mm; anthers c. 0.4 mm long. Ovary gla- brous or less often rather sparsely appressedly hairy and probably soon glabrescent; style glabrous or less often sparsely appressedly hairy in basal part; ovules 4. Fruit globular, up to 1.8 cm diam., dull, wrinkled, light greenish brown; pericarp soft, thin; pedicel mostly up to 7 mm, reddish. Distr. Malesia: Borneo (Sarawak, Tawau, N. Kalimantan). Ecol. Submontane rain-forest, 500—1200 m. Sabah, 18. Xanthophyllum subcoriaceum (CHODAT) MEU- DEN, Bot. J. Linn. Soc. 67 (1973) 120; Leiden Bot. Ser. 7 (1982) 85. — X. ellipticum var. subcoriaceum Cuopat in Merr. PI. Elm. Born. (1929) 134; MEUER, Bot. News Bull. Sandakan 7 (1967) 88. Shrub or small tree, up to 15 m, 20 cm dbh. Axil- lary buds 2 (or 3), (0.5—)1—2.7 mm long. Petiole 5—8.5 mm. Leaf-blade 5—12 by 1.5—5.5 cm, base sometimes more or less rounded, margin mostly strongly curved upwards when dry, apex cuspidate to acuminate; above sometimes slightly bullate between secondary nerves, dull, light greyish green to olive- green, secondary nerves mostly indistinct, venation mostly indistinct, sometimes scarcely visible, less often finely prominent; lower side yellowish green, secondary nerves (3 or) 4—6 pairs, forming a distinct intramarginal nerve, venation usually obscure; glands (0O—)2~12, situated at some distance from the POLYGALACEAE (van der Meijden) 509 midrib, 0.2—0.4 mm diam. IJnflorescences un- branched, + as long as the leaves; axes angular, light brown, sparsely appressedly shortly hairy; lowermost bracts sometimes leaf-like. Pedice/ 2.5—3.5 mm, mostly light brown, glabrous to sparsely minutely ap- pressedly hairy. Sepals (nearly) glabrous outside; outer ones 1.6—1.8 by 1.4—1.8 mm; inner ones 2—2.5 by 2—2.5 mm. Petals white, when dry orange, faintly ciliate apically, outside glabrous except at base, long- est one 8-10 mm. Stamens: anthers 0.5—0.7 mm long, sparsely hairy at base. Ovary 0.5—1.5 mm stipi- tate, subglabrous to rather densely appressedly hairy, rather soon partly glabrescent; style sparsely ap- pressedly hairy; ovules 4. Fruit at first + ellipsoid with a sharp beak because of subpersistent style, when mature globular, c. 1.7 cm diam., smooth, light green to brown, rather dull to shiny; pericarp thin; pedicel up to 5(—7) mm, mostly light brown. Distr. Malesia: Borneo (Sarawak, Sabah: Mt Kinabalu & Tawau). Brunei, 19. Xanthophyllum neglectum MEDEN, Bot. J. Linn. Soc. 67 (1973) 119; Leiden Bot. Ser. 7 (1982) 86. — X. palembanicum (non Mia.) KEITH, N. Born. For. Rec. 2 (1938) 225. Tree, up to 20m, 20 cm dbh. Axillary buds 2, ellip- tic to ovate-oblong, 1.5—4(—6) mm long, acute, light yellowish brown. Petiole 4—6.5 mm, glabrous to sparsely very shortly hairy especially in the upper groove. Leaf-blade 5—12 by 1.8—5.5 cm, margin un- dulate; above dull, greyish green; beneath rather dull, sometimes slightly waxy, secondary nerves 3—S pairs, forming a rather indistinct intramarginal nerve; glands 2—8, not in basal part, 0.2—0.3(—0.4) mm diam. J/nflorescences unbranched, 1.5—5S(—8) cm long, shorter than the leaves, bearing 3 or 4(—7) flowers; axes c. 0.5 mm diam., slightly flattened ba- sally, light brown, rather sparsely patently shortly hairy. Pedice/lc. 2mm, slightly grooved, rather dense- ly patently shortly hairy. Sepals: outer ones 2.2—2.5 by 2—2.3 mm; inner ones 2.8—3.5 by 2.2—3 mm. Pet- als white or yellowish, when dry light brownish, the longest one 7—10 mm; carina nearly glabrous to rather densely hairy outside, shortly hairy at both sides basally; other petals nearly glabrous except for some hairs at the base and apex. S/amens: filaments of abaxial 4 stamens basally widened and slightly thickened; anthers 0.3—0.4 mm long. Ovary patently hairy; ovules 4. Fruit (immature) ovoid, dull greyish green, hairy, slightly wrinkled when dry; pericarp rather soft; pedicel up to 4.5 mm, light brown. Distr. Malesia: Borneo (E. Sabah, E. Kaliman- tan). 20. Xanthophyllum pauciflorum Meupen, Bot, J. Linn. Soc, 67 (1973) 119; Leiden Bot. Ser. 7 (1982) 87. 510 FLORA MALESIANA [ser. I, vol. 10° Tree, up to 21 m, 25 cm dbh. Twigs + as thick as the petioles, only a few internodes long. Axillary buds 2, 0.5—1.3 mm. Petiole 3.5—4.5 mm. Leaf- blade 4.8—8(—9) by 1.2—2.2(—3.5) cm, apex cuspidate to acuminate; above olive- to dark-green, rather dull; beneath glaucous-papillose, secondary nerves 4—6 pairs, not forming an intramarginal nerve; glands 2—7, 0.1(—0.2) mm diam. /nflores- cences unbranched, much shorter than the leaves, bearing only 3—6 flowers; axes slender, slightly flat- tened, glabrous to sparsely very shortly patently hairy. Pedice/ 1.5—2.5(—3) mm, very shortly patently hairy. Sepals with a small apical tuft; outer ones 2—2.2 by 1.5—1.8 mm, very sparsely very shortly hairy outside; inner ones c. 2.7 by 3 mm, glabrous outside. Petals yellowish, when dry yellowish orange, very sparsely hairy to glabrous outside, apically slightly tufted and distinctly ciliate, basally rather densely hairy at both sides, the longest one 7.5—8.5 mm long. Stamens: anthers c. 0.4 mm long, with few hairs at base. Ovary appressedly hairy; ovules 4. Fruit (immature) ovoid-globular, olive- brown, smooth, with a distinct remainder of the style, roughly pubescent, glabrescent. Distr. Malesia: Borneo (Sarawak: Mersing Hill). 21. Xanthophyllum tardicrescens MEIDEN, Bot. J. Linn. Soc. 67 (1973) 120; Leiden Bot. Ser. 7 (1982) 87. Small tree, up to 6 m, 6 cm dbh. Twigs dull, bear- ing | or 2 (or 3) leaves per shoot. Axillary buds 2 (or 3?), smaller than 1 mm. Petiole 3.5—5 mm. Leaf- blade 7—16 by 2—5 cm, base rounded-truncate to -cordate; above dark greyish green, very dull, sec- ondary nerves slightly sunken, venation obscured; beneath olive-greenish, dull, secondary nerves 3 or 4 pairs, the basal nerves long, reaching often beyond the middle of the leaf, or forming an intramarginal nerve, venation not distinct; glands 6—12, scattered, 0.2—0.3 mm diam. /nflorescences unbranched, + as long as the leaves; axes grooved, not flattened, nearly glabrous, light brown; flowers often with 2 or 3 to- gether; bracts relatively long-persistent. Pedicel 3.5—4.5 mm, sparsely very shortly appressedly hairy. Sepals sometimes apically with tiny glandular spots; outer ones 2 by 2.1 mm; inner ones 2.3 by 2.3 mm. Petals white, the upper ones with a yellow spot, when dry yellowish orange, the longest one 7—8 mm; carina nearly glabrous outside; other petals glabrous. Stamens: filaments connate over 0.7—1 mm, very shortly hairy above base, further glabrous; anthers probably c. 0.7 mm long. Ovary very shortly stipi- tate, more or less appressedly hairy; ovules 4. Fruit (immature) smooth, slightly shiny, yellowish green, sparsely appressedly hairy, glabrescent; pericarp very thin. Distr. Malesia: Borneo (Sarawak: Semengoh). 22. Xanthophyllum parvifolium MEDEN, Bot. J. Linn. Soc. 67 (1973) 119; Leiden Bot. Ser. 7 (1982) 88. Tree, up to 25 m, 30 cm dbh. Twigs forming very short shoots bearing 2 or 3 leaves, the young ones very slender, c. 0.5 mm diam. Axillary buds 2, 0.7—1.2 mm long. Petiole 2—2.5 mm. Leaf-blade 1.6—5(—6) by 0.5—1.6(—1.9) cm, base rounded to cu- neate; above shiny, yellowish or greenish brown, midrib flat or slightly prominent, nervation finely prominent to obscure; beneath glaucous-papillose, yellowish to reddish brown, secondary nerves 1—3 pairs, rather indistinct, forming an indistinct intra- marginal nerve, venation rather indistinct; glands rather numerous, scattered, very small though rela- tively distinct, up to 0.1 mm diam. Flowers solitary in the upper leaf axils or twigs terminating in a 1—3-flowered inflorescence; this unbranched, shor- ter than to + as long as the leaves, up to c. 0.5 mm diam., up to 3.5 cm long, glabrous. Pedicel 10—11 mm, glabrous. Sepa/s purplish, (sub)glabrous out- side, outer ones c. 2 by 1.6 mm, inner ones 3—3.4 by 1.8—2.3 mm. Petals light orange, when dry orange brown, very sparsely hairy at base and at very apex, further glabrous, the longest one 10-11 mm. Stamens: filaments widened and slightly thickened above their base and there densely rather shortly patently hairy, further glabrous; anthers 0.35—0.4 mm long, with few short hairs at base. Ovary c. 1 mm stipitate, appressedly hairy; style rather sparsely ap- pressedly hairy in lower half, further glabrous; ovules 4. Fruit very shortly stipitate, globular, up to 1.1 cm diam., dull, light brown, very sparsely shortly appressedly hairy; pericarp rather thin; pedicel slender, 6.5—10 mm, completely glabrous. Distr. Malesia: Borneo (Sarawak: Sabal For. Res., Lambir hills). 23. Xanthophyllum philippinense CHOoDAT, Bull. Herb. Boiss. 4 (1896) 261; MerRR. En. Philip. 2 (1923) 387; MEUDEN, Leiden Bot. Ser. 7 (1982) 88, f. 10A. Twigs often with numerous adventitious buds on older nodes. Petioles 6-10 mm, often grading into the narrow leaf base, mostly in middle part with glands. Leaf-blade 7—14 by 2.5—6 cm, apex acutish; above shiny to rather dull; beneath + concolorous, rather dull, secondary nerves c. 5—7 pairs, apically hardly distinguishable; glands 1—6, 0.4—0.6 mm diam. /nflorescences unbranched or with one side- branch, up to 5 cm long, often several together in the leaf axils; axes rather dark, sparsely minutely hairy; lower bracts (sub)opposite. Pedicel c. 3.5 mm, red- dish brown, densely minutely hairy. Sepals: outer ones c. 2.5 by 2 mm; inner ones c. 3.5 by 3 mm. Petals orange when dry, the longest one c. 12 mm; carina faintly appressedly hairy outside near apex; other petals nearly glabrous. Stamens: filaments 1988] widened but hardly thickened above their base and there densely more or less woolly hairy, further gla- brous; anthers c. 1.2 mm long. Ovary c. 2 mm stipi- tate, appressedly hairy; style very long (c. 10 mm), exserting over c. 2 mm from the carina, faintly ap- pressedly hairy; ovules 4. Fruit c. 2 cm diam., smooth, light brown, faintly hairy; pedicel 2.5—6(—8) mm, minutely hairy. Seed/s) 1 or 2. Distr. Malesia: Philippines (Luzon, Mindoro, Mindanao). Ecol. In open flowers the stigma is exserted from the carina. This may mean that cross-pollination is obligatory. 24. Xanthophyllum ancolanum Mra. FI. Ind. Bat., Suppl. (1861) 394; CHopat, Bull. Herb. Boiss. 4 (1896) 261; Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 275, incl. f. angustifolia M1Q.; MEUDEN, Leiden Bot. Ser. 7 (1982) 89. — X. palembanicum Mig. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 277. — X. sumatra- num Mia. I.c. 275; BAKER, J. Bot. 62 (1924) Suppl. 7. Shrub or small tree, up to 4 m, 4 cm dbh. Twigs glabrous to densely short hairy. Axillary buds c. 1—2.5 mm long, hairy; older nodes often with nu- merous adventitious buds. Petiole (S—)7—15 mm, not transversely wrinkled, glabrous to densely short- ly hairy, apically sometimes with glands. Leaf-blade 10-26 by (2.5—)3.5—10.5 cm, apex shortly acuminate to cuspidate; above greyish green, rather dull; beneath green, secondary nerves 8—10 pairs, at least in apical part forming an intramarginal nerve; glands mostly rather numerous, scattered, 0.2—0.5 mm diam., basal glands often present. /nflorescences situated at end of young twigs, but also axillary and on old nodes (plant partly ramiflorous), 1 (or 2) in each axil, erect to strongly reflexed, unbranched, or sometimes with a side-branch; axes 1—11 cm long, dark, rather sparsely shortly hairy; lower bracts (sub)opposite. Pedicel 4.5-—6 mm, dark, rather sparsely minutely patently hairy. Sepals dark purple; outer ones 2.5—3.8 by 2.1—3.1 mm; inner ones 4.2—5.2 by 3—3.9 mm. Petals when dry light to dark orange, the longest one 13—19 mm long; carina long unguiculate, sparsely very shortly appressedly hairy outside; other petals very sparsely hairy outside to glabrous. Stamens: filaments connate over (0.7—)2—2.5 mm, slightly widened and hardly thick- ened above base; anthers 0.8—1.2 mm long, very shortly hairy at base. Ovary c. 2 mm stipitate, half- patently hairy; style exserted from the carina for less than 0.5 mm, sparsely hairy in 2 rows; ovules 4. Fruit (immature) shortly stalked, more or less globular, slightly beaked, brownish, hairy; pedicels 8—11 mm. Distr. Malesia: Sumatra (North: Gajolands: Mt Bandahara; Central: N. of Padang; South: Lampong Distr.: Muaradua). Ecol. Montane rain-forest, 500-1200 m. POLYGALACEAE (van der Meijden) = 25. Xanthophyllum brigittae MEUDEN, sp. nov. — Fig. 20. Gemmae axillares 2,5—3,5 mm longae. Folia basi cordata incurvata; glandulae sparsae; nervis laterali- bus 14—20-jugis; petiolo 15—18 mm longo. Petala in- aequalia, petalo longissimo 16 mm longo; carina subglabra, extus sparse appresse pilosa. Filamenta (subjlibera; antherae 1 mm longae. Ovarium sparse appresse pilosum; stylum appresse pilosus; ovula 4. Fructus ignotus. — Typus: W.J.J.O.DE WILDE & B.E.E.pE Wi_peE Duyryes 15060 (L). Shrub, c. 2 m high. Twigs glabrous. Axillary buds ovate-oblong, 2.5—3.5 mm long, hardly thickened basally, sparsely shortly hairy; older nodes with nu- merous small adventitious buds. Petiole 15—18 mm, distinctly transversally wrinkled, apically with a pair of indistinct glands. Leaf-blade 25—50 by 9—12 cm; base cordate with the margins curved upwards above the base of the petiole; apex slightly acuminate; above greyish green, rather dull; beneath green, sec- ondary nerves 14—20 pairs, in apical part forming a rather indistinct intramarginal nerve; glands indis- tinct, few, scattered, c. 0.1 mm diam., basal glands usually present, 0.3 mm diam. /nflorescences numer- ous in the upper part of young twigs and also on older nodes, 1 or 2 in each axil, patent or slightly re- flexed, unbranched or basally sometimes with a side- branch; axes 4—10 cm long, dark, rather sparsely shortly hairy; lower bracts subopposite. Pedicel 3—4 mm, brown, densely shortly appressedly hairy. Sepals purplish brown, shortly hairy; outer ones c. 3 by 2.5 mm; inner ones c. 4.5 by 3 mm. Petals whitish with pink to lilac tips when fresh, light to dark orange when dry, the longest one c. 16 mm; carina ungui- culate, sparsely shortly appressedly hairy outside; other petals (sub)glabrous outside. Stamens: filaments free or connate for 0.1 mm, hardly widened and not thickened above base; anthers 1 mm long, very shortly hairy at base. Ovary c. 1.5 mm stipitate, shortly appressedly hairy to near apex; ovules 4. Fruit unknown. Distr. Malesia: N. Sumatra (Gajolands: Mt Leu- ser National Park). Notes. Like 24. X. ancolanum, differing in the longer, distinctly transversally wrinkled petioles, the more numerous secondary nerves, the leaf base with its upturned margin (like in 32. X. adenotus) and its (nearly) free filaments. Half a century ago VAN STEENIS collected (also on Mt Leuser) a fragment of this species (VAN STEENIS 10075, BO!), thrown down by monkeys, with a single flower and a small part of a single leaf. | erroneously attributed this to 5/. X. erythrostachyum GAGNEP. noting, however, that the single flower possessed 4 instead of 11 ovules. Having now a fine collection at hand, it turned out that the original count of the ovules in VAN STeeNis’ collection was correct, 512 FLORA MALESIANA [ser. I, vol. 103 Fig. 20. Xanthophyllum brigittae MEIDEN. a. Habit, x 0.5; b. leaf base, x 1.5; c. gynoecium, x 3; d. ovary, longitudinal section, 6 (DE WILDE & DE WILDE-DUYFIJES 15728). The present species is named in honour of Ms. Bri- GITTA DE WILDE-DuyFJEs who ardently assisted her husband in the botanical exploration of the Leuser National Park in the Gajolands of N. Sumatra. 26. Xanthophyllum tenuipetalum MEUDEN, Bot. J. Linn. Soc. 67 (1973) 120; Leiden Bot. Ser. 7 (1982) 89, f. 9a. — X. affine (non MiQ.) Koorp. Minah. (1898) 344. Tree, up to 30 m, 40 cm dbh. Axillary buds 2 (or 3), 0.5—2.5 mm long, basally wrinkled, in apical part smooth or slightly keeled. Petiole 6—9 mm, often ap- pearing somewhat longer because of the attenuate leaf base, always with 2 rather distinct glands usually situated in the middle part or at the (very) base. Leaf- blade 9—20 by 3.5—11 cm; above slightly bullate be- tween the secondary nerves, shiny, dark to brownish green, nervation often very distinct; beneath slightly shiny, secondary nerves c. 6—8 pairs, apically dif- ficult to count, forming an irregular and fine intra- marginal nerve or ending in the venation; glands mostly very numerous, scattered, c. 0.3—0.5 mm diam. /nflorescences also in lower leaf axils, un- branched or rarely with one short branch, up to 7 cm long; axes rather slender, smooth, very densely shortly more or less appressedly hairy; flowers with 3 together or in the apical part solitary; lower bracts (sub)opposite. Pedicel 2-4 mm, very densely nearly appressedly shortly hairy. Sepals: outer sepals 1.8—2.9 by 1.9—3 mm; inner sepals 2.9—3.5 by 2.6—3.2 mm. Petals rather thin, white (or yellow?) when fresh, when dry light brown or orange, not cov- ering the stamens in anthesis, the longest one 10.5—12.5 mm; carina long-unguiculate, apically ciliate, rather sparsely to rather densely more or less appressedly woolly hairy outside in apical region and slightly so near the base, inside glabrous; other petals ciliate at very apex, near base sparsely to rather densely hairy on either side. Stamens 8, occasionally 9; filaments connate over 0.1—0.5 mm, rather dense- ly more or less appressedly woolly hairy in basal half, glabrous upwards; anthers (0.5—)0.6—0.7 mm, sparsely ciliolate along slits, sparsely and shortly hairy at base. Ovary appressedly hairy; style rather sparsely hairy; ovules 4. Fruit globular, 1.8—2 cm 1988] POLYGALACEAE diam., slightly shiny, light brown, faintly appressed- ly hairy; pericarp rather thin; pedicel up to 6 mm. Distr. Malesia: N. & S. Celebes (Minahassa; Ma- lili; Kendari; MunalI.), Moluccas (Taliabu & Kai Is.), West New Guinea (Vogelkop Peninsula). 27. Xanthophyllum impressum MeEvUDEN, Leiden Bot. Ser. 7 (1982) 90. Tree, up to 23 m, 20 cm dbh. Axillary buds, when resting, mostly more or less enclosed between the base of the petiole and a low ridge of the twig, 1—1.8 by 1.5—2 mm, for c. 1 mm of its length uncovered; scales strongly thickened, especially at base, but leav- ing a narrow scar. Petiole 10-14 mm, sometimes with glands. Leaf-blade 10—20 by 3.5—9 cm, apex acutish to shortly acuminate; above rather dull, grey- ish green; beneath light yellowish green, papillose, secondary nerves c. 8 or 9 pairs, not forming an intramarginal nerve; glands scattered, probably rather numerous but often seemingly absent, up to 0.2 mm diam., exceptionally larger. Inflorescences up to 20 cm long; axes reddish brown, densely mi- nutely appressedly hairy; lower bracts (sub)opposite. Pedicel 1.5—4 mm, grooved, appressedly shortly hairy. Sepals: outer sepals 2—2.5 by 2.6—3.3 mm; in- ner sepals 3.2—3.7 by 3.2—3.3 mm. Petals white, the upper ones with a yellow spot, when dry orange to dark red, sometimes with incrustations, the longest one 8.5—10.5 mm; carina densely more or less ap- pressedly hairy outside; other petals glabrous out- side. Stamens: filaments widened and thickened above base, there appressedly hairy, further gla- brous; anthers 0.6—0.7 mm long, hairy to (sub)gla- brous at base. Ovary nearly sessile, half-patently hairy; ovules 4. Fruit globular, c. 1.7 cm diam., dull, smooth, light brownish, appressedly hairy. Distr. Malesia: E. Borneo (E. Sabah, E. Kali- mantan), Philippines (Catanduanes). 28. Xanthophyllum griffithii Hoox./. ex A.W.BEN- NETT, FI. Br. India 1 (1874) 210; KinG, Mat. Fl. Mal. Pen. (1890) 136; Marmncay, Kew Bull. (1890) 114; Branpis, Indian Trees (1906) 45; GAGNeEP. in Desv. J. Bot. 21 (1908) 251; Rrptey, Fl. Mal. Pen. 1 (1922) 149; Burk. & Henp. Gard. Bull. S. S. 3 (1925) 346; HeEnp. ibid. 4 (1928) 222; Burk. Dict. (1935) 2269; Wyatt-Smitu, Mal. For. Rec. 17 (1952) 80, 362; Na, Tree Fl. Mal. 1 (1972) 357, f. 2, excl. var. curtisii et var. montanum ; Mal. For. 38 (1975) 85, f. 8.1 A—E, 8.2; Mevpen, Leiden Bot. Ser. 7 (1982) 92. — Banis- terodes griffithii (HooK f. ex A.W.BENNeTT) O. K. Rev. Gen. Pl. 1 (1891) 46, nom. illeg. — X. parvum Cuopat, Bull. Herb. Boiss. 4 (1896) 264. — X. gra- cile Cuopat, |.c. 256; K. & V. Bijdr. Booms. Java 5 (1900) 302; Backer, Schoolfl. Java (1911) 80; Koorp. Exk. Fl. Java 2 (1912) 454. — X. pseudosti- (van der Meijden) 513 pulaceum Merk. Philip. J. Sc. 10 (1915) Bot. 316; En. Philip. 2 (1923) 387. Tree, up to 27 m, 40 cm dbh. Twigs glabrous to minutely patently hairy. Axillary buds erect to half- patent, (1.5—)3—8 mm long, not thickened at base, glabrous to densely minutely hairy; enclosing a pair of nearly similar buds of second order (those at base of a new twig often half-patent). Petiole 4-12 mm, sometimes with | or 2 glands in apical part. Leaf- blade 4—12(—15) by 1—4.5(—9) cm, apex sometimes cuspidate; above dark green to brownish; beneath lighter coloured, smooth to glaucous-papillose, sec- ondary nerves 4—6 pairs, usually forming an indis- tinct intramarginal nerve in apical half; glands 4—20, scattered but often near midrib, 0.2—0.3 mm diam. Inflorescences up to 10 cm long, at very base with 2 side-axes or with a pair of buds of second order; axes often reddish brown, densely minutely patently hairy; lower bracts opposite. Pedicel 1—4.5 mm, grooved, densely minutely appressedly hairy. Sepals sometimes with 2 glands in apical part; outer sepals 1.6—2.5 by 2.1—2.7 mm; inner sepals 2.6—3.3 by 2.1—3 mm. Petals white, the upper ones with a yellow spot, when dry dark red to orange-red, the longest one (S—)7—8 mm long; carina densely more or less appressedly hairy outside; other petals gla- brous to appressedly hairy outside in apical part. Sta- mens: filaments widened above base and with a knob-like, densely hairy appendage at inner side, further glabrous; anthers (0.3—)0.4(—0.5) mm long. Ovary 0.5—2 mm stalked, more or less appressedly hairy; ovules 4. Fruit globular, up to 1.5 cm diam., more or less smooth brown, appressedly hairy; pedi- cel up to 4 mm. Distr. SE. Asia and Malesia. Note. VAN DER MEIDEN (/.c.) distinguished 3 subspecies, of which the typical one (from Burma, Mergui) occurs outside Malesia. KEY TO THE SUBSPECIES 1. Axillary buds half-patent, flat a. ssp. angustifolium 1. Axillary buds erect, flattened against the twig, basally: ‘convex: iii. 2998s ees b. ssp. erectum a. ssp. angustifolium (NG) Mevpen, Leiden Bot. Ser. 7 (1982) 94. — X. griffithii var. angustifolium Na, Fed. Mus. J. n.s. 13 (1971) 137. — X. parvum Cuopat. — X. gracile CHopat. — X. pseudostipula- ceum MERR. Axillary buds half-patent, elliptic to lanceolate, 1.5—8 mm long, at base often stalk-like constricted, more or less flat, wrinkled, glabrous or soon glabres- cent. Petiole 4—8(—9) mm. Leaf-blade 4—8(—10) by 1—4(—5) cm, in juvenile shoots up to 10 by 2.5 cm; beneath glaucous-papillose to (nearly) smooth and 514 FLORA MALESIANA [ser. I, vol. 103 not glaucous, secondary nerves 4 or 5 (or 6) pairs; glands 0.2—0.3 mm diam. Pedicel 1.5—4.5 mm. Petals: longest ones 6.5—7.8 mm. Stamens: anthers 0.4 mm long. Fruit c. 1.1 cm diam. Distr. Malesia: Central Sumatra (Indragiri), Malay Peninsula, Borneo (Sarawak, Sabah, E. Kali- mantan), Philippines (Luzon). Ecol. Usually in submontane rain-forest, up to 1400 m. b. ssp. erectum MEWDEN, Leiden Bot. Ser. 7 (1982) 94. — X. griffithii Hoox.f. ex A.W.BENNETT, excl. Mergui coll. Axillary buds erect or nearly so and flattened in their upper part against the twig, ovate to ovate- lanceolate, rarely elliptic, (3—)4—8 mm long, basally convex, slightly wrinkled, glabrous to densely mi- nutely hairy. Petiole 6-12 mm. _ Leaf-blade 5—12(-15) by 2—4.5(—9) cm; beneath glau- cous-papillose, secondary nerves c. 5 or 6 pairs; glands c. 0.2 mm diam. Pedicel 1—2(—3) mm. Petals: longest one (6.5—)7—8 mm. Stamens: anthers 0.4(—0.5) mm long. Fruit up to 1.5 cm diam. Distr. Malesia: Malay Peninsula (common). 29. Xanthophyllum monticolum MEuDEN, Leiden Bot. Ser. 7 (1982) 95. — X. griffithii var. montanum Noa, Fed. Mus. J. n.s. 13 (1971) 137; Tree Fl. Mal. (U972)9359). fa 2: Tree, up to 10 m, 20 cm dbh. Axillary buds often appressed against the petiole, (1—)1.8—2.9 mm long, base wrinkled, apex acute. Petiole 9-14 mm. Leaf- blade 8—16 by 2—5(—7.5) cm; above usually dark green, sometimes brownish; beneath glaucous- papillose, secondary nerves 6—8 (or 9) pairs, in apical part forming a weak intramarginal nerve; glands 4—16, mostly in basal half near midrib, 0.2—0.3 mm diam., basal glands sometimes present, slit-like, c. 0.8 mm long. /nflorescences up to 12 cm long; axes angular, basally flattened, grooved, very densely minutely patently yellowish brown hairy; flowers solitary or in basal part with 2 together; lower bracts (sub)opposite. Pedice/ 3.5—5 mm, slightly grooved, very densely appressedly hairy. Sepals: outer sepals 2.6—3 by 2.5—3 mm; inner sepals 4.4—5.5 by 3—4.5 mm. Petals whitish, when dry dark orange red, the longest one 10.5 mm long; carina rather densely ap- pressedly hairy outside, subglabrous inside; other petals outside in basal part shortly appressedly hairy, further glabrous. Stamens: filaments widened above base and with a knob-like, rather densely hairy ap- pendage at inner side, further glabrous; anthers 0.5—0.7 mm long. Ovary half-patently hairy, up to 1.5 mm stipitate, inserted on a rather wide, minutely hairy receptacle; ovules 4. Fruit globular, c. 1.7 cm diam., dark, shortly patently hairy; pericarp rather soft. Distr. Malesia: Malay Peninsula (Cameron Highlands, Fraser’s Hill, G. Benom). Ecol. Submontane rain-forest, 1000—1500 m. 30. Xanthophyllum vitellinum (BLUME) Dretr. Syn. Pl. 2 (1840) 1277; Watp. Rep. 1 (1842) 248; Hassk. Cat. Hort. Bog. (1844) 227; Pl. Jav. Rar. (1848) 296; Mia. Fl. Ind. Bat. 1, 2 (1858) 129; Hassk. in Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 193; Mia. Lc. 272; TEUSM. & Brinn. Cat. Hort. Bog. (1866) 218; CuHopaT, Monogr. I (1891) t. 9, f. 1, 2; t. 12, f. 4c—e; Burck, Wand. Bot. Tuin Btzg (1892) 31; WIESNER, Ann. Jard. Bot. Btzg, Suppl. 2 (1898) 97, t. 3; BoERL. Cat. Hort. Bog. (1899) 58; K. & V. Bijdr. Booms. Java 5 (1900) 294; VALETON, Icon. Bog. 1, 4 (1901) 9, t. 79; GAGNEP. in Desv. J. Bot. 21 (1908) 251; FI. Gén. I.-C. 1 (1909) 243; Backer, Schoolfl. Java (1911) 80; Koorp. Exk. Fl. Java 2 (1912) 453; MERR. En. Born. (1921) 326; BAKER, J. Bot. 62 (1924) Suppl. 7; DocTERS VAN LEEUWEN, Zoocecidia (1926) 273, 274; GAGNEP. FI. Gén. I.-C. Suppl. 1 (1939) 218; BACKER & BakH./f. Fl. Java 1 (1963) 200; MEIJDEN, Leiden Bot. Ser. 7 (1982) 97. — Jakkia vitellina BLuME, Cat. (1823) 17, 64; NEEs, Fl. Bot. Zeit. 8 (1825) 120 (‘Jackia’); BLUME, Bijdr. (1825) 61 (‘Jackia’); G.Don, Gen. Hist. 1 (1831) 368. — Jak- kia longifolia BLUME, Bijdr. (1825) 61 (‘Jackia’); G.Don, Gen. Hist. 1 (1831) 368 (‘Jackia’). — Mon- nina longifolia (BLUME) SPRENGEL, Syst. Veg. 3 (1827) 265. — Monnina vitellina (BLUME) SPRENGEL, l.c. 265; STEUDEL, Nom. ed. 2, 2 (1841) 157. — X. longifolium (BLUME) DietrR. Syn. Pl. 2 (1840) 1277; Hassk. Cat. Hort. Bog. (1844) 228; Mia. Fl. Ind. Bat. 1, 2 (1858) 129; Hassk. in Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 194. — Monnina macrophylla STEUDEL, Nom. ed. 2, 2 (1841) 157, nom. illeg. — X. paniculatum Mia. Fl. Ind. Bat., Suppl. (1861) 393; BaKER, J. Bot. 62 (1924) Suppl. 7. — X. flavescens (non Roxs.) F.-Vitt. Nov. App. (1880) 14; VIDAL, Sinopsis (1883) 13. — X. griffithii (non A.W.BEN- NETT) ROLFE, J. Bot. 23 (1885) 210; VipAL, Rev. PI. Vasc. Filip. (1886) 51; CERon, Cat. Pl. Herb. Manil- la (1892) 19. — X. hookerianum Kina, J. As. Soc. Beng. 59, ii (1890) 139; RrpLey, Fl. Mal. Pen. 1 (1922) 144; Burk. Dict. (1935) 2268; No, Tree FI. Mal. 1 (1972) 359. — X. kunstleri KinG, J. As. Soc. Beng. 59, ii (1890) 139; Ann. R. Bot. Gard. Calc. 5 (1896) 137, pl. 162; RmpLey, J. Str. Br. R. As. Soc. n. 33 (1900) 45; BurK. & HEND. Gard. Bull. S. S. 3 (1925) 346; Watson, Mal. For. Rec. 5 (1928) 249; Burk. Dict. (1935) 2268; Nc, Tree Fl. Mal. 1 (1972) 359. — X. curtisii Kinc, J. As. Soc. Beng. 59, ti (1890) 138; Ripley, Fl. Mal. Pen. 1 (1922) 146; Burk. Dict. (1935) 2269; Koripa, Gard. Bull. Sing. 17 (1958) 19, 51, f. 1; F. HALLE c.s. Trop. Trees & Forests (1978) 56; CoRNER, Gard. Bull. Sing. Suppl. 1 (1978) 146, 211. — Banisterodes longifolia (BLUME) 1988] POLYGALACEAE O.K. Rev. Gen. Pl. 1 (1891) 46, nom. illeg. — Banis- terodes vitellinum (BLUME) O. K. /.c. 46, nom. illeg. — X. robustum CHopDaAT, Bull. Herb. Boiss. 4 (1896) 262; MerR. En. Born. (1921) 326; En. Philip. 2 (1923) 387; MASAMUNE, En. Phan. Born. (1942) 381; MEuER, Bot. News Bull. Sandakan 7 (1967) 88. —X. robustum var. elmeri CHODAT in Merr. Pl. Elm. Born. (1929) 136; MASAMUNE, En. Phan. Born. (1942) 381. — X. griffithii var. curtisii (KING) NG, Tree Fl. Mal. 1 (1972) 359, f. 2. Shrub or tree, up to 30 m, 36 cm dbh. Axillary buds varying from narrowly triangular with strongly thickened base and then often 1.5—3 mm long, to rhomboid-ovate or ovate-oblong and then often 6—11 mm long. Petiole 8—14(—16) mm, very often with a pair of glands in apical half. Leaf-blade 8—20(—30) by 3.5—11 cm, sometimes a few leaves of a twig smaller; above greyish green to yellowish brown, midrib protruding to nearly flat in basal half; beneath: secondary nerves (6 or) 7—9(—11) pairs, in apical half forming an indistinct intramarginal nerve; glands mostly more than 10, near midrib or scattered, 0.2—0.4 mm diam., basal glands mostly present. Jnflorescences branched, 8—30 cm long, branches often in pairs in lower part; axes basally mostly strongly flattened, grooved, glabrous to densely shortly patently or less often appressedly hairy; in basal part flowers with up to 3 together, solitary in apical part; lower bracts nearly opposite. Pedicel 1.5—5.5 mm, very rarely longer, grooved, densely shortly patently (sometimes appressedly) hairy. Sepals basally often more or less thickened and wrinkled; outer sepals (1.7—)2—3.3(—3.9) by 1.9-4 mm; inner sepals (2.6—)3—5.3(—5.7) by (2.5—)3—4(—5) mm. Petals dark yellow to white, when dry orange to dark reddish and often with white incrustations, the longest one (7—)8—12, ex- ceptionally up to 15.5 mm; carina densely appressed- ly hairy outside; other petals glabrous outside or with a few hairs at apex. Stamens: 8, very rarely 9; fila- ments free or connate over up to 0.7 mm, widened above base and with a knob-like, rather densely hairy appendage at inner side, further glabrous; anthers 0.4—0.6(—0.7) mm long. Ovary subsessile or up to 1 mm stipitate, half-patently hairy; style hairy in basal half, little hairy upwards; ovules 4. Fruit globular, up to 1.8 cm diam., often wrinkled when dry, rather dull or rarely shiny, usually light brown, sometimes dark reddish brown, hairy; pericarp rather thin. Distr. Malesia: Sumatra (incl. Siberut & Simalur Is.), Malay Peninsula (incl. Penang), Java, Borneo, Philippines (Babuyan, Luzon, Mindanao). One of the most common species. Note. Three collections from Sumatra, Riouw District (bb 24833, 27509, 30108) and an (otherwise different) collection from Borneo (S 23996) have an unusual type of axillary buds. The buds are globular (van der Meijden) 515 to ovoid, 1.5—2.5 mm long, rather strongly thick- ened in the middle and apical part. Such buds also occur in the sterile collection SF 20520 from the Anambas Is., in which, however, most axillary buds are very large and flat, c. 10O—12 by 5S—6 mm, resemb- ling those of 40. X. heterophyllum. 31. Xanthophyllum incertum (BLUME) MEIDEN, Leiden Bot. Ser. 7 (1982) 99, f. 3A-e. — Guatteria in- certa BLUME, Fl. Java (1830) 100, t. 49B. — ? X. acuminatissimum Mig. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 276. — Monoon incertum (BLUME) Mia. ibid. 2 (1865) 19. Small tree, to 10 m. Axillary buds narrowly trian- gular to lanceolate, (2—)4.5—10(—11) mm long, more or less wrinkled; buds of second order rarely present. Petiole (6—)8—10(—12) mm. Leaf-blade 6—16(—22) by 2.3—5.5(—8) cm, apex cuspidate; above dark green, shiny; beneath green, secondary nerves 5 or 6 (or 7) pairs, in apical part forming an intramarginal nerve; glands few, 0.2 mm diam., basal glands some- times present. Jnflorescences branched or un- branched, up to 7 cm long; axes densely shortly hairy; in basal part flowers with 3 together; lower bracts (sub)opposite. Pedicel 1.5—2 mm, densely shortly more or less appressedly hairy. Sepals some- times with tiny glandular spots; outer sepals 2.8—3.5 by 3.1—4.1 mm; inner sepals 4.3—5.6 by 3.1—4.2 mm. Petals pinkish, reddish white or pale yellowish and purplish, when dry orange, the longest one 10.5—11.5 mm; carina shortly and rather sparsely ap- pressedly hairy outside; other petals more or less gla- brous or sometimes sparsely shortly hairy outside in apical part. Stamens: filaments free or connate over 0.5(—1) mm, widened above base and with a knob- like, shortly (half-)appressedly hairy appendage at inner side, further glabrous; filament of lateral alternipetalous stamens hairy to base in two rows; anthers 0.6—0.9 mm long. Ovary patently hairy; style nearly glabrous to rather densely appressedly hairy; ovules 4. Fruit globular, c. 1.5 cm diam., + shiny, brown, densely patently hairy; pedicel up to 3.5(—6) mm. Distr. Malesia: Central Sumatra (Toba Lands, Pajakumbuh, Mt Sago), West and Central Java. Ecol. Montane rain-forest, (200—)500—1300 m. 32. Xanthophyllum adenotus Mig. Fl. Ind. Bat., Suppl. (1861) 393; Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 275; Meupen, Leiden Bot. Ser. 7 (1982) 100. — X. cordatum Korru. ex Mig. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 274; Merr. En. Born, (1921) 325; Riptey, Kew Bull. (1925) 77; Cuopar in Merr. Pl. Elm. Born. (1929) 133, incl. /. aequale Cuopat; Kerr, N. Born. For. Rec. 2 (1938) 225; MASAMUNE, 516 FLORA MALESIANA [ser. I, vol. 10° En. Phan. Born. (1942) 379; MEER, Bot. News Bull. Sandakan 7 (1967) 87. — X. arsatii C.E.C.FISCHER, Kew Bull. (1932) 176; MASAMUNE, En. Phan. Born. (1942) 379. Shrub or small tree, up to 10 m, 25 cm dbh. Twigs glabrous or rarely minutely patently hairy. Axillary buds oblong or sometimes ovate-lanceolate, (1.3—)3—6(—10.5) mm long, the longer ones strongly thickened at base and usually suddenly widened and flattened upwards, usually glabrous, the smaller ones less distinctly flattened and usually rather densely shortly hairy; buds of second order often present. Petiole (8—)15—18(—23) mm, glabrous to rather densely minutely hairy, usually with 2 small, promi- nent glands. Leaf-blade (linear-)lanceolate, (9—)22—47 by (1.3—)5—10(—20) cm; base usually cordate with the margins curved upwards and con- nate above the apex of the petiole, or flat and round- ed to broadly cuneate, apex acutish; above usually slightly bullate between secondary nerves and intra- marginal nerve, greyish green to brown; beneath usually brownish, glabrous to minutely patently hairy all over, secondary nerves (9—)13—20 pairs, mostly forming a distinct, nearly complete intramar- ginal nerve; glands 2—6(—20), usually near the mid- rib (if few, only present in basal part), 0.3—0.4 mm diam. /nflorescences sometimes also axillary on the older nodes; axes slightly angular, slightly grooved, main axis basally usually sparsely minutely appres- sedly hairy, side axes and main axis in upper part more densely hairy; flowers solitary or very rarely with 2 together; lower bracts opposite. Pedicel (1—)1.5—2(—3.5) mm, more or less distinctly grooved, densely minutely appressedly to patently hairy. Sepals often with minute, rather distinct glands; outer sepals (2.1—)2.8—3.8(—4.1) by (2.4—)3—4.9 mm; inner sepals 3-—5.5 by (2.8—)3.4—4.6 mm. Petals pinkish to pale violet, the upper petals with a yellow spot, when dry dark red, the longest one (8.5—)9.5—12.5(—14.5) mm; carina rather densely appressedly hairy outside, inside sparsely minutely hairy in apical part only, further glabrous; other petals very sparsely minutely hairy above base outside, sparsely patently hairy outside near apex, inside glabrous to hairy up to + halfway. Stamens: filaments free or connate over 1(—2) mm, widened above base and especially those of abaxial stamens with a more or less distinct (half-)patently hairy knob-like thickening at inner side, further gla- brous; anthers (0.6—)0.7—0.9(—1) mm long, ciliate along slits. Ovary (half-)patently hairy; style (rather) sparsely half-patently hairy in basal half, very sparsely hairy in apical half, glabrous near apex; ovules 4. Fruit globular, 1.5—1.8 cm diam., rather dull, light to reddish brown, distinctly hairy; peri- carp thin, brittle. Distr. Malesia: Sumatra, Borneo. KEY TOT HE VARIETIES 1. Leaf-blade (4.5—)5—10(—20) cm wide. Secondary nerves 13—20 pairs, forming a nearly complete intramarginal nerve.......... a. var. adenotus 1. Leaf-blade 2—5.2 cm wide. Secondary nerves 9-14 pairs, forming a weak, irregular intramar- Binal nesvews2,..ccn eee eee b. var. lineare a. var. adenotus Axillary buds (1.8—)3—6(—10.5) mm long. Petiole (10—)15—21 mm, glabrous or hairy. Leaf-blade (15—)22—47 by (4.5—)5—15 cm, base cordate to cu- neate. Secondary nerves 13—20 pairs, forming a dis- tinct, nearly complete intramarginal nerve. Flowers: upper petals glabrous or hairy inside to about half- way. Distr. Malesia: Sumatra, Borneo. b. var. lineare MEIDEN, Leiden Bot. Ser. 7 (1982) 101. Axillary buds at upper side partly enclosed by a distinct ridge formed by the twig, 1.3—2.2 mm long. Petiole (8—)10—12(—18) mm, glabrous. Leaf-blade linear-lanceolate with more or less parallel sides over most of its length, (9Q—)13—30 by (1.3—)2—5.2 cm, base rounded to obtuse. Secondary nerves 9-14 pairs, forming a weak, irregular intramarginal nerve. Flowers rather small in all parts; upper petals shortly patently hairy inside to about halfway. Fruit unknown. Distr. Malesia: Borneo (Sabah). 33. Xanthophyllum palawanense Emer, Leafl. Philip. Bot. 5 (1913) 1673; MeEtDEN, Leiden Bot. Ser. 7 (1982) 101. — X. cordatum (non MiqQ.) MERR. En. Philip. 2 (1923) 386. Small tree, up to 8 m, 5 cm dbh. Twigs sparsely minutely hairy, glabrescent, older nodes often strongly thickened and with numerous adventitious buds. Axillary buds oblong, 6—9 mm long, basally narrowed and strongly thickened, sparsely shortly hairy, glabrescent. Petiole 15—18 mm, more or less densely shortly hairy, with 2(—4) more or less distinctly protruding small glands. Leaf-blade ovate- oblong to ovate-lanceolate, rarely elliptic, ¢. 20—40 by (6—)8—12(—15) cm, base cordate, the margins at base flat or only little upturned, apex gradually nar- rowed to shortly acuminate; above often slightly bul- late between secondary nerves and intramarginal nerve, dark green to greenish brown; beneath sparse- ly minutely hairy on the nerves in basal part, second- ary nerves 9—12 pairs, often irregular, forming a nearly complete, somewhat irregular, intramarginal nerve; glands few, situated in middle and basal part, 0.2(—0.4) mm diam. /nflorescences sometimes also on older shoots from adventitious buds, up to 22 cm 1988] POLYGALACEAE (van der Meijden) 517 long; axes angular, slightly grooved, densely shortly patently hairy; lower bracts opposite. Pedicel 2—3.5 mm, grooved, densely shortly half-patently hairy. Sepals: outer sepals 3.5—4.3 by 4.3—5 mm; inner sepals S—5.9 by 4.3—4.5 mm. Petals dark red when dry, the longest one 15—18.5 mm; carina rather densely shortly appressedly hairy outside, glabrous inside except at base; other petals minutely sparsely appressedly hairy in basal part out- and inside, fur- ther glabrous. Stamens: filaments widened and slightly thickened above base and only there rather densely half-patently hairy; anthers 1.1—1.2 mm long, ciliate along slits. Ovary stipitate for 1—1.5 mm, half-patently hairy; style densely hairy in basal part, upwards sparsely hairy to near apex; ovules 4. Fruit globular, c. 1.7 cm diam., dull brown, rather distinctly half-patently hairy; pericarp rather thin, brittle; pedicel up to S—6 mm. Distr. Malesia: Southern Philippines (Palawan, Sulu Is.: Tawitawi, Jolo). 34. Xanthophyllum ceraceifolium MEUDEN, Bot. J. Linn. Soc. 67 (1973) 117; Leiden Bot. Ser. 7 (1982) 102. Small tree, up to 15 m, 16 cm dbh. Axillary buds elliptic to oblong, S—7 mm long. Petiole (18—)25—30 mm. Leaf-blade 22—42 by 7—15.5 cm; above rather dull, greenish, beneath dull, concolorous, secondary nerves c. 8—10 pairs, little prominent, in apical part forming an indistinct intramarginal nerve, venation obscure; glands 2—8, 2 situated at the very base and 0.6—1 mm diam., the other ones (if present) scat- tered, sometimes close to midrib, 0.5 mm diam. Jn- florescences much shorter than the leaf; axes strong- ly flattened basally, grooved, brown, minutely hairy; lower bracts (sub)opposite. Pedicel 2.5—3.5 mm, grooved, densely shortly patently hairy. Sepals: outer sepals 2.8—3.5 by 3.6—4.4 mm; inner sepals 4.5—4.9 by 3.6—4.7 mm. Petals yellowish, when dry dark red with large incrustations, glabrous inside, the longest one 9—10.5 mm; carina appressedly hairy outside. Stamens : filaments widened above base and with a distinct, rather shortly (half-)appressedly hairy knob-like thickening at inner side, further gla- brous; anthers 0.6 mm long. Ovary nearly sessile, ap- pressedly hairy; ovules 4. Fruit unknown. Distr. Malesia: Borneo (Sarawak: Semengoh; Sabah). 35. Xanthophyllum petiolatum Mevupen, Leiden Bot. Ser. 7 (1982) 103. Tree, 14 m. Twigs minutely patently hairy. Axil- lary buds erect, oblong, c. 11-12 by 4 mm, base broad, rounded, apex rounded. Petiole 26-31 mm. Leaf-blade 6—13.5 by 4—7 cm, base rounded, apex rounded to slightly obtuse; above: midrib sunken in apical half, slightly prominent in basal half, second- ary nerves slightly sunken; beneath glaucous-papil- lose, secondary nerves 6—8 pairs, forming an indis- tinct intramarginal nerve in apical part, venation hardly protruding; glands 1—3, situated in middle and apical part, mostly c. 0.4mm diam., basal glands sometimes present, rather large. /nflorescences up to 15 cm long; axes dark, patently, extremely shortly hairy; lower bracts opposite. Pedice/ 4 mm, grooved, densely very shortly half-patently hairy. Sepals: outer sepals 2.9 by 3.3 mm, slightly pustulate; inner sepals 4 by 3.3 mm. Perals dark red when dry, the longest one 11.5 mm; carina densely appressedly hairy outside; other petals glabrous to sparsely short- ly hairy outside. Stamens: filaments connate over 0.5—0.8 mm between upper and lateral petals, con- nate over c. 1.5 mm between lateral petals and carina, the free parts constricted at very base and then widened and with a distinct densely hairy knob- like thickening at inner side, further glabrous; an- thers 0.7 mm long. Ovary c. 1.5 mm stipitate, appres- sedly hairy; ovules 4. Fruit unknown. Distr. Malesia: Borneo (Brunei: Andalau For. Res.). 36. Xanthophyllum clovis (STEEN. ex MEIJDEN) MEUDEN, Leiden Bot. Ser. 7 (1982) 103. — X. vitellinum var. clovis STEEN. ex MEUDEN, Bot. J. Linn. Soc. 67 (1973) 120. Tree, up to 14 m. Axillary buds with the form of a clove; scales 6.5—12 mm long, at base slightly en- larged and convex, distinctly enlarged at the rounded to + emarginate apex, and there with 2 more or less distinctly prominent knob-like appendages; buds of second order c. 5—6 mm long, hardly thickened at apex. Petiole 9-17 mm. Leaf-blade 8.5—18 by 3.5—6.5 cm; above greenish to brownish; beneath glaucous-papillose, secondary nerves c. 7 or 8 pairs, forming an indistinct intramarginal nerve in apical half; glands few to rather numerous, mostly near the midrib, c. 0.2—0.3 mm, basal ones up to 0.5 mm diam. Inflorescences up to 20 cm long; axes dark, minutely patently hairy; in basal part flowers in clus- ters of up to 7 together; lower bracts opposite. Pedicel 4.5mm, + grooved, densely very shortly, + appressedly hairy. Sepals: outer sepals 3 by 3.2 mm; inner sepals 4.1 by 4.1 mm, with tiny glandular spots at apex. Petals dark red when dry, the longest one 8.5 mm; carina appressedly hairy outside; other petals glabrous outside, the upper ones sparsely ciliate to halfway. Stamens: filaments widened above base and with a distinct densely appressedly hairy knob- like appendage at inner side, further glabrous; an- thers 0.5 mm long. Ovary subsessile, half-patently hairy; style hairy in two rows to near apex; ovules 4, Fruit unknown, Distr. Malesia: Borneo (Brunei, Sabah, Labuan 1.), 3 collections. 518 FLORA MALESIANA [ser. I, vol. 10° Fig. 21. Xanthophyllum bracteatum Cuopat. a. Habit, x 0.5; b. flower; c. flower, longitudinal section, gy- noecium removed; d. gynoecium, all x 3; e. ovary, longitudinal section, x6; /. carina, x4; g. base of leaf with glands, x 2.5 (EDANO BS 28512). 1988] 37. Xanthophyllum reflexum ME DEN, Leiden Bot. Ser. 7 (1982) 104. Small tree, up to 10 m, 12 cm dbh. Twigs minutely patently hairy. Axillary buds erect, appressed against twig, scales laterally flattened, triangular, 3—4.5 mm long, minutely densely patently hairy. Petiole 9-10 mm, densely minutely hairy. Leaf-blade 11-18 by 3.5—5.5 cm; above rather dull, dark greenish to yel- lowish brown, midrib + sunken, nervation rather obscure; beneath nearly concolorous, secondary nerves c. 6—9 pairs, not very distinct, in apical part forming an indistinct intramarginal nerve, venation rather obscure; glands 2 (or 3), situated near base, 0.4—0.7 mm diam. /nflorescences shorter than the leaves; axes dark, very densely more or less patently hairy; lower bracts opposite. Pedicel 2.5—3 mm, + grooved, very densely shortly patently hairy. Sepals: outer sepals 2.2—2.5 by 3.4—3.6 mm; inner sepals 3.8—3.9 by 3.6—4.9 mm. Petals yellowish white, when dry dark red, the longest one 13-14 mm; carina densely more or less appressedly hairy out- side; other petals glabrous. Stamens: filaments widened above base and with a knob-like shortly (half-)appressedly hairy appendage at inner side, fur- ther glabrous; anthers 0.7—0.8 mm long, hairy or nearly glabrous at base. Ovary subsessile, half- patently hairy; ovules 4. Fruit unknown. Distr. Malesia: Borneo (Sarawak: Semengoh). 38. Xanthophyllum angustigemma MEUDEN, Leiden Bot. Ser. 7 (1982) 104. Axillary buds erect or nearly so, mostly flattened against twig; scales ovate-lanceolate, 6—9 mm long, not thickened at base; buds of second order distinct, 3.5—5 mm long. Petiole 10-14 mm. Leaf-blade c. 6—12 by 2.5—5.8 cm; above greyish green to brown- ish green; beneath glaucous-papillose, secondary nerves c. 5—7 pairs, usually forming an indistinct intramarginal nerve in apical part; glands rather nu- merous, scattered, 0.1—0.2 mm diam. /nflorescences about as long as the leaves; axes dark, minutely pa- tently hairy; in basal part flowers with 3 together; lower bracts opposite. Pedicel 3.5—-4 mm, very densely whitish shortly patently hairy. Sepals: outer sepals 3.2—3.7 by 2.9—3.3 mm; inner sepals 3.7—5.4 by 2.3—3.2 mm. Petals dark red when dry, the longest one c. 13 mm; carina densely half-patently hairy outside; other petals faintly hairy along midrib outside. Stamens: filaments c. 2 mm connate, the free parts constricted at very base and then widened and slightly thickened, only there densely half- patently hairy; anthers 0.7 mm long. Ovary half- patently hairy; ovules 4. Fruit unknown. Distr. Malesia: Philippines (Luzon, Samar), 2 collections. POLYGALACEAE (van der Meijden) 519 39. Xanthophyllum bracteatum CuHopatT, Bull. Herb. Boiss. 4 (1896) 258; MERR. En. Philip. 2 (1923) 386; MEUDEN, Leiden Bot. Ser. 7 (1982) 105, f. 15. — Fig. 21. Axillary buds (ob)ovate-oblong to linear-lanceo- late, 7-20 by 1.5—6 mm, basally slightly wrinkled, acute, more or less flat. Petiole 5—11(—14) mm, sometimes with glands. Leaf-blade 9.5—25 by 3-8 cm, rounded-attenuate to cordate; above (yellowish) green, midrib nearly flat to distinctly prominent; be- neath glaucous-papillose, secondary nerves 10—12 pairs, forming a rather distinct intramarginal nerve; glands scattered, numerous, 0.1—0.2 mm diam. /n- florescences up to 10 cm long; axes (rather) densely minutely patently hairy, smooth to pustulate, angular, reddish; flowers solitary or in basal part with up to 3 together, sometimes turned upside- down; lower bracts opposite. Pedicel 5—7 mm, slen- der, very densely patently shortly whitish hairy. Sepals: outer sepals c. 3 by 2.1 mm; inner sepals c. 5—5.5 by 4—4.5 mm. Petals dark red when dry, the longest one c. 14—17 mm; carina rather densely more or less appressedly hairy outside in middle and apical part; other petals nearly glabrous outside. Stamens: filaments connate over 1—3 mm, widened and hardly thickened above base, basally rather densely whitish hairy in 2 rows; anthers c. 1—1.2 mm. Ovary more or less patently whitish hairy; ovules 4. Fruit (im- mature) globular, yellowish brown; pericarp thin. Distr. Malesia: Philippines (Luzon). 40. Xanthophyllum heterophyllum MEuDEN, Leiden Bot. Ser. 7 (1982) 107. — X. pseudostipulaceum (non MERR.) MEUER, Bot. News Bull. Sandakan 7 (1967) 87; WEBERLING, Beitr. Biol. Pfl. 50 (1974) 279, f. 1, 1h Tree, up to 33 m, 70 cm dbh. Axillary buds elliptic to oblong, (8—)11—20(—30) by 6—12(—14) mm; scales flat, wrinkled, indistinctly nerved, more or less shiny, sometimes in middle part with 1—4 rather in- distinct glands, base shortly attenuate, apex rounded to obtuse. Petiole 7—15 mm. Leaf-blade (3—)4.5—12 (—19) by (1.8—)2.5—5(—7.5) cm, apex shortly acuminate to cuspidate; above + shiny, brownish green; beneath yellowish brown, secondary nerves 7—10(—12) pairs, not forming an intramarginal nerve; glands few to many, mostly situated in middle and apical part, 0.2—0.4 mm diam. /nflorescences branched; axes densely patently shortly hairy; lower bracts opposite. Flowers unknown. Fruit globular, up to 1.7 cm diam., shiny, brown, slightly hairy; pericarp rather soft; pedicel 1.5—2.5 mm, densely shortly patently hairy. Seed 1; 3 ovules abortive. Distr. Malesia: Borneo (Sarawak, Brunei, Sabah). Note. Although quite a number of collections are known, these are either vegetative or in fruit. 520 FLORA MALESIANA [ser. I, vol. 103 41. Xanthophyllum korthalsianum Mig. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 277; MeuDEN, Leiden Bot. Ser. 7 (1982) 107. Tree, up to 21 m, 23 cm dbh. Axillary buds insert- ed (1.5—)3—15 mm above the axils on 1—2 mm long stalks; scales elliptic to linear-lanceolate, 6—18 by 1.5-8 mm long, faintly nerved. Petiole 7-10 mm, glands present or not. Leaf-blade 8—14 by 2.5—5 cm; above often slightly bullate between the secondary nerves, mostly dark green, secondary nerves finely prominent to obscure, venation obscure to finely prominent; beneath glaucous-papillose, secondary nerves 6—8 pairs, forming a + distinct intramarginal nerve; glands either not numerous, mostly situated near midrib, and c. 0.3 mm diam., or numerous, scattered, and 0.1—0.2 mm diam. Jnflorescences shorter to much longer than the leaves, the lower branches distinctly supra-axillary, (sub)opposite; axes densely minutely hairy, more or less grooved, angular. Pedicel 1.5-2 mm, grooved, densely patently minutely hairy. Sepals glabrous inside ex- cept for a few hairs at the very base; outer sepals c. 2 by 2mm; inner sepals c. 3.5 by 3 mm. Petals incom- pletely known, carina and lateral petals unknown; upper petal probably c. 8.5 mm long, sparsely hairy at apex. Stamens: unknown. Ovary patently whitish hairy (short and long fine hairs mixed); style and stig- ma unknown; ovules 4. Fruit unknown. Distr. Malesia: Central Sumatra, Borneo (Sara- wak, SE. Kalimantan), 4 collections. 42. Xanthophyllum discolor CHopatT, Bull. Herb. Boiss. 4 (1896) 257; RrpLey, Fl. Mal. Pen. 1 (1922) 147; Watson, Mal. For. Rec. 5 (1928) 249; Burk. Dict. (1935) 2268; WyaTT-SMITH, Mal. For. Rec. 237 (1963) f. 8; NG, Tree Fl. Mal. 1 (1972) 356, f. 1; ME- DEN, Bot. J. Linn. Soc. 67 (1973) 118, incl. ssp. ma- cranthum MEIDeEN; Leiden Bot. Ser. 7 (1982) 108, f. 3A-q. — X. macranthum CHODAT ex ELMER, Leafl. Philip. Bot. 5 (1913) 1674, nomen. — X. hypoleucum Merk. PI. Elm. Born. (1929) 135 (excl. BS 44034); KeitH, N. Born. For. Rec. 2 (1938) 225; MASAMUNE, En. Phan. Born. (1942) 380; MEER, Bot. News Bull. Sandakan 7 (1967) 88. — X. flavo- virens ELMER, Leafl. Philip. Bot. 10 (1939) 3776, nom. inval. (anglice). Very low shrub or small tree, 1-10 m, up to 10 cm dbh. Axillary buds narrowly triangular, c. 2.5-—6mm long, basally and centrally thickened, acute, shiny. Petiole 3—6(—7) mm, often rather shiny. Leaf-blade 3.8—25 by 1.6—10 cm, base obtuse to cordate or cu- neate, apex acutish; above rarely faintly bullate be- tween the secondary nerves, light to dark greyish green, midrib mostly slightly sunken, sometimes a little prominent, venation rather indistinct; beneath glaucous-papillose to nearly smooth, secondary nerves (5 or) 6—13 pairs, forming a mostly rather in- distinct intramarginal nerve; glands numerous, scat- tered, c. 0.1 mm diam., the basal ones often some- what larger. Inflorescences unbranched or rarely with one branch, much shorter than to three times as long as the leaves; axes mostly very slender, mostly less than 1 mm thick, (rather) sparsely minutely hairy; in basal part flowers usually with 3 together; bracts small, either with 2 large glands (in Bornean material) and then rather long-persistent, or eglan- dular (in Malayan and Philippine collections) and then soon caducous; lower bracts (sub)opposite. Pedicel 2—25 mm, slightly grooved, appressedly to patently, sparsely to rather densely hairy, rarely gla- brous. Sepals rarely glabrous outside; outer sepals (1.5—)2—4 by (1.3—)2—3.5 mm, without or with (in most Bornean material) very distinct glands; inner sepals 3—6 by 4—6 mm. Petals white or pinkish, when dry brownish to dark reddish, nearly glabrous, api- cally with few hairs, basally inside slightly hairy, the longest one 11—23 mm. Stamens: filaments free or connate over up to 2 mm; anthers 2—2.5 mm long, minutely hairy all over. Ovary sessile to distinctly stipitate, appressedly whitish hairy; ovules 8—15. Fruit globular, up to 1.8—3 cm diam., dull, light brownish; pericarp thin, rather brittle. Distr. Malesia: Malay Peninsula, Borneo, Phil- ippines. KEY TO THE SUBSPECIES 1. Inflorescence less than half as long as the leaves. Inner sepals 3—3.8 mm long. Longest petals LS amines Nan eee Se a. ssp. discolor 1. Inflorescence 0.5—3 times as long as the leaves. In- ner sepals 5—6 mm long. Longest petals 15—23 MMA Reece is bE oe b. ssp. macranthum a. ssp. discolor — X. discolor CHopaT. — X. hypo- leucum MERR. Very low shrub or small tree, up to 10 m. Second- ary nerves (5 or) 6 or 7(—9) pairs. Inflorescences mostly much less than (rarely up to) half as long as the leaves, often few-flowered. Pedice/ 2—8.5 mm. Sepals: outer sepals (1.5—)2—2.5 by (1.3—)2—2.5 mm; inner sepals 3—3.8 by 4—4.5 mm. Longest petals 11-15 mm. Fruit up to 1.8 cm diam. Distr. Malesia: Malay Peninsula (Johore; Singa- pore), Borneo. b. ssp. macranthum MEUDEN, Bot. J. Linn. Soc. 67 (1973) 118; Leiden Bot. Ser. 7 (1982) 109, f. 3A-q. Low shrub or small tree up to 8 m. Secondary nerves 6—13 pairs. Inflorescences half as long to three times as long as the leaves, many-flowered. Pedicel 10—25 mm. Sepals: outer sepals 2.8—4 by 2.2—3.5 mm; inner sepals 5—6 by 4.2—6 mm. Long- est petals 15—23 mm. Fruit up to 3 cm diam. 1988] POLYGALACEAE Distr. Malesia: throughout the Philippines. 43. Xanthophyllum penibukanense HEINE, Mitt. Bot. Staatssamml. Miinchen 6 (1955) 215; Pfl. Cle- mens Kinab. (1953) 50; MEER, Bot. News Bull. San- dakan 7 (1967) 88; MEIDEN, Leiden Bot. Ser. 7 (1982) 110, f. 9b. Small shrub or tree, up to 12 m, 30 cm dbh. Axil- lary buds ovate-oblong, 3.2—7 mm long, acute, flat, but basally strongly thickened, there often with exu- berant cork-forming which may hide the scale com- pletely, greyish to cream-coloured, more or less shiny. Petiole 8—15 mm, often seemingly much longer because of the long-attenuate leaf base. Leaf- blade (S—)7—28 by 2.3—10 cm, base long-attenuate, above dark green, shiny, midrib slightly sunken to slightly prominent basally, further + prominent, secondary nerves and venation very distinct, some- times even more distinct than beneath; glaucous- papillose beneath, secondary nerves c. 4—6 pairs, first nerves reaching to halfway or further, intra- marginal nerve in apical part rather distinct; glands very numerous, scattered, 0.1—0.2 mm diam. Jnflo- rescences unbranched, shorter than the leaves; axes minutely rather sparsely appressedly hairy to nearly glabrous; in basal part flowers with up to 3 together. Pedicel 2.5—6 mm, more or less smooth, minutely sparsely to densely, appressedly hairy. Sepals sparse- ly hairy along the midrib to (nearly) glabrous out- side, often some with rather distinct glandular spots; outer sepals 2.7—2.9 by 2.1—2.4 mm; inner sepals 3.1—3.3 by 3—3.2 mm. Petals creamish white to light purplish, the upper ones with a yellow spot, when dry orange, glabrous outside, apically and_ basally sparsely ciliate, the longest one 11—13 mm. Stamens: filaments connate over up to 2.5 mm; anthers c. 1.2—1.8 mm long, often minutely hairy all over. Ovary 1\—2 mm stipitate, glabrous to densely appres- sedly whitish hairy; style thinly appressedly hairy in basal part, further glabrous; ovules 8—12. Fruit sometimes distinctly stipitate, globular, c. 1.5 cm diam., + shiny, brownish, glabrous or nearly so; pericarp thin; pedicel up to 10 mm. Distr. Malesia: Borneo (Sarawak, E. Kaliman- tan). Ecol. Mostly found 500—1500(—1800?) m. Note. Very variable in the indumentum of the ovary. in mountain forests, 44. Xanthophyllum pseudoadenotus Mevupen, Leiden Bot. Ser. 7 (1982) 110. — X. stapfii Cuovat, Bull. Herb. Boiss. 4 (1896) 260, p.p., pro specim. HAVILAND 1620. Small tree, up to 9 m, 12 cm dbh. Axillary buds oblong, 3—6 mm long, basally strongly thickened, obtuse, probably glabrous; secondary and adven- (van der Meijden) 521 titious buds often present, numerous. Pefiole (8—)10.5—14 mm, smooth. Leaf-blade 13-32 by 5—10.5 cm, base attenuate to cordate, apex acutish; above dark green, often slightly bullate between the secondary nerves, midrib prominent, often with a groove from halfway down; beneath glaucous- papillose, secondary nerves 11—14 pairs, forming a rather distinct complete intramarginal nerve; glands numerous, smaller than 0.1 mm diam. /nflorescences 1.5—6 cm long, also arising from adventitious buds on older nodes, unbranched; axes minutely appres- sedly hairy (hairs 0.1 mm long). Pedicel S—6 mm, minutely appressedly hairy (hairs 0.1 mm long). Sep- als: outer sepals 2 by 2 mm; inner sepals 3.2 by 2.7 mm. Peta/s subglabrous, brownish orange when dry, the longest c. 15 mm. Stamens: filaments c. 1.5 mm connate; anthers 2 mm long, ciliate along slits. Ovary minutely appressedly hairy (hairs 0.1—0.2 mm long); style glabrous in apical part; ovules 9—11. Fruit globular, c. 1.5 cm diam., minutely appressedly hairy; pericarp thin, brittle; pedicel up to 7 mm. Distr. Malesia: Borneo (Sarawak, Sabah). Note. Resembling 45. X. pulchrum ssp. stapfii, differing in the shorter, appressed hairs of inflores- cence and flowering parts, in the greater number of secondary nerves which form a distinct intramarginal nerve, in the longer pedicel, and in the smaller sepals. 45. Xanthophyllum pulchrum Kina, J. As. Soc. Beng. 59, ii (1890) 141; Ann. R. Bot. Gard. Calc. 5 (1896) 138, pl. 164; GaGNEP. in Desv. J. Bot. 21 (1908) 252; RimpLey, Fl. Mal. Pen. 1 (1922) 146; Burk. & HEND. Gard. Bull. S. S. 3 (1925) 346; No, Tree Fl. Mal. 1 (1972) 361, f. 3; CorNER, Gard. Bull. Sing. Suppl. 1 (1978) 27, 147, 211; MEpDEN, Leiden Bot. Ser. 7 (1982) 111. — X. stapfii CHopat, Bull. Herb. Boiss. 4 (1896) 260 (exc/. HAVILAND 1620); in E. & P. Nat. Pfl. Fam. 3, 4 (1896) 345 (‘stapferi’); Merk. En. Born. (1921) 326; MASAMUNE, En. Phan. Born. (1942) 381. — X. densiflorum Cuopat, Bull. Herb. Boiss. 4 (1896) 256; Merr. En. Born. (1921) 325; MASAMUNE, En. Phan. Born. (1942) 379. Small shrub to small tree, up to 8 m. Twigs gla- brous to minutely patently hairy. Axillary buds ovate, 1.8—3.5(—5?) mm long, very thick (mostly not especially basally), obtuse, light brown and often more or less reddish, often irregular because of cork- forming. Petiole 4—9 mm, the young ones nearly smooth, not transversely wrinkled, glabrous to min- utely densely patently hairy all round, the older ones soon becoming transversely cracked, more or less corky; glands often present, mostly rather distinct. Leaf-blade (5.5—)7.5-30 by 2.4—11.5 cm; base rounded-cordate, rarely rounded, obtuse, or cu- neate-rounded, apex acutish, rarely rounded; above rarely bullate between midrib and secondary nerves, greyish green, midrib slightly sunken to flat, rarely a22 FLORA MALESIANA indistinctly prominent; beneath glaucous-papillose, secondary nerves (6 or) 7—12 (or 13) pairs, forming mostly an indistinct intramarginal nerve in apical half; glands numerous, scattered, c. 0.1—0.2 mm diam. /nflorescences at end of young twigs but also axillary, not rarely on old nodes, unbranched, shor- ter than the leaves, many-flowered; axes stiff, minutely patently hairy; in basal part flowers with up to 3 together. Pedicel 2.5—3.5(—4.5) mm, finely grooved, very densely minutely patently hairy. Sepals often with rather distinct glands; outer sepals 2.1—5 by 2.7—4.2 mm;; inner sepals 3.2—6 by 3.2—4.9 mm. Petals pink or whitish, when dry red or brown- ish orange, slightly hairy apically and basally inside, further glabrous, the longest one 13—18 mm. Sta- mens: anthers (1.3—)1.7—2.5(—3.6) mm long, faintly hairy at base, sparsely ciliate along slits. Ovary up to 2 mm stipitate, patently light brownish pubescent; style glabrous in apical part; ovules 12—16. Fruit globular, up to 2 cm diam.; pericarp thin. Distr. Malesia: Sumatra, Malay Peninsula, Borneo. KEY TO THE SUBSPECIES 1. Secondary nerves 6 or 7 pairs. Longest petals 12—16 mm. Anthers 1.3—1.8 mm long a. ssp. pulchrum 1. Secondary nerves 7—13 pairs. Longest petals 15—18 mm. Anthers (1.7—)1—3.6 mm long b. ssp. stapfii a. ssp. pulchrum — X. pulchrum KING. Petiole 4—6.5 mm. Secondary nerves (6 or) 7 pairs. Outer sepals 2.1—4 by 2.7—3.9 mm, inner sepals 3.2—4.7 by 3.4—4.3 mm. Longest petals 12-16 mm. Anthers 1.3—1.8 mm long. Distr. Malesia: Sumatra, Malay Peninsula (incl. Penang I.). b. ssp. stapfii (CHODAT) MEUDEN, Leiden Bot. Ser. 7 (1982) 112. — X. stapfii CHopat. — X. densiflo- rum CHODAT. Petiole (4—)5—9 mm. Secondary nerves 7—12 (or 13) pairs. Outer sepals 3.4—5 by 3.1—4.2 mm, inner sepals 3.6—6 by 3.2—4.9 mm. Longest petals 15—18 mm. Anthers (1.7—)2.1—2.5(—3.6) mm long. Distr. Malesia: Borneo (Sarawak, Kalimantan). 46. Xanthophyllum beccarianum CuopatT, Bull. Herb. Boiss. 4 (1896) 257; Monogr. I (1891) t. 9, f. 3; MerrR. En. Born. (1921) 325; MASAMUNE, En. Phan. Born. (1942) 379; MeER, Bot. News Bull. Sandakan 7 (1967) 87; MEIDEN, Leiden Bot. Ser. 7 (1982) 112. Tree, up to 12 m, 17 cm dbh. Twigs very densely patently hairy (hairs up to 1 mm). Axillary buds tri- [ser. I, vol. 10° angular, 0.5—4.5 mm long, basally strongly thick- ened, acutish. Petiole 5—6 mm, very densely hairy. Leaf-blade 9.5—19 by 4—8 cm, base cordate, apex acutish; above dark green; beneath glaucous-papil- lose, rather sparsely hairy, midrib rather densely hairy, secondary nerves c. 6—8 pairs, forming an in- distinct intramarginal nerve in apical part; glands nu- merous, scattered, c. 0.1 mm diam. J/nflorescences unbranched, shorter than the leaves; axes densely minutely hairy (hairs up to 0.4 mm); in basal part flowers with up to 3 together, sometimes turned upside-down. Pedicel 7.5—9.5 mm, densely patently minutely hairy (hairs up to 0.25(—0.4) mm). Sepals nearly glabrous (very shortly hairy); outer sepals 2.5—2.7 by 3—3.6 mm, with rather distinct glandular spots; inner sepals 3.2—4 by 2.8—3.4 mm. Petals orange-red when dry, glabrous except for the ciliate base, the longest one c. 16—16.5 mm long, minutely hairy at base along margin, further glabrous. Stamens: anthers 2.2 mm long. Ovary patently hairy; style glabrous in apical half; ovules 13. Fruit (immature) apically pointed; pedicel c. 10-12 mm. Distr. Malesia: Borneo (Sarawak). 47. Xanthophyllum pedicellatum MEuDEN, Leiden Bot. Ser. 7 (1982) 113. Shrub to tree, 3.5—23 m, 60 cm dbh. Twigs densely patently brownish hairy, glabrescent. Axillary buds narrowly triangular, 1.5—3.5 mm long, hairy. Pet- iole 1.5—2.5(—3) cm, very densely patently hairy. Leaf-blade (S—)9—11 by (1—)1.5—3(—4) cm, base cu- neate to rounded or slightly cordate, apex usually acutish; above dark green, shiny, beneath glaucous- papillose, hairy mainly on midrib, secondary nerves c. 7 or 8 pairs, forming an indistinct intramarginal nerve or not; glands very numerous, scattered, c. 0.1 mm diam. /nflorescences unbranched, as long as the leaves; axes densely minutely patently hairy, some hairs up to 0.5 mm. Pedice/l (9—)10—15 mm, minute- ly patently hairy. Sepals very sparsely minutely hairy outside, glabrous inside except at very base; outer sepals c. 2—2.5 by 2 mm; inner sepals c. 3—3.5 by 2—2.4mm. Petals pinkish, when dry orange-red, gla- brous except for ciliate base, the longest one 12.5 mm. Stamens: anthers 1.5—1.6 mm long, sparsely minutely hairy at base. Ovary nearly sessile, half-patently brownish pubescent; style glabrous in apical part; ovules 9-11. Fruit globular, c. 2.2 cm diam., light brownish, hairy; pericarp thin. Distr. Malesia: Borneo (E. Sabah). 48. Xanthophyllum purpureum RIDLEY, Kew Bull. (1938) 114; MepDEN, Leiden Bot. Ser. 7 (1982) 114. — X. molle Riwiey, Kew Bull. (1938) 114. Shrub or small tree, up to 5 m, 10 cm dbh. Twigs very densely patently hairy. Axillary buds narrowly triangular, (1.5—)3—5 mm long, basally thickened. 1988] POLYGALACEAE (van der Meijden) 523 Petiole c. 5 mm, densely hairy. Leaf-blade (6—)10—20 by (1.5—)2.5—9 cm, base cordate to rounded-attenuate, rarely cuneate, apex acutish; above green, midrib slightly sunken to fiat; beneath glaucous-papillose, more or less densely hairy all over or only on nerves, secondary nerves (5 or) 6 or 7 pairs, not or only in apical part forming an intra- marginal nerve; glands numerous, scattered, c. 0.1 mm diam. /nflorescences unbranched, shorter than the leaves, often curved downwards; axes sparsely minutely hairy (hairs up to 0.2 mm long); in basal part flowers with 3 together. Pedice/ 2.5—5 mm, minutely densely hairy (hairs up to 0.2—0.3 mm). Sepals rather densely minutely hairy outside (hairs 0.1 mm), subglabrous inside, often with tiny, rather indistinct glandular spots; outer sepals 1.8—3 by 2.1—2.6 mm; inner sepals 2.7—4.3 by 2.7—3.5 mm. Petals (light) purple to rosa-violet, when dry orange- red, ciliate at base and apex, further glabrous, the longest one 11—12(—14?) mm. Stamens: filaments free or 0.4 mm connate; anthers 0.9—1.4 mm long, glabrous to shortly hairy at base. Ovary subsessile or c. 1.5 mm stipulate, patently hairy; style glabrous in apical half; ovules 8—14. Fruit globular, 1.2—1.5 cm diam., usually with remnant of style, hairy; pericarp thin; pedicel curved. Distr. Malesia: Borneo (Sarawak, Sabah, NE. Kalimantan). 49. Xanthophyllum reticulatum CHoDAT in Merr. PI. Elm. Born. (1929) 136; MEUDEN, Leiden Bot. Ser. 7 (1982) 114. Small tree, 2.5—15 m. Twigs very densely patently hairy (hairs up to I mm long). Axillary buds narrow- ly triangular, 4—6(—7.5) mm long, hairy. Petiolec. 5 mm, densely pubescent. Leaf-blade 7—19 by 3—5.5 cm, base obtuse to rounded, apex acutish; above dark green, midrib, secondary nerves and part of finer nervation sunken, midrib hairy at very base; be- neath green, smooth or indistinctly papillose, hairy on midrib and on basal part of nerves, secondary nerves c. 8 pairs (difficult to count), tertiary nerves strongly protruding, blade bullate in-between; finer nerves not strongly prominent; glands numerous, scattered, c. 0.1 mm diam. Flowers unknown. In- fructescences 0.8—4.5 cm long, unbranched; axes shortly sparsely hairy (hairs up to 0.25 mm long). Fruit globular, c. 1.5 cm diam., sessile, with remnant of style, hairy; pedicel S—10.5 mm, minutely patently hairy (hairs up to 0.2 mm long). Seed 1; abortive ovules 11—13. Distr. Malesia: Borneo (Sabah). 50. Xanthophyllum trichocladum CHopart in Merr. Pl. Elm. Born. (1929) 137; Masamune, En. Phan. Born. (1942) 382; Mever, Bot. News Bull. Sandakan 7 (1967) 87; Mevpen, Leiden Bot. Ser. 7 (1982) 115. Shrub or small tree, up to 12 m, 13 cm dbh. Twigs very densely patently hairy. Axillary buds ovate- oblong, (1.5—)2.5—5(—6) mm long, densely hairy. Petiole c. 4—7 mm, very densely hairy. Leaf-blade 11—31 by 3—9 cm, base cordate, covering upper side of petiole, apex acutish; above green, dull, hairy on the midrib; midrib distinctly sunken, rarely flat, sec- ondary nerves and intramarginal nerve faintly sunken, rarely slightly prominent, venation little prominent; beneath glaucous-papillose, pubescent all over, secondary nerves c. 9(—12) pairs, forming a distinct intramarginal nerve; glands very numerous, scattered, c. 0.1 mm diam. Jnflorescences un- branched, shorter than to as long as the leaves or sometimes with one side-branch at very base; axes very densely brownish patently pubescent (most hairs 0.5—0.8 mm long); flowers often turned upside- down. Pedicel 5—7 mm, very densely brownish hairy (hairs up to 1 mm long); pedicels of flower buds at first curved downwards, of open flowers turned up- wards and often half-twisted, rarely straight, pedi- cels of fruits curved downwards again. Sepals very densely brownish pubescent outside (hairs up to 1 mm long); outer sepals 3.2—3.8(—4) by 2.7—3.2 mm; inner sepals 3.2—4.2(—5.6) by 3.2—4.2 mm. Petals pink, the upper ones with a yellow spot, when dry dark reddish, the longest one 13(—16) mm; carina glabrous to sparsely appressedly hairy outside along central veins; other petals glabrous except for a few hairs at base, sometimes sparsely ciliate in basal part. Stamens: filaments free or 0.1—0.5(—1.5) mm con- nate; anthers 2.2—3 mm long, faintly hairy at base, ciliolate along slits. Ovary patently hairy; ovules 11— 16. Fruit globular, c. 1.5 cm diam., densely hairy; pericarp rather thin; sepals subpersistent in fruit. Distr. Malesia: Borneo (Sarawak, E. Sabah, Sa- marinda). 51. Xanthophyllum erythrostachyum GAGNEP. in Desv. J. Bot. 21 (1908) 250; Bull. Soc. Bot. Fr. 56 (1909) 36; MeWDEN, Leiden Bot. Ser. 7 (1982) 115, excl. STEENIS 10075. — X. forbesii BAKER, J. Bot. 62 (1924) Suppl. 7, nom. superfl. Axillary buds 0.8—1.5 mm long, minutely hairy, glabrous. Petiole 3-4 mm. Leaf-blade 10-—20.5 by 4—5.5 cm, base cordate to cordate-truncate, apex distinctly acuminate; upper side dull, greyish green, midrib prominent, nervation rather obscure; beneath greenish, secondary nerves 8—10 pairs, rather indis- tinct, forming an indistinct intramarginal nerve in upper part, venation rather obscure; glands numer- ous, 0.2—0.3 mm diam. /nflorescences unbranched or with one side-branch, up to 8 cm long; axes angu- lar, orange, rather sparsely appressedly minutely hairy. Pedicel 4—4.5 mm, rather densely appressedly minutely hairy. Sepals: outer sepals 2.7-3 by 1.8—2.2 mm, with rather distinct glands; inner sepals 524 FLORA MALESIANA [ser. I, vol. 10° 3.5—4 by 2.7-3.2 mm. Petals whitish tinged with rosa, when dry reddish orange, the longest one 13 mm; carina outside glabrous to rather sparsely patently minutely hairy near base, near apex very sparsely appressedly hairy, further glabrous; other petals glabrous. Stamens: filaments 0.3 mm con- nate: anthers 1—1.2 mm long, glabrous at base, cil- iate along slits. Ovary appressedly rather shortly brownish hairy; ovules 11. Fruit unknown. Distr. Malesia: S. Sumatra (Lampong Distr.), one collection. 52. Xanthophyllum laeve MevDEN, Bot. J. Linn. Soc. 67 (1973) 118 (‘leavis’); Leiden Bot. Ser. 7 (1982) 116. Shrub or small tree, 3—6 m. Axillary buds c. 1 mm long. Petiole 5-7 mm. Leaf-blade 4.5—13.5 by 1.9—5.3 cm, apex cuspidate; above greenish to red- dish brown, dull, midrib sunken, hardly visible, nerves obscure to slightly protruding; beneath light greenish to reddish brown, secondary nerves c. 5 pairs, rather indistinct to slightly protruding, form- ing an indistinct intramarginal nerve, venation indis- tinct; glands rather few, near midrib, 0.3—0.4 mm diam. Jnflorescences branched or unbranched, shorter than the leaves; axes glabrous, more or less smooth. Pedicel 8—15 mm, grooved, dark, glabrous. Sepals: outer sepals 1.8—2.1 by 2.4 mm; inner sepals 2.8—3 by 2.8 mm. Petals white with red spots, when dry reddish orange, the longest one 11—13 mm; carina shortly sparsely appressedly hairy outside, shortly hairy inside; other petals sparsely hairy out- side near apex. Stamens: anthers c. 0.5 mm long. Ovary glabrous or with a few hairs; style very sparse- ly more or less appressedly hairy; ovules 8. Fruit un- known. Distr. Malesia: NE. Sumatra (Sibolangit), 2 col- lections. 53. Xanthophyllum retinerve MEWDEN, Leiden Bot. Ser. 7 (1982) 117. Tree, up to 12 m, 20 cm dbh. Twigs sparsely short- ly appressedly hairy, glabrescent; axillary region shortly densely appressedly hairy. Axillary buds mostly 3, inconspicuous, shortly densely appressedly hairy, the upper one 1—2 mm supra-axillary, some- times more distinct and up to 0.8 mm long. Petiole 6-16 mm, appressedly shortly hairy. Leaf-blade 7-14 by 2.5—5 cm, apex shortly acuminate to cuspidate; above (greenish) brown, midrib sunken to slightly prominent; beneath reddish brown, some- times glaucous, sparsely shortly hairy, secondary nerves 5—7 pairs, sometimes forming an indistinct intramarginal nerve; glands 0—2, mostly close to midrib, 0.2(—0.4) mm diam. Jnflorescences up to 10 cm long, with 2 opposite branches directly above base; axes densely appressedly shortly hairy; lower bracts of side axes opposite. Pedicel 2—2.5 mm, densely appressedly shortly hairy. Sepa/s sometimes with tiny glands; outer sepals 1.4—1.5 by 1.3—1.4 mm; inner sepals 2.1—2.2 by 2.2—2.3 mm. Petals white, when dry orange-red, glabrous inside, the longest one c. 6.5 mm; carina appressedly hairy out- side; other petals sparsely hairy apically. Stamens: anthers 0.3—0.4 mm long. Ovary (sub)sessile, faintly ribbed, densely shortly appressedly hairy; ovules 4. Fruit globular, up to 2.2 cm diam., smooth, rather dull, brown, sparsely appressedly hairy; pericarp c. 4 mm thick, rather hard; pedicel up to 4 mm. Distr. Malesia: Malay Peninsula (Perak; Treng- ganu; Fraser’s Hill). 54. Xanthophyllum eurhynchum Mig. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 277; Kine, J. As. Soc. Beng. 59, ii (1890) 137; GAGNEP. in Desv. J. Bot. 21 (1908) 252; BAKER, J. Bot. 62 (1924) Suppl. 7; MEu- DEN, Leiden Bot. Ser. 7 (1982) 117, f. 3A-a,b. — X. maingayi Hook.f. ex A.W.BENNETT, FI. Br. India 1 (1874) 210; KiNG, Mat. Fl. Pen. (1890) 136; GAGNEP. in Desv. J. Bot. 21 (1908) 252; Burk. Gard. Bull. S. S. 3 (1923) 35; WyatTt-SmitH, Mal. For. Rec. 17 (1952) 362; BALAN MENON, ibid. 19 (1956) 34; Na, Tree Fl. Mal. 1 (1972) 360, f. 3; CornNER, Gard. Bull. Sing. Suppl. 1 (1978) 146, 147, 211. — Banisterodes maingayi (A.W.BENNETT) O. K. Rev. Gen. Pl. 1 (1891) 46, nom. illeg. — X. verrucosum CHODAT, Bull. Herb. Boiss. 4 (1896) 263; RipLey, Fl. Mal. Pen. 1 (1922) 147; HEND. Gard. Bull. S. S. 4 (1928) 222; Watson, Mal. For. Rec. 5 (1928) 249; WyaTrT- SmiTH, ibid. 17 (1952) 364. — X. palembanicum (non Mig.) KinG, J. As. Soc. Beng. 59, ii (1890) 137; Gac- NEP. in Desv. J. Bot. 21 (1908) 251; RmpLey, Fl. Mal. Pen. 1 (1922) 149; Burk. & HEND. Gard. Bull. S. S. 3 (1925) 346; Watson, Mal. For. Rec. 5 (1928) 249; Crap, Fl. Siam. En. 1 (1931) 105; Burk. Dict. (1935) 2269. Shrub or tree, 3—20 m, up to 20 cm dbh. Twigs gla- brous to minutely patently hairy, mostly soon gla- brescent; axillary region shortly densely patently hairy. Axillary buds 2—4, usually less than 0.5 mm long, mostly very densely patently hairy, the upper one sometimes slightly supra-axillary. Petiole 3—9(—11), exceptionally up to 14 mm, glabrous to shortly rather densely hairy in the upper groove, rarely (in some Sumatran coll.) shortly hairy all round; glands mostly indistinct. Leaf-blade rarely linear-lanceolate, (2.5—)3.5—15(—18) by (1—)2—5 (—7) cm, apex acuminate to cuspidate; above sometimes slightly bullate between the secondary nerves, greyish green, midrib slightly sunken or sometimes flat to slightly prominent; beneath rather dull, mostly glabrous, yellowish green, secondary nerves 3—5 (or 6) pairs, forming a rather distinct intramarginal nerve; glands (0—)2—7(—11), 0.1—0.3 1988] (—0.4) mm diam. /nflorescences solitary or with 2 to- gether, mostly unbranched or sometimes with a few short side axes at base; axes angular to terete, minutely hairy; flowers solitary or in basal part with 3 together; bracts and bracteoles relatively long- persistent. Pedicel (1—)2—4(—7.5) mm, densely minutely hairy. Sepals: outer sepals 1.6—2.3 by 1.4—2.3 mm; inner sepals 2.6—3.6 by 2—3.3 mm. Petals: white, when dry yellowish, the longest one 7—8.5(—9) mm; carina mostly bent outwards in open flowers, thus exposing the stamens, rather densely appressedly hairy outside, inside minutely hairy in apical part, ciliate in basal half; other petals glabrous to sparsely hairy outside at apex. Stamens: anthers (0.4—)0.5 mm long. Ovary (sub)sessile, more or less ribbed, more or less appressedly hairy with hairs of different length; ovules 4. Fruit globular to broadly ellipsoid, sometimes irregularly 2—4-sulcate, 1.2—1.8 cm diam., more or less smooth to strongly warty, light brown, velvety to densely appressedly hairy. Distr. S. Thailand; Malesia: Malay Peninsula (also Penang and Singapore), Sumatra. KEY TO THE SUBSPECIES 1. Fruit with distinct warts arranged in longitudinal PPE Sa Oe SO STEIN a. ssp. eurhynchum 1. Fruit more or less smooth, transversely wrinkled or finely rugose or irregularly grooved b. ssp. maingayi a. ssp. eurhynchum MEuDEN, Leiden Bot. Ser. 7 (1982) 119, f. 3A-b. — X. eurhynchum Mig. — X. verrucosum CHODAT. Fruit globular, 1.5—1.8 cm diam., strongly warty, densely appressedly hairy between the warts. Distr. S. Thailand; in Malesia: Malay Peninsula (incl. Penang I.), Sumatra. b. ssp. maingayi (Hook.f. ex A.W.BENNETT) MEU- DEN, Leiden Bot. Ser. 7 (1982) 119, f. 3A-a. — X. maingayi Hook.f. ex A.W.BENNETT, FI. Br. Ind. 1 (1874) 210. — X. palembanicum (non Mig.) KING. Fruit broadly ellipsoid, often irregularly formed, 1.2—1.5 cm diam., sometimes 2—4-sulcate, surface at lateral sides usually with depressions of irregular size or transversely wrinkled, sometimes finely rugose, very densely shortly velvety. Distr. S. Thailand; in Malesia: Malay Peninsula (incl. Singapore). 55. Xanthophyllum wrayi Kino, J. As. Soc. Beng. 59, ii (1890) 138; Ann. R. Bot. Gard. Calc. 5 (1896) 138, pl. 164; Gaonep. in Desv. J. Bot. 21 (1908) 251; Riptey, Fl. Mal. Pen. | (1922) 148; Burk. & HeND. Gard. Bull. S. S. 3 (1925) 346; Henp. ibid. 4 (1928) 222; Watson, Mal. For. Rec. 5 (1928) 249; Burk. POLYGALACEAE (van der Meijden) 525 Dict. (1935) 2268; No, Tree Fl. Mal. 1 (1972) 363, f. 5; CORNER, Gard. Bull. Sing. Suppl. 1 (1978) 147, 211; MEWDEN, Leiden Bot. Ser. 7 (1982) 119, f. 3A-c. — X. puberulum Rip ey, J. Str. Br. R. As. Soc. 7. 73 (1916) 139; HEND. Gard. Bull. S. S. 4 (1928) 222. Shrub to small tree, up to 10 m. Twigs shortly densely patently to appressedly hairy, rarely glabrous except for the densely hairy area above the leaf axil. Axillary buds 2 or 3, densely hairy, up to c. 0.5 mm long. Petiole (S—)7—11(—14) mm, usually not trans- versely wrinkled, densely minutely hairy all round, rarely hairy only in the upper groove, often with rather distinct prominent glands. Leaf-blade 10—30 by 3.5—13 cm, base rarely cordate; above often slightly bullate between the secondary nerves, mostly greyish green, midrib deeply sunken and mostly hairy in basal part; beneath yellowish green, rather dull, minutely hairy or rarely glabrous, secondary nerves 8—15 pairs, forming a distinct intramarginal nerve; glands (2—)4—14, often (very) close to the midrib, sometimes present only in upper part, (0.3—)0.5—0.7 mm diam. /nflorescences shorter than to + as long as the leaves; axes strongly ribbed, flat- tened at base, densely minutely hairy; in basal part flowers with 3 together; bracts and bracteoles rela- tively long-persistent. Pedicels 2.5—7 mm, densely minutely patently hairy. Sepals: outer sepals 2.1—3 by 1.5—2.1 mm, sometimes with small glands; inner sepals 2.8—4 by 2—2.5 mm. Petals white to lilac, the upper ones with a yellow spot, when dry yellowish, the longest one 5.8—7.5(—8) mm; carina densely min- utely hairy in apical part outside and inside; other petals sparsely hairy outside in apical part, lateral petals minutely hairy inside above insertion of fila- ments, upper petals rather densely patently hairy in- side up to apex. Stamens: filaments connate over 0.5—2 mm or sometimes triadelphous, sparsely min- utely hairy in basal part to densely more or less pa- tently hairy in middle part; anthers 0.4—0.5 mm long. Ovary patently to appressedly hairy; style pa- tently hairy in basal part, in apical half sparsely hairy to glabrous; ovules 4. Fruit more or less globular, up toc. 1.5 cm diam., verrucately ribbed to strongly tu- berculate-warty, apically rounded or with the style- scar sunken; pedicel 2-6 mm. Distr. Peninsular Thailand; in Malesia: Malay Peninsula (incl. Penang I.). 56. Xanthophyllum venosum Kino, J. As. Soc. Beng. 59, ii (1890) 139; Riptey, Fl. Mal. Pen. 1 (1922) 222; Meupen, Leiden Bot. Ser. 7 (1982) 120, f. 3A-d. Shrub to small tree, up to 10 m. Twigs rather densely hairy in axillary area, further glabrous to sparsely minutely hairy, Axillary buds 2—4, up to 0,5 mm long, hairy. Petiole (12~)15—21(-—27) mm, gla- brous to sparsely minutely hairy, not transversely 526 FLORA MALESIANA [ser. I, vol. 103 wrinkled, often with glands. Leaf-blade 12.5—40 by 4.5—12(—13) cm, base cuneate to rounded or cor- date; above often slightly bullate between the sec- ondary nerves, greenish to yellowish brown, midrib deeply sunken; beneath yellowish green, mostly gla- brous, secondary nerves 12—20 pairs, forming a dis- tinct intramarginal nerve; glands mostly numerous, scattered, 0.1—0.2 mm diam. /nflorescences some- times also on older nodes, shorter than the leaves; ax- es strongly flattened basally, ribbed, densely minute- ly hairy; in basal part flowers with 3 together; bracts and bracteoles relatively long-persistent. Pedicel 2—4 mm, minutely hairy. Sepals: outer sepals 2.3—3.3 by 2.1—3.3 mm, often with glands; inner sepals 2.8—4 by 2.5—4 mm. Petals light purple, when dry pale yel- low, the longest one 8.5—11.5 mm; carina sparsely minutely appressedly hairy outside, glabrous inside; other petals slightly hairy basally, further glabrous. Stamens: filaments connate over (0.5—)1—3 mm, rarely some filaments free; anthers 0.5—0.9 mm long. Ovary (sub)sessile, ribbed, appressedly hairy; ovules 4. Fruit ovoid, c. 2 by 1.5 cm, apically shortly but distinctly beaked, strongly verrucately ribbed; pedicel 3—5 mm, minutely hairy. Distr. Malesia: Malay Peninsula. 57. Xanthophyllum malayanum MEeEDEN, Bot. J. Linn. Soc. 67 (1973) 118; Nec, Tree Fl. Mal. 1 (1972) 365; CORNER, Gard. Bull. Sing. Suppl. 1 (1978) 146; MEUDEN, Leiden Bot. Ser. 7 (1982) 121. Tree, up to 10 m, 10 cm dbh. Twigs very densely brownish patently hairy with a mixture of very small and long hairs. Axillary buds nearly completely cov- ered by indumentum, narrowly triangular, possibly up to 2.5 mm long. Petiole 4—10 mm, very densely patently pubescent. Leaf-blade 5—24 by 1.5—8.5 cm, base obtuse to slightly cordate, apex obtuse to cuspi- date; above greyish green to light brownish, midrib and nerves sunken, venation obscure; beneath con- colorous, rather densely patently pubescent, second- ary nerves 5—8 pairs, forming a distinct intramar- ginal nerve; glands 2—8, up to 0.1 mm diam. /nflo- rescences unbranched or basally with a pair of side axes, shorter than to as long as the leaves; axes strongly, flattened and ribbed at base, densely rather shortly hairy, mixed with longer patent hairs; bracts and bracteoles small, relatively long-persistent. Pedi- cel 2.5—5.5 mm, ribbed, minutely densely patently hairy. Sepals: outer sepals 2.4—3 by 2.1—3.1 mm; in- ner sepals 3.4—3.8 by 2.9—3.5 mm. Petals purplish, when dry brownish orange, the longest one 7.5—10 mm; carina minutely hairy outside, inside minutely appressedly hairy in apical and basal part; other pet- als minutely patently hairy in apical part outside, up- per petals inside glabrous to rather densely patently hairy. Stamens: filaments free or 1 mm connate, ex- serted from the carina in open flowers; anthers c. 0.5—0.7 mm long. Ovary more or less patently pu- bescent; ovules 4. Fruit (immature) shortly beaked or apically rounded, verrucately ribbed. Distr. Malesia: Malay Peninsula (Pahang, Jo- hore). 2b. Subsection Eystathes Twigs and inflorescence axes sometimes with minute nodal appendages. Axillary buds (2 or) 3—5(—7), often all distant and supra-axillary. Seed(s) 1(—4); testa without a hard inner layer; albumen very thin; embryo without flattened areas near the base, radicle not exserted. 58. Xanthophyllum novoguineense MEUDEN, Leiden Bot. Ser. 7 (1982) 122. Tree, up to 30 m, 40 cm dbh. Axillary buds 0.8—1.6 mm long, faintly keeled. Petiole 6—10 mm. Leaf-blade 4—13.5 by 1.3—6.5 cm; above green; be- neath light green, papillose, secondary nerves c. 6—8 pairs, forming an indistinct intramarginal nerve or not; glands rather numerous, scattered, c. 0.2—0.4 mm diam., basal glands c. 0.6—1.5 mm diam. Jnflo- rescences up to c. 8 cm long; axes light brownish, slender, rather thinly minutely hairy; in basal part flowers with 3 together. Pedice/ 2.5 mm, grooved, rather densely minutely more or less patently hairy. Sepals: outer sepals 2.4 by 2.1 mm; inner sepals 2.9 by 2.5 mm. Petals yellowish orange when dry, the longest one c. 7 mm; carina woolly hairy outside in apical half; other petals outside with a few hairs at apex and at base. Stamens: anthers 0.35—0.4 mm long. Ovary appressedly hairy; ovules 4. Fruit unknown. Distr. Malesia: New Guinea (Sorong in W, Sepik in Central), 4 collections. Note. Part of the flowers of the type collection have abnormally developed ovules; these vary in number from 1—3 and they are placed basally in the ovary and have a distinct funiculus. The majority of the ovaries, however, contained 4 laterally inserted, sessile ovules. 59. Xanthophyllum ngii MEUDEN, Bot. J. Linn. Soc. 67 (1973) 119; No, Tree Fl. Mal. 1 (1972) 365, f. 5; MEUDEN, Leiden Bot. Ser. 7 (1982) 122, f. 3B-a. Tree, up to 35 m, 65 cm diam. at 5 m. Nodal ap- pendages extremely small, present on very short 1988] POLYGALACEAE straight ridges adjoining the insertion of the petiole. Axillary buds 0.5—1(—1.5) mm long, blackish, rather densely hairy. Petiole 7.5—10 mm. Leaf-blade 6—17 by 1.7—6.5 cm, base rounded to narrowly cuneate, margin slightly undulate, apex acutish to shortly acu- minate; above yellowish green or brownish, midrib nearly flat to slightly protruding, sometimes at very base a little sunken; beneath glaucous-papillose, sec- ondary nerves 5—8 pairs; glands 6—12, situated half- way between midrib and margin or near midrib, 0.3—0.5 mm diam., the basal ones mostly larger. Jn- florescences about as long as the leaves; axes flat- tened at base, blackish, rather densely shortly hairy; lower bracts (sub)opposite. Pedice/ 3—4.5 mm, very densely more or less appressedly shortly hairy. Sepals shortly rather sparsely appressedly hairy outside, (sub)glabrous inside; outer sepals 2.2—2.9 by 2—2.8 mm; inner sepals 3.1—3.6 by 2.5—3.2 mm. Petals brownish orange when dry, the longest one 10—12.5 mm; carina shortly appressedly hairy outside along median veins, further glabrous outside. Stamens: anthers 0.3—0.4 mm long, sparsely hairy at base. Ovary subsessile, shortly appressedly hairy; style sparsely appressedly hairy at very base only, further glabrous; ovules 4. Fruit more or less apple-shaped, up to 8 cm diam., the very short pedicel enveloped by the pericarp; pericarp very hard, in mature fruit up to 3. cm thick when dry. Seed 1 (or ‘1—more’ accord- ing to NG, /.c.), up to 2 cm diam. Distr. Malesia: Southern half of Sumatra, Malay Peninsula. 60. Xanthophyllum lanceatum (Miq.) J.J.Smiru, Ic. Bogor. 4 (1912) 109, t. 334; Gorter, Indische Mer- cuur 34 (1911) 410 (‘lanceolatum’); Tropenfl. 16 (1912) 50; Hea1, Fl. Mitteleur. 5-1 (1925) 87 (‘lanceo- latum’); HEYNE, Nutt. Pl. (1927) 901; Burk. Dict. (1935) 2268; MEUDEN, Leiden Bot. Ser. 7 (1982) 124, f. 1OA. — Skaphium lanceatum Mia. FI. Ind. Bat., Suppl. (1861) 357; Kurz, J. As. Soc. Beng. 40, ii (1871) 46; Scuerrer, Nat. Tijd. Ned. Ind. 34 (1874) 105. — X. glaucum Watt. [Cat. (1831) 4199] ex Hassk. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 193; Kurz, J. As. Soc. Beng. 42, ii (1873) 80; A.W.BEN- NETT, FI. Br. India 1 (1874) 209; Kurz, For. FI. Br. Burma | (1877) 81; Kinc, Mat. Fl. Mal. Pen. (1890) 136; Gace, Rec. Bot. Surv. India 3 (1904) 24; WiitaMs, Bull. Herb. Boiss. II, 5 (1905) 219; BRAN- pis, Indian Trees (1906) 44; Gaonep. in Desv. J. Bot. 21 (1908) 251; Fl. Gén. I.-C. 1 (1909) 245; Riptey, J. Str. Br. R. As. Soc. n. 59 (1911) 73; ibid. n. 73 (1916) 140; Fl. Mal. Pen. | (1922) 147; Warson, Mal. For. Rec. 5 (1928) 249; Crevost & PétreLot, Bull. Econ. Indochine (1929) 138; Cram, FI. Siam. En. 1 (1931) 105; Burk. Dict. (1935) 2268; Gaonep. Fl. Gén. L.- C. Suppl. 1 (1939) 219; Henn. J. Mal. Br. R. As. Soc. 17 (1939) 36; STADELMAN, For. Southeast Asia (van der Meijden) 527 (1966) 186; No, Tree Fl. Mal. 1 (1972) 357, f. 1. — Banisterodes glaucum (WALL. ex Hassk.) O. K. Rev. Gen. Pl. 1 (1891) 46, nom. illeg. — X. microcarpum CuHopatT, Bull. Herb. Boiss. 4 (1896) 263. Low shrub or small tree, 3—12 m, 18—20 cm dbh. Twigs often sparsely appressedly hairy when young, soon glabrescent. Nodal appendages bluntly conical to triangular, up to 0.2 mm long but often smaller. Axillary buds 2—3, up to 2 mm long, sessile or the up- per very shortly stipitate, acutish, often distinctly keeled, more or less densely shortly hairy. Petiole 3-5 mm, + indistinctly transversely wrinkled, brownish, often sparsely thinly hairy when young, glabrescent. Leaf-blade 5—14 by 1.4—4(—S.5) cm, margin more or less distinctly undulate, often a little incurved, apex acutish, rarely shortly acuminate; above yellowish green to brown, midrib flat to slight- ly prominent in apical half, in basal half with a cen- tral groove; beneath light yellowish or brownish, papillose, midrib glaucous or sometimes sparsely ap- pressedly hairy, secondary nerves mostly hardly dis- tinct from finer veins, c. 8—14 pairs, not forming an intramarginal nerve; glands mostly numerous, 0.2—0.3 mm diam., but sometimes larger (up to 0.8 mm) and then of irregular form. /nflorescences mostly longer to much longer than the leaves, basally branched but sometimes seemingly unbranched if lower bracts resemble normal leaves; axes mostly dis- tinctly flattened basally, rust-brown, more or less densely patently shortly hairy; in basal part flowers with 3—5 together, solitary in apical part. Pedicel 2—3.5mm, + densely patently shortly hairy. Sepals: outer sepals 1.7—2.2 by 1.5—2 mm; inner sepals 2.2—2.7 by 2—2.9 mm. Petals pinkish or white, the upper ones often with a yellow spot, when dry yel- lowish, the longest one 6—9 mm long; carina rather densely (woolly) hairy outside; other petals hairy at very apex. Stamens 8, rarely in some flowers 9; an- thers 0.3—0.5 mm long. Ovary appressedly hairy; ovules 4. Fruit broadly ellipsoid to globular, 1.2—3.5 cm diam., mostly greyish brown; pericarp often wrinkled when dry, rather thick, soft. Seed(s) | or 2. Distr. Continental SE. Asia (Bangla Desh, Bur- ma, Thailand, Cambodia, Laos, S. Vietnam); in Malesia: S. Sumatra (Palembang), Malay Peninsula (incl. Langkawi Is.). Ecol. Confined to streambanks and swamps. Note. A common species with the largest distri- butional area within this subsection, and little varia- tion in its characters except in the size of the ripe fruit. 61. Xanthophyllum lateriflorum Mig. Ann. Mus, Bot. Lugd.-Bat. 1 (1864) 318; Mevpen, Leiden Bot. Ser. 7 (1982) 129. Shrub or small tree, 5-6 m. Axillary buds (2 or) 3(—5), distant, the upper one 3—5(—15) mm supra- 528 FLORA MALESIANA [ser. I, vol. 103 axillary, usually shortly stipitate (stalk up to 3 mm), + oblong, c. 1—2 mm long (excl. stalk), more or less acute. Petiole 3.5—4.5(—6) mm. Leaf-blade 4—8 by 1.5—3.5 cm, apex acuminate to cuspidate; above dark green; beneath glaucous-papillose, secondary nerves c. (5—)7 or 8 pairs, not forming an intra- marginal nerve; glands c. 6—8(—10), usually near midrib, 0.1—0.3 mm diam., basal ones somewhat lar- ger. Flowers unknown. /nfructescences shorter than the leaves, unbranched; axes minutely hairy. Fruit (immature) globular, rather sparsely minutely appressedly hairy; pedicel 3—3.5 mm, minutely ap- pressedly hairy. Seed (immature) 1; abortive ovules ar Distr. Malesia: S. Sumatra (Palembang, Lam- pong Distr.), 5 collections; insufficiently known spe- cies. 62. Xanthophyllum virens Roxs. Pl. Corom. 3 (1820) 81, t. 284, f. 1; SPRENGEL, Syst. Veg. 2 (1825) 219 (‘virescens’); WALL. Cat. (1831) 4197; Roxs. FI. Ind. ed. Carey 2 (1832) 221; DreTR. Syn. Pl. 2 (1840) 1277 (‘virescens’); WiGHT, Ill. Ind. Bot. 1 (1840) 49, 50, t. 23, f. 10 (sub X. flavescens, sphalm.); DRuRY, Handb. Indian FI. 1 (1864) 56; BEpp. Fl. Sylv. Anal. Gen. 3 (1869) xix, pl. III, f. 2, 1—3; Kurz. J. As. Soc. Beng. 42, ii (1873) 79, 80; Prelim. Rep. For. Pegu (1875) 26; For. Fl. Br. Burma 1 (1877) 81; GAGNEP. in Desv. J. Bot. 21 (1908) 251; Crarp, Fl. Siam. En. 1 (1931) 107; GaGnep. FI. Gén. I.-C. Suppl. 1 (1939) 219; PURKAYASTHA in Chowdhury & Ghosh, Indian Woods 1 (1958) 60; MEpDEN, Leiden Bot. Ser. 7 (1982) 130, f. 10A. — X. flavescens var. virens (Roxs.) A.W.BENNETT, FI. Br. India 1 (1874) 209; CraIs, Bull. Misc. Inf. Kew (1911) 14. — X. affine (non Mig.) Ripiey, J. Fed. Mal. St. Mus. 10 (1920) 82. Tree, up to 30 m, 1 m dbh. Nodal appendages c. 0.1 mm long. Axillary buds 3—7, 0.5—1.6 mm long, the upper one (2—)3—10(—20) mm supra-axillary. Petiole (S—)6—12 mm, often in apical part with small, usually not protruding glands. Leaf-blade (6.5—)10—23 by 2.5—7.5 cm, margin slightly un- dulate, often somewhat irregular, apex acutish; above dark or greyish green, midrib prominent to flat, in basal part with a central groove; beneath usually brownish to yellowish green, smooth to papillose, secondary nerves 7—10 pairs, forming an indistinct intramarginal nerve; glands either 0—3 (or 4) and often of an irregular form, or numerous and 0.1—0.2 mm diam., basal glands larger. /nflores- cences usually 2 per leaf axil, 5-20 mm supra- axillary, as long as or longer than the leaves, much- branched, the basal branches |—3 together; axes flat- tened basally, sparsely minutely hairy at the nodes with minute appendages; flowers 1—7 together; lower bracts opposite. Pedice/ 2.5—5.5 mm, minutely patently to appressedly hairy, sometimes subgla- brous. Sepals: outer sepals 1.4—2.2 by 1—1.9 mm; inner sepals (1.9—)2.3—3.3 by 1.6—3.5 mm. Petals white or pinkish, the upper ones with a yellow spot, when dry yellowish orange, the longest one 6.5—11 mm; carina sparsely to densely appressedly hairy outside, inside sparsely hairy in apical part; other petals glabrous or sparsely hairy at apex. Stamens: anthers 0.3—0.5 mm long. Ovary appressedly hairy; style rarely only basally hairy; ovules 4. Fruit globu- lar, c. 1.5 cm diam., smooth, dull, greyish, appres- sedly hairy apically; pericarp rather thick; pedicel 4—6 mm (see note). Distr. Continental SE. Asia (Bangla Desh, Bur- ma, Thailand); in Malesia: Malay Peninsula (Kelan- tan, Perak). Ecol. Outside Malesia in (usually submontane) monsoon forest. Notes. The Malayan collections are either sterile or only in fruit; as they have longer pedicels, they may not belong to this species. In some collections all flowers examined had 7 sta- mens only; instead of 2 carinal stamens only a single one is present in those flowers. II. Subgenus Coriaceum MEUDEN, Leiden Bot. Ser. 7 (1982) 133. Nodal glands distinct, c. 0.3 mm diam. Axillary buds indistinct when resting. Leaf-blade: ter- tiary nerves coarsely reticulate, sometimes partly scalariform. Jnflorescences inserted in older nodes, unbranched, few-flowered, axes up to 1.5 cm, with minute nodal glands. Sepals glabrous except for ciliate margin. Petals glabrous in apical half out- and inside, lateral petals and carina spoon-shaped, upper petals narrower. Stamens monadelphous. Ovary glabrous; style glabrous; stigma peltate; ovules 8—12. Fruit indehiscent, stipitate, + globular, apically pointed, 1.2 cm diam. Seed 1; testa 2-layered, less than 0.1 mm thick inner layer hard; albumen forming a rather distinct, thin layer which is very thin at lateral sides of cotyledons; embryo more or less globular, green, plumule undifferentiated, radicle exserted. 1988] POLYGALACEAE (van der Meijden) 529 a Le: WZ Zs ft, ft V4; Fig. 22. Xanthophyllum ramiflorum MeEuDEN. a~b. Habit, x 0.7; c. base of leaf with glands, x 2; d. part of twig with axillary bud; e. flower; /. flower, longitudinal section, gynoecium removed; g. lower petal; A. gynoecium; all x4; /. ovary, longitudinal section, * 8 (S 16051). 530 FLORA MALESIANA 63. Xanthophyllum ramiflorum MevupEN, Blumea 18 (1970) 392; Leiden Bot. Ser. 7 (1982) 133, f. 16. — X. spec.: ANDERSON, Gard. Bull. Sing. 20 (1963) 152; Wuit. Trop. Rain For. Far East (1975) 147. — Fig. 22. Tree, up to 30 m, 27 cm dbh. Axillary buds 2, very indistinct when resting, apparently sunken into the tissue of the twig, the upper one when bursting with 2 broadly ovate c. 1—1.5 mm long persistent scales. Petiole (S—)8—12 mm. Leaf-blade coriaceous, 7—20 by 3—8 cm, apex obtuse to subacute; above brown- ish, midrib slightly sunken to flat; beneath pale glaucous-papillose, secondary nerves c. 8 pairs, not forming an intramarginal nerve; glands numerous, c. 0.3 mm diam. /nflorescences solitary or up to 9 to- gether in the axils of the lower leaves and those of the already fallen ones, 1—10-flowered; axes up to 1.5 cm long, thin, glabrous; nodal glands very indistinct; bracts small, scale-like. Pedice/ 8-10 mm, glabrous. [ser. I, vol. 103 Sepals dark reddish when dry; outer sepals c. 4—4.2 by 2.6 mm; inner sepals 4.5—4.8 by 3—3.3 mm. Pet- als white, the upper ones with a purple mark, when dry yellowish, minutely patently hairy in basal half outside and inside, further glabrous; carina like the lateral petals but a little shorter; lateral petals 7.5—8.5 by 7 mm; upper petals up to 7 by 2 mm. Sta- mens 8, exceptionally 7, up to c. 5 mm long; fila- ments connate over c. 1.5—2 mm, densely shortly pa- tently hairy up to about halfway; anthers c. 0.7—0.8 mm long, minutely ciliate, very shortly hairy at base. Ovary light brownish, glabrous; style c. 5 mm, gla- brous; stigma peltate, oblique, rather large; ovules 8-12. Fruit stipitate, + globular, up to 1.2 cm diam., pustulate, dull, reddish brown, the style-scar more or less protruding and excentric. Distr. Malesia: Borneo (Sarawak, Brunei). Ecol. Confined to the lowland ‘padang’ peat- swamp forest on a very poor, sandy, wet soil. III. Subgenus Triadelphum MEIDEN, Leiden Bot. Ser. 7 (1982) 135. Nodal glands usually distinct, 0.3—0.7 mm diam. Axillary buds seemingly single, small, the scales usually not fully covering the bracts of young inflorescences, 0.4—1(—1.5) mm long. Leaf- blade: tertiary nerves coarsely reticulate. Inflorescences unbranched, axes slightly angular, dark, glabrous to sparsely hairy, with usually distinct nodal glands. Sepals glabrous outside, usually minutely hairy inside. Peta/s (sub)glabrous outside, lateral petals and the somewhat short carina spoon-shaped, upper petals narrower. Stamens triadelphous, connate parts c. 3—4 mm high. Ovary usually black, glabrous; style glabrous or hairy; stigma peltate; ovules 8—14. Fruit indehis- cent, usually stipitate, 1-2 cm diam. Seed 1; testa 2-layered, c. 0.2 mm thick, inner layer c. 0.1 mm thick, hard; albumen copious, (nearly) separated into 2 halves; embryo flat, nerved, green, plumule undifferentiated, radicle exserted. 64. Xanthophyllum ellipticum Kortu. ex Mig. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 276; A.W.BENNETT, Fl. Br. India 1 (1874) 211; Kina, Mat. Fl. Mal. Pen. (1890) 140; CHopatT in E. & P. Nat. Pfl. Fam. 3, 4 (1896) 344; RipLey, J. Str. Br. R. As. Soc. n. 33 (1900) 45; GAGNEP. in Desv. J. Bot. 21 (1908) 253; Merr. En. Born. (1921) 326; CHopDAT in Merr. PI. Elm. Born. (1929) 133, excl. var.; FISCHER, Kew Bull. (1933) 487; KeitH, N. Born. For. Rec. 2 (1938) 225; MASAMUNE, En. Phan. Born. (1942) 379; MEu- ER, Bot. News Bull. Sandakan 7 (1967) 88; Fox, Sabah For. Rec. 7 (1970) 65; Nc, Fed. Mus. J. n.s. 13 (1971) 137; Tree Fl. Mal. 1 (1972) 357, f. 3; Cor- NER, Gard. Bull. Sing. Suppl. 1 (1978) 146, 211; MEUDEN, Leiden Bot. Ser. 7 (1982) 135, f. 3A-p, 17. — Banisterodes ellipticum (KoRTH. ex Miq.) O. K. Rev. Gen. Pl. 1 (1891) 46, nom. illeg. — X. citrifolium CHODAT, Bull. Herb. Boiss. 4 (1896) 255; in’ B. ‘&/PsoNat. Pfl. “Ram: 3, 4° 896)" 345; MASAMUNE, En. Phan. Born. (1942) 379; ANDERSON, Gard. Bull. Sing. 20 (1963) 152. — X. kingii Cuopat, Bull. Herb. Boiss. 4 (1896) 255; RmDLey, FI. Mal. Pen. 1 (1922) 143; Watson, Mal. For. Rec. 5 (1928) 249; Crars, Fl. Siam. En. 1 (1931) 106; Burk. Dict. (1935) 2268; Wyatt-SmiTH, Mal. For. Rec. 17 (1952) 80, 361; ibid. 237 (1963) iii, 12, 57. — Fig. 23. Shrub or tree, up to 30 m, 40 cm dbh. Twigs gla- brous to minutely hairy. Petiole (4—)5—7 mm, gla- brous to minutely hairy. Leaf-blade 5—20 by 2-7 cm, margin often irregular, sometimes shallowly cre- nate because of glands; above usually brownish, midrib narrowly sunken; beneath more or less con- colorous, secondary nerves 5—9 pairs, forming a rather distinct intramarginal nerve; glands rather nu- merous, c. 0.4—0.8 mm diam., scattered but at least 8 present on the leaf margin itself, other glands 1988] POLYGALACEAE (van der Meijden) 531 3 \ ZW Saks S&S ROM SS \\ \, Wy x N “ Y \ » S\N Fig. 23. Xanthophyllum ellipticum Kortu. ex Mig. a. Habit, «0.5; b. flower and inflorescence axis; c. flower, longitudinal section, gynoecium removed; d. carina with two stamens enclosed; e. gynoecium; all x 3; J. ovary, longitudinal section, x6; g. part of twig with young inflorescence, * 4.5 (S 25564). 532 FLORA MALESIANA (4—)6—10. Inflorescences often several together on the secondarily thickened nodes, shorter than the leaves. Pedicel c. 3—4 mm, rather sparsely to rather densely minutely (woolly-)hairy. Sepals mostly nigrescent, glabrous to minutely hairy inside; outer sepals 3—3.8 by 1.8—2.2(—2.7) mm; inner sepals 3.8—4.8(—6) by 2.3—3.2(—4.5) mm. Petals white to light yellow, when dry light brown to dark orange, inside minutely hairy only above base; carina 6—7(—8) mm long; other petals 8—9 mm long, lateral petals 3—4 mm wide, upper petals 1.5—2 mm wide. Stamens: filaments glabrous; anthers 0.4—0.6 mm long, shortly hairy at base, ciliolate along slits, often cohering around the stigma. Ovary subsessile; style glabrous; ovules 8—14. Fruit sessile, globular, 1.5—2.2 cm diam., smooth, dark reddish, somewhat shiny; pericarp thin, brittle; pedicel 3—6(—8) mm long. Seed often (sub)apical. Distr. S. Thailand; in Malesia: Malay Peninsula (incl. Singapore, Penang I.), Sumatra (incl. Simalur, Nias; Riouw: Karimun Is.), Borneo. 65. Xanthophyllum celebicum MEDEN, Leiden Bot. Ser. 7 (1982) 137. Tree, 25 m. Older nodes usually with a cluster of adventitious buds. Petiole 5—7 mm, not transversally wrinkled, more or less smooth. Leaf-blade 8—15 by 3—6 cm; above green, midrib sunken; beneath con- colorous, secondary nerves 5 or 6 pairs, not forming an intramarginal nerve; glands present only in apical half of the leaf-blade, c. 0.2 mm diam., scattered but up to 6 present on the leaf margin itself, other glands c. 8—12. Inflorescences up to 4 cm long; nodal glands rather distinct. Pedicel 5-7 mm, rather sparsely minutely woolly hairy. Sepals sparsely minutely hairy inside; outer sepals 4 by 2.5 mm; inner sepals 4.9 by 2.4mm. Petals brownish orange when dry, the longest one 7.5 mm long; carina inside in middle part shortly appressedly sparsely hairy; other petals inside densely appressedly shortly hairy. Stamens triadel- phous; filaments densely rather shortly hairy; an- thers 0.6 mm long, sparsely hairy at base, ciliolate along slits. Ovary shortly stipitate, slightly pustulate; style rather densely shortly more or less patently (+ woolly) hairy; ovules 13. Fruit globular, c. 1.5 cm diam., smooth, dark reddish; pericarp thin; pedicel 7-—9.5 mm. Distr. Malesia: Central Celebes (Malili), one col- lection. 66. Xanthophyllum montanum MeEpDEN, Leiden Bot. Ser. 7 (1982) 137, f. 10B. Tree, up to 30 m, 60 cm dbh. Nodal glands elliptic, 0.3—0.6 mm long. Petiole 3.5—5 mm, indistinctly transversely wrinkled. Leaf-blade (4—)6—9 by 1—2.5(—3.5) cm, apex gradually acuminate; above brownish green, midrib sunken; beneath concolor- [ser. I, vol. 103 ous, secondary nerves c. 6—8 pairs, forming an intra- marginal nerve; glands (2—)4—8(—15), present in middle and apical part, absent from margin itself but (in Sumatran coll.) rather close to the margin, or (in Bornean coll.) rather closely together and not near the margin, 0.3—0.6 mm diam. /nflorescences up to 3 cm long; nodal glands elongate, usually very dis- tinct. Pedicel c. 5 mm, sparsely minutely woolly hairy. Sepals minutely hairy inside; outer sepals c. 3.1 by 2.3 mm; inner sepals 3.5—4.3 by 2.3—2.8 mm. Petals yellowish brown when dry, the longest one 6—6.5 mm long; carina inside minutely hairy above base, upper petals minutely hairy inside. Stamens: filaments minutely hairy in basal part; anthers 0.6 mm long, glabrous. Ovary 0.5—0.7 mm stipitate, gla- brous; style glabrous or only at base very sparsely minutely hairy; ovules 8—12. Fruit globular, up to 0.9 cm diam., yellowish to greenish brown, smooth, dull; pericarp thin; pedicel 4—5.5 mm. Distr. Malesia: Sumatra (near Lake Toba), Borneo (Sabah: Mt Kinabalu). Ecol. Submontane rain-forests, 900—1600 m. 67. Xanthophyllum contractum MEDEN, Leiden Bot. Ser. 7 (1982) 138. Tree, flowering on older twigs from adventitious axillary buds. Nodal glands indistinct, elongated. Petiole 9-10 mm. Leaf-blade 14—20 by 6.5—8 cm, apex obtuse to very shortly acuminate; above yellow- ish green, midrib protruding; beneath: secondary nerves 8 or 9 pairs, not forming an intramarginal nerve; glands c. 12—18, mostly at 2—5 mm from the margin, some scattered, 0.2—0.3 mm diam. /nflores- cences up to 4cm long; nodal glands present but very indistinct. Flowers unknown. Fruit (immature) c. 3 mm stipitate, ovoid, c. 2 by 1.2 cm, more or less fleshy, finely pustulate, light reddish, glabrous; peri- carp hard; pedicel 5-6 mm, glabrous. Seed 1 (sub)apical, developing from one of the 12 opposite ovules which are situated only in the apical half of the young fruit. Distr. Malesia: Borneo (Sarawak, Brunei). 68. Xanthophyllum hildebrandii MEIsDEN, Leiden Bot. Ser. 7 (1982) 139, f. 10B. Petiole c. 6 mm, not transversally wrinkled. Leaf- blade c. 15 by 5—7 cm, papery thin; above dull, mid- rib sunken; beneath reddish brown, secondary nerves c. 7 pairs, in apical half forming a rather indistinct intramarginal nerve; glands numerous, mostly situ- ated very close to midrib and a few scattered, 0.5—1.1 mm diam. /nflorescences | or 2 together; nodal glands distinct; axes up to 10 cm long, sparsely shortly woolly hairy. Flowers unknown. Fruit (very young) c. 2.5 mm stipitate, ovoid, apically with gland-like pustules, black, glabrous; pedicel 7—10 mm, dark, minutely woolly hairy. Seed (immature) 533 1988] POLYGALACEAE (van der Meijden) Note. Dedicated to the late Mr. F.H.HILDE- BRAND who cleverly recognized many inadequate specimens of the genus by means of macroscopical characters of the wood of the twigs. 1, (sub)apical, developing from one of c. 12 ovules situated in apical 2/3 part of the fruit. Distr. Malesia: Borneo (Sabah: Mt Kinabalu, Dallas), one collection only. IV. Subgenus Exsertum MEUDEN, Leiden Bot. Ser. 7 (1982) 139. Presence of nodal glands uncertain. Axillary buds 2(—4), usually seemingly single, black when dry. Leaf-blade: tertiary nerves coarsely reticulate. Jnflorescences unbranched, up to 6—8 cm long, axes slightly angular, black when dry, rather sparsely woolly hairy, sometimes with in- distinct nodal glands. Sepals nigrescent. Petals (sub)equal, spathulate-lanceolate, apex flat, (sub)glabrous outside, densely lanately ciliate. Stamens: filaments free, longer than petals, rather densely woolly hairy. Ovary black when dry, hairy inside, outside glabrous or hairy; stigma small, bilobed; ovules 8—16. Fruit indehiscent, globular, 2—6 cm diam., black when dry. Seeds c. 4—12, more or less bean-shaped, testa with rather thick and soft, fibrous outer layer and a hard inner layer; albumen copious, (nearly) separated into 2 halves; embryo flat, elliptic, nerved, plumule undifferentiated, radicle exserted. 69. Xanthophyllum suberosum C.T.Wuirte, J. Arn. Arb. 10 (1929) 229; MEuDEN, Leiden Bot. Ser. 7 (1982) 141, f. 18. — Fig. 24. Tree, up to 25 m, up to 70 cm dbh. Petiole 6—7 mm, black, sometimes glaucous. Leaf-blade 4.5—11 by 2—5.5 cm; above olive-green, midrib narrowly sunken, nervation mostly rather indistinct; beneath reddish olive-green, secondary nerves c. 7—9 pairs, forming an indistinct intramarginal nerve, glands 10—14, sometimes more, mostly + halfway between midrib and margin, or near the margin, 0.1—0.3 mm diam., basal ones c. 0.3—0.4 mm diam. Pedicel 8—11 mm, rather sparsely shortly lanate. Sepals sparsely minutely hairy outside, rather densely shortly lanate. Sepals sparsely minutely hairy outside, rather dense- ly shortly hairy inside; outer sepals 3.5—5.5 by 3.5—4 mm; inner sepals 5—7.5 by 4—5 mm. Petals 14—16 mm long, white, when dry dark reddish, rather sparsely hairy outside, subglabrous inside. Stamens: filaments 17~—22 mm, glabrous at very base, further densely woolly hairy; anthers 1—1.3 mm long, hairy from base to apex. Disk densely hairy, with a small number of patent hairs on upper and lower lobes. Ovary very densely (velvety) hairy; style rather sparsely woolly hairy; ovules 10-16. Fruit (im- mature) c. 2cm diam., shortly stipitate, finely pustu- late, dark, distinctly bluish-waxy, hairy; pericarp c. 1—4 mm thick; pedicel slender, c. 15-18 mm. Distr. Malesia: New Guinea (incl. Meos Num, Biak & Japen Is.). 70. Xanthophyllum amoenum Cuopar, Bull. Herb. Boiss. 4 (1896) 259; in E. & P. Nat. Pfl. Fam. 3, 4 (1896) 344; GAGNEP. in Desv. J. Bot. 21 (1908) 252; MASAMUNE, En. Phan. Born. (1942) 379; WyatTrT- SmitH, Mal. For. Rec. 17 (1952) 80, 363; ibid. 237 (1963) f. 8; ANDERSON, Gard. Bull. Sing. 20 (1963) 152; MEUER, Bot. News Bull. Sandakan 7 (1967) 87; Na, Tree Fl. Mal. 1 (1972) 356, f. 1; MEUDEN, Leiden Bot. Ser. 7 (1982) 141, f. 3A-n. — X. stipitatum var. nitidum Cuopat in Merr. Pl. Elm. Born. (1929) 137; MASAMUNE, En. Phan. Born. (1942) 381. — X. stipitatum var. pachyphyllum Cxopat in Merr. Pl. Elm. Born. (1929) 137; MASAMUNE, En. Phan. Born. (1942) 381. Tree, up to 35 m, up to 80 cm dbh. Axillary buds 2 or 3, the upper one up to 2 mm long. Petiole 4.5-10.5 mm. Leaf-blade 4—14 by (1—)2—7 cm; above deep brown to reddish brown, midrib narrow- ly sunken, nervation rather obscure to rather dis- tinct; beneath sometimes waxy, secondary nerves 5—7, not forming an intramarginal nerve; glands 6—10(—20), scattered, 0.1—0.4(—0.5) mm diam. Pedicel 8—15 mm, rather densely minutely lanate. Sepals: outer sepals 2.5—3 by 1.8—2.2 mm, minutely rather densely hairy at both sides; inner sepals 4—4.5 by 1.8—2.4 mm, hairy outside along midrib, inside densely hairy at base, further more or less glabrous. Petals (8—)9—11(—12) mm long, white (‘3 lower with yellow centre’; HAVILAND 2112), when dry dark red- dish, glabrous outside, inside lanate especially in ba- sal and apical part. Stamens: filaments 11—13(—16) mm; anthers 0.7—0.9 mm long, glabrous or with a few hairs at base. Disk glabrous. Ovary stipitate, (glabrous? or) densely lanate; style lanate; ovules 8~—16. Fruit globular or less often ovoid, up toc, Scm 534 FLORA MALESIANA [ser. I, vole | WY {| | Hil } Weg y, Wa ' <~_> Fig. 24. Xanthophyllum suberosum C.T.Wuirte. a. Habit, x 0.7; b. part of twig with axillary buds; c. flower; d. flower, longitudinal section, gynoecium removed; e. lower petal; f. gynoecium; all x2; g—h. ovary, longitudinal and cross section, X4 (VAN ROYEN 3201). 1988] POLYGALACEAE diam., sometimes bluish waxy, hairy or apparently glabrous; pericarp 1—10 cm thick. Seeds 6 or more. Distr. Malesia: Malay Peninsula, Central Suma- tra, Borneo. 71. Xanthophyllum stipitatum A.W.BENNETT, FI. Br. India 1 (1874) 210; Kinc, Mat. Fl. Mal. Pen. (1890) 140; CHopaT in E. & P. Nat. Pfl. Fam. 3, 4 (1896) 345; Riptey, Fl. Mal. Pen. 1 (1922) 145; HEND. Gard. Bull. S. S. 4 (1928) 222; Watson, Mal. For. Rec. 5 (1928) 249; CHopatT in Merr. Pl. Elm. Born. (1929) 137, incl. var. borneense CHODAT; Burk. Dict. (1935) 2268; Wyatt-SmitH, Mal. For. Rec. 17 (1952) 81, 361; BALAN MENON, ibid. 19 (1956) 34; MeEvER, Bot. News Bull. Sandakan 7 (1967) 87; Na, Tree Fl. Mal. 1 (1972) 363, f. 5; Cor- NER, Gard. Bull. Sing. Suppl. 1 (1978) 147; MEUDEN, Leiden Bot. Ser. 7 (1982) 142. — Banisterodes stipitatum (A.W.BENNETT) O. K. Rev. Gen. Pl. 1 (1891) 46. — X. amoenum (non CuopatT) KEITH, N. Born. For. Rec. 2 (1938) 225. Tree, up to 50 m, 1.20 m dbh. Axillary buds 2(—4), c. 2 mm long. Petiole 3-8 mm. Leaf-blade (2.5—)4—13 by (1—)2—7 cm; above mostly greyish brown to dark brown, sometimes reddish brown, midrib slightly sunken, nerves mostly obscure; be- neath concolorous, sometimes more or less waxy, secondary nerves 5 or 6 pairs, not forming an intra- marginal nerve, finely prominent to rather obscure; glands few, 0—2(—4) per leaf, near the base and in the middle, 0.1—0.2 mm diam. Pedicel c. 8—12 mm, dark, minutely lanate. Sepals densely minutely hairy inside at base, further almost glabrous; outer sepals 1.8—3 by 1.8—2 mm; inner sepals 3—3.5 by 2—2.5 mm. Petals 7—7.5(—8) mm long, dark reddish when (van der Meijden) 555 dry, glabrous outside, hairy inside at base and sometimes also in apical part. Stamens: filaments 11—13 mm, shortly rather densely lanate, glabrous in apical part, free or (in var. glabrum) forming a ‘tube’ by their intertwined hairs; anthers c. 0.8 mm long, glabrous. Disk glabrous. Ovary densely lanate or rarely (sub)glabrous; style glabrous in upper half; ovules 8—12. Fruit c. 2—6 cm diam., often apparent- ly sterile, shortly to long-stipitate, black, often waxy, glabrous or hairy; pericarp 0.5—1.5 cm thick; pedicel up to c. 1.5 mm. Seeds c. 4-12. Distr. Malesia: Malay Peninsula, Sumatra, Borneo. KEY TO THE VARIETIES 1. Filaments forming a ‘tube’ by their intertwined hairs. Ovary (sub)glabrous .... b. var. glabrum 1. Filaments free, hairy. Ovary densely lanate a. var. stipitatum a. var. stipitatum — X. stipitatum A.W.BENNETT, incl. var. borneense CHODAT. Stamens: filaments free, hairy. Ovary densely lan- ate; style densely hairy basally. Distr. Malesia: Malay Peninsula (incl. Singa- pore), Central Sumatra, Borneo. b. var. glabrum MEUDEN, Leiden Bot. Ser. 7 (1982) 144. Stamens: filaments forming a ‘tube’ by their inter- twined hairs. Ovary glabrous or with a few appressed hairs; style thinly hairy basally. Distr. Malesia: E. Borneo (Kalimantan: Long Petah). V. Subgenus Brunophyllum MEUDEN, Bot. J. Linn. Soc. 67 (1973) 117; Leiden Bot. Ser. 7 (1982) 144. Nodal glands usually distinct, c. 0.3—0.5 mm diam. Leaf-blade: tertiary nerves coarsely reticu- late. Inflorescences unbranched. Sepals glabrous to minutely hairy outside, usually woolly-ciliate, usually minutely hairy inside. Pefa/s unequal with the carina boat-shaped, or equal, (sub)glabrous outside. Stamens 8 (—10), filaments free or up to 6 mm connate, densely woolly hairy. Ovary gla- brous to hairy; style glabrous or hairy at base; stigma peltate; ovules 8—18(—23). Fruit indehis- cent, pear-shaped to ovoid, large. Seeds 8—more, large; testa 2-layered, outer layer usually thick (thin in 76. X. chartaceum), soft, inner layer hard; embryo broadly triangular-ovoid and with lit- tle albumen, to flattened-ovoid and covered by much albumen; cotyledons cordate, usually seem- ingly peltate (except in 72. X. brevipes), plumule usually differentiated into a number of small scales, radicle fully enveloped by the cotyledons or just exserted at base. Nn Ww ON FLORA MALESIANA [ser. I, vol. 103 Fig. 25. Inflorescence of Xanthophyllum brevipes MEISDEN, collected in Arboretum, Semengoh For. Res., Sarawak (S 26838) (Photogr. P.Sre, 1971). 72. Xanthophyllum brevipes MEUDEN, Bot. J. Linn. Soc. 67 (1973) 117; Leiden Bot. Ser. 7 (1982) 144. — Fig. 25. Tree, up to 35 m, 40 cm dbh. Twigs (sometimes?) all curved downwards, pendent. Nodal glands often indistinct. Petiole 1.5—3 mm. Leaf-blade 2.6—7.5 by 0.7—2.6 cm, apex acuminate to cuspidate; above brown to olive-green, midrib sunken; beneath olive- greenish brown, papillose, secondary nerves c. 10 pairs, little more distinct than finer nerves, forming an indistinct intramarginal nerve; glands more than 10, in a row between margin and midrib, c. 0.1 mm diam. Inflorescences 2—5-flowered; axes hardly thicker than pedicel, glabrous. Pedicel c. 7 mm, gla- brous. Sepals minutely ciliate, further glabrous; outer sepals c. 3.5 by 3.5 mm; inner sepals c. 4 by 4.5 mm. Petals unequal, white, when dry orange-brown, faintly ciliate, the longest one 15—16 mm; carina un- guiculate, boat-shaped, ciliate only at base; lateral petals narrowly boat-shaped; upper petals more or less straight, with cup-shaped apex. Stamens: fila- ments c. 0.5 mm connate, rather densely shortly hairy in basal part; anthers c. 1 mm long, glabrous. Ovary shortly stipitate, orange-brown, glabrous; style glabrous; ovules 18. Fruit pear-shaped (to broadly ovoid?), up to 4cm diam., strongly wrinkled when dry, shiny, brown; pericarp probably very fleshy; pedicel 12—15 mm, blackish, shiny. Seeds more than 10, flattened-ovoid, c. 1 cm long; albumen nearly absent from lateral sides of cotyledons, at the broad side of these forming a layer equal in thickness to each cotyledon; embryo elliptic in side view, trans- versally flattened, cordate at base; cotyledons thick- ened; plumule not differentiated; radicle slightly ex- serted. Distr. Malesia: Borneo (Sarawak, Brunei). Note. ASHTON mentioned in a fieldnote that the habit of the tree is very reminiscent of old specimens of Salix babylonica because of its pendant twigs. 73. Xanthophyllum obscurum A.W.BENNETT, FI. Br. India 1 (1874) 211; Kinc, Mat. Fl. Mal. Pen. (1890) 141; RipLey, J. Str. Br. R. As. Soc. n. 33 (1900) 45; Fl. Mal. Pen. 1 (1922) 144; HEyNE, Nutt. Pl. (1927) 902; HEND. Gard. Bull. S. S. 4 (1928) 222; Watson, Mal. For. Rec. 5 (1928) 249; WyattT- SmiTH, Mal. For. Rec. 17 (1952) 81, 363; BALAN MENON, ibid. 19 (1956) 34; WyatTT-SMITH, ibid. 237 (1963) f. 8; NG, Tree Fl. Mal. 1 (1972) 361, f. 3; Mal. For. 38 (1975) 89, f. 8.1 F—J, 8.3, 8.4; CorNER, Gard. Bull. Sing. Suppl. 1 (1978) 147, 211; MEDDEN, Leiden Bot. Ser. 7 (1982) 145, f. 3B-b, 10B. — X. in- signe A.W.BENNETT, Fl. Br. India 1 (1874) 211; KinGc, Mat. Fl. Mal. Pen. (1890) 144. — X. scor- techinii Kinc, J. As. Soc. Beng. 59, ii (1890) 140; Ann. R. Bot. Gard. Calc. 5 (1896) 138, pl. 163; GAGNEP. in Desv. J. Bot. 21 (1908) 253; Rm ey, FI. Mal. Pen. 1 (1922) 143; HENbD. Gard. Bull. S. S. 4 1988] (1928) 222; Watson, Mal. For. Rec. 5 (1928) 249; Burk. Dict. (1935) 2268; Wyatr-SmitTH, Mal. For. Rec. 17 (1952) 81, 361; BALAN MENON, ibid. 19 (1956) 34; Wyatt-SmitH, ibid. 237 (1963) f. 5; No, Tree Fl. Mal. 1 (1972) 363, f. 3. — Banisterodes in- signe (A.W.BENNETT) O. K. Rev. Gen. | (1891) 46, nom. illeg. — Banisterodes obscurum (A.W.BEN- NETT) O. K. /.c., nom. illeg. Tree, up to 47 m, 70 cm dbh. Twigs often strongly thickened on the nodes and with adventitious buds. Nodal glands usually distinct, sometimes elongate, c. 0.5 mm diam. Petiole S—11(—15) mm. Leaf-blade (4—)7.5—17 by (1.5—)3.5—9 cm, apex rounded to ob- tuse Or sometimes very shortly acuminate; above dark or greyish redbrown, rarely green, midrib flat to protruding; beneath concolorous or darker than above, midrib prominent or flat, secondary nerves c. (3—)6—9 pairs, sometimes in upper part forming an intramarginal nerve; glands 2—16, usually situated near or on the margin of the leaf, (0.2—)0.5—0.7 (—1.2) mm diam. /nflorescences sometimes also on the older nodes, shorter than the leaves; axes angu- lar, black, glabrous to sparsely shortly lanate. Pedicel 3—11 mm, glabrous to sparsely hairy. Sepals black; outer sepals 2.8—5.5 by 2.8—6 mm; inner sep- als 4—7.5 by 3.5—7 mm. Peta/s unequal, white or purple, the upper ones with a yellow or green spot, when dry black, inside glabrous or rather densely woolly hairy above insertion of filaments and at apex, the longest one 14-19 mm; carina boat- shaped, 9.5—16 mm long; lateral petals more or less spathulate, distinctly longer than upper petals and carina; upper petals more or less linear, flat to slight- ly channelled, curved upwards. Stamens 7.5—12 mm long; filaments connate for (0.1—)1—3 mm, glabrous in basal part, free parts of filaments lanate in basal part, hairs often intertwined, thus forming a fila- mental ‘tube’, glabrous upwards; anthers 0.7—1.7 mm long, glabrous to minutely hairy, free or at- tached to each other around the stigma. Ovary black, glabrous; style black, glabrous; ovules 8—18. Fruit globular, very large, the largest up to 14 cm diam., dull pinkish brown, with numerous small dark spots; pericarp 0.5—2 cm thick. Seeds 8—16, sticking to- gether in drying like a ball, each c. 2—6 cm wide; testa 2-layered, outer layer thick, soft, fibrous, inner layer c. 0.1 mm thick; albumen nearly absent, usually vis- ible only at base; embryo thick, triangular in side view, plumule and radicle situated in the very centre of the embryo, the plumule differentiated into a number of decussate scales; cotyledons seemingly peltate, peripherically with numerous vessel-like ele- ments forming a regularly reticulate pattern, Distr. Southernmost Thailand; in Malesia: Ma- lay Peninsula (incl. Singapore & Penang I.), Su- matra, Borneo. POLYGALACEAE (van der Meijden) 537 74. Xanthophyllum papuanum WHITM. ex MEIDEN, Bot. J. Linn. Soc. 67 (1973) 119; Wuitm. Guide For. Br. Sol. Is. (1966) 92, 151; PatmMans, Land Research Ser. 29 (1971) 107; VERSTEEGH, Meded. Landb. hogesch. Wageningen 71-19 (1971) 63; EDDOwEs, Commerc. Timbers P.N.G. (1977) 46 (‘Boxwood’); MEUDEN, Leiden Bot. Ser. 7 (1982) 147, f. 3B-c, 19. — X. affine (non Mia.) K.ScH. & HOLtR. FI. Kaiser Wilhelm Land (1889) 68; K.Scu. & Laut. Fl. Schutz- geb. Stidsee (1901) 388. — Fig. 26. Tree, up to 43 m, up to 1.1 m dhb. Petiole 5—9 mm. Leaf-blade 5—14 by 2.8—9.3 cm; above greyish mid-green, rarely yellowish brown, midrib mostly sunken, sometimes flat, rarely prominent; beneath concolorous, secondary nerves 5 or 6 pairs; glands 6—10(—14), in middle part close to the midrib, in up- per part often along margin and midrib, 0.4—0.5 mm diam. /nflorescences + as long as to longer than the leaves; axes angular, brown, rather densely shortly lanate; in basal part flowers with 3(—7) together. Pedicel 3—4 mm, ribbed, densely shortly lanate. Sep- als brownish, shortly lanate outside; outer sepals 2.5—3.5 by c. 2mm; inner sepals 3—4.5 by c. 2.5mm. Petals (sub)equal, obovate-lanceolate, 7—8.5 mm long, with slightly cupped apex, white, when dry orange-brown, outside with a few woolly hairs along midrib, woolly-ciliate especially at apex, inside wool- ly hairy. Stamens 8 or 9, the middle carinal stamen rather often developed; filaments free or 0.1 mm connate, densely lanate to apex; anthers 0.7—0.8 mm long, sparsely woolly hairy. Ovary 0.5—0.7 mm stipi- tate, inside glabrous to sparsely hairy, outside dense- ly shortly lanate on median rib, further glabrous; style basally shortly lanate, upwards nearly glabrous; ovules 14—16. Fruit pear-shaped, c. 5-15 cm long, 3—12 cm diam., reddish brown to greyish greenish brown, often with darker spots; pericarp c. 0.5 cm thick, hard; pedicel up to 5 mm. Seeds mostly more than 8, rarely less, sticking to the pericarp in drying, broadly triangular-ovoid, up to 5 cm wide when dry; testa 2-layered, outer layer thick, soft, not fibrous, inner layer less than 0.1 mm thick, hard; albumen nearly absent, usually visible only at the base; em- bryo thick, triangular in side view, plumule and rad- icle situated below the centre of the embryo, plumule differentiated into a number of decussate scales; co- tyledons seemingly peltate, peripherically without vessel-like elements; top of radicle hardly exserted at base. Distr. Solomon Islands; in Malesia: New Guinea (incl. Fergusson I. in E, and Salawati, Numfoor, Japen & Mios Waar Is. in W), Moluccas (Ceram), Central Celebes (Palopo). Note. In a number of collections all flowers ex- amined have 9 stamens: 3 instead of 2 stamens are placed before the adaxial petal. 538 FLORA MALESIANA [ser. I, vol. 10° Ss SS SSSA SEN, Fig. 26. Xanthophyllum papuanum Wuitn. ex MEISDEN. a. Habit, x 0.7; b. flower and part of inflorescence axis; c. flower, longitudinal section, gynoecium removed; d. lower petal with three stamens; e. gynoecium; all x4; f. ovary, longitudinal section, x 8 (HOOGLAND 5072). 1988] POLYGALACEAE (van der Meijden) 539 75. Xanthophyllum ecarinatum CuHopatT, Bull. Herb. Boiss. 4 (1896) 254; in E. & P. Nat. Pfl. Fam. 3, 4 (1896) 344; MerrR. En. Born. (1921) 325; Masa- MUNE, En. Phan. Born. (1942) 379; MEER, Bot. News Bull. Sandakan 7 (1967) 87; MEIDEN, Leiden Bot. Ser. 7 (1982) 148, f. 3B-d. — X. kalimantanum MEUDEN, Bot. J. Linn. Soc. 67 (1973) 118. Tree, up to 25 m, 16 cm dbh. Petiole (1.5—)4—6.5 mm. Leaf-blade (ovate-)oblong, (3—)7—17 by (1—)2.8—7 cm, apex acuminate to cuspidate; above dark reddish brown to dark olive-green, midrib slightly prominent to flat, or sunken in basal part; beneath concolorous, secondary nerves S—7 pairs, forming a rather indistinct intramarginal nerve; glands 0-8, situated in middle and apical part, (0.1—)0.2-0.4 mm diam. IJnflorescences (much) shorter than the leaves; axes lanate; flowers solitary or in basal part with up to 3 together. Pedicel (1.5—)3—4 mm, dark, sparsely lanate. Sepals black when dry; outer sepals (3—)4—5.5 by (1.5—)2.5—3.5 mm; inner sepals (3.5—)6—7 by (1.5—)2.5—4 mm. Petals subequal, 9.5—12 mm long, white, the upper ones with a yellow spot, when dry nearly black, cil- jate to apex, inside hairy above insertion of fila- ments. Stamens 8(—10); filaments connate over c. 5—6 mm, glabrous at base, upwards densely (woolly) hairy; anthers 0.5—0.65 mm long, glabrous to sparse- ly woolly hairy at base. Ovary stipitate for 2.5—3 cm, dark, glabrous or with a few rather long hairs; style glabrous to rather sparsely lanate; ovules 12—18 (—23). Fruit ellipsoid, up to 11 by 6 cm, attenuate at base and apex, often more or less smooth, orange to dark brown; pericarp rather soft, c. 0.5 mm thick. Seeds 8 or more, like those of 74. X. papuanum, but albumen forming a thin layer along outer side of the cotyledons. Distr. Malesia: Borneo (Sarawak, Sabah, Kali- mantan). 76. Xanthophyllum chartaceum MevpEN, Bot. J. Linn. Soc. 67 (1973) 118; Leiden Bot. Ser. 7 (1982) 149. — X. spec. E, NG, Tree Fl. Mal. 1 (1972) 366. Tree, up to 30 m, | m dbh. Petiole 3—4 mm. Leaf- blade 4.5—11.5 by 2—4.2 cm, papery thin, base near- ly cordate to cuneate, apex acuminate to cuspidate; above very dark green, midrib flat to slightly promi- nent; beneath + glabrous, concolorous, secondary nerves c. 5—7 pairs; glands very few, up to 3, c. 0.1 mm diam., very indistinct. Flowers unknown. /n- fructescences short; axes black, glabrous. Fruit glob- ular to more or less pear-shaped, up to 8.5 cm long, strongly wrinkled, when dry black, probably gla- brous; pericarp rather thick, hard. Pedicel c. 5 mm, black, glabrous. Seeds 8 or more, like those of 74. X. papuanum, but both layers of the testa much thin- ner, and albumen forming a thin layer along the outer side of the cotyledons. Distr. Malesia: Malay Peninsula, Sumatra (Indragiri). Note. Because of its thin nigrescent leaves with few laminar glands, its habit is atypical in Xantho- phyllum. As its flowers may be atypical too (i.e. in being possibly ‘regular’), flowering material may have been collected already but identified wrongly. Central Excluded Xanthophyllum glaucescens Mig. Fl. Ind. Bat., Suppl. (1861) 394; Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 274, has been identified by HILDEBRAND and KOSTERMANS as Litsea insignis (BLUME) BOERL. (Lau- raceae). Xanthophyllum hebecarpum Cuopart, Bull. Herb. Boiss. 4 (1896) 263; RrpLey, J. Str. Br. R. As. Soc. n. 73 (1916) 139; Fl. Mal. Pen. 1 (1922) 149, was identified by SyMINGTON, Kew Bull. (1937) 318 as Ryparosa kunstleri KinG (Flacourtiaceae); SLEUMER, Fl. Males. I, 5 (1954) 48. Xanthophyllum subglobosum Emer, Leafl. Philip. Bot. 5 (1913) 1676, incl. var. longifolium ELMER, was identified by MerriLL, En. Philip. 2 (1923) 485 as Siphonodon celastrineus GRIFFITH (Celastraceae); DinG Hou, FI. Males. I, 6 (1964) 395. Excluded Semeiocardium Zou. Nat. Tijd. Ned. Ind. 17 (1858) 245, the type of a monospecific genus, was by ZoL- LINGER assigned to Balsaminaceae, but later referred to Polygalaceae by HASSKARL (in Miq. Ann. Mus. Bot. Lugd.-Bat. 1, 1863, 142). Still later Cuopat reduced it to Polygala triphylla (Monogr. 1, 1891, 41). BACKER (Gard. Bull. S. S. 9, 1938, 70) showed that ZOLLINGER was correct. » el all 1 aw vies, ast ah Jel. wi) se CF ae LDS hi}. thew. i) ne grt rte dia trc. ay _ , Be wn -- a. 7 td ie.) a ban ee ei ts ue Purell j Hagina’* Mwy , ; hal A1on (3 reN h re sadn haa 7 ‘lei tb oe I pa] Wg sail ye i } > : ; 4 A oye eet aS aa ia 's erreta i oe pene \ v7) . 1% > 1 | eet ae * iad aA, ( f) > ih fj U Bf) = } a iit w- he » bh Ff ol i ihe Fpl uae aie 4 ~ FY s 1 4 ave = te » Nyiket ® wo > Lied al yy Didagil ith poe) Servo yi ootta im —yiun swiins yu “ a 4a is hee a « ie Ste P ut! Or) Ae ibaa Louie x tier ee uae | p am ae a a) ita =. ce Oe ee ee CRUCIFERAE (B. Jonsell, Stockholm) Herbs, sometimes subshrubs. Leaves spirally arranged, basal ones often in a ro- sette, exstipulate, petiolate to sessile and amplexicaul, entire to variously divid- ed. Inflorescences terminal or sometimes axillary racemes, in flower mostly con- densed and often corymbose, in fruit elongate, usually ebracteate. Flowers bi- sexual, actinomorphic or slightly zygomorphic, hypogynous, cyclic, tetra- merous, heterochlamydous. Sepals 4, free, usually equal, spathulate to clawed, imbricate or contorted. Stamens 6, tetradynamous (rarely 4 or 2), episepalous usually free; anthers usually 2-thecous opening lengthwise. Nectarial glands variously arranged at the filament bases. Ovary superior, sessile or stipitate, of seemingly two united carpels, secondarily divided into two locules by a thin membranous septum (sometimes transversely locular by intrusions from the fruit wall); placentation parietal, ovules usually many, anatropous or campylo- tropous; stigma bifid or connate. Fruit a bivalved dehiscent siliqua or silicula (see key), sometimes a nutlet, lomentaceous or otherwise constricted. Seeds vir- tually devoid of endosperm, with cotyledons incumbent, accumbent or various- ly folded. Distribution. A cosmopolitan family with about 380 genera and more than 3000 species, especially diversified in the Mediterranean and the Irano-Turanian regions as well as in parts of Southern Africa, North America and montane South America. The family is comparatively sparse in the tropics, mainly confined to montane and arid areas. The family includes a number of important crops and spices, notably in the genera Brassica, Sinapis, Raphanus, Crambe, Lepidium, Rorippa, some now cultivated in montane parts of the tropics. Several cosmopolitan weeds belong to the family (within Lepidium, Coronopus, Capsella, Thlaspi, Arabis, Sisymbrium, Rorippa, Camelina, and others) and may appear as casual aliens virtually anywhere. In Malesia there are 6 genera with a total of 24 species. The genus Papuzilla (3 spp.), regarded as a New Guinean endemic, is here included within Lepidium. In each of the genera Cardamine and Rorippa 3 spp. are endemic to New Guinea and some adjacent areas, and Rorippa backeri is an endemic of Java. The affinities of the endemic Rorippa species is with Australia and New Zealand, while the Car- damine species belong to an alliance of tropical montane species around the widespread C. africana. The remaining 14 species may all have been introduced by man and partly naturalized. Ecology. The native species are montane (some secondarily at lower levels). Natural occur- rences are open forests, rocky outcrops, riverbanks, mostly places without a closed vegetation cover. Many are favoured by clearings, fires, road building, efc. Dispersal. A\l Malesian species (except Raphanus sativus) disperse by small seeds without special devices. Some seeds are mucilaginous (get sticky with water), which may facilitate trans- port with birds and other agents. Morphology. In spite of its size, the family is remarkably uniform. Lignification is rarely substantial except in basal parts, the hypocotylar region and roots. The herbaceous life-form seems to be primary, at least so far as many woody groups seem to have evolved from herbaceous forms. Annuals are common in many genera. In floral parts little specialization is to be found, but not rarely reductions as to stamens, petals and nectaries. The position of the nectaries varies in relation to the filament bases (and is of taxo- nomic importance) but no particular elaboration is found. (541) 542 FLORA MALESIANA [ser. I, vol. 10° Variation and specialization are important and often drastic as to fruits and diaspores, which means that without developed fruits many species are not determinable. The seeds are variously ornamented, often winged. The testa is rather uniformly built from one- cell-thick layers: epidermis (which may contain mucilage), subepidermis (which may be absent), palissade layer (usually with pigmented wall thickenings) and a parenchymatous pigmented layer. The ripe seed is almost filled up by the embryo; endosperm is reduced to a one-cell-thick ‘aleurone’ layer adjacent to the testa. The folding of the cotyledons and their position in relation to the radicle is of taxonomic importance. Vegetative anatomy. Wood anatomy is quite variable within the family and largely asso- ciated with stem width. The xylem in thin-stemmed species forms a closed cylinder and the cam- bium forms a continuous ring. In thicker stems sclerenchymatous tissue separates the xylem bundles and the cambium cylinder is mostly discontinuous. In woody species medullary rays may also separate the xylem bundles. The variation largely reflects degrees of adaptations to arid con- ditions and occurs within genera and species as well as intra-individually according to age. Even in strongly lignified species the wood elements are of little advanced types, making it probable that woodiness has evolved comparatively recently in the family. Secretory cells containing myrosinase (myrosin cells) are widely distributed in the family, but in various organs in various genera. Their distribution has been regarded to be of some taxonomic value. Hairs are always unicellular, but shape varies from unbranched to forked, stellate, T-shaped, etc. and is of considerable taxonomic interest at various levels. The stomata are mostly of a characteristic type with one small and two larger subsidiary cells: so-called anisocytic or cruciferous stomata. This type is, however, by no means restricted to the Cruciferae. References: CARLQUIST, Aliso 7 (1971) 365—384; INAMDAR & Rao, Feddes Rep. 94 (1983) 183—190; IvERSEN, BAGGERUD & BeEIsvAaG, Z. Pflanzenphysiol. 94 (1979) 143-154; Kowa & CUTLER, Kew Bull. 30 (1975) 503—507; METCALFE & CHALK, Anatomy of the Dicotyledons 1 (1950) 79—87; PANT & Kipwal, Ann. Bot. (N.S.) 31 (1967) 513—521; Rao & INAMDaR, Bot. Mag. Tokyo 96 (1983) 15—28. Palynology. As with macromorphology, the pollen morphological variation within Cruciferae is small in proportion to the number of genera and species. Shape of the grains ranges from suboblate to subprolate, but prolate and perprolate shape is easily attained by invagination of the long colpi under dry conditions (e.g. during transport through the air). The grains are small to medium-sized (10—45 ym), rarely somewhat larger. The apertural system is nearly always 3-colpate. Seldom also 2-, 4-, 5-, 6-, or 7-colpate grains occur. ERDTMAN (1952) records the rare occurrence of inaperturate pollen, which probably repre- sents a kind of tritenuate apertural system. Exine stratification is mostly distinct, sexine and nexine being clearly distinguishable. A reticulate tectum with up to 1 um broad muri is usually present. Columellae may be distinct or not, which mainly depends on their size. On the strength of extensive material (200 out of about 380 genera) CHIGURIAEVA (1973) estab- lished 5 main pollen types within Cruciferae. Most characters used in this subdivision are quanti- tatively determined and the various pollen types occur together in most tribes of the family. Be- sides, compared with the pollen morphological range in the whole family, relatively much varia- tion appeared to be present within single species. Thus, pollen morphology seems of little signifi- cance for taxonomy in Cruciferae. As to be expected, cruciferous pollen keys out very difficultly (ERDTMAN c.s., 1963). References: CHIGURIAEVA, Pollen morphology of Cruciferae, in Kuprijanova, Pollen and spores morphology of the recent plants. Proc. III Int. Palyn. Conf. (1973) 93—98; ERDTMAN, Pollen morphology and plant taxonomy, Angiosperms (1952) 133—134; ERDTMAN, PRAGLOWSKI & Nixsson, An introduction to a Scandinavian Pollen Flora II (1963). — R.W.J.M. VAN DER HAM. 1988] CRUCIFERAE (Jonsell) 543 Phytochemistry. The family is rather uniform and highly characteristic also from the chemical point of view. Particularly the seeds but also other organs contain glucosides with sulphur and nitrogen in their molecules, so-called mustard oil glucosides, or glucosinolates, com- pounds unique to the family. The enzyme myrosinase, localized in particular cells, will split the glucosinolates when cell walls are crushed in the presence of water into three compounds, among those the pungent mustard oils. They are either isothiocyanates of usually pleasant flavour or thiocyanates with a strong, often garlic-like odour (e.g. in Lepidium and Thlaspi). Very important in the seeds are also lipid acids, particularly unsaturated ones with 18, 20 or 22 carbon atoms. Particular for the family and very widespread is the erucic acid, which because of unliked properties should be kept at a minimum in strains of e.g. Brassica cultivated as oil seed crops. Others of those fatty acids (oleic, linolenic and linoleic acids) are of utmost economic im- portance. Among alkaloid-like compounds sinapin, a protoalkaloid of bitter taste, is very com- mon in the family and concentrated in the seeds. Proteins are of importance in the seeds, while starch is lacking. References: HEGNAUER, Chemotaxonomie der Pflanzen 3 (1964) 586—607; VAUGHAN c.s. (ed.), The biology and chemistry of the Cruciferae (1976) 191—330. Chromosomes. By far the most widespread basic number is x=8, but dysploidy occurs within several genera (e.g. Brassica) and may account for x=7, which prevails in a few genera (e.g. Thlaspi, Sisymbrium); x=5 is known from Arabidopsis, the southern hemispheric tribes Stenopetaleae and Heliophileae, and on polyploid level in Crambe. Polyploidy, often combined with aneuploidy, is extensive in most of the large genera, e.g. Cardamine. The chromosomes are small and do not readily lend themselves to structural studies. References: FEDorov (ed.), Chromosome numbers of flowering plants (1969); MANTON, Ann. Bot. n.s. 1 (1932) 438—462. Taxonomy. The uniformity of this large family makes subdivision difficult. A number of tribal classifications have been proposed, most of them ending up with 15—20 tribes, 9 of those distinct and largely restricted to limited parts of the southern hemisphere. They include few genera and mostly also few species. Among widespread tribes, rich in genera and species, Brassiceae (with Brassica and Raphanus in the Malesian flora) is the only one really distinct, and not been dis- agreed upon as to its circumscription. Most of the other tribes have been rather schematically de- limited and may contain a nucleus of closely related genera to which others seem to have been more arbitrarily added. According to SCHULz’s system the Malesian genera should be included within Lepidieae (Lepidium and Capsella) and Arabideae (Cardamine and Rorippa). References: HEDGE in Vaughan c.s. (ed.), The biology and chemistry of the Cruciferae (1976) 1—45; Scuutz, Cruciferae, in Engler & Prantl (ed.), Die nattrlichen Pflanzenfamilien ed. 2, 17b (1936) 227-658. Cultivated species. The following species have been reported as cultivated only, and are not treated separately: Aromoracia rusticana G.M. & SCHERB. See BACKER & BAKH./. Fl. Java 1 (1963) 188. — Brassica chinensis L. See HEYNE, Nutt. Pl. Ned. Indié (1927) 677; OcHsE & BAKH. Indische Groenten (1931) 162; Stone, Fed. Mus. J. 26 (1981) 81. — B. napus L. See HEYNE, Nutt. Pl. Ned. Indié (1927) 677; Ocuse & Baku. Indische Groenten (1931) 168. — B. oleracea L. See Koorp. Meded. Lands Plantentuin 19 (1898) 341; Exk. Fl. Java 2 (1912) 284; Hocur. Candollea 2 (1925) 367; Heyne, Nutt. Pl. Ned. Indié (1927) 677; Ocuse & BAKH. Indische Groenten (1931) 169; DocTeRs VAN LEEUWEN, Verh. Kon. Akad. Wet. Amsterdam sect. II, 31 (1933) 166; BACKER & Baku. Fl. Java 1 (1963) 188. — B. rapa L. (B. campestris L.). See Koorp. Exk. Fl. Java 2 (1912) 284; Hocur. Candollea 2 (1925) 367; Heyne, Nutt. Pl. Ned. Indié (1927) 678; Ocuse & Baku. Indische Groenten (1931) 174; Backer & Baku./. Fl. Java | (1963) 188; Git, Ann. Naturhist. Mus. Wien 83 (1980) 429. — B. rugosa PrRain. See Koorp. Exk. Fl. Java 2 (1912) 286; Heyne, Nutt. Pl. Ned. Indié (1927) 678. — Cochlearia officinalis L. See BACKER, Schoolfl. Java (1911) 57; Koorp. Exk. Fl. Java 2 (1912) 283. — /beris amara L. See BACKER & Baku. Fl. Java 1 (1963) 188. — J. umbellata L. See Backer & Baku /. Fl. Java 1 (1963) 188. — Lobularia 544 FLORA MALESIANA [ser. I, vol. 10? bs b ut: i acy. \% Fig. 1. Brassica juncea (L.) CZERN. 1. Habit, x 0.6; 2. flower, x 6; 3. petal, x 6; 4. siliqua, X 2; 5. seed, x 12 (1-3 DRUMMOND & HEMSLEY 3360; 4, 5 Conrabs 230; drawn after African material). Courtesy FL slrop..e. Afr., Crucif. (1982) 4, f. 1. 1988] CRUCIFERAE (Jonsell) 545 maritima (L.) Desv. See BACKER & BAKH./. Fl. Java 1 (1963) 192. — Matthiola incana (L.) R.Br. See BACKER & BaAkH./. Fl. Java 1 (1963) 192. — Sinapis alba L. See Koorp. Meded. Lands Plantentuin 19 (1898) 342; Exk. Fl. Java 2 (1912) 284. In addition the following species has been reported, probably as occasional, but it is in L represented only by indeterminable material: Barbarea vulgaris R.Br. See BACKER, Schoolfl. Java (1911) 55; Koorp. Exk. Fl. Java 2 (1912) 288; Back. & Baku./f. Fl. Java 3 (1968) 643. KEY TO THE GENERA 1. Fruit a siliqua, /.e. less than three times as long as broad. Ovary ovoid to pyriform. 2. sitienle with two seeds, orbicular, elliptic in outline............6..--seceeecee reece 3. Lepidium 2. Silicule with more than two seeds, triangular in outline ....................2.000e00:- 4. Capsella 1. Fruit a siliqua, i.e. more than three times as long as broad. Ovary + cylindrical. Pe COrcY-SNOneV: NON-CehiSCent . 5.4 ¢ a «ce « 3.5. eines kaw k ok wile ene ee 2. Raphanus 3. Siliqua opening by two valves. 4. Siliqua narrowing into a conical beak (rostrum), terminated by the style............... 1. Brassica 4. Siliqua without a rostrum, + abruptly terminated by the style. 5. Siliqua with a thickened replum (i.e. thickened edge of the septum of the siliqua, on which the seeds pam expiosively, dehiscent by coming Valve ix. «jap iu « since viciy a.0.8 wivista otis sisian.oacisis oka 5. Cardamine 5. Siliqua with a thin replum; valves opening without coiling .......................+-. 6. Rorippa 1. BRASSICA LINNE, Sp. PI. (1753) 666; Gen. PI. ed. 5 (1754) 299; ENDL. Gen. PI. (1839) 882; B. & H. Gen. Pl. 1 (1862) 84; Barton, Hist. Pl. 3 (1871) 248; PRANTL in E. & P. Nat. Pfl. Fam. III, 2 (1891) 177; Scnutz, Pflanzenr. IV-105, Heft 70 (1919) 21; in E. & P. Nat. Pfl. Fam. ed. 2, 17b (1936) 321. — Fig. 1. Annual, biennial or perennial herbs, rarely subshrubs, glabrous or with hispidulous simple hairs. Leaves undivided to pinnatipartite. Racemes terminal, usually ebracteate, corymbiform to paniculate, with many rather large flowers, in fruit elongate, lax. Sepals erect, connivent or spreading, the inner larger and somewhat saccate. Petals yellow or white, spathulated to clawed. Stamens 6, the lateral ones often ascending, filaments linear. Lateral nectaries prismatic or re- duced, median ones semiglobose to filiform, often large. Ovary cylindrical, sometimes on a gynophore; stigma semiglobose to slightly bilobed, ovules numerous, usually uniseriate. Fruit a siliqua, readily dehiscent, terete or slightly laterally compressed, in transverse section + biconvex, attenuate into a conical to filiform beak with 0—3 seeds; valves rather thick, convex, with prominent midnerve and inconspicuous side-nerves. Seeds usually globose, reticulate, not mucilaginous. Distr. As indigenous mostly Mediterranean; a number of the nearly 40 spp. have widely spread as weeds, some of them in connection with cultivation. Ecol. Natural habitats are open, rather dry places like seacliffs, etc. In the tropics only as weeds of cul- tivation, roadsides, waste places, efc. 1. Brassica juncea (L.) Czern. Consp. Pl. Chark. Scnurz, Pflanzenr. 1V-105, 70 (1919) 55; Ocuse & (1859) 8; Backer, Fl. Batavia (1907) 49; Mere. Fl. Baku. Ind. Groenten (1931) 164, f. 99-101; Burk. Manila (1912) 214; Koorp. Exk. Fl. Java 2 (1912) —_ Dict. (1935) 361; Backer & BAKH/. FI. Java 1 (1963) 286; Mere. Int. Rumph. Herb. Amb. (1917) 240; =: 188; Henry, Harmful Pl. PNG, Bot. Bull. Lae 12 546 FLORA MALESIANA [ser. I, vol. 10° (1980) 43. — Sinapis juncea LINNE, Sp. Pl. (1753) 668. — Sinapis timoriana DC. Syst. Nat. 2 (1821) 616; Mia. Fl. Ind. Bat. 1, 2 (1858) 93; DECNE, Herb. Timor. Descr. (1835) 97; Bogert. Handl. Fl. Ned. Ind. 1, 1 (1890) 59; Quis. Medic. Pl. Philip. (1951) 332. — B. besseriana ANDRZ. ex TRAUTV. Bull. Soc. Nat. Moscou 33 (1860) 134; Koorp. Meded. Lands Plantentuin 19 (1898) 341. — B. integrifolia (WEsT) Rupr. FI. Ingrica 1 (1860) 96; Merr. En. Philip. 2 (1923) 207. — Fig. 1. Annual herb, glabrous or somewhat hispid at bases of stem and petioles, somewhat glaucous. Stem erect, usually 30—150 cm high, with long ascending branches in upper part. Basal and lower /eaves pet- ioled, not auriculate, with obovate attenuate blade up to 20 by 10 cm, rather coarsely and irregularly dentate, and 1—3 pairs of small lateral lobes; median and upper leaves + distinctly petioled, not auricu- late, obovate and obtuse or oblanceolate and acute, usually 5—10 cm long, irregularly dentate. Racemes corymbiform, rather loose, with numerous flowers; in fruit lax, up to 50 cm long; pedicels ascending, 5—12 mm long. Sepals green, oblong, 4—6 mm long. Petals bright yellow, clawed, with obovate blade, 6—10 mm long. Anthers 1.5—2.5 mm long. Ovary with 10—20 ovules. Siliquae linear, often somewhat torulose, sometimes inflated, 25—75 by 2—3.5 mm, attenuate into a narrowly conical seedless beak, 6—12 mm long; valves with distinct + keeled midnerve. Seeds dark brown, globose, 1—1.5 mm in diameter, with a distinct fine reticulum. Distr. Wild origin unknown, perhaps E. Europe or SW. Asia; widely spread with cultivation in Asia, S. Europe, Africa and America; in Malesia: Malay Peninsula, Java, Borneo (Brunei, Sarawak), Cele- bes, Moluccas, Philippines, New Guinea. Ecol. Weed of waste land, open places, cultiva- tions (sometimes in gardens); mostly at 2000—3000 m. Note. Brassica juncea has in later years become the world’s most important mustard crop. 2. RAPHANUS LINNE, Sp. Pl. (1753) 669; Gen. Pl. ed. 5 (1754) 300; ENDL. Gen. PI. (1839) 886; B. & H. Gen. Pl. 1 (1862) 101; PrRanti in E. & P. Nat. Pfl. Fam. III, 2 (1891) 179; BaILLon, Hist. Pl. 3 (1871) 250; ScHuiz, Pflanzenr. IV-105, 70 (1919) 194; in E. & P. Nat. Pfl. Fam. ed. 2, 17b (1936) 347. Annual to perennial herb, + hispid with simple hairs. Leaves lyrate-pinnati- partite to pinnatifid. Racemes terminal, ebracteate, + corymbiform, with many rather large flowers, in fruit elongate, lax. Sepals erect, the inner larger and somewhat saccate. Petals white, yellow or + violet, clawed. Stamens 6; fila- ments linear; anthers linear, obtuse. Lateral nectaries very small, median semi- globose or filiform. Ovary biarticulate; lower segment very short, upper seg- ment long with 2—20 ovules; stigma capitate. Fruit a transversely articulate sili- qua; lower segment usually stalk-like, very short, seedless; upper segment in- dehiscent, cylindrical, in transverse section circular, + constricted between the seeds, sometimes lomentaceous; beak narrow, seedless. Seeds ovoid to globose, reticulate to smooth. Distr. About 8 spp., native in the Mediterranean area; 2 spp. are almost cosmopolitan weeds. 1. Raphanus sativus LINNE, Sp. Pl. (1753) 669; BACKER, Fl. Batavia (1907) 50; Merr. FI. Manila (1912) 214; Koorp. Exk. Fl. Java 2 (1912) 287; HEyYNE, Nutt. Pl. Ned. Ind. (1927) 679; OcHSE & BakH. Ind. Groenten (1931) 178; Burk. Dict. (1935) 1566; BACKER & Baku .f. Fl. Java 1 (1963) 188; Quis. Medic. PI. Philip. (1951) 336; SroNE, Fed. Mus. J. 26 (1981) 80. — R. caudatus LINNE, Mant. Pl. (1767) 95; Mia. Fl. Ind. Bat. 1, 2 (1858) 93. Annual or usually biennial herb with thickened napiform to cylindrical taproot, hispid especially in lower parts. Stem erect, usually branched, 15—150 cm high. Basal and lower cauline /eaves lyrate- pinnatipartite to -pinnatisect, petioled, not auricu- late, up to 15 cm long; terminal lobe large, at least half as long as the whole leaf, elliptic to ovate, acute, + cuneate, + coarsely dentate; lateral lobes in up to 7 pairs (but usually much fewer), small, oblong or narrowly triangular, + dentate; upper leaves shortly to indistinctly petiolate, undivided, oblong to lan- 1988] CRUCIFERAE (Jonsell) 547 ceolate, acute, cuneate, serrate to dentate. Racemes loosely corymbiform to paniculate with many flow- ers on 10—20 mm long pedicels, in fruit elongate, lax with up to 30 mm long pedicels. Sepals lanceolate, obtuse, 5—10 mm long. Petals white, yellow or lilac, + dark-veined, 15—20 mm long with obovate, emar- ginate to rounded blade, S—6 mm broad. Anthers + 2.5 mm long. Siliqua 20—90 by 7—15 mm; lower seg- ment much reduced; upper segment of + spongy to corky consistency, not or slightly constricted be- Seeds brownish, ellipsoid to globose, 1.5—4 mm long, finely reticulate. Distr. Only known as a cultivated species, which has probably originated from hybrids between forms of R. raphanistrum L. In Malesia as a weed (prob- ably escaped from cultivation) at least in Java, prob- ably elsewhere. Cultivated also in the Malay Penin- sula and the Philippines, probably elsewhere. Ecol. Cultivated land, waste places, roadsides; usually 1800—2200 m altitude. tween the 1—12 seeds; beak conical, 10—20 mm long. 3. LEPIDIUM LINNE, Sp. PI. (1753) 644; Gen. Pl. ed. 5 (1754) 291; ENDL. Gen. Pl. (1839) 879; B. & H. Gen. PI. 1 (1862) 87; BAILLon, Hist. Pl. 3 (1871) 284; PRanTr in E. & P. Nat. Pfl. Fam. III, 2 (1891) 160; THELL. Die Gattung Lepidium (1906); ScHutz in E. & P. Nat. Pfl. Fam. ed. 2, 17b (1936) 407; HEwson, Brunonia 5 (1982) 73. — Papuzilla Rwxey, Trans. Linn. Soc. Lond. II, Bot. 9 (1916) 17; Scuu1z in E. & P. Nat. Pfl. Fam. ed. 2, 17b (1936) 410; P.RoYEN, Mt. Fl. New Guinea 3 (1982) 2014. — Fig. 2. Annual or perennial, small to medium-sized herbs or subshrubs with taproot, often finely pubescent with simple hairs. Stems erect or ascending, sometimes procumbent, freely branching. Leaves membranous or coriaceous, simple to imparipinnatipartite. Racemes terminal or axillary, ebracteate, densely corym- bose, with small to minute flowers, in fruit narrowly cylindrical, contracted or elongate but usually rather dense. Sepals elliptic, not saccate, greenish. Petals usually whitish, somewhat longer or shorter than the sepals, or often lacking. Stamens 6, 4, or 2, median and/or lateral ones reduced; filaments usually linear to subulate. Nectarial glands usually 6 or 4, adjacent to the bases of the lateral filaments. Ovary flat, dorsiventrally compressed, elliptic in outline, ovules 2. Fruit a silicula, dehiscent, flat, strongly compressed, narrowly septate, usually orbicular to elliptic in outline, winged (especially distally) or not, with the short style as an apical notch. Seeds single in each locule, very finely reticulate, muci- laginous, usually reddish brown, rather flat, elliptic to obovate in outline, some- times narrowly and unilaterally winged. Distr. Worldwide; a number of the c. 150 spp. are cosmopolitan weeds. Ecol. Largely open, wet or dry ground without closed vegetation, in elevated regions. Very often in dis- turbed places. KEY TO THE SPECIES 1. Inflorescences terminal with numerous flowers. Spines absent. 2. Leaves deeply divided. Siliculae with prominent wings ........ 66600 eee eee eee eens 1. L. sativum 2. Leaves undivided, serrate. Siliculae winglesS .........0ceccceeeeeeeseseeveneuces 2. L. virginicum 1. Inflorescences very short with 5 or fewer flowers inserted at base of spines. $;/Subshrub; more than 20.cm high. 605.0 5 Ooee ev ee Peeves eeu aed seed vEnvete 5. L. maccowagei 3. Cushion forming, rhizomatous, partly woody herbs, less than 15 cm high. 548 FLORA MALESIANA [ser. I, vol. 103 4. Cushions over 5 cm thick. Leaves mostly serrate or incised with up to 4 pairs of lobes 4. L. minutiflorum 4. Cushions less than 5 cm thick. Leaves mostly deeply pinnatifid with up to 7 pairs of lobes 1. Lepidium sativum LINNE, Sp. Pl. (1753) 644; Heyne, Nutt. Pl. Ned. Ind. (1927) 676; Burk. Dict. (1935) 1331; BACKER & BAKH./f. Fl. Java 1 (1963) 188. Annual herb with slender taproot, up to 70 cm high, erect, simple or much branched, glabrous or with scattered minute hairs. Leaves membranous, petioled, very sparsely pubescent, up to 10 cm long, imparipinnati- or bipinnatipartite, with 2—4 pairs of lateral lobes; lobes linear, lanceolate or oblanceolate, up to 3 cm long; uppermost leaves sometimes simple, serrate. Racemes terminal, with rather conspicuous whitish flowers, in fruit loose, up to 25 cm long; pedicels in fruit 3—6 mm long, ascending to almost erect, straight, glabrous. Sepals green with membra- nous margins, elliptic, 1—1.5 mm long. Petals white or violet, spathulate to slightly clawed, with rounded apex, 1.7—3 mm long. Stamens 6, with a single small nectary between each of them. Ovary elliptic, emar- ginate, with projecting style. Si/iculae elliptic in outline, 4.5—6.6 mm long, 3—4 mm broad, with prominent apical wings, deeply emarginate; style dis- tinct, with stigma completely within or just level with the top of the sinus. Seeds wingless, reddish brown, 2—3 mm long, c. 1.5 mm broad. Distr. Probably native in Ethiopia and neigh- bouring countries to the west and north, and in the Near East; spread by cultivation or inadvertently to many parts of the world; in Malesia introduced: Java, Flores, probably elsewhere. Ecol. Occurs as an occasional escape from culti- vation; uncertain whether naturalized anywhere in the area. 2. Lepidium virginicum LINNE, Sp. Pl. (1753) 645; Merk. En. Philip. 2 (1923) 207; BACKER & BaAKH./. Fl. Java 1 (1963) 187. Annual or short-lived perennial herb with taproot, 20—80 cm high with one to few erect-ascending stems, branching above, puberulent with thin falcate hairs. Leaves membranous, lowest ones lyrate-pinnatifid, up to7 cm long, median and upper ones mostly oblan- ceolate, acutely serrate. Racemes terminal with minute flowers, rather dense in fruit, up to 15 cm long; pedicels in fruit 3.5—5.5 mm long, patent, straight, glabrous. Sepa/s greenish, c. 1 mm long. Petals white, longer or shorter than sepals, very nar- row. Stamens 2 (median) with triangular nectaries. Ovary elliptic, emarginate, style not projecting. Siliculae wingless, suborbicular, 3—3.5 by 2.7—3.5 mm, rather widely and deeply emarginate; style short with stigma completely contained within sinus. Seeds narrowly winged, red-brown, c. 1.5 by 0.9 mm. 3. L. laeteviride Distr. Native of North America; widespread as a weed; in Malesia known from E. Java (Mt Tengger) and the Philippines (Luzon). Ecol. In Java naturalized in young forest at c. 1500 m. Elsewhere a weed in settled areas. 3. Lepidium laeteviride (P.ROYEN) HEwson, Brunonia 5 (1982) 74. — Papuzilla laeteviridis P.RoyEN, Nova Guinea, Bot. 19 (1964) 430; Mt FI. New Guinea 3 (1982) 2014, f. 615, pl. 154. — Fig. 2. Perennial, mat- or cushion-forming, rhizomatous herb with persistent old leaf bases. Stems up to 3.5 cm long, sparingly branched, puberulent, and with a few leaf-opposed, up to 1 cm long spines. Leaves coriaceous, sessile; basal ones in a rosette, sparsely puberulent, bipinnatifid with up to 7 pairs of lobes, acute, without spines; cauline leaves obovate, entire or acutely serrate. Racemes axillary at the base of spines with 1—3 minute flowers, in fruit hardly elon- gating, up to 10 mm long; pedicels 2—3 mm long, pa- tent, sparsely pubescent. Sepals green with hyaline margins, c. 1 mm long. Petals white, shorter than sepals, 0.6—0.8 mm long. Stamens 2 (median), with 4 linear nectaries, one on either side of each stamen. Ovary elliptic with inconspicuous style. Siliculae in outline broadly ovate to obovate, 3—3.5by2.5—3mm, with very narrow wing in upper third, shallowly emar- ginate; style very short, stigma sessile or nearly so. Seeds wingless, dark brown, 1.5—2 by 0.8—1.3 mm. Distr. Malesia: Papua New Guinea (Morobe District, Sarawaket Range, Mt Bangeta). Ecol. On overgrown screes and among boulders at c. 4100 m. 4. Lepidium minutiflorum (RIDLEY) HEwson, Brunonia 5 (1982) 74. — Papuzilla minutiflora Riptey, Trans. Linn. Soc. Lond. II, Bot. 9 (1916) 16, f. 7-14, pl. 1; MERR. & Perry, J. Arn. Arb. 24 (1943) 207; P.RoYEN, Nova Guinea, Bot. 19 (1964) 428; Mt. Fl. New Guinea 3 (1982) 2014, f. 614. Perennial, mat- or cushion-forming rhizomatous herb with persistent old leaf bases. Stems up to at least 12 cm, intricately branched, puberulent, with numerous leaf- or bract-opposed, up to 1.2 cm long spines. Leaves coriaceous, sessile, up to 25 mm long, glabrous to sparsely puberulent, impari- (or rarely bi-)pinnate with 2—4 pairs of lobes, or undivided, acutely serrate or entire. Racemes axillary at the base of spines with 1—3 minute flowers, in fruit little elon- gating, to 5 mm long; pedicels 2—4 mm long, straight, patent, puberulent. Sepals green with hya- line margins, 1—1.5 mm long. Petals white, shorter than or equalling sepals, linear. Stamens 2 (median) 1988] CRUCIFERAE (Jonsell) 549 Fig. 2. Lepidium laeteviride (P.ROYEN) HEwson. a. Habit, x 4; b. flower, x 12; c. ditto, calyx removed; d. petals; e. ovary; f. ovary, lateral view; all x 16; g. fruit; A. seeds; i. fruit valve; all x 4 (P. Roven 20026). with 4 shield-like nectaries, one on either side of each stamen. Ovary elliptic with very short style. Siliculae broadly to cordate in outline, 2.5—5 by 2—4 mm, with narrow wing in upper third, not or indistinctly narrowly and shallowly emarginate; style 0.5—1 mm long, stigma exserted from the sinus. Seeds wingless, dark brown, 1—2 by 0.5—1.3 mm. Distr. Malesia: West New Guinea (Mts Car- stensz & Wilhelmina), Papua New Guinea (W. Sepik District: Mt Capella). Ecol. Screes, wet open places, tussock grassland, heaths; 3100-4350 m. 5. Lepidium maccowagei Hewson, Brunonia 5 (1982) 75. — Papuzilla minutiflora auct. non Riptey: P.Roven, Mt. Fl. New Guinea 3 (1982) 2014, p.p., as to Papua New Guinea specimens. Subshrub with taproot, 20—35 cm high or more, woody, branched, puberulent, with numerous leaf- opposed, up to 1.3 cm long spines. Leaves coria- ceous, sessile, up to 14 by 5 mm, glabrous to puber- ulent, ovate to obovate, acutely serrate. Raceme re- duced to a single flower at the base of a spine; pedi- cels 3—S mm long, straight, patent, puberulent. Sepals green with hyaline margins, 0.7—1 mm long. Petals white, shorter than or equalling sepals, linear. Stamens 2 (median) with 4 linear nectaries, one on either side of each stamen. Ovary elliptic with very short style. Siliculae broadly elliptic to obovate, 3.5—6.5 by 2.5—3.8 mm, with narrow wing in upper third, narrowly and shallowly emarginate; style c. 0.3 mm long, just level with the top of the sinus or slightly exserted. Seeds wingless, red-brown, 2—2.5 by 1.1—1.5 mm. Distr. Malesia: Papua New Guinea (Southern Highlands: Mt Giluwe, Kaguba). Ecol. Screes, stream-edges, efc., at lower alti- tudes along roadsides; 2750-3500 m. 4. CAPSELLA Mepicus, Pflanzengatt. 1 (1792) 85; ENDL. Gen. PI. (1839) 878; B. & H. Gen. Pl. 1 (1862) 86; BaiLton, Hist. Pl. 3 (1871) 287; Scnuz in E. & P. Nat. Pf. Fam. ed. 2, 17b (1936) 453. 550 FLORA MALESIANA [ser. I, vol. 10° Annual to short-lived perennials, small to medium-sized with taproot. Hairs simple, or forked, or branched. Racemes terminal, ebracteate, corymbose, with small flowers, in fruit loose. Sepals somewhat spreading, not saccate. Petals white, sometimes reddish or yellowish, + clawed, rarely absent. Stamens 6; filaments linear; anthers ovoid, obtuse. Nectaries as semicircular glands at each side of each lateral stamen. Ovary dorsiventrally flattened, obovate in outline, with 10—40 ovules in each locule. Fruit a silicula, dehiscent, flat, with narrow septum, obtriangular to obcordate, valves keeled. Seeds numerous, not mucila- ginous. Distr. About 5 spp., mainly European; doubtfully congeneric species in other parts of the world; in Malesia introduced. 1. Capsella bursa-pastoris (L.) MEpicus, Pflanzen- gatt. 1 (1792) 85; Backer, Schoolfl. Java (1911) 58; Koorp. Exk. Fl. Java 2 (1912) 291; Merr. En. Philip. 2 (1923) 208; BACKER & BAKH./. Fl. Java 1 (1963) 192; EVERAARTS, Weeds Veget. Java (1981) Sle sis Soe Annual to short-lived perennial herb, 5—50 cm high. Stems 1—few, erect or ascending, simple or sparsely branched. Rosette /eaves numerous, petiolate, imparipinnatipartite to coarsely dentate, oblanceolate in outline, 3—15 cm long; cauline leaves rather few, smaller, oblong to lanceolate, undivided (lower leaves sometimes lyrate-pinnatifid), sagittate and clasping, coarsely to finely distantly dentate or entire. Stem and leaves with simple and branched hairs. Racemes dense, with numerous small flowers, pedicels spreading to divaricate, 5-20 mm long. Sepals green, mostly pubescent, oblong (sometimes with a tinge of purple), c. 1.5 mm long. Petals white, clawed, distinctly exceeding sepals, 1.8—3 mm long. Silicula obtriangular, with straight or slightly con- cave margins, + emarginate, 5-10 mm long, 2.5—8 mm broad; style c. 0.3 mm long. Seeds reddish brown, oblong, 0.8—1 by c. 0.4 mm, minutely verru- cose. Distr. Almost cosmopolitan as a weed, probably native in Europe, introduced in Malesia, recorded from highlands of E. Java, Philippines (Luzon) and E. New Guinea; probably more widespread. Ecol. Weed of cultivated soil in mountain areas; 2000—2500 m. Vern. Shepherd’s purse (EB). in fruit elongate, loose, sparsely hairy or glabrous; 5. CARDAMINE LINNE, Sp. PI. (1753) 654; Gen. Pl. ed. 5 (1754) 295; ENDL. Gen. Pl. (1839) 865; B. & H. Gen. Pl. 1 (1862) 70; PRANTL in E. & P. Nat. Pfl. Fam. UI, 2 (1891) 184; BaILLon, Hist. Pl. 3 (1871) 234; Scuuiz, Bot. Jahrb. 32 (1903) 280; in E. & P. Nat. Pfl. Fam. ed. 2, 17b (1936) 527. — Pteroneurum DC. Syst. Nat. 2 (1821) 269. — Fig. 3. Annual, biennial or perennial, small to large herbs, with a taproot and often with creeping or tuberous rhizomes. Hairs often present, simple. Sfems erect, ascending, sometimes basally rooting. Racemes mostly terminal, ebracteate, densely to laxly corymbose, with a few to numerous small to medium-sized flowers, in fruit at least basally elongate and lax. Sepals not saccate, not spread- ing; margins hyaline. Petals white, cream, violet or purple, spathulate to clawed, sometimes lacking. Stamens 6—4, rarely fewer; filaments linear to subulate. Nectaries horseshoe-shaped or a close ring-shaped tissue around the lateral stamen and usually as a conical gland outside each median pair of sta- mens. Ovary cylindrical, usually with a short narrower style; stigma flat; ovules 1988] CRUCIFERAE (Jonsell) 551 4—40. Fruit a siliqua, dehiscent, linear, straight, tapered towards the narrow style; stigma flat; fruit disrupting explosively by the valves coiling from the base upwards; replum swollen; valves without prominent nerves. Seeds uniseriate, flattened, usually broadly elliptic to oblong, minutely rugose to colliculate, not mucilaginous. Distr. In subarctic, temperate, and montane tropical areas all over the world; c. 130 spp., of which 6 spp. in Malesia. Ecol. Most species grow in moist places, many in or along running water, in marshes, efc. Quite a few form undergrowth in rather dense forests. Very few species are widely spread as weeds, though many are local- ly favoured by human interactions (clearings, ditches, efc.). Note. Specific delimitation is notoriously difficult in this genus through frequent hybridization and occur- rence of polyploids. KEY TO THE SPECIES 1. Leaves tripartite or undivided. 2. Leaves undivided or terminal leaflet at least 6 times as long as the lateral ones ....... 1. C. papuana 2. Leaves tripartite, terminal leaflet not more than 3 times as long as the lateral ones.... 2. C. africana 1. Most leaves with two or more pairs of lateral leaflets. 3. Leaflets oblanceolate in outline. Plants usually very slender with internodes much longer than leaves 3. C. altigena 3. Leaflets ovate to orbicular in outline. Plants not very slender, with most internodes equalling or shorter than leaves. aeeersiswet least:6 mim long’. 2....2--. 25.65. f-sk 4. Petals not exceeding 4 mm. bes ia «cM ecnsie te Se RSE Os See tae 4. C. keysseri 5. Pedicels of siliquae patent at an angle with stem exceeding 45° (usually c. 60°). Seeds at least 1.1 mm OUL ea See eS i ai ae ae rs ae ht Ria nes bee eae ee 5. C. flexuosa 5. Pedicels of siliquae erect to erecto-patent (angle with stem c. 45° or less). Seeds shorter than 1.1 mm 1. Cardamine papuana (LAutT.) O.E.ScuHutz, Bot. Jahrb. 55 (1918) 271, f. 1E; Merr. & Perry, J. Arn. Arb. 24 (1943) 207; P.RoveN, Mt. Fl. New Guinea 3 (1982) 2021. — C. africana L. ssp. borbonica (PERS.) O.E.ScHuLz var. papuana Laut. in K.Sch. & Laut. Fl. Deut. Schutzgeb. Siidsee, Nachtr. (1905) 271; Putte, Nova Guinea 8 (1911) 641. — Fig. 3. Perennial, rhizomatous, glabrous (rarely pubes- cent) herb with 15—SO cm high, erect or ascending shoots, appearing solitarily or with 2 or 3 together. Stems unbranched or with a few rather long branches in the upper part. Leaves tripartite or unifoliate (side-lobes reduced); basal leaves (often not present) up to 12 cm long with 9 cm long petioles; cauline leaves rather few, up to 8 cm long, without or with only slightly auriculate, 2~8 cm long petioles; termi- nal leaflet ovatotriangular, acute to acuminate, at base cuneate or rounded (rarely truncate), distantly serrate-dentate, 3—12 by 1—4 cm on 5—20 mm long petiolules (in partite leaves); lateral leaflets ovate, acute, sparsely serrate or dentate, 0.5—3 by 0.2—2. cm on 1—4 mm long petiolules. Racemes terminal (but often surpassed by upper leaves) with 5—15 flowers on 0.5—5 cm long peduncles, in fruit condensed or 6. C. hirsuta loose, 1—6 cm long; pedicels straight, erecto-patent, 5—12 mm long. Sepals green with hyaline margins, oblong, 2.5—4 mm long. Petals white, broadly spathulate, usually notched, 2 or 3 times longer than sepals, 8-12 mm long, + half as broad. Stamens 6, with linear filaments; anthers c. 0.8 mm long. Ovary cylindrical with indistinctly demarcated style and somewhat expanded stigma. Si/iqua straight, linear, 20—40 by 1.5—2 mm; style 0.5—1.7 by 0.3—0.5 mm. Seeds dark to light brown, oblong in outline, 1.7—2 by 1—1.3 mm, smooth. Distr. Malesia: New Guinea (throughout), rather widely distributed. Ecol. Wet cliffs, rocky banks of streams in forests; 500-3100 m. 2. Cardamine africana Linn&é, Sp. Pl. (1753) 655; Scuutz, Bot. Jahrb. 32 (1903) 414; Koorp. Exk. Fl. Java 2 (1912) 290; BotpinGcu, Zakfl. Landb. Java (1916) 67; Koorp. Fl. Tjibodas 3 (1923) 80; RipLey, J. Mal. Br. R. As. Soc. nm. 87 (1926) 52; Henty & PrircHarp, Weeds New Guinea, Div. Bot. Lae, Bot. Bull. 7 (1973) 96. — C. borbonica Pers, Syn. Pl. 2 (1807) 195, — Pteroneurum javanicum BLUME, 552 FLORA MALESIANA [ser. I, vol. 103 Vi pp Fig. 3. Cardamine papuana (Laut.) O.E.Scuutz. a. Habit, x 0.6; b. siliqua, x2; c. seed, scale bar 1 mm (SCHODDE 5540) Bijdr. (1825) 51; ZoLi. & Mor. Syst. Verz. (1845/46) 35; Mia. FI. Ind. Bat. 1, 2 (1858) 93. — C. javanica (BLUME) Mra. Ill. Arch. Ind. (1871) pl. X; Borrt. Handl. Fl. Ned. Ind. 1, 1 (1890) 58; Backer, School- fl. Java (1911) 56. — C. africana L. ssp. borbonica (PERS.) SCHULZ, Bot. Jahrb. 32 (1903) 414; DocTERs VAN LEEUWEN, Verh. Kon. Akad. Wet. Amsterdam, . Del. K.G. Forss. sect. II, 31 (1933) 167; BACKER & BakH./f. Fl. Java 1 (1963) 191; STEEN. Mt. Fl. Java (1972) pl. 13/6; P.RoyeEN, Mt. Fl. New Guinea 3 (1982) 2021; STREI- MANN, Canberra Nat. Bot. Gard. (1983) 100 (‘var. borbonica’). Perennial rhizomatous herb with 10—80 cm high, erect or ascending shoots appearing solitary or clus- 1988] CRUCIFERAE (Jonsell) 558 tered. Stems often basally rooting, unbranched or with a few branches, most from the lower parts, gla- brous or basally puberulent. Leaves tripartite (rarely with one additional small pair of leaflets), glabrous or hispidulous above; basal leaves up to 15 cm long with up to 8 cm long petioles, soon disappearing; cauline leaves few to numerous, 5—20 cm long, in lower leaves with proportionally very long (up to 12 cm), slightly auriculate petioles; terminal leaflet broadly to narrowly ovate or subtriangular, acute to acuminate (but the tip often obtuse with a short point), at base truncate or rounded (in upper leaves also cuneate), often oblique, crenate, serrate or in- cised (especially plants from New Guinea), 1.5—7.5 by 0.8—4 cm on 2—20 mm long petiolules; lateral leaflets of similar shape as the terminal, slightly to considerably smaller, 0.5—6.5 by 0.2-3 cm on 0.5—10 mm long petioles. Racemes terminal with 5—20 flowers on 1—25 cm long peduncles, in fruit condensed or loose, 2—25 cm long; pedicels straight, erecto-patent, 4—30 mm long. Sepals green with hya- line margins, elliptic, 1.4—4.5 mm. Petals white to pale violet, spathulate, broadly obtuse, at least twice as long as sepals, 3—12 mm long, + 1/3 as broad. Stamens 6 with subulate filaments; anthers 1—1.5 mm long. Ovary cylindrical with short narrower style and somewhat expanded stigma. Si/iqua straight, lin- ear, 20—55 by 1.3—2.5 mm, tapered towards style, which is 0.6—2.5 by c. 0.4 mm. Seeds red or dark brown, broadly elliptic or oblong in outline, 1.5—2.5 by 1—1.5 mm, smooth. Distr. Ubiquist in tropical montane areas. In Africa from Ethiopia south to South Africa (Cape Province) and westwards to Mt Cameroun; also in Fernando Pdéo, S40 Tomé, Comores, Madagascar and Réunion. In America in many Caribbean is- lands, Central America and NW. South America; in Asia in India and Sri Lanka; in Malesia: Sumatra (northern parts: W., E. coast, Tapanuli, Atjeh), Java (W., E.), Lesser Sunda Islands (Bali, Lombok, Sumbawa), Ceram, Papua New Guinea. Ecol. Undergrowth in montane forests, often also in clearings, along roads and paths. Usually in damp, shady places; 200—2400 m. 3. Cardamine altigena O.E.Scuutz, Bot. Jahrb. 62 (1929) 479; Merr. & Perry, J. Arn. Arb. 24 (1943) 207; HooGLaAnpD, Blumea Suppl. 4 (1958) 227; P.Roven, Mt. Fl. New Guinea 3 (1982) 2021, f. 616, pl. 154. Perennial, slender herb with 20—50 cm long stems, ascending to largely prostrate and running with root- ing nodes from which more stems may arise. Stems glabrous with few, slender branches. Leaves gla- brous or sparsely ciliate, imparipinnate with 1—3 pairs of lateral leaflets (rarely leaflets lacking); basal leaves 2—7 cm long with 1.5—3 mm long petioles, soon disappearing; cauline leaves distantly inserted, rather few, up to 6 cm long with non-auriculate, up to 2 cm long petioles; terminal leaflets trifid, orbicu- lar or reniform, in upper leaves oblanceolate, obtuse, 0.5—1.5(—3) by 0.1—1.2(—2) cm, on 0—8 mm long petiolules; lateral leaflets trifid to oblanceolate, 0.2—0.7 by 0.1—0.5 cm, petiolules c. 1 mm. Racemes terminal, with 3—10 flowers, often very slender; pedicels straight, erect to erecto-patent, 8-15 mm long. Sepals green to reddish white with hyaline mar- gins, oblong, 1.8—2.7 mm long. Petals white, spathulate, obtuse, 3.5—6 mm long, + 1/3 as broad. Stamens 6, tetradynamous, with subulate filaments; anthers 0.3—0.5 mm long. Ovary cylindrical with in- distinct style and flat stigma. Siliqua straight linear, 13—35 by 1.3—1.8 mm; style 0.4—2 by c. 0.5 mm. Seeds red-brown, oblong in outline, 1.4—1.7 byc. 1.1 mm, smooth. Distr. Malesia: New Guinea (throughout), wide- spread in high mountain areas. Ecol. Preferably in or near running water; along streams, in wet grassy slopes; 1700—4000 m. 4. Cardamine keysseri O.E.ScHuLz, Bot. Jahrb. 62 (1929) 480; P.RoyEN, Mt. Fl. New Guinea 3 (1982) 2025,9f:7617, (pl 156: Annual to probably short-lived perennial herb with 10—40 cm high, ascending main stem, basally prostrate and rooting at nodes, from which one to several new stems may arise. Stems in lower parts mostly hispidulous without or with rather few, usual- ly arcuate branches. Leaves glabrous or with a few scattered hairs, imparipinnate with 2—4 pairs of lateral leaflets; basal leaves 4—8 cm long with 1—4 cm long petioles, soon disappearing; cauline leaves often numerous, up to 9 cm but mostly much smaller with non-auriculate petioles; leaflets trifid, acute (often with a point), at base attenuate, cuneate (rarely trun- cate), 0.3—2.5 by 0.2—1.4 mm; terminal leaflet often larger and proportionately narrower than the lateral; petiolules 0-15 mm. Racemes terminal with 5—10 flowers on up to 4 cm long peduncles, in fruit up to 10 cm long; pedicels straight, erecto-patent, 4—10 mm long. Sepals green to reddish with hyaline mar- gins, oblong, 2.5—3.5 by c. 1.2 mm. Petals white, spathulate, obtuse or emarginate, 2—3 times longer than sepals, 7—8 mm long, + half as broad. Stamens 6 of nearly equal length with subulate filaments; an- thers c. 0.8 mm long. Ovary bottle-shaped, with short distinct style and slightly expanded stigma. Sili- quae straight, linear, 20-38 by 1-2 mm; style 0.5—1.4 by ec. 0.5 mm. Seeds red-brown, elliptic in outline, 1.4—1.8 by 1.2—1.4 mm, smooth. Distr. Malesia: New Guinea (throughout), rather widespread. Ecol. Sandy or gravelly streambanks, moist forest clearings, moist eroded soil, in gardens; 900-4000 m. 554 5. Cardamine flexuosa WirTH. Arr. Brit. Pl. ed. 3, 3 (1796) 578; Scuutz, Bot. Jahrb. 32 (1903) 473; Koorp. Exk. Fl. Java 2 (1912) 290; BoLDINGH, Zakfl. Landb. Java (1916) 67; RmLey, J. Fed. Mal. St. Mus. 8, 4 (1917) 15. — Pteroneurum decurrens BiuME, Bijdr. (1825) 51. — C. decurrens (BLUME) Zou. & Mor. Syst. Verz. (1845/46) 35; Mia. Fl. Ind. Bat. 1, 2 (1858) 93; Ill. Arch. Ind. (1871) pl. X; BoeERL. Handl. Fl. Ned. Ind. 1, 1 (1890) 58; BACKER, Voorl. Schoolfl. (1908) 11; Schoolfl. Java (1911) 56. — C. regeliana Miq. Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 73; MerR. En. Philip. 2 (1923) 208; RiDLey, J. Mal. Br. R. As. Soc. n. 87 (1926) 52. Mostly perennial, diffusely branching herb; from the base usually many erect to ascending, curved or flexuous stems, in lower parts sometimes prostrate and rooting, 10—50 cm high with few to numerous scattered hairs. Basal /eaves rarely persistent; cauline leaves nearly glabrous to sparsely pubescent, mostly numerous, imparipinnate with a terminal and 2 or 3 pairs of lateral leaflets, 2—10 cm long; petiole non- auriculate; leaflets narrowly to broadly ovate or or- bicular in outline, obtuse, truncate to cuneate, usual- ly crenate and often shallowly trifid; terminal leaflets 0.7—3.5 by 0.7—2 cm on distinct petiolules; lateral leaflets 0.4—1.5 by 0.3—1.5 cm on 1.5—7 mm long petiolules. Racemes terminal with 5—20 flowers, in fruit rather lax, up to 12 cm long; pedicels in fruit pa- tent (angle to stem over 45°), 6—12 mm long with pa- tent to erect siliquae, angled upwards at junction with the pedicel. Sepals green with narrow hyaline margins, oblong, obtuse, c. 1.5 mm long. Petals white, spathulate, 2.5—4 mm long. Stamens 6, dis- tinctly tetradynamous; anthers 0.3—0.4 mm long. Ovary cylindrical with a rather short style, narrower than the ovary, and a flat stigma. Siliqua straight, linear, 18—30 by 0.8—1.2 mm; style 0.7—2 by c. 0.3 mm. Seeds reddish brown, often with a vestigial wing, broadly oblong in outline, 1.1—1.4 by 0.8—1.1 mm, nearly smooth. Distr. Circumpolar in northern temperate region. Also in some montane areas further south. In Malesia: N. Sumatra (Atjeh), Java, Celebes (Mena- do), Philippines (Mindanao, Luzon), Moluccas (Buru, Ceram). Ecol. Moist places in forests, among rocks and boulders, at streams, in clearings; 1000—2200 m. 6. Cardamine hirsuta LINNE, Sp. Pl. (1753) 655; BoERL. Handl. Fl. Ned. Ind. 1, 1 (1890) 58; Koorp. Nat. Tijd. Ned. Ind. 62 (1902) 234; ScHuLz, Bot. Jahrb. 32 (1903) 464; Backer, Schoolfl. Java (1911) 56; Koorp. Exk. Fl. Java 2 (1912) 290; RIDLeEy, Trans. Linn. Soc. Lond. II, Bot. 9 (1916) 17; Koorp. FLORA MALESIANA [ser. I, vol. 103 Fl. Tjibodas 3 (1923) 80; Hocur. Candollea 2 (1925) 371; BurK. & HEND. Gard. Bull. S. S. 3 (1925) 345; DocTERS VAN LEEUWEN, Verh. Kon. Akad. Wet. Amsterdam, sect. II, 31 (1933) 168; BACKER & BakH./. Fl. Java 1 (1963) 191; STEEN. Mt. Fl. Java (1972) pl. 13/5; Henty & PrircHARD, Weeds New Guinea, Div. Bot. Lae, Bot. Bull. 7 (1973) 96, fig.; Gitu, Ann. Naturhist. Mus. Wien 83 (1980) 429; EvERAARTS, Weeds Veget. Java (1981) 53, f. 39. — C. flexuosa auct. non WiTH.: STONE, Fed. Mus. J. 26 (1981) 80. Usually annual, branching from the base but not or very sparsely above (occasionally perennial and then branched above and devoid of leaf-rosette). Stems erect or in tall specimens curving, 5—30 cm high, glabrous or with scattered hairs. Basal leaves forming a distinct rosette, cauline leaves rather few; all leaves imparipinnate, with a terminal and 2—6 pairs of lateral leaflets, 1.5—10 cm long (including petiole); petiole non-auriculate, with a few straight hairs at base; leaflets rhombic, ovate, orbicular or reniform, truncate to broadly cuneate (in upper parts also narrowly cuneate), entire to deeply crenate, oc- casionally with secondary lobes, glabrous or with scattered hairs; terminal leaflet 0.5—1.5 by 0.3—2 cm, distinctly petioled, the lateral ones oblique, 0.2—1 by 0.2—1.1 cm, nearly sessile or with a petiole up to the length of the leaflet. Racemes terminal, very dense, with several small whitish flowers, in fruit rather lax (up to c. 10 cm long); pedicels in fruit erect to suberect (angle to stem usually c. 45° or less), 3—8 mm long with erect siliquae overtopping the flowers. Sepals green with narrow white margins, oblong, ob- tuse, c. 1.5 mm long. Petals white, narrowly spathu- late, usually 2-3 mm long (rarely absent). Stamens 4—6; anthers c. 0.4 mm long. Ovary cylindrical with a very short style narrower than the ovary, and a flat stigma. Siliqua glabrous, linear, 12—27 by c. 1 mm; style 0.5—1 by c. 0.4 mm. Seeds reddish brown with a vestigial wing, broadly oblong to suborbicular in outline, c. 1 by 0.8 mm, smooth. Distr. Native at least in Europe, and probably in some tropical mountain areas (Ethiopia, E. Africa, Cameroun). Widespread as a weed in most parts of the world and probably naturalized in many areas; in Malesia: possibly native in some mountain areas, elsewhere a weed or naturalized; Malay Peninsula (incl. also Singapore), Sumatra (N. parts, Ben- coolen), Java, Borneo (Sabah), Philippines (Luzon), New Guinea (throughout). Ecol. Moist, open ground in montane forests, clearings, efc.; 850-3000 m. Weed in gardens, flow- erbeds, also at low altitudes. 1988] CRUCIFERAE (Jonsell) 555 6. RORIPPA Scop. Fl. Carniol. (1760) 520; JonsELL, Symb. Bot. Upsal. 197 (1968); StucKEY, Sida 4 (1972) 279. — Nasturtium R.Br. in Ait., Hort. Kew. ed. 2, 4 (1812) 109; ENDL. Gen. Pl. (1839) 863; B. & H. Gen. Pl. 1 (1862) 68; PRANTL in E. & P. Nat. Pfl. Fam. III, 2 (1891) 184; BarLton, Hist. Pl. 3 (1871) 232; ScHutz in E. & P. Nat. Pfl. Fam. ed. 2, 17b (1936) 551. — Trochiscus Gii1, Ann. Naturhist. Mus. Wien 83 (1980) 430. — Fig. 4. Annual to perennial, small to rather tall herbs with a taproot, sometimes also with root-runners or adventive roots from the stem-nodes, practically glabrous to pubescent with simple, pointed hairs. Stems prostrate to erect, usually much branched. Leaves membranous, usually pinnatifid to pinnate, often auriculate. Racemes terminal, less often axillary, with or without bracts, nearly corymbose, with numerous, rather small to small flowers, in fruit cylindrical and loose. Sep- als greenish, elliptic, not saccate, spreading. Petals yellow or white, equalling or upto + twiceas long as the sepals. Stamens 6, distinctly tetradynamous, with usually linear filaments; anthers ovate to oblong, yellow, blunt. Nectarial tissue + surrounding the lateral filament bases, usually also extending outside the me- dian ones but sometimes lacking there. Ovary spheroid to narrowly cylindrical, with numerous ovules. Fruits terete, very variable in shape (globose to narrowly linear), without or rarely with a very short gynophore, usually with a short but distinct style and flat to slightly bifid stigma, dehiscent by 2 valves with or without very fine midnerve. Seeds many to numerous, not mucilaginous, finely and variously sculptured, yellowish to dark brown, ellipsoid to spheroid, wing- less, 0.5—1.5 mm diameter. KEY TO THE SPECIES Seeecenties bracteate: yo.) )5.4/))c Hoes Miles sR OBIS... ERTS A ee ee 8. R. benghalensis 1. Racemes ebracteate (rarely with a few bracts at the lowest pedicels). 2>Siliquae Hot more than\1':5:mm thick. is(.\) 0022... oes dines 40 1 SORE aenie 7. R. heterophylla 2. Siliquae more than 1.5 mm thick. 3. Pedicels of fruits straight. Ae Seeds notiover:0:8 mim long 2 oi) V3 Wire «0.6 okt. CITT ee 9. R. micrantha 4. Seeds longer than 0.8 mm. 5. Cauline leaves numerous; seeds distinctly reticulate (c. 150 areoles per face)........ 2. R. backeri 5. Cauline leaves absent or few; seeds very finely reticulate (over 300 areoles per face) 3. R. peekelii 3. Most pedicels of fruits curved. 6, Petals white. Seeds coarsely reticulate’... 3.0... see Tei ole eae an 1. R. nasturtium-aquaticum 6. Petals yellow. Seeds with very fine surface patterns. 7. Ripe siliquae inflated, semighobose............csseeecceeeeeneessevenenecees 6. R. schlechteri 7. Ripe siliquae + ellipsoid to sausage-shaped, not inflated. 8. Seeds with warty processes. Very tall and stout plants ........... sees eeees 5. R. hybosperma 8. Seeds without processes. Moderately high plants ..........60 cece cece eee eens 4. R. palustris 1. Rorippa nasturtium-aquaticum (L.) Hayek, 2, 4 (1812) 110; Mig. Fl. Ind. Bat, 1, 2 (1858) 73; Sched. FI. Stir. Exs. 3-4 (1905) 22. — Sisymbrium — Boert. Handl. Fl. Ned. Ind. 1, 1 (1890) 58; Koorp. nasturtium-aquaticum Linne#, Sp. Pl. (1753) 657. — Meded. Lands Plantentuin 19 (1898) 342; Backer, Nasturtium officinale R.Br. in Ait., Hort. Kew. ed. — Schoolfl. Java (1911) 55; Koorp. Exk. Fl. Java 2 556 FLORA MALESIANA [ser. I, vol. 102 Fig. 4. Harvest of Rorippa nasturtium-aquaticum (L.) HAYEK (Photogr. W.F. WINCKEL, 1918). (1912) 288; Merr. En. Philip. 2 (1923) 208; HEYNE, Nutt. Pl. Ned. Ind. (1927) 680; OcHsE & Baku. Ind. Groenten (1931) 176, f. 108; Burx. Dict. (1935) 1534; Quis. Medic. PI. Philip. (1951) 335; BACKER & BakH./. Fl. Java 1 (1963) 191; Gru, Ann. Naturhist. Mus. Wien 83 (1980) 430; LENCH & OsBoRNE, Fresh- water Pl. Papua New Guinea (1985) 105. — Rorippa officinalis (R.BR.) P.ROYEN, Mt. Fl. New Guinea 3 (1982) 2029. — Fig. 4. Perennial, practically glabrous herb. Stems juicy and hollow, prostrate to ascending, usually rooting at the lower nodes, richly branched, 10—100 cm long. Leaves petiolate, auriculate, pinnatisect, oblong in outline, up to 10 cm long; lateral leaflets in 2—9 pairs, sessile, narrowly obovate, elliptic or nearly or- bicular, entire to faintly dentate; terminal leaflet of similar shape, often somewhat larger. Racemes mostly terminal, ebracteate with numerous small white flowers, in fruit rather loose but short, usually c. 10 cm long; pedicels spreading to somewhat re- flexed, 8—12 mm long. Sepals green, elliptic, c. 2mm long. Petals white, often with a violet tinge, obovate, c. 4mm long. Anthers c. 0.7 mm long. Ovary nar- rowly ellipsoid, with short but distinct style. Siliqua broadly linear, often curved and torulose, 10—18 by 2—2.5 mm, 7—12 times as long as broad, with seeds arranged in two distinct rows; style 0.8—1.8 by c. 0.5 mm, with a slightly expanded stigma; valves rather thin, bulging when seeds are ripe. Seeds shining, dark red-brown, nearly orbicular in outline, 1—1.3 mm long, covered by a distinct rather high reticulum, which forms 25—SO areoles at each side of the seed. Distr. Native in Europe and West Asia, perhaps also in Ethiopia; widely cultivated (‘watercress’) and spread with European settlements to temperate and montane areas throughout the world; in Malesia: Java (especially West, rarer in Central and East), Lesser Sunda Islands (Flores), Borneo (Sabah), Philippines (Luzon), Papua New Guinea. Probably naturalized in many more areas than so far recorded. Ecol. Along streams, ditches and all sorts of run- ning water, on shores or floating in shallow water; c. 350—2500 m. Forms often large mats which very rarely produce flowers. 2. Rorippa backeri (O.E.ScHULZ) JONSELL, Bot. Notis. 132 (1979) 536. — Nasturtium backeri O.E.Scuutz, Notizbl. Bot. Gart. Berlin 9 (1925) 84; BACKER & BAKH./. Fl. Java 1 (1963) 191; STEEN. Mt. Fl. Java (1972) pl. 12/7. 1988] Annual or short-lived perennial herb with taproot, glabrous, 30—150 cm high. Stems erect, usually soli- tary, unbranched or with few erecto-patent bran- ches. Basal and lower cauline /eaves originally nu- merous (in older specimens lacking), 40—150 by 10—30 mn, petiolate, lyrato-pinnate with 1—4 pairs of oblong 5—12 by 2—10 mm lateral lobes; terminal lobe elliptic to ovate, 20—35 by 9-30 mm; cauline leaves numerous, 3—10 by 0.7—2 cm, lyrato- pinnatifid with long petioles, auriculate, with usually one pair of side-lobes; terminal lobe 1.5—5 by 0.7—2 cm; margins serrate to distantly crenate. Racemes ebracteate, terminal, sometimes numerous from up- per branches, in flower much condensed with 30—70 flowers, in fruit up to 80 cm long; pedicels divaricate at + 90°, straight, stout, 5-8 mm long. Sepals oblong, 2.5—4 mm long. Petals white, broadly spathulate, 3.5—7 mm long. Stamens 6; anthers 0.6—0.8 mm long. Ovary narrowly cylindrical with c. 0.5 mm long style. Sil/iquae linear, straight or fairly curved, 30—50 by 1.7—2.2 mm; valves rather firm; style 0.8—1.2 by c. 0.4 mm; stigma flat, slightly broader than style. Seeds dark brown, broadly oblong to subspheroid in outline, 1.2—1.4 by 0.9-1.1 mm, regularly reticulate-foveolate. Distr. Malesia: Java (eastern part: Mts Merba- bu, Wilis, Jang & Idjen: Suket). Ecol. In burnt forests, thickets, locally common; 1600—3000 m. Fi. fr. throughout the year. 3. Rorippa peekelii (O.E.ScHULZ) P.ROoYEN, Mt. Fl. New Guinea 3 (1982) 2032. — Nasturtium peekelii O.E.Scuut1z, Bot. Jahrb. 55 (1918) 270, f. 1D; von Ma. in Fedde, Rep. 41 (1937) 295; Scuutz in E. & P. Nat. Pfl. Fam. ed. 2, 17b (1936) 555, f. 337D; VELDKAMP, Blumea 28 (1982) 166. — Nasturtium novo-guineense Gitul, Ann. Naturhist. Mus. Wien 83 (1980) 429. — Fig. 5. Annual to shortlived perennial herb with taproot, glabrous, 20-60 cm high. Stems one to few, + straight, spreading, sometimes prostrate-ascending, usually with few erecto-patent branches from the lower parts. Basal /eaves mostly numerous, in a + dense rosette, 4-13 by 1—5 cm, petiolate, lyrato- pinnate with 1—3 pairs of oblong, lateral lobes, 0.4—2.5 by 0.2—1.5 cm; terminal lobe elliptic-ovate, 1.5—5 by 1—3 cm; margins sinuate to irregularly ser- rate; cauline leaves similar but smaller, the upper- most without side-lobes, petiolate, without or with rather small auricles, sometimes in axillary rosettes; margins crenate to acutely serrate. Racemes ebrac- teate, terminal, in flower very condensed with 20—50 flowers, in fruit very prolonged, up to 30 cm long; pedicels divaricate at 60—90°, straight, rather thin, 4-13 mm long. Sepals elliptic-oblong, 2—3.7 mm long. Petals white, broadly spathulate, 3-4 mm long. Stamens 6; anthers c. 0.7 mm long. Ovary nar- CRUCIFERAE (Jonsell) 557 rowly cylindrical with c. 0.3 mm long style. Siliquae linear, straight, 23—30 by 1.4—2 mm; valves rather firm; style 0.7—2.5 by 0.4—0.5 mm; stigma + flat, slightly broader than style. Seeds red-brown, subglo- bose to oblong, very finely reticulate-foveolate, 1.1—1.3 by 0.9—1.1 mm. Distr. Malesia: E. Timor (Mts Mutis & Tatamai- lau), Papua New Guinea (W., E. & S. Highlands), Bismarck Archipelago (New Ireland). Ecol. Along streams on rocks and beaches, a weed in gardens and along damp tracks. In New Guinea 1350—3500 m altitude, in Timor 1600—2850 m. 4. Rorippa palustris (L.) Bess. Enum. Pl. Volhyniae (1822) 27; JonseLL, Symb. Bot. Upsal. 19? (1968) 157. — Sisymbrium amphibium L. var. palustre LINNE, Sp. Pl. (1753) 657. — Nasturtium palustre (L.) DC. Syst. Nat. 2 (1821) 121; Biume, Bijdr. (1825) 50; Mig. Fl. Ind. Bat. 1, 2 (1858) 93; BACKER, Schoolfl. Java (1911) 55. — Rorippa islandica (OED.) Bors. Bal. Fl. (1900) 392; BACKER & BAKH./. Fl. Java 1 (1963) 190. — Nasturtium homalosper- mum O.E.SCHULZ, Bot. Jahrb. 55 (1918) 270, f. 1C, incl. var. macrocarpum; in E. & P. Nat. Pfl. Fam. ed. 2, 17b (1936) 551, f. 337C. — Nasturtium in- dicum auct. non DC.: K.ScH. & Laut. Fl. Deut. Schutzgeb. Stidsee (1901) 334, p.p. Annual or shortlived perennial herb with taproot, glabrous, 10-110 cm high. Stems erect, solitary or a few together, unbranched or branched in upper parts. Basal /eaves lyrate-pinnatisect, usually evanes- cent; cauline leaves few to numerous, 2.5—12 by 0.7—3 cm, lyrato-pinnatisect with 2—6 pairs of lan- ceolate lateral lobes, shortly to indistinctly petiolate, auriculate; terminal lobe 1/3 to 1/2 of total leaf length; margin irregularly serrate-crenate. Racemes ebracteate, terminal, often branched, and from up- per leaf axils, in flower much condensed with numer- ous flowers, in fruit up to 50 cm long and rather dense; pedicels somewhat curved, patent to deflexed, 3.5—7 mm long. Sepals oblong, 1.6—2.4 mm long. Petals pale yellow, spathulate, 1.6—2.6 mm long. Anthers 0.3—0.6 mm long. Ovary cylindrical with distinct style and not or slightly expanded stigma. Si- liquae sausage-like, often curved, 5-12 by 1.7—3 mm; valves rather firm; style 0.4—1 by c. 0.3 mm with not or slightly broader stigma. Seeds brown to light brown, spheroid, 0.6—0.9 mm, finely collicu- late. Distr. Over the whole north temperate area; widespread in many montane regions in subtropics and tropics; introduced to south temperate areas and elsewhere. In Malesia: Java (W., E.), Lesser Sunda Is. (Alor), Moluccas (Tanimbar), New Guinea (West: Vogelkop; East: Sepik, S. Highlands). Ecol. Moist or marshy places, ditches and other 558 FLORA MALESIANA [ser. I, vol. 103 Fig. 5. Rorippa peekelii (O.E.Scuutz) P.ROYEN. a. Habit, x 0.7; b. fruiting raceme, x 0.7; c. siliqua, x2; d. seed, scale bar 1 mm (a Brass 30519, b Savers NGF 213 16, c, d WOMERSLEY NGF 14272). Del. K.G. Forss. 1988] kinds of disturbed moist ground; from near sea-level to c. 2100 m. 5. Rorippa hybosperma (O.E.ScHULZ) JONSELL, comb. nov. — Nasturtium hybospermum O.E. SCHULZ, Bot. Jahrb. 55 (1918) 268, f. 1B; in E. & P. Nat. Pfl. Fam. ed. 2, 17b (1936) 551, f. 337b. — Nas- turtium indicum auct. non DC.: K.ScH. & HOLiR. Fl. Kaiser Wilh. Land (1889) 49; Wars. Bot. Jahrb. 13 (1891) 317; K.Scu. & Laut. Fl. Deut. Schutzgeb. Stidsee (1901) 334, p.p. Annual herb with taproot, glabrous, 50—150 cm high. Stem solitary, erect or in basal part ascending, shining, mostly rather stout, with many long erecto- patent branches. Leaves petiolate, auriculate, 3—17 by 1—8 cm, lyrato-pinnate with 1—3 pairs of oblong- elliptic lateral lobes, 0.8—4 by 0.3—1.5 cm; terminal lobe elliptic-ovate, 1.5—7 by 0.8—2.5 cm; margins crenulate-serrulate. Racemes terminal on stem and side-branches, ebracteate or with a few bracts in the lowest part, in flowers very condensed, in fruit rather loose, sometimes branched, up to 25 cm long; pedi- cels patent, the lower somewhat curved, S—10 mm long. Sepals elliptic, 1.5—1.8 mm long. Petals yellow, spathulate, 2—2.5 mm long. Stamens 6; an- thers c. 0.5 mm long; filaments narrowly subulate. Ovary cylindrical with a c. 0.3 mm long style. Sili- quae sausage-shaped, often curved, 12—17 by 1.5—2.7 mm; valves thin; style 0.9—1.5 by 0.2—0.3 mm, with a broader stigma. Seeds yellow brown, spheroid, finely colliculate and with prominent ir- regular warts around the margin, c. 1.2 mm long. Distr. Malesia: Philippines (N. Mindanao: Dulawan), New Guinea (West; East: Nuru, Sepik, Lae). Ecol. Along streams and rivers and in marshes in the lowland. Also a weed in moist places. 6. Rorippa schlechteri (O.E.ScHULZ) P.RoYEN, Mt. Fl. New Guinea 3 (1982) 2031. — Nasturtium schlechteri O.E.Scuutz, Bot. Jahrb. 55 (1918) 266, f. 1A;inE. & P. Nat. Pfl. Fam. ed. 2, 17b (1936) 554, f. 337A; VeLpKaAmp, Blumea 28 (1982) 166. — Trochiscus macrocarpus Gitu1, Ann. Naturhist. Mus. Wien 83 (1980) 430. Annual to short-lived perennial with taproot, 15—50 cm high. Stem solitary, erect, unbranched or with rather few, comparatively long branches. Leaves indistinctly petiolate, auriculate, 1.5—15 by 0.9—3.5 cm, lyrato-pinnate with 1—3 pairs of oblong lateral lobes, 0.2—1.5 by 0.1—0.6 mm; terminal lobe elliptic-ovate, 0.8—4 by 0.3—2.5 mm; margins ir- regularly dentate. Racemes terminal and on side- branches, ebracteate, in flower very condensed, in fruit rather loose, up to 15 cm long; pedicels patent to reflexed, 3—~10 mm long. Sepals oblong, 1.5—2 mm long. Petals yellow, spathulate, 1.7—-2.5 mm CRUCIFERAE (Jonsell) 559 long. Stamens 6; anthers c. 0.5 mm long; filaments linear. Ovary ellipsoid with c. 0.2 mm long style. Sili- quae inflated, semiglobose to broadly ellipsoid, S—9 by 3—6 mm; valves thin; style 0.6—0.9 by c. 0.3 mm, with flat, slightly broader stigma. Seeds red-brown, spheroid, 0.8—1 mm, minutely foveolate. Distr. Malesia: Papua New Guinea. Ecol. Moist or marshy places, various kinds of disturbed moist ground; c. 200—2600 m. Vern. Heleki, Okapa, heyenka, Tairora, kemu kembili, Kaugel. 7. Rorippa heterophylla (BLUME) WILLIAMS, FI. Tri- nidad & Tobago 1 (1929) 24. — Nasturtium hetero- phyllum Buume, Bijdr. (1825) 50; Mia. Ill. Arch. Ind. (1871) pl. IX; Boeri. Handl. Fl. Ned. Ind. 1, 1 (1890) 58; BAcKErR, Fl. Batavia (1907) 47; Schoolfl. Java (1911) 55; Koorp. Exk. Fl. Java 2 (1912) 288; HeEyYNE, Nutt. Pl. Ned. Ind. (1927) 680; OcHsE & Baku. Ind. Groenten (1931) 175, f. 107; Burk. Dict. (1935) 1534. — Cardamine sublyrata Mig. Ann. Mus. Bot. Lugd.-Bat. 2 (1848) 178. — Nasturtium indicum (L.) DC. var. javana BLUME, Bijdr. (1825) 50; Mia. Fl. Ind. Bat. 1, 2 (1858) 93. — Nasturtium diffusum auct. non DC.: Mig. Fl. Ind. Bat. 1, 2 (1858) 93; Boer. Handl. Fl. Ned. Ind. 1, 1 (1890) 58. — Nasturtium indicum auct. non (L.) DC.: Boer. Handl. Fl. Ned. Ind. 1, 1 (1890) 58; BACKER, Fl. Batavia (1907) 47; Schoolfl. Java (1911) 55; Merr. Fl. Manila (1912) 213; Koorp. Exk. Fl. Java 2 (1912) 289; RrpLey, Trans. Linn. Soc. Lond. Il, Bot. 9 (1916) 16; Merr. Interpr. Rumph. Herb. Amb. (1917) 240; Ripiey, Fl. Mal. Pen. 1 (1922) 119; Merk. En. Philip. 2 (1923) 208; Koorp. FI. Tjibodas 3 (1923) 80; Hocur. Candollea 2 (1925) 370; HEYNE, Nutt. Pl. Ned. Ind. (1927) 680; Burk. Dict. (1935) 1534; H.J.LAM, Blumea 5 (1945) 570; Quis. Medic. Pl. Philip. (1951) 335; BACKER & BAkHu/. Fl. Java 1 (1963) 190. — R. indica (L.) HteRN var. apetala (Lour.) Hocur. Candollea 2 (1925) 370. — R. dubia (Pers.) HARA, J. Jap. Bot. 30 (1955) 196; BACKER & Baku./. Fl. Java 1 (1963) 190. Annual herb with taproot, nearly glabrous. Stems erect or ascending, one or more from the base, 5—50 cm high, not or little branched. Leaves in a rosette and along the stem, the lower 1.5—8 cm long with 8—60 mm long petioles, narrowly obovate in outline, undivided or lyrate-pinnatipartite with 1—3 pairs of small lateral lobes; terminal lobe ovate, 1—7 by 0.8—4 cm, dentate, obtuse, basally cuneate or trun- cate; upper leaves rather few, undivided, narrowly ovate. Racemes terminal, and on short patent side- branches, ebracteate, with many small, very con- densed flowers, in fruit usually rather dense, 2—10 cm long; pedicels spreading, straight, 2—8 mm long. Sepals green, elliptic, c. 2 mm long. Petals often re- duced or lacking, sometimes up to 3.2 mm long, nar- 560 rowly spathulate. Stamens up to 3 mm long with c. 0.7 mm long anthers. Ovary narrowly cylindrical, with very short style and flat expanded stigma. Sili- qua linear, straight, 14—25 by 0.7—1.3 mm; valves rather thin; style narrow, 0.5—1.3 by c. 0.3 mm; stigma broader than style. Seeds uniseriate, red- brown, roundedly quadrangular in outline, c. 0.7 mm long, very finely colliculate. Distr. Indigenous in East Asia, at least from Japan to Burma; introduced in other tropical areas, also in Africa and America; in Malesia: Sumatra, Malay Peninsula, Java, Lesser Sunda Islands (Timor), Philippines (Luzon, Mindanao), Moluccas (Ambon, Buru, Ceram), Celebes, Papua New Guinea. Ecol. Open, moist ground, often cultivated places; 250—1500 m. 8. Rorippa benghalensis (DC.) Hara, J. Jap. Bot. 49 (1974) 132. — Nasturtium benghalense DC. Syst. Nat. 2 (1821) 198; ScHuLz in Fedde, Rep. 33 (1934) 281. Annual, pubescent herb, with taproot. Stems erect or ascending, up to 50 cm high, branched at least in upper parts. Leaves obovate in outline, auriculate, lyrato-pinnatipartite, irregularly serrate-dentate, up to 15 by 3 cm, successively transgrading into smaller undivided bracts; terminal lobe ovate, 1.5—5S by 1—3 cm; lateral lobes in 1—4 pairs. Racemes terminal or axillary, at least in lower and middle parts bracteate, with numerous small light yellow flowers, in fruit rather dense, up to 7 cm long; pedicels spreading, straight, 3—7 mm long. Sepals green, elliptic, c. 1.5 mm long. Petals pale yellow, spathulate, 1-2 mm long, about as long as the sepals. Anthers c. 0.5 mm long. Ovary cylindrical with very short style and flat expanded stigma. Si/iqua linear, straight, 6—17 by 1—1.5 mm; valves rather thin; style 0.3—0.8 by c. 0.4 mm; stigma flat, broader than style. Seeds biseriate, FLORA MALESIANA [ser. I, vol. 103 dark brown, spheroid, c. 0.5 mm long, minutely fo- veolate. Distr. India to Vietnam; in Malesia: Java, Lesser Sunda Islands (Sumbawa). Ecol. Weed in settled areas, perhaps only inci- dental. 9. Rorippa micrantha (ROTH) JONSELL, Svensk Bot. Tidskr. 68 (1974) 384; Fl. Trop. E. Afr., Cruciferae (1982) 55, f. 19. — Sisymbrium micranthum RortH, Nov. Pl. Sp. (1821) 324. Annual herb with taproot. Stems erect or ascend- ing, up to 70 cm high but usually much shorter, branched at least in upper parts, somewhat pubes- cent especially basally. Basal /eaves lyrate-pinnati- partite in an evanescent rosette; cauline leaves petio- late, auriculate, nearly amplexicaul, broadly oblong- elliptic in outline, imparipinnatisect to lyrate-pinna- tipartite; lateral lobes usually in 2—S pairs, sessile, el- liptic to lanceolate, serrate, up to 3cm long. Racemes terminal and axillary, ebracteate, with numerous small yellowish flowers, in fruit rather loose, up to 20 cm long; pedicels spreading, straight, 2.5—5 mm long. Sepals light to yellowish green, elliptic, 1.5—2.5 mm long. Petals pale yellow, sometimes with a violet tinge, spathulate, 2—3 mm long, usually longer than the sepals. Anthers c. 0.7 mm long. Ovary ellipsoid to cylindrical, with very short style and flat expanded stigma. Siliqua linear, straight, 14—25 by 1.8—2.5 mm; valves rather firm; style 1—2.2 by 0.5—0.8 mm; stigma broader than style and slightly bifid. Seeds red-brown, finely orbicular to broadly elliptic in out- line, 0.5—0.8 by 0.4—-0.8 mm, ffinely reticu- late-foveolate. Distr. India, widespread in Africa from Egypt to Zaire and South Africa, also in Madagascar; in Malesia: collected a few times in E. Java (Pasuruan, Besuki, efc.). Ecol. Weed in settled areas, perhaps only inci- dental. MAGNOLIACEAE (H.P. Nooteboom, Leiden) Trees or shrubs, glabrous or with an indumentum of single hairs. Leaves spirally arranged, simple, entire or 2—10-lobed, penninerved, evergreen or deciduous; stipules present, at first enclosing and protecting the innovations, later caducous and leaving an annular scar around the node. Flowers terminal or pseudo- axillary on a short shoot in the axils of the leaves, bisexual, rarely unisexual, pedunculate. Peduncle bearing 1 or more caducous spathaceous bracts which leave annular scars. Perianth spiral or spirocyclic, simple or differentiated in calyx and corolla, perianth members 6 or more, free, imbricate. Stamens numerous, free, spirally arranged; filaments short or more or less elongated; an- thers linear, 2-locular, dehiscing introrsely, latrorsely or rarely extrorsely; con- nective usually more or less produced into an appendage. Gynoecium sessile or stipitate (a gynophore present); carpels numerous to few (rarely one), spirally arranged (except in Pachylarnax), free or sometimes concrescent; ovules 2 or more, biseriate on the ventral suture. Fruit apocarpous, sometimes syncarpous; fruiting carpels opening along the dorsal and/or ventral suture, or circum- scissile, rarely indehiscent. Seed(s) 1 or more in each fruiting carpel, large, in dehiscent carpels hanging from the elongated spiral vessels of the funiculus, with arilloid testa, rarely, when fruit indehiscent adherent to the endocarp; endosperm copious, oily; embryo minute. Distribution. Seven genera in temperate and tropical SE. and E. Asia and from North America southward through the West Indies and Central America to S. Brazil. Ecology. Several of the tropical species are found at low altitudes, many grow at medium and high altitudes, in Malesia up to c. 3500 m. Pollination. — THIEN (1974) studied the floral morphology of North American Magnoliaceae. They are all protogynous. The flowers are pollinated by beetles which enter at all stages from large bud to open flower and feed on stigmas, pollen, nectar, and the secretions of the petals. They become covered with pollen when feeding. Because the pollen is shed after the stigmas cease to be receptive, the first flowers that open often are not pollinated. There are indications that if all the flowers blossom in one flush, as rarely happens after a severe winter, no seed is set (TRESEDER, 1978). The timing of the flowers, including opening and closing of petals, the appression of stig- mas to the gynoecium (in some species), large quantities of food, and protogyny all suggest that the flowers are highly specialised for exclusive pollination by beetles. Despite the sharing of polli- nators few hybrids have been found in nature. Not enough is known about isolating mechanisms in Magnolia to explain fully the lack of natural hybrids. Between several species artificial hybrids easily can be obtained (THIEN, 1974). References: Tuten, Amer. J. Bot. 61 (1974) 1037-1045; Treseper, Magnolias (1978). Morphology. — Vegetative morphology. All Magnoliaceae are trees or shrubs with decid- uous or persistent stipulate leaves. The stipules are first connate, thus forming a cap for the ter- minal bud and then rupture longitudinally and fall, leaving a circular scar round the twig. They can be free from the petiole or (partly) adnate to it. Growth of the twigs often is faster in the begin- ning of the season, thus first producing longer internodes than later. In tropical species, however, there may be no conspicuous difference in length of internodes throughout the twig. In Lirioden- droideae and in tribe Magnolieae of Magnolioideae growth is sympodial, and when a flower ter- minates a twig, the axillary bud of the uppermost leaf or a lower leaf gives rise to the next vegeta- tive shoot which in turn eventually will be terminated by a flower. In tribe Michelieae growth is monopodial, the flowers grow from the axil of a leaf and are mostly borne on a brachyblast that consists of two or several internodes. (561) 562 FLORA MALESIANA [ser. I, vol. 10° Flower structure and floral anatomy. — The flowers are borne on a pedicel which mostly is the ultimate internode of a brachyblast (axillary in E/merrillia and Michelia; terminal in the other genera). The flower bud is often protected by the stipules of the uppermost leaf, especially in the genera with terminal flowers. In temperate species these stipules often are leathery and/or hairy. The brachyblast, if present, bears one to several spathaceous bracts which also surround and pro- tect the flower bud. Essentially these bracts consist of a petiole (reduced lamina) with its stipules adnate to it, together forming a cap like the one surrounding the vegetative buds. The uppermost spathaceous bract, sometimes called bracteole, surrounds the base of the pedicel. This pedicel can be very short or (nearly) absent or long. The perianth is either spiral throughout, such as in Mag- nolia stellata, or consists of some ternate whorls. Ontogenetic studies have shown that the latter are initiated successively, although they form two tiers. Thus there is a clear transition from the spiral phyllotaxy to a whorled phyllotaxy in perianths of the Magnoliaceae (TUCKER, 1960; ERBAR & LEINS, 1982, 1983). While there is often no clear differentiation of calyx and corolla such a sep- aration may exist in some cases, e.g. in Liriodendron, although UEDA (1986) argues for the regular presence of a true calyx. Vascular system. — Two vascular systems, a stelar and a cortical system of vascular bundles, are present in the floral axis of the Magnoliaceae (Sk1PWORTH, 1970). The cortical system arises below the flower, either directly from the stele or as branches of leaf and stipule traces. In general it provides the lateral traces of the flower appendages, while the stelar system gives rise to the me- dian traces. This is true of bracts, perianth members, stamens, and carpels. The stamens are spirally arranged and develop either directly after the tepals and in the same 8 (7—10) parastichies like in Magnolia stellata (ERBAR & LEINS, 1982), or after a relatively long interval like in Magnolia denudata (l.c.) and Michelia fuscata (TUCKER, 1960). In Magnolia denudata the spiral of the stamens is somewhat irregular. A number of tropical species has broad 3-veined stamens with two pairs of linear sporangia which are deeply embedded in their surface. They are comparable to those of the Degeneriaceae and Himantandraceae. CANRIGHT (1952) describes several trends of specialisation in the stamens of the Magnoliaceae such as: (1) elongation of the apices; (2) differentiation of a filament; (3) re- duction in number of veins from three to one; and (4) the transition from a laminal to a marginal position of the sporangia. Teratological stamens often occur, which range from broad petaloid stamens to narrow filamented ones. These different forms were found in the same flower of Michelia champaca. Also carpels with varying degrees of fertility were found among the stamens. The carpels are arranged spirally on the lengthened receptacle. Their phyllotaxy was studied in Magnolia stellata (ERBAR & LEINS, 1982) and Michelia fuscata (TUCKER, 1961). They arise in the same spirals as stamens and tepals, i.e. 8 (sometimes 7, 9) parastichies. In Michelia different divergence angles were found to occur in different flowers. In contrast to the more primitive Ranalean families, the carpels of the Magnoliaceae and the related Himantandraceae (CANRIGHT, 1960) appear strongly advanced. The most significant specialisations are: (1) the closure of the fertile part of the carpel; (2) the restriction of the external stigmatic surfaces to the margins of the conduplicate style; and (3) conspicuous changes in the vascularisation of the carpels. There is also a tendency towards a reduction in the number of ovules (to two in many Magnoliaceae) and to lateral adherence or actual concrescence of crowded carpels. In summary the carpels of Magnoliaceae exhibit more or less advanced stages of phylo- genetic modification. The carpels, like the stamens, are supplied by three veins. The dorsal traces commonly originate from the stelar system while the two ventral traces originate from the cortical system. CANRIGHT (1960), however, found in cleared material from Michelia champaca that ‘not uncommonly, the basal carpels of a gynoecium are vascularized entirely by the cortical system, and the apical carpels entirely by the stelar system.’ VAN HEEL (1981, 1983) suggests that the carpels are basically cupuliform. His SEM studies show that carpels arise as cup-shaped (ascidiate) or partly cup-shaped organs lateral or terminal on the flower apex. According to CANRIGHT the margins of the carpels become adnate to the floral 1988] MAGNOLIACEAE (Nooteboom) 563 axis. According to VAN HEEL’s observations there is no later fusion, but already in an early state there is no abaxial margin present. On the side of the floral axis the cup is deepened secondarily, and the ovules originate in it on the inside margin close to the floral axis when an ovary and a style have not yet differentiated. Fruits. — The fruits of Magnolioideae principally consist of several to many (in Michelia mon- tana sometimes one) free carpels spirally arranged around the receptacle. The fruiting carpels are woody and entirely free in some taxa while they are, secondarily, more or less connate in others. The young carpels are always free when initiated, and only relatively late in ontogeny they may become concrescent. Dehiscence of the mature carpels is mostly along the dorsal suture, often also along the ventral suture. Sometimes the carpels become bivalved, the 2 valves only adnate to the central axis, shedding their seeds. Sometimes the carpels form a pseudosyncarp, becoming more or less concrescent. But only the outer layers of the exocarpium are really concrescent, giving the fruit the appearance of a syncarp. In the latter case the dorsal (abaxial) parts of the carpels finally fall away leaving their base, which is inbedded in the receptacle, exposed with the seeds mostly hanging from their placenta on the elongate funicle which in that stage consists of spirally thick- ened vessels. In some taxa the abaxial parts of the carpels when falling also dehisce along the dor- sal suture (‘7soongiodendron’, but also in some ‘Talaumas’ and in ‘Manglietiastrum’). In Mag- nolia nitida and M. kachirachirai the carpels are essentially connate. When maturing they tear apart and dehisce along the dorsal suture thus giving the appearance of acommon Magnolia fruit. This condition probably exists in many species with crowded carpels. The concrescence of the car- pels has apparently developed independently in different lineages of Magnoliaceae. Species with free and concrescent carpels sometimes are very closely allied, judging from the other characters. In Liriodendron the fruits are free, indehiscent, samaroid (produced at the apex into a wing-like beak) and, contrary to Magnolioideae, caducous. Seed. — In Magnolioideae the seeds are exposed, hanging from the dehiscent carpels. In Lirio- dendron the testa is adherent to the endocarp and the seed falls with the fruit. The seed coat is organised chiefly from the outer integument while the inner integument is represented by a layer of crushed cells. In a ripe seed the testa consists of an outer fleshy region (sarcotesta) comprising the epidermis of the outer integument, 2 or 3 layers of tangentially elongated cells, a 10—12 layered fleshy zone, 2 or 3 layers of tangentially compressed cells, and an inner stony region of 3 or 4 layers of lignified cells. In Liriodendron no sarcotesta is formed. The same tissue remains thin-layered and becomes partly sclerotic. The differences in seed coat structure between Magnolia and Liriodendron are clearly associated with differences in seed dispersal (BOUMAN, 1977). The seed has such primitive features as the multiplicative sarcotesta, stomata, ligneous sclerotesta (endotesta sensu CORNER), massive chalaza, copious endosperm, and minute embryo. It is advanced in the apparently functionless nature of the tegmen, perhaps in loss of rumination, and in the substitution of the aril by the sarcotesta. The specialities of the seed of Magnolioideae are the pink pellicle of the sarcotesta, the nature of the cells composing the multiple endotesta, and the tubular lignified sheath around the chalaza. The lignified cells of the sclerotesta are pecu- liar and need study by electron microscopy. A system of lignified filaments pervades each cell as if the endoreticulum were lignified (CORNER, 1976). TiFFNEY (1977) studied the sclerotesta of many living and fossil species and found that it is possible to recognise morphological distinctions between the genera and in many cases between species within a genus. Embryology. — The anther wall at the microspore mother cell stage comprises epidermis, endo- thecium, 2—4 middle layers, and a bilateral glandular tapetum. By the time the cytokinesis is com- pleted in the microspore mother cells, a large number of Ubisch granules line the inner walls of the tapetum. In a fully mature anther the papillate epidermis and endothecium along with 2 or 3 middle layers persist. After meiosis II in the microspore mother cell the cytokinesis takes place by furrowing, resulting in tetrahedral or isobilateral tetrads. The mature pollen is shed at the 2-celled stage. The generative cell is surrounded by a thin sheath of finely granular cytoplasm and a delicate membrane. 564 FLORA MALESIANA [ser. I, vol. 103 The ovules are anatropous, bitegmic, and crassinucellate; the outer integument is vascularised. The hypodermal archesporium is multicellular and of hypodermal origin although ultimately only one cell functions. The primary parietal cell divides repeatedly to form the parietal tissue so that the megaspore mother cell is buried deep in the nucellus. At the end of meiosis II linear or T- shaped megaspore tetrads are formed. The chalazal megaspore functions, giving rise to the Polygonum type of embryo sac. The synergids and antipodal cells are ephemeral. The endosperm is cellular from the beginning, and embryogeny conforms to the Myosurus variation of the Onagra type or is irregular (HAYASHI, 1964, 1966; BHANDARI, 1971; Davis, 1966). References: BHANDARI, J. Arn. Arb. 52 (1971) 1—40, 285—304; Bouman, Acta Bot. Neerl. 26 (1977) 213—223; CANRIGHT, Amer. J. Bot. 39 (1952) 484—497; ibid. 47 (1960) 145—155; CoRNER, Seed of Dicotyledons (1976) 177; Davis, Systematic embryology of the Angiosperms (1966) 167; ERBAR & LEINS, Beitr. Biol. Pflanzen 56 (1982) 225—241; Bot. Jahrb. 103 (1983) 433—449; HayasHl, Sci. Rep. Tohoku Univ. ser. IV Biol., 30 (1964) 89—98; ibid. 32 (1966) 111—118; vAN HEEL, Blumea 27 (1981) 499—522; ibid. 28 (1983) 231—270; SkipwortH, Phytomorphology 20 (1970) 228—235; TIFFNEY, Bot. J. Linn. Soc. 75 (1977) 299—323; TucKER, Amer. J. Bot. 47 (1960) 266—277; ibid. 48 (1961) 60—71; UEDA, Bot. Mag. Tokyo 99 (1986) 333-349. Anatomy. — Vegetative anatomy. The epidermal cells are irregular in form, less often poly- gonal and with sinuous, curved or nearly straight walls which are often thickened. Many members of the family have a modified subepidermal layer (hypodermis) beneath the upper epidermis, some have such a layer associated with both upper and lower epidermis, and few have it only with the lower epidermis. Many species lack specialised subepidermal cells entirely. Hairs may be present on both the upper and lower epidermis, or only on the lower. In some species the leaves lack hairs altogether. They are uniseriate, consisting of one to several or many cells. Several types of hair base have been distinguished by BARANOVA (1972), of which the most primitive occurs in Manglietia while the most advanced are found in El/merrillia, Michelia (Tsoon- giodendron), and in some scattered species of Magnolia. The stomata are paracytic in Magnolioideae (but in rare cases NONG VAN TieP, 1980: 519, found also anomocytic stomata in Manglietia) and paracytic as well as anomocytic in Liriodendroideae. They are confined to the lower surface. Significant thickenings on the walls of the epidermal and subsidiary cells, as well as strongly developed cuticular thickenings on the outer wall of the guard cell, occur mainly in the tropical sections of Magnolia subg. Magnolia as well as in many Asian species of Magnolia subg. Talauma. Foliar sclereids are present in certain taxa. They vary widely in form, size, wall thickness, pitting, and degree of ramification. They may be found in four tis- sues or cell assemblages: First, as idioblastic sclereids in the petiole and along the midrib, especial- ly in evergreen leaves; less frequently in the mesophyll, in dermal layers and in the vein sheath system including veinlet endings. Taxonomically the mentioned characters are of different value. Each genus shows a particular range of features, some of which can be used diagnostically to separate taxa above the genus level. The coriaceous texture of tropical leaves is achieved in various ways arising from the variable form and diverse distribution of the sclereids. Magnolia subg. Talauma species characteristically have the entire vein system of the leaf encased in sclerenchymatous sheaths, and the veinlets terminate in sclerified elements. Manglietia species in contrast have parenchymatous vein sheaths near the terminations and lack sclerified terminal cells, but most of the mesophyll and epidermal cells are sclerified. Manglietia leaves are characterized by sclerified epidermal and hypodermal layers, sclerified arm parenchyma in the mesophyll, unlignified lobate vein-sheath cells, and an absence of either sclerified veinlet terminal cells or a sclerified leaf margin. No other genus combines all those features, although some do show one or more. Sterile specimens of Manglietia can easily be recognized on the basis of cleared leaf segments. Magnolia subg. Magnolia and subg. Talauma and Michelia exhibit in many of their members sclerified terminal veinlet cells, thick sclerified leaf margins, and stellate sclereids along the mid- rib. Rarely those features may be diagnostic for species, but in most cases they are too common 1988] MAGNOLIACEAE (Nooteboom) 565 to be useful. The foliar characteristics of Magnolia subg. Talauma include veinlets terminating in thick-walled stellate sclereids and thinner walled tracheary elements. These cell types are typi- cally absent from temperate zone Magnolia sections and therefore are possibly an adaptation to climatic conditions. In all the Asian species of subg. Talauma examined there is a thick sclerified margin, usually including a vein. In the American species this is absent but in few of them a weak approach to this condition is seen. The massive marginal ridge helps to identify certain species of Magnolia subg. Talauma and subg. Magnolia, Manglietia (3 spp.), and Michelia (3 spp.). In Alcimandra, Liriodendron, ‘Paramichelia’, and most Elmerrillia species foliar sclereids are rare. This may indicate that these taxa have not developed vegetative modifications different from those of the related large genera (mainly after BARANOVA, 1972 and TUCKER, 1977). Wood anatomy (largely from H. GoTTWALpD, a manuscript of a lecture given for the Interna- tional Association of Wood Anatomists in Amsterdam, which he kindly put at my disposal). Wood anatomy is homogeneous throughout the family. The secondary xylem shows a well de- veloped fibrous ground tissue of thin to medium thick-walled fibres, with diameters from 15 to 25 um depending on the species. The vessels are mostly evenly distributed, except for the temper- ate species which show (semi-)ring-porous tendencies. Vessel diameter ranges from 50 to 180 um on average. The rays are nonstoried, mostly 3 cells wide and 0.5—1 mm high. The axial paren- chyma is mainly banded and marginal (bands 1—2(—12) cells wide). CHowpHuRY (1964) estab- lished the annual periodicity of these parenchyma bands, even in the tropical Michelia champaca. For other tropical species the periodicity of the parenchyma bands remains to be studied, and is probably not annual. Only in Magnolia subg. Talauma the bands tend to fork. The paratracheal parenchyma is usually scanty or in some species virtually absent. The parenchyma cells do not contain any kind of crystals, nor are there any phenolic substances found in the vessels. There are some striking anatomical details occurring only in groups of individual taxa. Firstly the primitive vessel pitting varying from purely scalariform to opposite. The vessel perforations are mostly scalariform with only | to 15 bars. In the advanced taxa Liriodendron and sect. Yulania of Magnolia also simple vessel perforations occur, always in connection with opposite pits. A rare feature is the occurrence of amorphous silica as a coating on the cell walls including the tyloses, or as solid occlusions completely occluding the lumina. Complete solid plugs of silicium dioxide occur in all species of sect. Blumiana of Magnolia subg. Talauma and in the monotypic sect. Lirianthe of Magnolia. The estimated volume percentage of silica is up to 8%, based on air-dried volume; sometimes almost every vessel is filled with silicium dioxide. This means that these taxa belong to the woody species with the highest silica content known in the vascular plants. Spiral thickenings on the vessel walls are a further special feature of limited occurrence in Magnoliaceae. They are only weakly pronounced and hard to detect. Some peculiar idioblastic parenchyma cells are found mainly between the marginal cells of the rays. They are thin-walled, have mostly amorphous brownish contents, and are often referred to as ‘oil cells’, although their chemical nature is still unknown. These enlarged cells are of irregular distribution so that their taxonomic value is limited. Silica bodies which occur in the ray cells of a few species in sect. Maingola of Magnolia and in sect. Blumiana of Magnolia subg. Talauma, point to their close relationship. Crystalline inclusions are found in fine wood-splits of Magnolia subg. Magnolia and subg. Talauma; they consist of calcium carbonate and are always of traumat- ic origin. The last special item concerns the occurrence of tyloses in the fibres, besides those in the vessels. This rare feature was observed in 24 species, belonging to 7 genera and subgenera. The homogeneity of characters enhances a mixture of overlapping structural details which is detrimental to the internal classification of the family. The largest genus, Magnolia (without subg. Talauma) includes almost all structural traits which are otherwise distributed over the remaining genera. There are only few taxa which can be distinguished anatomically such as the genus Lirioden- dron, which has neither spirals in the vessels nor tyloses in the fibres. Magnolia sect. Yulania is 566 FLORA MALESIANA [ser. I, vol. 10° characterized by simple perforations, and Magnolia subg. Talauma sect. Blumiana by its silica- bearing vessels and the pronounced heterogeneity of the rays. Magnolia subg. Talauma sect. Aro- madendron differs from the rest of subg. Talauma in that the pores are almost twice the size of the investigated Asian species of subg. Ta/lauma. They are also the largest in the whole family (CANRIGHT, 1955). Differences are also apparent in the distribution of wood parenchyma and of characteristics of fibre tracheids. In this respect E/merrillia falls within the range of Michelia whereas Kmeria is in- distinguishable from many tropical Magnolia species. There is little difference in the wood anatomy of Manglietia and many temperate Magnolia species. Although neither the primitive nor the advanced wood anatomical characters are confined to any one genus, there are many indica- tions that the woods of the temperate species of Magnolia and Liriodendron are the most special- ized. Magnolia subg. Talauma exhibits the largest assemblage of primitive characters. The tropical Magnolia paenetalauma from Hainan exhibits a great number of primitive features. Also in the external morphology the species of sect. Gwillimia, to which this species belongs, resemble the species of subg. Talauma so closely that they cannot be distinguished in absence of fruits. References: BARANOVA, Taxon 21 (1972) 447—469; CANRIGHT, J. Arn. Arb. 36 (1955) 119—140; CHOWDHuRY, J. Indian Bot. Soc. 43 (1964) 334—342; GorTwaLpb, Wood Sci. Technol. 6 (1972) 121—127; METCALFE & CHALK, Anatomy of the Dicotyledons ed. 2, 3 (1987) 24—33; NoNG VAN TrEP, Feddes Rep. 91 (1980) 497—576; TuckER, Bot. J. Linn. Soc. 75 (1977) 325—356. Chromosomes. The basic chromosome number is x = 19. Polyploidy has been demonstrated in Magnolia. In wild species diploidy as well as tetraploidy is found. TRESEDER (1978: 208) reports for cultivated Magnolias besides diploid, also triploid, tetraploid, pentaploid, hexaploid, hepta- ploid, and octoploid numbers, while aneuploid numbers also are recorded. The chromosomes are characteristically small, short rods (WHITAKER, 1933). The chromosome base number of all Angiosperms is 7, the higher numbers are derived from it by palaeopolyploidy (EHRENDORFER C.S., 1968). The same basic chromosome number of 19 is found in Myristicaceae (also 21) and Monimiaceae (also 22). References: EHRENDORFER C.S. Taxon 17 (1968) 337—468; TRESEDER, Magnolias (1978); WHITAKER, J. Arn. Arb. 14 (1933) 376—385. Phytochemistry. Chemical characters of the family were summarized and analyzed by HEGNAUER (1969). A general occurrence of essential oils deposited in large idioblasts and of alka- loids of the biosynthetic pathway resulting in the so-called benzyltetrahydroisoquinoline family of alkaloids comprising among others benzylisoquinolines, bisbenzylisoquinolines, aporphines, oxoaporphines, and protoberberines was stressed. Leaf phenolics were represented mainly by kaempferol, quercetin, rhamnetin, procyanidins, and caffeic acid. At the same time a total lack of flavonols with trihydroxylated B-ring, of ordinary flavones and of galli- and ellagitannins had been demonstrated. Other characters mentioned are strongly silicified leaves in many taxa, oil- rich seeds without starch, and a strong tendency to store cyclitols in Liriodendron (liriodendritol), Magnolia subg. Magnolia (pinitol) and subg. Talauma (quercitol). The sporadic occurrence of lignans (lirioresinol), neolignans (magnolol), coumarin glycosides (magnolioside), cyanogenic compounds (not identified), and of the sesquiterpene lactones costunolide and parthenolide was noted. The totality of known chemical characters was interpreted as being highly typical of Polycarpicae. In the meantime much additional phytochemical research was performed with members of the genera Alcimandra, Elmerrillia, Liriodendron, Magnolia subg. Magnolia and Talauma, and Michelia. In each instance alkaloids were isolated, some being new compounds and some already known from other members of Policarpicae. All, however, belong to the phenyltetrahydroiso- quinoline class. The oxoaporphine liriodendrine and the quaternary aporphine magnoflorine seem to be nearly ubiquitous. New features are the rather general occurrence of biologically active sesquiterpene lactones belonging to the germacranolide, eudesmanolide, elemanolide, and guaianolide groups of these constituents, and of lignans and neolignans (together called ligna- 1988] MAGNOLIACEAE (Nooteboom) 567 noids). Hitherto sesquiterpene lactones were isolated from species of Liriodendron, Magnolia, and Michelia. All species investigated for lignanoids were shown to contain such metabolites. To- day many individual compounds are known from members of Liriodendron, Magnolia subg. Magnolia and Talauma, and Michelia; they represent at least 12 different structural types. Cyano- genic compounds were detected in Liriodendron tulipifera (taxifolin and triglochinin) and in Magnolia sprengeri cv. ‘Diva’ (taxifolin). They belong to the tyrosine-derived class of cyanogens. From the taxonomic point of view the secondary metabolism of Magnoliaceae can be con- sidered as highly characteristic of Polycarpicae and closely related orders such as Piperales and Aristolochiales. They all have essential oils in idioblasts and comprise members synthesizing ben- zylisoquinolines. Similar lignanoids are presently known, among others, from Aristolochiaceae, Eupomatiaceae, Lauraceae, Myristicaceae, Piperaceae, and Trimeniaceae. Sesquiterpene lac- tones occur also in Chloranthaceae and Lauraceae. They form one of the arguments for the pro- position of an evolutionary line Polycarpicae — Rutales — Umbelliferales s.str. — Asterales. — R. HEGNAUER. Palynology. (After PRAGLOWSKI, 1974). The pollen grains of Magnoliaceae are 1-(ana)col- pate, bilateral, heteropolar. Aperture simple, with markedly thin, frequently slightly undulated margins. Colpus usually longer, occasionally as long as, or rarely shorter than the longest axis. Colpus invagination usually absent or insignificant. Exine structure in tectate grains (about 95%) consists of a continuous tectum perforatum supported by columellae which have no geometrical connection with the tectum or with the supratectal elements. In rarely occurring semitectate grains the exine structure is microreticulate, without sculpturing. Sculpturing rugulose or absent (tectum smooth). Sexine at the proximal face thicker than nexine, usually twice or more. Columellae mi- nute, usually indistinct. Elmerrillia pollen grains show partly similarity with pollen grains of Michelia, but the micro- reticulate structure of the grains of E. tsiampacca ssp. mollis is rather unique showing no resemblance to Michelia pollen. The reticulum is considerably coarser than that of Pachylarnax pollen grains. Michelia, Paramichelia, and Tsoongiodendron possess pollen grains that are very similar. Pachylarnax possesses pollen with microreticulate exine structure consisting of thin muri encom- passing very minute lumina. This rather peculiar exine structure is more delicate than that in other Magnoliaceae. Similarly, the rather symmetric ellipsoidal shape of the pollen makes it quite dif- ficult to include them among typical magnoliaceous pollen. Alcimandra pollen possesses an exine without sculpturing which shows sometimes resemblance with Talauma pollen. Magnolia pollen grains show a rather large morphological variation; nearly all the pollen types of subfamily Magnolioideae are found in Magnolia. Pollen grains of Elmer- rillia and Michelia show slight differences to that of Magnolia. Manglietia pollen grains show a high morphological similarity to those of Magnolia. Talauma pollen grains show features similar to those encountered in Magnolia subg. Magnolia. Aromadendron pollen grains show similarity to Alcimandra pollen grains, but also with those of Magnolia sect. Maingola. On palynological evidence the joining of Michelia, Tsoongiodendron, and Paramichelia is strongly supported. The same holds for Manglietia, Magnolia, and Talauma, while the joining of Aromadendron with Magnolia is not contradicted. Reference: PRaGLowski, World Pollen and Spore Flora 3 (1974). The family can easily be divided into two subfamilies, Magnolioideae and Liriodendroideae, the latter not in Malesia. 568 FLORA MALESIANA [ser. I, vol. 10° SUBFAMILY MAGNOLIOIDEAE Leaves entire or occasionally two-lobed at the apex; stipules free from the petiole or adnate to it. Anthers introrse or latrorse. Fruiting carpels longitudin- ally dehiscent or circumscissile, at least the base remaining adnate to the torus, free or concrescent into a syncarp. Testa free from the endocarp, externally aril- loid. In Magnolioideae two tribes can be recognized clearly. KEY TO THE TRIBES 1. Growth sympodial. Flower buds arising terminal on the twigs................. 1. Tribus Magnolieae 1. Growth monopodial. Flower buds arising on brachyblasts in the axils of the leaves 2. Tribus Michelieae 1. Tribus Magnolieae Growth sympodial. Flower buds arising terminal on the twigs, the latter growing in length from an axillary bud of one of the upper leaves. Genera: Magnolia (incl. Talauma, Alcimandra, Manglietiastrum and Aromadendron), Maneglie- tia, Kmeria (not in Malesia), and Pachylarnax. KEY, TO THRE GENERA 1. Fruit consisting of few to many separate free or connate carpels along the torus. 2. Ovules 4 or more in each carpel. The hair base replaces a normal epidermal cell, so that the loss of a hair leaves a pore in the cutical membrane. Leaf anatomical characters: sclerified epidermal and hypodermal layers, sclerified arm parenchyma in the mesophyll, unlignified vein-sheath cells, absence of either scleri- HeEGAVEIMlen tenmInal ices OF a SClerined leah MALOU ce oo onus sioueo os ordre oyna mp2 ofS es 2. Manglietia 2. Ovules 2 in each carpel (4 in Magnolia kachirachirai), sometimes 4 in the lower carpels. The hair base con- sists of at least two epidermal cells. The hair does not leave a pore when falling........ 1. Magnolia 1. Fruit a woody loculicidal capsule composed of few (2—8) concrescent carpels. Tepals 9—15, subequal. Ovules about 4—8 in each carpel. Stipules free from the petiole..................... 3. Pachylarnax 1. MAGNOLIA LINNE, Sp. Pl. (1753) 535; Gen. Pl. ed. 5 (1754) 240; Danby, Kew Bull. (1927) 259; in Hutch. Gen. FI. Pl. 1 (1964) 55; in Treseder, Magnolias (1978) 29; Noor. Blumea 31 (1985) 83; ibid. 32 (1987) 343. — Type: M. virginiana LINNE, E. United States. Talauma Juss. Gen. PI. (1789) 281; Danby, Kew Bull. (1927) 259; in Hutch. Gen. Fl. Pl. 1 (1964) 55. — Magnolia sect. Talauma BatLLon, Adansonia 7 (1866) 3, 66; Noor. Blumea 31 (1985) 83. — Magnolia subg. Talauma PIERRE, Fl. For. Cochinch. 1 (1881) sub t. 1. — Violaria Post & O. K. Lexic. Gen. PI. (1903) 588, p.p. — Type: T. plumierii (SCHWARTZ) A.DC. (Magnolia plumierii SCHWARTZ). Aromadendron BuiuMgE, Bijdr. (1825) 10; Fl. Java Magnol. (1829) 25 (‘Aro- madendrum’); DANDY, Kew Bull. (1927) 259; in Hutch. Gen. FI. Pl. 1 (1964) 55. 1988] MAGNOLIACEAE (Nooteboom) 569 — Violaria Post & O. K. Lexic. Gen. Pl. (1903) 588, p.p. — Talauma sect. Aromadendron Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 70 (excl. T. vrie- seana). — Type: A. elegans BLUME. Blumia NEEs, Flora 8 (1825) 152, non Blumea DC., nom. conserv. — Type: Talauma candollei BLUME. Alcimandra Danvy, Kew Bull. (1927) 260. — Type: A. cathcartii (HooK./f. & THomMs.) DANDy. — Fig. 1-6. Trees or shrubs. Stipules adnate to or free from the petiole. Flowers terminal, solitary, bisexual. Tepa/s 9-21, 3—5-merous, subequal or more rarely the outer whorl forming a true calyx. Anthers introrse to latrorse, connective produced into a longer or shorter appendage or rarely unappendaged. Gynoecium sessile or in some species stipitate; carpels many to few, usually free but connate in sect. Talauma, sect. Gynopodium, and in some other species. Fruiting carpels free, crowded, dehiscent along the dorsal suture, in subg. Talauma the basal parts of the ripe carpels staying adnate to the torus, the apical parts falling and thus ex- posing the seeds which hang from the lengthened funiculus, sometimes the apical parts during or before falling dorsally dehiscing; in some species of sect. Gynopodium the carpels connate but tearing apart when maturing and de- hiscing dorsally. Ovules generally 2 in each carpel, rarely 3 or 4 in the lower car- pels, in few species up to 4 or 5 in all carpels. Ripe seeds often hanging from the funicles which lengthen through uncoiling of the spiral vessels. Distr. About 120 spp. of which about one third in the New World from SE. North America to S. Brazil, the remainder in temperate and tropical SE. Asia from the Himalayas to China, Japan, Taiwan and Malesia. In Malesia represented by sect. Maingola of subg. Magnolia, and sect. Blumiana and sect. Aromadendron of subg. Talauma. Note. Two species of subg. Magnolia are commonly cultivated in Malesia, M. coco (Lour.) DC. and M. grandiflora L. Magnolia coco (of sect. Gwillimia) differs from the species in sect. Maingola because the stip- ules are adnate to the petiole. The species can be distinguished from sect. Blumiana, especially from M. can- dollii, by the midrib being not prominent on the uppersurface. Magnolia grandiflora (of sect. Theorodon) has petals of 7 cm long or even longer while those in sect. Maingola never exceed 5 cm. The stamens in M. grandiflora are 2—3 cm long and the densely appressedly pubescent brachyblast 8 mm diam. or more. KEY TO THE SECTIONS (only in Malesia) 1. Stipules free from the petiole. Midrib not prominent above. Penaowith free carpels ..5 5.355% 6 vs' do Pe Pa Pie Ve Sis been ees nny 1. Sect. Maingola Zeretn COnnate carpels <).....)2:s!0'0) vs Sic doles elilen oo ava iahele ete ee ete 2. Sect. Aromadendron 1. Stipules adnate to the petiole. Midrib prominent above ..........5:00e ce eeeeeees 3. Sect. Blumiana To make identification of collections without fruits possible, besides the keys to the species of each section a separate key is given to the sections Maingola and Aromadendron together. KEY TO THE SPECIES OF SECTIONS MAINGOLA AND AROMADENDRON 1. Young twigs and underside of leaves hairy. Carpels free in fruit. 2. Gynoecium hairy. 3. Young twigs pubescent or pilose. Scars of perianth and stamens along 5~9 mm of the torus under the fruit. Stamens 7.5—12 mm, the 2—3 mm long connective appendage not included... 1. M. macklottii 570 FLORA MALESIANA [ser. I, vol. 103 3. Young twigs densely woolly hairy. Scars of perianth and stamens along 10—15 mm of the torus. Stamens 6—9 mm long, the 1.5—2 mm long connective appendage not included............ 3. M. maingayi 2. Gynoecium glabrous. 4. Scars of perianth and stamens along S—9 mm of the torus under the fruit. Stamens 7.5—12 mm, the 2—3 mm long connective appendage not included. Outer tepals 3.5—4 cm long ........ 1. M. macklottii 4. Scars of perianth and stamens along 10—20 mm of the torus under the fruit. Stamens c. 6 mm long, the c. 5 miniwlone, connective appenGare NOWMCUGEGn None tien ata ie ois nissan 4. M. uvariifolia 1. Young twigs mostly and underside of leaves always glabrous. Carpels free or connate in fruit. 5. Tepals c. 18 or more. Carpels connate in fruit. Connective appendage 10—15 mm. Petiole 8—25 mm. Ped- icle 0-10 mm. Alveoles in general more than 0.5 mm diam. Reticulation on the upper surface quite distinct. 6. Leaves mostly narrowly elliptic; primary nerves in 11—16 pairs. Pedicle 5—10 m. Outer tepals 4, 4.5—7 cm, inner tepals 15—33. Connective appendage 12-15 mm ....................002- 6. M. elegans 6. Leaves mostly obovate; primary nerves in 8—12 pairs. Pedicle absent. Outer tepals 3, c. 2 cm long, inner tepalsic- sla COnnechIVe.aADDendaAge LO — Osi arte nk aati nin aoe Bie Ae eile ae 7. M. bintuluensis 5. Tepals at most 12. Carpels free or connate in fruit. Connective appendage 0.5—15 mm. 7. Stamens c. 10, 4 mm long, the connective appendage c. 7 mm. Carpels c. 6. Pedicle absent, scars of SiamensianGapenianthaat mostel mime ees see . Gaye ae ee Sipe ash © prot bye epee 10. M. pahangensis 7. Stamens more than 10 and longer than 4 mm. Carpels in general many. 8. Petiole 2.5—5 cm. Pedicle 7—8 mm. Gynophore 5—10 mm. Carpels connate in fruit. 8. M. ashtonii 8. Petiole 3—17 mm. Pedicle 0-5 mm. Gynophore 0—5 mm. Carpels free or connate in fruit. 9. Outer tepals 4, c. 4.5 cm long. Stamens 9—14 mm, the 10—15 mm long connective appendage not in- cluded. Scars of tepals and stamens under the fruit along c. 5 mm of the torus. Carpels connate in THAW we bree eagiestvere oy atene Rave ee cnet NCA Cac kero RIC cece cere ase heen on aca ears eel ads 9. M. borneensis 9. Outer tepals 3, 1.5—4 cm long. Stamens 5—12 mm, the 0.5—3 mm long connective appendage not in- cluded. Scars of tepals and stamens under the fruit along c. 1-9 mm of the torus. Carpels free in fruit. 10. Stamens (S—)7.5—12 mm, the 2—3 mm long connective appendage not included. Scars of tepals and stamens along S—9 mm of the torus. Carpels many (rarely less than 15) ...... 1. M. macklottii 10. Stamens 5—10 mm, the 0.5—3 mm long connective appendage not included. Scars of tepals and sta- mens under the fruit along 1—3 mm of the torus. Carpels 1—15. 11. Stamens c. 5 mm, 0.5—1 mm long connective appendage not included. Scars of perianth and sta- MIGHS: AlOnOMIATINMOl Ne LOMISE at smatiaetins, «4: Ses 6G Antec eis eee Set ee 5. M. phaulantha 11. Stamens 5S—10 mm, the 1—1.5 mm long connective appendage not included. Scars of perianth and SiaIMensalonee: —5 MinOlAthe LOUISH ia. 4Geu- ie ss.cee Somaeaas. Ak ee fo eee 2. M. carsonii I. Subgenus Magnolia Ripe fruits consisting of free carpels which dehisce along the dorsal suture. Anthers dehiscing in- trorsely. Flowers neither precocious nor with a much reduced calyx-like whorl of outer tepals. Leaves in Asia evergreen. Only one section indigenous: I. Section Maingola Danby, Curtis Bot. Mag. 155 (1948) sub t. 16; Noort. Blumea 31 (1985) 88; ibid. 32 (1987) 346. KEY, TO THE SPECIES 1. Carpels many, or at least 15 (rarely in M. macklottii fewer). Scars of perianth and stamens in fruit along 5—20 mm of the torus. 2. Twigs (long) yellowish pubescent, pilose, or glabrous. Scars of perianth and stamens in fruit along 5—9 MAMIE eMeT ONUSey Pipe. wrk: OLE Atytee. Cle Rasilee 128 2 thes kee ee Tis sete 1. M. macklottii 2. Twigs woolly hairy when young. Scars of perianth and stamens in fruit along 10—20 mm of the torus. ee 1988] MAGNOLIACEAE (Nooteboom) 571 3. Fruits hairy, 5—8 cm long. Scars of perianth and stamens in fruit along 10(—15) mm of the torus. Brachy- Bestel — wom... 6439.20 3 5 Se erro es * diet aiaess s ae ee ie 3. M. maingayi 3. Fruits glabrous, 7—12 cm long. Scars of perianth and stamens in fruit along 10—20 mm of the torus. raaehublast 4:5—hliem < -kie «'sthedlevelo ety eelnece tn 1b. M. glauca var. sumatrana 2. Peduncles glabrous. 3. Underside of leaves densely minutely reddish-brown appressedly hairy. Nerves in 10—15 pairs 3. M. calophylla 3. Underside of leaves glabrous. Nerves in 14—20 pairs. ............ccceeeeeeevees 4. M. sabahensis 1. Stipules higher adnate to petiole, stipular scar clearly present. 4. Innovations, peduncles, stipules and underside of leaves rufous woolly pubescent .. 2. M. lanuginosa 4. Twigs and stipules glabrous or nearly so. Hairs if present minute and appressed. 5. Fruits cylindrical, 2—3 times as long as wide; carpels mostly opening along the dorsal suture first. Ovary Sarticabior cylindrical 3:5 osi56i 5 secld dv eo ncateels » oo pb see sltiilelee pele ee a 5. M. dolichogyna 5. Fruits ellipsoid or ovoid, only slightly longer than wide ............005. 1. M. glauca (var, glauca) 1. Manglietia glauca BLume, Verh. Bat. Gen. K. W. — b/umei Pranti in E. & P. Nat. Pfl. Fam. 3, 2 (1888) 9 (1823) 150; Bijdr. (1825) 8; Fl. Java Magnol. (1829) 16. — Type: Biume, Salak (”.v.) 22, t. 6; Kortu. Ned. Kruidk. Arch. 2, Versl. (1851) M. macklottii auct. non Korru.: Mig. Fl. Ind. 97; Mio. Fl. Ind. Bat. 1, 2 (1858) 15; Ann. Mus. Bot. Bat. 1, 2 (1858) 15, p.p. — M. oortii auct. non Lugd.-Bat. 4 (1868) 71; K. & V. Meded. Lands Plan- Kortu.: Mig. Suppl. (1860) 153. — Type: Teus- tent. 17 (1896) 150; Backer, Schoolfl. Java (1911) MANN HB 466, Haleban. 14; Koorp. Exk. Fl. Java 2 (1912) 238, f. 47; M. sumatrana Mig. Suppl. (1860) 153, nomen; Koorp.-Scuum. Syst. Verz. 1, Fam. 95 (1913) 1; — ibid. (1861) 367; Ann. Mus, Bot. Lugd.-Bat. 4 (1868) Backer & Baku. Fi. Java | (1963) 97; Noor. 71. — M. oortii (non Kortu.) Mig. Fl. Ind, Bat. 1, Blumea 31 (1985) 92. — Michelia doltsopa auct. non —_2: (1858) 15, p.p. — M. glauca var, sumatrana Dan- Bucn.-Ham. ex DC.: Serena. Syst. Verz. 4,2(1827) py, Kew Bull, (1928) 188. — Type: TevsMANN HB 217; Hassk. Cat. Hort. Bog. (1844) 178.— Magnolia 468, Bukit Silit (L; BO), 590 FLORA MALESIANA [ser. I, vol. 10° Fig. 7. Fruits of a. Manglietia glauca BLuME and b. M. dolichogyna DaNDy ex Noot., both x 1 (a KOORDERS 4449, b CLEMENS 26279). M. pilosa P.PARMENT. Bull. Sc. Fr. Belg. 27 (1896) 217, 292. — M. singalanensis AGostIN1, Atti Com. Accad. Fisiocrit. Siena, ser. IX, 7 (1926) sep. 22. — Type: BEccarRi PS 334 (L, FI), Sumatra, Mt Singa- lan. — Fig. 7a. Tree up to 35 m by 122 cm diam. Twigs warted, with many ring-formed scars of the fallen stipules, glabrous or with some hairs at the apex, in var. suma- trana appressedly hairy in innovations. Stipules gla- brous or with some hairs on the apex, adnate to the petiole, c. 15S—60(—90) mm. Leaves glabrous above, minutely appressedly hairy beneath, sometimes the hairs microscopically small and the leaves seemingly glabrous, glaucous, (long-)elliptic to slightly obo- vate, 10—35 by 5—12 cm; acumen 3—15 mm; base acute to slightly acuminate; midrib much prominent; nerves in (9—)11—16(—18) pairs, decurrent along the midrib, prominent on both sides, straight, anasto- mosing at some distance from the margin and merg- ing into the venation, sometimes an intramarginal vein distinct; reticulation fine, prominent on both surfaces. Petiole glabrous, 1.5—3(—4.5) cm, stipular scar c. 3—10(—15) mm, less than one third of its length, rarely slightly longer, sometimes even up to above halfway its length. Peduncle glabrous, or ap- pressedly hairy in var. sumatrana, without scars or with one scar 15—40 mm, pedicel glabrous, or ap- pressedly hairy in var. sumatrana, 2—8 mm; spatha- ceous bracts at least one towards the base of pedicel and one at base of peduncle or higher, sometimes a third at the middle of peduncle or lower. Outer fepals 3, c. 5—6.2 by 2 cm, inner tepals 6, smaller than the outer tepals. Stamens many, filament c. 3 mm, an- thers c. 6mm, connective appendage very acute, c. 4 mm; gynoecium ovoid to orbicular, c. 15 by 7 mm. Carpels 20—c. 50, nearly entirely connate when young, c. 3 mm long along the dorsal suture, dorsal face polygonous; style free, 1.5—2 mm long. Fruits ellipsoidal or ovoid, 3.5—8(—9.5) by 2.5—5.5(—6.5) cm. Seeds flat-orbicular to ellipsoidal, often irregu- larly shaped, 5—8 by 4—5 mm. a. var. glauca Peduncle and pedicel glabrous. Fruits at most 8 by 5.5 cm. Dorsal face of carpels from 1—1.5 cm in the upper to at most 2.5 cm in the lower carpels. Twigs in innovations glabrous. Distr. Malesia: Sumatra (Aceh, G. Ketambe; —E———————— 1988] MAGNOLIACEAE (Nooteboom) 59] Sumatera Utara, E. Coast and Tapanuli; W. coast; Lampung, G. Tanggamus), Java (common in the mountains of W. Java, less so in Central and E. Java), Lesser Sunda Is. (Bali, Sumba, Flores), Sula- wesi (Central, near Malili; North, Mt Nokilalaki). Ecol. In forest; 500—2400 m; /1. fr. Jan.—Dec. Uses. Good timber, used for building houses. Vern. Sumatra: antuang, bungo, madang bustak, m. kaladi, m. tjampago, tjampago; Java: baros, tempoko baros, tjepoko kantil; Bali: tjim- paka; Sulawesi: manglid. Field notes. Flowers cream, rarely white or yel- lowish green. Fruit (reddish) green to red brown. b.var. sumatrana Danby, Kew Bull. (1928) 188; Noort. Blumea 31 (1985) 93. — M. sumatrana Mig. — M. pilosa P.PARMENT. Peduncle and pedicel appressedly hairy. Fruits c. 9.5 by 6.5 cm. Dorsal face of carpels from 2 cm in up- per to 3.5 cm in lower carpels. Twigs in innovations appressedly hairy. Stipules often adnate to the very base of the petiole only. Distr. Malesia: W. Sumatra (G. Singalan, G. Talang, G. Merapi, G. Silit and Padang Panjang). Ecol. In forests; 600—1300 m; f/. Sept.—Oct., fr. June. Vern. Sikibus. Field notes. Flowers yellow or white, fruit black. 2. Manglietia lanuginosa (DANDy) Noor. Blumea 31 (1985) 94. — M. glauca var. lanuginosa DANDY, Kew Bull. (1928) 187. — Type: FRI bb 8531 (BO). Tree up to 40 m high and | m diam. Innovations and peduncles, stipules, young petioles and under- sides of leaves rufous woolly pubescent. Leaves (nar- rowly) elliptic, sometimes slightly obovate, 15—26 by 7—11 cm; nerves in 14—20 pairs. Petiole 2.5—4 cm, with stipular scar only towards the base. Further as M. glauca. Distr. Malesia: Sumatra (around lake Toba: Deli, Simelungun & Karolands, Tapanuli). Ecol. Primary forest; 1000—1500 m; /7. Feb., /r. Feb.—Nov. Vern. Aduwang, antuang, modang sanggar. Field notes. Flowers cream or green, sweet scented; fruit grey-green becoming brown. 3. Manglietia calophylla DANpy, J. Bot. 66 (1928) 46; Noor. Blumea 31 (1985) 94. — M. glauca (non Biume) Riptey, J. Fed. Mal. St. Mus. 8, 4(1917) 14. — Type: Ropinson & Kioss 200 (BM; SING), Su- matra, Korinchi Peak. Tree to at least 30 m by | m. Stipules appressedly hairy to nearly glabrous, only adnate to the very base of the petiole. Leaves ovate, covered by a dense indu- mentum of very small glistening reddish brown hairs beneath (often only to be seen with a magnification of more than x50), 10—20 by 4—8 cm with faintly acuminate apex, recurved, bony margin and acute to nearly rounded base; midrib much prominent be- neath, grooved; nerves prominent on both sides, in 10—15 pairs, anastomosing and meeting in an intra- marginal vein which is hardly distinct from the finely netted reticulation on both surfaces. Petiole 2—2.5 cm. Peduncle glabrous. Flowers as in M. glauca but smaller, the tepals as far as known not longer than c. 2.5 cm. Fruits as the smaller fruits in M. glauca c. 4—5.5 by 3.5—5 cm, number of carpels c. 20. Seeds c. 4 from each carpel, flat ovoid, c. 7 by 4 mm. Distr. Malesia: Sumatra (W. Coast, G. Kerinci). Ecol. Primary mountain forest; 2000—2200 m; fl. Oct., fr. May—Aug. Field notes. Flowers white or yellow. 4. Manglietia sabahensis DANDy ex Noor. Blumea 31 (1985) 95. — ‘Sabah Manglietia’ DANDY ex MEUER, Bot. Bull. Herb. Sabah 11 (1968) 11, nomen. — M. ‘sabahensis’ DANDY ex COCKBURN, Trees of Sabah 2 (1980) 56, nomen. — Type: CLEMENS 34192 (L; A, BO), Kinabalu, Colombon basin, 4500 ft. — Fig. 8. Tree. Stipules glabrous, c. 7 cm long, adnate to the very base of the petiole only. Leaves glabrous, the undersurface in young leaves with an occasional very small hair, elliptic to obovate, 12—22 by 6—9 cm; base cuneate, apex acute to rounded, usually not or only very shortly acuminate; midrib sulcate above, very prominent and grooved below; nerves in 14—c. 20 pairs, meeting in an intramarginal vein which is slightly more prominent than the densely netted re- ticulation. Petiole 23—30 mm, stipular scar hardly visible. Outer fepals 3, c. 3 by 1—1.5 cm; inner tepals 6, narrower. Stamens c. 1 cm long; ovary ellipsoid, c. 15 by 10 mm. Fruits ellipsoid to ovoid, c. 6 by 4 cm, the carpels opening along the dorsal suture only or both along dorsal and ventral suture. Peduncle slender, c. 3 cm long, pedicel slender, c. 2 cm long. Seeds c. 4 in each carpel, 5—8 by 3—4 mm. Distr. Malesia: Borneo (Sabah, Kinabalu). Ecol. In forest; c. 1300 m; fr. June—Aug. 5. Manglietia dolichogyna Danby ex Noor. Blumea 31 (1985) 95. — M. glauca (non BLuME) KiNG, J. As. Soc. Beng. 58, ii (1889) 370; Ann. Bot. Gard. Calc. 3 (1891) 213, t. 56, excel. fr.; Riptey, Fl. Mal. Pen. 1 (1922) 14, p.p.; Burk. Dict. (1935) 1407. — ‘Long- pistillate Manglietia’ Mever, Bot, Bull. Herb. Sabah 11 (1968) 10. — M. ‘dolichogyna’ Danpy ex Cock- BURN, Trees of Sabah 2 (1980) 56, nomen. — Type: SAN 41051 (L; SAN). — Fig. 7b. Tree up to 21 m and 60 cm diam. Stipular scar on petiole to c. halfway. Flowers: peduncle c, 2—3 cm, pedicel 4—7 mm, spathaceous bract glabrous, outer tepals 3, c. 3.5 by 14 mm, inner tepals 6, smaller. 592 FLORA MALESIANA [ser. I, vol. 102 / BN | as iu in OX CO. wih 54 ran PeeN EAS O means ee) RN eat” Sale rx Fig. 8. Manglietia sabahensis DANDY ex Noor. a. Habit, x 0.6; b. fruit, x 0.6; c. ovary, x 0.9; d. anther, x 3.8 (a, b CLEMENS 40979, c, d CLEMENS 40769). 1988] Stamens many, c. 10 mm, incl. the c. 3 mm long acute connective appendage. Ovary conical or cylin- drical, dorsal face of carpels 2—3 mm along suture, style c. 2 mm long. Ripe fruits cylindrical 4—7.5 by 2—c. 2.5 cm, 2 to 3 times as long as wide, carpels c. 60, c. 1 cm along the dorsal suture, opening along the dorsal suture first and sometimes later also along the ventral suture. Seeds flat ellipsoidal, c. 6 by 4 mm. Otherwise as M. glauca. Distr. Malesia: Malay Peninsula (Perak, Tai- ping, road to Fraser’s Hill, near Gap Valley), Borneo (Sabah, Tenom, Tambunan, Kinabalu). MAGNOLIACEAE (Nooteboom) 593 Ecol. Primary forest; 450—1500 m; fl. March— May, fr. June—Feb. Field notes. Outer tepals dark purple on green, inner tepals lemon with purple tinge; flowers pale yellow. Excluded species Manglietia ? minahassae K. & V. Meded. Lands Plantent. 19 (1898) 328, nomen; Koorp.-SCHUM. Syst. Verz. 3, 1 (1914) 41, nomen = Madhuca bur- ckiana (KoorpD.) LAMK (Sapotaceae). 3. PACHYLARNAX DaAnpDy, Kew Bull. (1927) 260; in Hutch. Gen. FI. Pl. 1 (1964) 55; Noor. Blumea 31 (1985) 97. — Type: P. praecalva DaNnvby. — Fig. 9. Trees. Stipules free from the petiole. Flowers terminal, solitary, bisexual. Tepals 9—15, 3—5-merous, subequal. Anthers introrse; connective produced in- to a short appendage. Gynoecium sessile; carpels few (2—8), concrescent; ovules about 4—8 in each carpel. Fruit a thick-walled woody loculicidal capsule, the carpels dehiscent along the dorsal suture and sometimes separating towards the apex. Distr. There are 2 spp., of which one in Assam and one in Indochina and in Malesia (Sumatra and the Malay Peninsula). 1. Pachylarnax praecalva DANDY, Kew Bull. (1927) 260, excl. plantae ex Annam; H.KENG, Tree FI. Malaya 2 (1973) 289, t. 4; Noor. Blumea 31 (1985) 97, f. 4,5. — Type: HanirF 4067 (K; SING), Penang. — Fig. 9. Tree 10—60 m by 30—100 cm, glabrous in all its parts. Stipules free from the petiole. Leaves glossy above, less so beneath, (narrowly) elliptic to obovate, 7-16 by 3—6.5 cm; apex obtuse or rounded; base cuneate or attenuate; margin recurved; midrib prom- inent beneath, slightly so above; nerves in 12—15 pairs, prominent on both surfaces, curved upwards and meeting in a looped intramarginal vein which is rather indistinct from the coarsely netted venation. Petiole without stipular scar, slender, 1.5—3 cm long. Flowers terminal on an incrassate peduncle of 0.5—20 mm (sometimes the peduncle becoming much longer); 1—3 (rarely many) spathaceous bracts under each flower, pedicel very short. Tepals 9(—10), + similar, the outermost oblanceolate to oblong, 2.5—3.5 cm long. Stamens c. 17-20 mm long with an acute connective appendage; gynoecium elongate- obovoid; carpels 2—4; ovules 4—8. Fruit 3.5—6 cm long, + orbicular before opening, loculicidal, split- ting into 2—4 valves, the carpels more or less separ- ating from each other later, in the centre a columella with the attached fruits persistent. Seeds black with enveloping pink aril. Distr. Annam (Bana near Tourane); Malesia: Sumatra (W. coast), Malay Peninsula (Kedah, Pe- nang, Selangor). Ecol. In primary forest; 360-1800 m; fl. Jr. probably the whole year round, 2. Tribus Michelieae Law Yuu-wu, Acta Phytotax. Sin. 22 (1984) 89. Growth monopodial. Flower buds arising on brachyblasts in the axils of the leaves. Genera: Michelia (incl. Paramichelia and Tsoongiodendron) and Elmerrillia. 594 FLORA MALESIANA [ser. I, vol. 103 Nes es ICME Fig. 9. Pachylarnax praecalva Danpy. a. Habit, x 0.7; b. fruit, x 0.7; c. ovary, x 2.2; d. anther, x 3 (CurTIS 3012). 1988] MAGNOLIACEAE (Nooteboom) 595 KEY TO THE GENERA ERE SUT SCSSHC! <5. s\.05 . scigrnt dee eo SOOM aut ook. ORR as & < Se ee ere 4. Elmerrillia Pe eumMeISHINGEY. SHDItAten 2. 25....!5 Gavi k «x So. veh. «Betis dake s, Sasa coe 5. Michelia 4. ELMERRILLIA DANDY, Kew Bull. (1927) 261; in Hutch. Gen. FI. Pl. 1 (1964) 56; Noor. Blumea 31 (1985) 100. — Type: FE. papuana (SCHLTR) DANDY. Elmerrillia sect. Pseudoaromadendron Danpy in Praglowski, World Pollen & Spore Flora 3 (1974) 5. — Type: E. ovalis (Mig.) DANDY. — Fig. 10, 11. Trees. Stipules free from petiole. F/owers terminal on axillary brachyblasts, solitary or sometimes 2—3-nate, bisexual, growth monopodial. Sepals 4 (or 5); petals 5—c. 10, 3—5-merous, subequal. Anthers introrse; connective produced into a short appendage. Gynoecium sessile; carpels many, with the base sunken in the torus, free or concrescent; ovules 2—6 in each carpel. Fruiting carpels basally sunken in the torus, free, crowded, and dehiscent along the dorsal su- ture, finally 2-valved, or concrescent to form a fleshy syncarp. In the latter case the carpels hexagonal, the apical parts falling away, often in irregular masses, thus shedding the seeds, or carpels tearing apart towards the outside and de- hiscing longitudinally. Distr. There are 4 spp., all in Malesia. KEY TO THE SPECIES (based on flower and fruit characters) 1. Tepals 12. Carpels free (not seen in EF. platyphylla). Per tos ATU SLIDES KIXITY =. sc iSc ce sco ates + cs05 > vebugn's> vaste ole eee 4. E. tsiampacca PmnOs ARCS. SLIDES PIADTONUS . oc es ss owe so %:o:ceule cca wane © Kinbninte Seiko neeTeE 2. E. platyphylla 1. Tepals (12—)17. Carpels free or concrescent. For New Guinea plants with 12 tepals and fruiting carpels free, see under E. tsiampacca. 3. Undersurface of leaves glaucous (sometimes a dense indumentum of appressed hairs obscuring the glau- cousness), hairy (in New Guinea sometimes glabrous: var. glaberrima). Fruiting carpels free, dorsally de- MORIN cls o'r go ccs a Fk ws 2 3.clm © ole aie. o Wo atR ARG GTks, seca sak ahve eee a orate ree 4. E. tsiampacca 3. Undersurface of leaves not glaucous, glabrous or hairy. Fruiting carpels concrescent. 4. Twigs glabrous or yellowish villous, soon glabrescent, rarely pubescent. Nerves in (10—)14—21 pairs 1. E. ovalis 4. Twigs densely fulvously pubescent or tomentose, later glabrescent. Nerves in 20-24 pairs 3. E. pubescens KEY TO THE SPECIES (based on fruit characters) 1. Fruit a syncarp, ripe carpels shedding their apical parts. 2. Young twigs and stipules glabrous or yellowish villous, soon glabrescent, rarely pubescent, Nerves in CE Bed |) es As ee ere i ee 1. E. ovalis 2. Twigs densely fulvously pubescent or tomentose, later glabrescent. Nerves in 20-24 pairs 3. E. pubescens 1. Fruit apocarpous, ripe carpels longitudinally dehiscing. 3. Twigs and leaves glabrous. Brachyblast glabrous ..........00cccccceuceeeeeeeeee 2. E. platyphylla 3. Twigs and leaves glabrous or hairy. Brachyblast hairy (except rarely in New Guinea: E. tsiampacca var. WIADOTITINGY «60-000 heed oleh LHI 0 UVite waa oes. o Vevialisind «on sake eee baie miele 4. E. tsiampacca 596 1. Elmerrillia ovalis (M1ig.) DANDY, Kew Bull. (1927) 261; Noor. Blumea 31 (1985) 101. — Talauma ovalis Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 69; Koorp. Meded. Lands Plantent. 19 (1898) 331; Suppl. Fl. N.O. Celebes 2 (1922) t. 8. — Type: For- STEN s.n. (L; BO), Celebes near Tondano. Talauma yvrieseana Mia. Ann. Mus. Bot. Lugd.- Bat. 4 (1868) 70. — Magnolia vrieseana (MiqQ.) BAILLON ex PIERRE, FI. For. Cochinch. | (1881) sub t. 2. — E. vrieseana (Mi1Q.) DANDY, Kew Bull. (1927) 262. — Type: DE VRIESE & TEIJSMANN S.n. (L), N. Celebes. Tree to 45 m high and 100(—150) cm diam. Twigs glabrous or yellowish villous, soon glabrescent, rare- ly pubescent. Stipules glabrous to yellowish villous. Leaves glabrous or underside minutely sparsely ap- pressedly hairy, rarely sparsely patently pubescent, elliptic, 7—36 by 4—16 cm; acumen less than 10 mm; base cuneate (to rounded), usually attenuate; nerves in (1O— )l14—21 pairs, intramarginal vein often rather inconspicuous in the prominent fine reticulation. Petiole glabrous or with same indumentum as twigs, (1—)2.4—4.5(—7) cm. Brachyblast glabrous or yel- lowish villous; spathaceous bracts (sparsely) pubes- cent to glabrous. Flowers (creamy) white; tepals c. 16 in 2 or 3 whorls, more or less coriaceous, the longest 25—35 mm. Stamens 8—14 mm long; carpels puberu- lous, 4(—6)-ovuled, concrescent. Fruit ellipsoidal, the ripe carpels disintegrating, losing their apical portions and thus shedding the 1—4 seeds. Distr. Malesia: Sulawesi (incl. Muna), Moluccas (Morotai, Ambon). Ecol. In forests at low and medium altitudes, up to 1000 m; fl. fr. Jan.—Dec. Uses. The timber is very durable and amongst others used for house-building. The trees are culti- vated for timber (see KoorDERS, 1898). 2. Elmerrillia platyphylla (MERR.) Noort. Blumea 31 (1985) 102. — Michelia platyphylla MERR. Philip. J. Sc. 13 (1918) Bot. 11; Enum. Philip. 2 (1923) 153. — Type: FB 26866 (K), Leyte. Tree. Twigs glabrous. Stipules glabrous. Leaves el- liptic, glabrous, 23—30 by 9—13 cm; acumen shorter than 10 mm; base cuneate, shortly attenuate; nerves in 18—23 pairs meeting in an intramarginal vein; reticulation rather fine. Petiole 2—3 cm. Brachyblast glabrous, 4—5 cm long; spathaceous bracts glabrous. Flowers white; tepals c. 12, the outer ones c. 25—35 mm long. Stamens c. 12 mm; ovary glabrous. Fruits unknown as yet. Distr. Malesia: Philippines (Leyte, Agusan; Mindanao, Zamboanga). Ecol. Secondary forest at low altitudes. 3. Elmerrillia pubescens (MERR.) DANDY, Kew Bull. (1927) 261; Noor. Blumea 31 (1985) 102. — Talauma FLORA MALESIANA [ser. I, vol. 103 pubescens MeErRR. Philip. J. Sc. 3 (1908) Bot. 133; Enum. Philip. 2 (1923) 152. — Type: CLEMENS 686 (iso BO), Mindanao. Tree c. 1S—40 m high and up to 80 cm diam. Twigs densely fulvously pubescent or tomentose, glabres- cent. Stipules densely fulvously long tomentose or pubescent. Leaves pubescent beneath, elliptic to ovate, 15—30 by 8—14 cm; acumen less than 1 cm; base nearly rounded, slightly acuminate; nerves in 20—24 pairs, meeting in a looped intramarginal vein; reticulation fine. Petiole with same indumentum as twigs, 20—25 mm. Brachyblast densely fulvously to- mentose or pubescent, 3—5 cm; spathaceous bracts densely pubescent. Flowers white; tepals c. 15, glabrous, the outer ones 27—35 mm, the inner ones gradually shorter and narrower. Stamens c. 10 mm; carpels many, 3—4 ovuled, pubescent. Fruit 3—6 by c. 2 cm; carpels c. 8 mm long, concrescent, tearing apart towards the outside when mature, longi- tudinally dehiscing, 1—4-seeded. Distr. Malesia: Philippines (Mindanao, Lanao lake, Mt Katanglad, Mt Apo). Ecol. Altitude c. 2000 m; fl. fr. Jan.—Dec. Vern. Hangilon, Bukidnon. 4. Elmerrillia tsiampacea (LINNE) DANDy in Prag- lowski, World Pollen & Spore Flora 3 (1974) 5; Noor. Blumea 31 (1985) 103, f. 6—8. — Michelia tsiampacca LINNE, Mant. (1767) 78; Miq. Fl. Ind. Bat. 1, 2 (1858) 18, p.p.; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 73; MerRR. Interpr. Rumphius (1917) 224. — Michelia champacca auct. non LINNE: LINNE, Syst. ed. 10, 2 (1759) 1082, p.p. — Michelia celebica Koorp. Meded. Lands Plantent. 19 (1898) 328, 631, nom. ill.; Suppl. Fl. N.O. Celebes 2 (1922) t. 9. — E. celebica (KOORD.) DANDY, Kew Bull. (1927) 261. — Type: Sampacca sylvestris RuMPH. Herb. Amb. 2 (1741) 202, t. 68. Talauma papuana SCHLTR, Bot. Jahrb. 50 (1913) 70. — E. papuana (SCHLTR) Danby, Kew Bull. (1927) 261; Crorr in Womersley, Handb. Papua New Guinea (1978) 130, t. 65. — Type: SCHLECHTER 19001 (not seen). Michelia forbesii Baker f. J. Bot. 61 (1923) Suppl. 2. — Type: ForseEs 442a (K; P), Sogeri. Michelia arfakiana AGostTINi, Atti Com. Accad. Fisiocrit. Siena, ser. IX, 7 (1926) sep. 25. — Type: BEcCARI PP 890 (FI), Mt Arfak. E. mollis Danby, Kew Bull. (1928) 184. — Michelia mollis (DANDY) McLAuGHLIN, Trop. Woods 34 (1933) 36. — Type: ENDERT 5252 (K; BO, L), Borneo, W. Kutei. E. papuana var. adpressa DaNpy, Kew Bull. (1928) 185. — Type: LEDERMANN 13089 (K; L), Sepik region. E. papuana var. glaberrima DaANDy, /.c. — Type: LEDERMANN 9505 (K), Sepik. MAGNOLIACEAE (Nooteboom) $97 1988] v, | )) Sat) * SSS Z c S > a " ” x) p) y a. Habit, * 0.6; b- npaca . (star Fig. 10. Elmerrillia tsiampaca (Linnt) DANDY var x2; e. young fruit, x 2; /. ditto, longitudinal section, 598 E. sericea C.T.Wuirte, J. Arn. Arb. 10 (1929) 212. — Type: Brass 661 (A; K), Sogeri. — Fig. 10, 11. Tree to 60 m high and 150(—200) cm diam. Twigs (densely) ferrugineously or fulvously (woolly) pubes- cent or tomentose when young, rarely glabrous. Stip- ules with same indumentum. Leaves often appressed- ly pubescent above in innovations, soon glabrescent, hairy beneath but glabrous in var. glaberrima, some- times glaucous, (narrowly) elliptic, rarely ovate (often so in Sulawesi), 10—46 by 4—15 cm; acumen 5—17 mm; base cuneate to rounded, rarely subcor- date; nerves in 11—28 pairs, meeting in an often hardly distinct intramarginal vein; reticulation rather fine to very fine. Petiole 7-35 mm. Brachyblast densely pubescent (glabrous in var. glaberrima), 15—40(—60) mm. Flowers white to yellow; fepals (10—)12—c. 15, glabrous or the outer ones pubescent, 20—35 mm long. Stamens 10—14 mm; carpels many (c. 50), pubescent or puberulous, glabrous in var. glaberrima. Fruit cylindrical, 4—9 by 1.5—2 cm. Distr. Malesia: Sumatra (Mentawai I., Siberut), Borneo, Central & N. Sulawesi, Moluccas (Ambon, Buru), New Guinea (incl. Biak & Japen), New Brit- ain. Ecol. On fertile soil in forest; 0O—1800 m; fl. fr. Jan.—Dec. KEY TO THE INFRASPECIFIC TAXA 1. Leaves 10—30 cm; nerves in (11—)14—22(—24) pairs. Hairs underneath the leaf blade not curved towards their base. Tepals (12—)c. 15. Sulawesi, Moluccas, New Guinea..... a. ssp. tsiampacca 2. Young twigs and stipules hairy; ovary puberu- TOUS ke doce sc eee eee al. var. tsiampacca 2. Entire plant glabrous..... a2. var. glaberrima 1. Leaves 16—46 cm; nerves (14—)17—18 pairs. Hairs underneath the leaf blade usually (uncinately) curved towards the base. Tepals (10—)12. Su- THOU Oe AOT COLE. eee ore b. ssp. mollis a. ssp. tsiampacca. — All synonyms except E. mollis DANDY. FLORA MALESIANA [ser. I, vol. 10% Tree to 60 m high and 150(—200) cm diam. Leaves beneath very densely, often minutely appressedly or patently hairy, 10-30 by 3—15 cm. Petiole 12—35 mm. Distr. Malesia: Central & N. Sulawesi, Moluccas (Ambon, Buru), New Guinea (incl. Biak & Japen), New Britain. al. var. tsiampacca. — All synonyms except E. pa- puana var. glaberrima DANDY. — Fig. 10. Distr. As the subspecies. Ecol. On fertile soil in forest; to 1400 m; //. fr. Jan.—Dec. Uses. The most valuable and most demanded timber in N. Sulawesi. Already becoming scarce in 1898 (KooRDERS). Very durable. Vern. Sulawesi: tjempaka utan, t. u. aloes, uru tanduk, wasian, w. rintek, w. sela; New Guinea: ba- lamtalogo, Mooi lang., boska, Manikiong lang., arimot, pui, Biak, hui, Sogeri, wasau, wuka, Morobe, kaule, Kainantu, bibau, Hattam lang., war- mei, Waskuk lang., pubitza, Garaina lang., rap, Sepik, biendjung. a2. var. glaberrima (DANDY) Noor. Blumea 31 (1985) 107, f. 6c. — E. papuana var. glaberrima DANDY. Distr. Malesia: New Guinea (Kebar Valley, Idenburg R., Sepik R., Morobe Distr.). Ecol. Primary forest; up to 1200 m. Vern. Mamer, mamier, Kebar lang. b. ssp. mollis (DANDY) Noor. Blumea 31 (1985) 108, f. 6d, 8. — E. mollis DANDy. — Fig. 11. Tree to 40 m high and 80 cm diam. Leaves beneath villous, pubescent, or puberulous. Petiole 7—25 mm. Carpels with 2 ovules. Distr. Malesia: Sumatra (Mentawei I., Siberut), Borneo. Ecol. In forest at low and medium altitude. In Sabah 1500—1800 m. F/. fr. Jan.—Dec. Vern. Sumatra: minjaran; M; Borneo: arau, miharo, Dyak. 5. MICHELIA LINNE, Sp. Pl. (1753) 536; Gen. Pl. ed. 5 (1754) 240; Danby in Hutch. Gen. FI. Pl. 1 (1964) 56. — Champaca ADANS. Fam. PI. 2 (1763) 365, 537. — Sampacca O. K. Rev. Gen. Pl. 1 (1891) 6. — Type: M. champaca LINNE. Liriopsis Spacu, Hist. Natur. Veget., Phanerog. 7 (1839) 460, non Liriopsis REICHB. (1828). — Magnolia sect. Liriopsis BAILLON, Hist. Pl. (1868) 142, note 4. — Type: L. fuscata (ANDR.) SPACH. 1988] MAGNOLIACEAE (Nooteboom) : : Fig. 11. Elmerrillia tsiampaca (LINNE) DANDY var. mollis (DANDY) Noor. Twig with flower, deflorated flow- er, and bud showing the stipular nature of the spathaceous bract; natural size (NOOTEBOOM 4518A). Paramichelia H.H.Hu, Sunyatsenia 4 (1940) 142. — Type: P. baillonii (PIERRE) Hu. — Fig. 12, 13. For further synonymy see Noor. Blumea 31 (1985) 108. Trees or shrubs. Stipules adnate to or free from petiole. Flowers bisexual. Tepals 6—21, 3—6-merous, subequal or rarely the outer whorl different. An- thers latrorse or sublatrorse (to introrse). Gynoecium stipitate, carpels many to few (rarely 1), free or connate; ovules 2 to many; fruiting carpels free or concrescent. Distr. About 30 spp., in Southeast Asia from India and Sri Lanka eastwards to S, Japan and Taiwan and : cultivated ] southeastwards into Indonesia (not in Sulawesi and New Guinea). In Malesia 6 spp., 2 other spp. commonly KEY TO THE SPECIES Petiole 3—5 mm long, stipular scar present for nearly its whole length. Leaves glabrous, 4.5-6.5 by 2-3 cm. Tepals 6 8. M. figo 600 FLORA MALESIANA [ser. I, vol. 103 ( ’ ‘ 917 Vest A) VY CXS { A) ih Wy \ VK f v) iy " , Vp h, Wa WHA hy Wy LY if Vy ¥ We, yy We vy, SSK SY Fig. 12. Michelia scortechinii (KiNG) Danpy. a. Habit, x 0.7; b. young fruit, x2; c. anther, x 4; d—e. ovary, x 4 (a, c-e SF HoittuM 31244; b GRASHOFF 335). 1988] MAGNOLIACEAE (Nooteboom) 601 1. Petiole longer than 5 mm. The other characters different or the tepals more than 6. 2. Young twigs glabrous. Terminal buds hairy at the apex only. Carpels 1—4. Stipules free from the 15—35 mm long petiole. Leaves glabrous, 9—35 cm long Cts oS ER tao Se ee eo eee 4. M. montana 2. Young twigs hairy, at least directly under the terminal bud. Carpels 8 or more. 3. Stipules adnate to the petiole for one third or more than one third of its length. 4. Leaves 8—12 by 2.5—3.5 cm. Petiole 6—16 mm 4. Leaves 10—35 by 4—11 cm. Petiole 14—50 mm. 1. M. scortechinii 5. Tepals 15, from light yellow to orange, 20—45 mm long. Carpels c. 30, fertile. Leaves 10—30 cm 3. M. champaca 5. Tepals c. 12, white, 30—55 mm long. Carpels c. 10, sterile. Leaves 15-35 cm ........ 7. M. alba 3. Stipules free from petiole or adnate to its base only (in M. salicifolia sometimes up to one third). 6. Leaves 15—35 by 5.5—11 cm. Petiole 15—50 mm. Tepals c. 12, white, 30—55 mm. Carpels sterile, c. 10 7. M. alba 6. Leaves 4—16 by 2.3—6 cm. Petiole 5—20 mm. Tepals white to yellow, 10—40 mm. Carpels fertile. 7. Brachyblast 10-17 mm. Outer tepals 3, membranous, inner tepals 6, coriaceous. Twigs and stipules puberulous to nearly glabrous. Leaves glabrous beneath, 6—16 by 3—6 cm. Carpels c. 10 2. M. koordersiana 7. Brachyblast 3—7 mm. All the tepals the same, 9-17. Twigs and stipules pubescent or puberulous. Leaves glabrous or hairy beneath, 4—13 by 1.5—4 cm. Carpels 8—16 or c. 30. 8. Tepals 20—40 mm. Leaves 9—13 by 2.3—4 cm, often glaucous beneath. Carpels c. 30 5. M. salicifolia 8. Tepals 10-15 mm. Leaves 4—12 by 1.5—3.5 cm, not glaucous beneath. Carpels 8—c. 16 1. Michelia scortechinii (KING) DaNpy, Kew Bull. (1927) 262; H.KENG, Tree Fl. Malaya 2 (1973) 288; Noort. Blumea 31 (198%) 111, f. 9. — Manglietia scortechinii KiNG, J. As. >oc. Beng. 58, ii (1889) 370; Ripey, Fl. Mal. Pen. 1 (1922) 14. — Paramichelia scortechinii (KING) Danby in Praglowski, Pollen & Spore Flora 3 (1974) 21. — Type: SCORTECHINI 764 (BM, K, SING). — Fig. 12. Tree to 37 m high and c. 50 cm diam. Twigs zigzag, ferrugineously pubescent to tomentose when young. Stipules adnate to the petiole for at least half its length, to 30 mm long. Leaves (densely) ferrugin- eously pubescent or puberulous beneath, disti- chously arranged, elliptic, 8—12 by 2.5—3.5 cm; apex shortly acuminate; base cuneate; nerves in 12—18 pairs, with the very fine reticulation prominent on both sides. Petiole 6-16 mm. Brachyblast 6-8 mm; spathaceous bracts densely ferrugineously pu- bescent. Flowers white; tepals c. 12, more or less the same, 12—18 mm long. Stamens c. 8 mm, incl. the c. 1 mm long connective appendage; carpels c. 20, densely ferrugineously pubescent. Fruiting carpels concrescent. Distr. Malesia: Sumatra (W. Coast, Bengkulu, Palembang), Malay Peninsula (Perak, Pahang, Cameron Highlands). Ecol. Primary forest; 650-1300 m; fl. fr. April- May. 2. Michelia koordersiana Noor. Blumea 31 (1985) 111, f. 10. — Fig. 13. Tree to 32 m high and 62 cm diam. Twigs finely ap- pressedly puberulous when young, or only so directly under the terminal bud, soon glabrescent, often zig- 6. M. philippinensis zag. Stipules with same indumentum to nearly gla- brous, not adnate to petiole, 10O—15 mm. Leaves gla- brous, more or less elliptic, distichously arranged, 6—16 by 3—6 cm; apex shortly acuminate, acumen (O—)3—8 mm; base cuneate; nerves in 7—13 pairs, with the fine reticulation prominent on both sides. Petiole 10-20 mm. Brachyblast appressedly pubes- cent, 10-17 mm. Flowers (orange) yellow; outer fep- als 3, membranous, 12—22 mm; inner tepals 6, coria- ceous. Stamens incl. the 0.5 mm long appendage 5—7 mm; carpels c. 10, with the c. 5 mm long gynophore minutely tomentellous or puberulous. Fruiting car- pels 2—10. Distr. Malesia: Sumatra (W. coast, Padang; E. coast, Palembang), Malay Peninsula (Selangor). Ecol. Primary forest; 0O—1000 m; f/. fr. probably Jan.—Dec. 3. Michelia champaca LinNE, Sp. Pl. (1753) 536; BiumE, Bijdr. (1825) 7; Fl. Java Magnol. (1829) 9, t. 1; Bianco, FI. Filip. (1837) 462; Kortu. Ned. Kruidk. Arch. 2 (1851) 96; Mig. Fl. Ind. Bat. 1, 2 (1860) 101, 153; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 72, excl. var.; K. & V. Bijdr. 4 (1869) 159; Atlas 4 (1918) t. 799; Koorp. Exk. Fl. Java 2 (1912) 241; Mere. Enum. Born. (1921) 251; Enum. Philip, 2 (1923) 152; Backer, Schoolfl. Java (1911) 16; Riptey, Fl. Mal. Pen. 1 (1922) 15; Burk. Dict. (1935) 1465; Corner, Wayside Trees (1940) 434; Backer & Baku /. Fl. Java | (1963) 98; Noor, Blu- mea 31 (1985) 113. — Type: HERMANN FI, Zeyl. 144 (BM). M. suaveolens Pens. Syn. 2 (1806) 94, p.p. excl. syn. Rueepe ef Rumen, — M. blumei Steup. 602 FLORA MALESIANA [ser. I, vol. 103 Fig. 13. Michelia koordersiana Noor. a. Habit, x 0.6; b. fruit, x 0.6; c. ovary, x 4; d. anther, X 6 (VAN DER ZWAAN for THORENAAR 1345). 1988] Nomencl. ed. 2, 2 (1841) 139. — M. tsiampacca L. var. blumei Moritz1 in Zoll. Syst. Verz. (1846) 36. — Type: RumpH. Herb. Amb. 2 (1741) t. 67. M. tsiampacca BuiuME, Bijdr. (1825) 7, non M. tsiampacca LINNE (1767). — M. velutina BLUME, FI. Java Magnol. (1829) 17, non M. velutina DC. (1824); Mia. FI. Ind. Bat. 1, 2 (1858) 17; K. & V. Bijdr. 4 (1896) 162; BacKER, Schoolfl. Java (1911) 16; Koorp. Exk. Fl. Java 2 (1912) 241; H.KENG, Tree Fl. Malaya 2 (1973) 288 pro coll. KEP/FRI 6943. — Champacca velutina O. K. Rev. Gen. Pl. 1 (1891) 6. — M. pilifera Baxu.f. Blumea 12 (1963) 61; BACKER & Baku-/f. Fl. Java | (1963) 98. — Type: BLUME s.n. (L). M. pubinervia BLumE, Fl. Java Magnol. (1829) 14, t. 4. — M. rufinervis BLuME, Bijdr. (1825) 8, non M. rufinervis DC. (1817). — M. champaca Mia. FI. Ind. Bat. 1, 2 (1858) 16, p.p., non M. champaca LINNE (1753); BisscHop GREVELINK, Pl. Ned. Ind. (1882) 277, p.p. — M. champaca LINNE var. pubinervia (BLUME) Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 72. — Type: BLUME 670 (L; BO). Talauma villosa Mig. forma celebica Miq. Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 70. — Type: DE VRIESE & TEUSMANN s.n. (L; BO). M. montana auct. non BLUME: BAKER /. J. Bot. 62 (1924) Suppl. 2. Huge forest tree to 50 m high and 183 cm diam. Twigs (appressedly) pubescent, glabrescent. Stipules pubescent, adnate to petiole at least for one third of its length. Leaves spirally arranged, (long) elliptic or ovate, pubescent below especially on midrib and nerves, often glabrescent, 10-30 by 4—10 cm; acumen 7—13(—25) mm; base cuneate to more often rounded; nerves in 14—23 pairs, intramarginal vein often hardly more prominent than fine reticulation. Petiole 14—36(—40) mm. Brachyblast densely pu- bescent, (S—)10—18(—25) mm long; spathaceous bracts pubescent. Flowers light yellow becoming dark orange; tepals 15, 20—45 mm long. Stamens 6—8 mm, incl. the up to | mm connective appendage; carpels c. 30, the c. 3 mm long gynophore densely pu- bescent. Distr. From India to SW. China and Indochina; in Malesia: Sumatra, Malay Peninsula, Java, Lesser Sunda Islands. KEY TO THE VARIETIES 1. Leaves ovate with cuneate-attenuate base; the acu- men often quite long. Petiole with a stipular scar up to shortly below its middle to up to its apex. Tree to c. 30 m high and 50 cm diam, a. var, champaca 1. Leaves more or less elliptic with cuneate to round- ed base, the acumen often rather short, oblique. Petiole with a stipular scar from 0,3 up to 0.7 of MAGNOLIACEAE (Nooteboom) 603 its length. Tree to 50 m high and 180 cm diam. b. var. pubinervia a. var. champaca. — All synonyms except those under var. pubinervia. Distr. Commonly cultivated throughout the tropics. Probably originally from India, where it is cultivated on the temple ground of Jains and Hindus. Uses. See BuRKILL, Dict. ed. 2 (1966) 148. Vern. 7jampacca ((India: cempaka) or t. kuning is the common name in most of its area. Further: djeumpa, Aceh, djempa, Gajo, kantil, lotjari, pet- Jari, J, kembang konéng, tjampacca mera, Mad., t. barak, t. warangan, Bali, hépaka, képaka, Sawu, sampacca, s. modarag, t. mariri, Ald., Cel., bunga edja, Mak., bunga matjela, Bug., kupa haja, Ceram, kupa pokur, papokur, pupukuljo, walatol, Ulis, hapaka, tjapaka, N. Halmaheira, ¢. goratji, Ternate, Tidore. b. var. pubinervia (BLUME) Mia. Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 72; Noor. Blumea 31 (1985) 115. — M. pubinervia BLUME. — M. tsiampacca BLUME. — M. montana auct. non BLUME: BAKER /. Distr. Malesia: Sumatra (Aceh; Bengkulu, Lake Ranau), Malay Peninsula (Kedah, Langkawi Is.; Bt. Kaju Hitam; Kelantan, Cameron Highlands), Java (common in the mountains), Lesser Sunda Ils. (Sumbawa). Ecol. Evergreen primary forest on fertile soil; 250—1500 m, in Java mostly between 1000 and 1200 m. Fi. fr. Jan.—Dec. Uses. The wood is highly esteemed for building and furniture. The properties are at least the same as of Tectona grandis. Because of its nice structure the value for furniture is higher than that of the wood of M. montana. In the beginning of this century its ex- tinction was already feared by Koorpers because of the use made of it. The trees can easily be cultivated and reach a height of c. 27 m and a diameter of c. 55 cm in 27 years (K. & V., 1896). Vern. Sumatra: kemait, M; Java: baros, manglis, S, kadjeng sekar, J, kadju kempheug, Md, lungjung, J; Sumbawa: fengkel. 4. Michelia montana BLume, Verh. Bat. Gen. K. W. 9 (1823) 153; Bijdr. (1825) 7; Fl. Java Magnol. (1829) 15, t. 5; Mig. Fl. Ind. Bat. 1, 2 (1858) 17; Suppl. 1 (1860) 153; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 73; Backer, Schoolfl. Java (1911) 15; Koorp, Exk. Fl. Java 2 (1912) 241; Riptey, Fl. Mal. Pen. 1 (1922) 15; Burk. Dict. (1935) 1491; Corner, Wayside Trees (1940) 434; Backer & Baku /. Fl. Java 1 (1963) 98; Noor. Blumea 31 (1985) 116, f. 11. — Sampacca montana O. K. Rev. Gen, Pl. 1 (1891) 6. — Lecto- type: BLume 575 (L, NY). M. ecicatrisata Mig. Fl. Ind. Bat. Suppl. [(1860) 604 153, nomen] (1861) 368. — M. montana BLUME var. subvelutina Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 73. — Lectotype: TEUSMANN HB 4457 (L; BO). Tree becoming over 40 m tall and over 2 m diam. Twigs glabrous. Stipules pubescent at the apex only, exceptionally entirely pubescent, free from petiole. Leaves glabrous, spirally arranged, more or less el- liptic, 9—30(—35) by 4—13(—20) cm; acumen 2—20 mm; base attenuate; nerves in 9—15 pairs, conspic- uous, intramarginal vein rather conspicuous; reticu- lation dense, prominent on both surfaces. Petiole often thickened towards its base, 15—35 mm. Brachyblast 5—20 mm, glabrous to (rarely) densely pubescent; spathaceous bracts glabrous except the margins towards the apex. Outer fepals 3, rather thin, 15S—30 mm, greenish or greenish white; inner tepals creamy or white, 6, thick, coriaceous, 14—40 mm. Stamens incl. the c. 2 mm long connective ap- pendage 10—13 mm; carpels 1—4, with the 4-8 mm long gynophore reddish puberulous when dry, green in vivo. Fruiting carpels free. Distr. Malesia: Sumatra (Aceh; W. coast; Lam- pong; Palembang; Banka), Malay Peninsula (Perak; Pahang; Cameron Highlands), Borneo (Sabah; E. Kalimantan), Java (common), Bali. Fig. 14. la Zs = - ee 6 ities | end | oe ee: Je? o2 | . *¢ ® on a eee ° 7 -§ “8 as os : ws é pe Se NG A ma een Yea Gera asl nM je Y \ 3 es pai “tA | <— sat tS an Be a) he as By ea a = ae , = S Wyn EN 2 GY ( } a j % \. MUR athnitenntt aa : Sp Miley ye.) UN Be, 100 Noel . 7 Loe } og tyes Cong Sir nn alelettal seh (arnt in a ca ee ee a a ayers ner mae LINACEAE (A.M.N. van Hooren & H.P. Nooteboom, Leiden) In this work Linaceae sensu lato have been split into three families: Linaceae, Ixonanthaceae and Ctenolophonaceae, among which the latter deviates most. In order to elucidate distinction of the two segregated families of Linaceae sensu lato VAN HooreEN & Nooresoom (Blumea 29, 1984, 550) prepared the following diagnoses: Linaceae — Lianas, trees, shrubs, or herbs. Tufted hairs absent. Stipules lateral. Leaves spirally or distichously arranged. Flowers hypogynous. Petals caducous, contorted. Disk absent (or traces of an extrastaminal disk present, /.c. 556 sub Philbornea). Filaments basally connate in a tube. Styles 3—5(—6), simple. Fruit a drupe or a capsule (sometimes with indehiscent mericarps). Seed not persistent, with slightly or not developed arillode. — Stomata paracytic. Ixonanthaceae — Trees. Tufted hairs absent. Stipules lateral. Leaves spirally arranged. Flowers perigynous. Petals persistent in fruit, imbricate (extra-Mal. also contorted). Disk intrastaminal. Filaments free, inserted outside and against the disk. Style 1, simple. Fruit a capsule. Seed not persistent, with an obvious basal wing or suprahilar arillode. — Stomata paracytic. Ctenolophonaceae — Trees. Tufted hairs present. Stipules interpetiolar. Leaves opposite. Flowers hypogynous. Petals caducous, contorted. Disk extrastaminal. Filaments free, inserted halfway on inside of disk. Style 1, apically bifurcate, with 2 stigmas. Fruit a capsule. Seed persis- tent on the columella after the valves have been shed, with hairy-papillose arillode. — Stomata anomocytic. Linaceae sensu stricto are distinguished by several flower characters from the other two fami- lies, viz. disk absent or almost so, filaments connate at base, styles 3—5, and arillode absent or hardly developed. The distinction of these families is also sustained by wood and anatomical re- search (HEIMSCH & TSCHABOLD, 1972) and pollen morphological studies (SAAD, 1962; OLTMANN, 1971). NARAYANA & RAo (1978) concluded that on the basis of floral morphology and embryology Linaceae are related to Erythroxylaceae and Humiriaceae, in addition to showing affinity with Ctenolophonaceae and Ixonanthaceae. In his studies on seeds, CORNER (1976) opposed an affinity with Geraniaceae, but suggested Malpighiaceae and possibly also Oxalidaceae as closer relatives. Within Linaceae sensu stricto there are two distinct subfamilies, Linoideae and Hugonioideae. They can be distinguished as follows: 1. Linoideae — Erect herbs or small shrubs. Petals usually long-clawed. Stamens as many as pet- als, alternating with the same number of staminodes. Ovary 6—10-celled. Fruit usually a cap- sule. — Almost entirely confined to the northern hemisphere. 2. Hugonioideae — Trees or lianas with hooks, rarely shrubs, all ligneous. Petals not or hardly clawed. Stamens twice as many as petals. Ovary 3—S5-celled. Fruit a drupe, rarely splitting fi- nally in indehiscent mericarps. — Pantropical, but hardly on the northern hemisphere except in southern Southeast Asia. In Malesia only Hugonioideae occur, and have not seldom been distinguished as a separate family Hugoniaceae (e.g. by Exe, & MENDONGA, 1951; TAKHTAJAN, 1969; DAHLGREN, 1975; CRONQUIST, 1981). CORNER (/.c.) dwelt extensively on the anatomical structure of their seeds. In his opinion the simple tegmen of the Linoideae may be derived from the mesotestal construction in the Hugonioideae. The genus Indorouchera of the Hugonioideae may yield the most primitive pollen type in Linaceae. References: CORNER, The seeds of dicotyledons | (1976); Cronguist, An integrated system of classification efc. (1981); DAHLGREN, Bot. Notis. 128 (1975) 119-147; Exet, & MENDONGA, Con- spectus florae Angolensis 1, 2 (1951) 242—249, 390-392; Hemmscu & TscHABOLD, Bot. Gaz. 133 (1972) 242—253; NARAYANA & Rao, J. Ind. Bot. Soc. 57 (1978) 258-266; OLTMANN, Pollen- morphologisch-systematische Untersuchungen innerhalb der Geraniales. Diss. Bot. 11 (1971); SAAD, Pollen et Spores 4 (1962) 65—82; TAKHTAJAN, Flowering plants, Origin and dispersal (1969) 226. (607) 608 FLORA MALESIANA [ser. I, vol. 10° SUBFAMILY HUGONIOIDEAE Trees or lianas, rarely shrubs. Hairs, if present, simple, uniseriate, or multi- seriate with multicellular, glandular heads. Stipules lateral, caducous. Leaves simple, spirally or distichously arranged, pinnately nerved. /nflorescences an axillary or terminal compound panicle, raceme, corymb, or cyme, or an axile fascicle, few- to many-flowered, rarely 1-flowered. Flowers actinomorphic, pentamerous, bisexual, hypogynous, sometimes heterodistylous. Sepals basally shortly connate or free, quincuncially imbricate, subequal or unequal, flabel- lately nerved, persistent, often slightly indurated and enlarged in fruit. Petals free, rarely basally clasping, shortly or not clawed, contorted, flabellately nerved, caducous. Disk absent. Stamens 10, alternately shorter and longer; fila- ments basally connate in a tube; anthers dorsoversatile, 2-celled, introrse. Extrastaminal nectary glands adnate to staminal tube, up to 5, or absent. Ovary superior, 3—5(—8)-celled; styles 3—5(—8), free or basally connate; stigma capi- tate. Ovules 2 per cell, axile, collateral, pendulous, epitropous. Fruit a drupe, rarely splitting in pyrenes. Seed(s) 1 or 2 per cell, arillode hardly or not devel- oped. Endosperm dry or fleshy, copious or scanty. Embryo straight or slightly curved. Distribution. Pantropical; 5 genera, of which 2 small (Hebepetalum, Roucheria) confined to tropical South America, | (Philbornea) confined to Malesia, 1 Indo-Malesian (Jndorouchera), and | in the Old World tropics (Hugonia, incl. Durandea), including Madagascar, with some three dozen species, eastwards extending as far as the Solomon Islands, Queensland, New Caledonia, and Fiji. In Malesia in all 5 spp. Ecology. Tropical, everwet rain-forest, mostly in the lowland but locally ascending to 1500 m on Mt Kinabalu. As to latitude up to c. 20° N and 25° S (Madagascar). The South American representatives are trees, the Old World ones predominantly lianas, rarely shrubs in New Caledonia. Sometimes the lianoid species may remain shrubs or small trees as long as no support is available to climb. As soon as a support is present, the plants climb by means of stout, woody, curved climbing hooks (branch metamorphosis), which wind around the support and considerably thicken afterwards. Vegetative anatomy. The Hugonioideae are leaf anatomically very strictly defined by their shared possession of subsidiary cells which are lobed underneath the stomatal guard cells and cris- tarque cells (cells with Ca-Oxalate crystals and a unilaterally thickened, sclerified cell wall); para- cytic stomata are another constant feature of the group. This very unusual combination of charac- ters induced VAN WELZEN & Baas (1984) to advocate family status for Hugoniaceae, because the Linoideae (or Linaceae) lack this combination of characters. Most Hugonioideae have lignified guard cell pairs, but Jndorouchera constitutes an exception and can be separated from the other two Malesian genera on account of its unlignified cells. The wood anatomy of the Malesian Hugonioideae is incompletely known. /ndorouchera has been recorded to have exclusively scalariform perforations; in Hugonia they are simple. The fibres have distinctly bordered pits like in all members (families or subfamilies) of the Linaceae sensu lato. References: HEmscH, Lilloa 8 (1942) 83—198, pl. 1—17; HEmmscH & TSCHABOLD, Bot. Gaz. 133 (1972) 242-253; METCALFE & CHALK, Anatomy of the dicotyledons 1 (1950) 268—279; VAN WELZEN & Baas, Blumea 29 (1984) 453—479. — P. Baas. Palynology. Pollen grains in Linaceae are suboblate to prolate, and measure from 20 to 90 1988] LINACEAE (van Hooren & Nooteboom) 609 um. The apertural system is mostly tricolpate or tricolporate. In Linum also pantocolpate and pantocolporate pollen occur. Anisadenia and Reinwardtia pollen is always pantoporate. Jn- dorouchera has inaperturate grains, which, however, actually represent a tritenuate condition. In most genera the exine shows differentiation in sexine and nexine. In Jndorouchera the exine is a thin, homogeneous sheet, covered by small verrucae. Otherwise, the sexine is granulate or baculate. Sometimes the tips of the bacula are fused to form a columellate-tectate sexine. The macromorphologically distinct subfamilies Linoideae and Hugonioideae can be separated in a pollen morphological way too. Pollen of Linoideae is tricolpate, pantocolpate, or panto- porate, and has a more or less granulate sexine, while that of Hugonioideae is tricolporate (Hugonia, Philbornea) or inaperturate (Jndorouchera), and mostly has a more or less baculate or a columellate-tectate sexine (SAAD, 1962). Generally speaking, subfamily Linoideae has more primitive pollen characters than subfamily Hugonioideae, although Indorouchera may yield the most primitive pollen type in Linaceae. Following Saap (/.c.) the pollen of the Linaceae is primitive within the Geraniales. Relationship of the Linaceae with [xonanthaceae, as well as with Erythroxylaceae and Humiriaceae, is sup- ported by pollen morphology (OLTMANN, 1971). However, pollen also indicates the distinctness of the Linaceae within this group of families. Pollen of Ctenolophonaceae appeared to be dissimilar to that of Linaceae. References: OLTMANN, Pollenmorphologisch-systematische Untersuchungen innerhalb der Geraniales. Diss. Bot. 11 (1971); SAAb, Pollen et Spores 4 (1962) 65—82. — R.W.J.M. VAN DER Ham. Phytochemistry. See HEGNAUER, Chemotaxonomie der Pflanzen 4 (1966) 393. KEY TO THE GENERA 1. Indumentum present, at least on the calyx. Styles 5(—6). Ovary 5(—6)-celled, locules all distinct in the fruit. Peet 2 4(—5) Geveloped SEEdS .. . 2. ies ee acbe cde scans wee Come ieee enna 1. Hugonia 1. All parts glabrous. Styles 3—4(—5). Ovary 3(—4)-celled, only one, rarely 2 locules distinct in the fruit. Drupe usually with one developed seed. 2. Resin absent. Leaves spirally arranged. Flowers in rather lax racemes or panicles...... 2. Philbornea 2. Buds and stipules often covered by resin. Leaves distichously arranged. Flowers in axillary fascicles, sub- tended by often densely packed, imbricate bracteoles ..............020eeeeeeeeees 3. Indorouchera 1. HUGONIA LinnE, Gen. Pl. ed. 5 (1754) 305; Sp. Pl. (1753) 675; Srapr in Hook. Ic. PI. (1906) t. 2822; HALL.f. Beih. Bot. Centralbl. 39, 2 (1923) 43; Hus. WINKLER in E. & P. Nat. Pfl. Fam. ed. 2, 19a (1931) 108; vaN HooreNn & Nooresooo, Blu- mea 29 (1984) 553, map. — Durandea PLANcu. in Hook. Lond. J. Bot. 6 (1847) 594, nom. cons.; HUB. WINKLER in E. & P. Nat. Pfl. Fam. ed. 2, 19a (1931) 108. — Hugonia sect. Durandea (PLANCH.) BaAILLon. Hist. Pl. 5 (1874) 48. — Fig. Ss. Indumentum present at least on calyx. Stipules palmatifid-laciniate, pinnati- lobed or simple, entire or dentate. Leaves spirally arranged. Flowers in axillary or terminal more or less densely flowered racemes or leafy or leafless panicles, or axillary, in few-flowered cymes or solitary, rarely ramiflorous. Bracts and bracteoles present or not. Sepals free, unequal, rarely subequal. Petals shortly clawed or thickened at base. Nectary glands present or not. Ovary 5(—6)- 610 FLORA MALESIANA [ser. I, vol. 10° Zl th Fig. 1. Hugonia costata Mia. a. Habit, x 0.5; b. climbing hooks, x 0.5; c. stipules from inside, x 3; d. flower bud; e. innermost sepal; /. pistil (in bud), all x 5; g. fruit, x 1.5; A. fruitin CS, x 1.5; i. seed, x 3; 7. embryo, x 3 (a, c—j FORBES 2978, b ForBEs 2814). 1988] LINACEAE (van Hooren & Nooteboom) 611 loculed; styles 5(—6). Drupe (pseudo-)indehiscent or splitting into 5 pyrenes; endocarp woody, provided with more or less prominent longitudinal ribs, either compact and forming one stone with 5(—6) fertile locules alternating with as many interlocular sterile cavities or divided into 5 thin, more or less obviously ribbed pyrenes which are separated by a fleshy mesocarp. Seeds 1 per locule, 2—4(—5) developed per fruit, apically attached. Embryo with fleshy endosperm. Distr. About 40 spp., in the Old World reaching from Senegal in the west to Fiji in the east and from Bombay in India at 20° N as far as Madagascar at c. 25° S; in Malesia 2 spp., one in the west (Sumatra, Borneo) and a second one in the east (Moluccas, New Guinea). They belong to two allopatric sections: sect. Hugonia possesses some 30 spp. in Africa, Mauritius, and 4 spp. in SE. Asia (Sri Lanka, Indonesia, Malesia); sect. Durandea (PLANCH.) BAILLON has 4 spp. in E. Malesia, Queensland, Solomons, New Caledonia (2 or 3 endemic), and Fiji. Ecol. Mixed tropical forests; 0—1500 m (Mt Kinabalu). KEY TO THE SECTIONS AND THE SPECIES 1. Young parts, twigs, stipules, petiole and nerves densely tomentose. Stipules digitately laciniate or pinnati- lobed. Sepals S—15 mm long. Petals 12—30 by 4-10 mm. Drupe usually indehiscent, sometimes finally sep- ree MRE EIU COMI «3.5 yn.sysasoudlivss i kaleicie os mieeAee oiedb-n ol aipundngs se aintalnguacaamceiaiaaaek an eens ae 1. H. costata 1. Young parts, stipules, petiole and nerves glabrous or with some very small scattered hairs. Stipules simple, entire or dentate. Sepals up to 3.5(—4) mm long. Petals 2.5—3.5 by 1.5—2.5 mm. Drupe splitting into 5 SemeeeeCh. OUTANGEH ..~ . <.- <0-.n.. liens» = «oO 1. Hugonia costata Mra. Illustr. (1871) 67; BorRL. Hand. Fl. Ned. Ind. 1 (1890) 139; Ha... Beih. Bot. Centralbl. 39, 2 (1923) 45; van HooREN & NOOTE- BOOM, Blumea 29 (1984) 555. — Fig. 1. Liana. Tomentum appressed to patent, hairs up to 1 mm. Twigs densely tomentose when young, gla- brescent. Climbing shoots often patent, some grow- ing out to a leafy, sometimes fertile branch; first in- ternodium 3—12 cm long; hooks (1—)2, subopposite or opposite, apically on the first internodium, 2.5—5 cm long, later thickening; at apex often with cata- phylls. Stipules narrowly ovate to obovate, 5—10 by 2.5—3.5 mm, digitately laciniate up to 1/3, with 7—10 lobules. Leaves sparsely to densely tomentose, narrowly elliptic to obovate, sometimes asymmetric, (4—)6—18 by 2.5—7 cm; base obtuse to acute or cu- neate; margin glandular-crenulate to crenate, glands caducous, conical; apex acuminate to cuspidate, acute to obtuse, acumen up to 1.5 cm; nerves 11—17 pairs, arching upwards and ending in the dense re- ticulation; petiole 3-14 mm, the margin with 1-3 conical glands on each side. /nflorescence an axillary (and terminal?) 3-flowered cyme, 1—1.5 cm, some- times reduced to | flower, densely tomentose; bracts caducous, narrowly ovate to obovate, 10—15 by 2—3.5 mm, deeply digitately laciniate into 8 lobules. Pedicel 2—5 mm, articulate below the calyx. Flowers only known in bud. Buds ovoid, c. 8 by 5.5 mm. Sep- als leathery, the 3 or 4 outer ones tomentose, boat- shaped, ovate, 9-10 by 5—6.5(—8.5) mm, inner ones 1—2 mm shorter. Petals boat-shaped, broadly ovate, Poo s 6 nein 0's win 48's 90 re ane oe 2. H. jenkinsii at least 10 by 7 mm. Staminal tube + thickened at base, sometimes persistent in fruit; filaments + hairy. Ovary 5- (or 6-)loculed, glabrous, cylindric; styles 5 (or 6), exceeding the stamens, of different length. Drupe broadly obovate to globular, 18—23 by 20—24 mm; base with vascular openings absorbed in- to a navel with slightly prominent margin; mesocarp thick-fleshy, hard and strongly irregularly wrinkled when dry, adnate to the stone; endocarp woody, very hard, with thick radial septa, 5 (or 6) narrowly ellip- tic to narrowly ovate (in CS), seminal cavities and as many alternating, always empty, cordate, sterile cavities, 1.7 mm diam. Seed/(s) | or 2, exarillate, ven- trally attached, flattened obovoid, 11.5 by 4.5 mm, c. 1 mm thick; hilum not obvious; testa thin, 2-layered. Embryo straight; cotyledons obovate, 7 by 4mm; radicle cylindric, 3 by 0.8 mm. Endosperm co- pious, slightly granular. Distr. Malesia: Sumatra (Palembang, Padang), Malay Peninsula (Lobis For. Res.), Borneo (Sabah, Kutai, E. Kalimantan). Fig. 2. Ecol. Typically matures in late building gaps, well drained ridges, and flat upland areas; in Malaya on gently undulating ground in lowland forest at 100 m. Although rarely collected, according to LEIGHTON (coll. no. 1010) locally relatively common, 8 mature individuals/km’; on Mt Kinabalu at 1500 m. 2. Hugonia jenkinsii F.v.M. Fragm. 5 (1865) 7; VAN Hooren & Nooresoom, Blumea 29 (1984) 555 (with complete synonymy). — Durandea pallida K.Scn. in FLORA MALESIANA [ser. I, vol. 103 Fig. 2. Range of Hugonia costata Miq. (stars) and H. jenkinsii F.v.M. (dots). K.Sch. & Hollr. Fl. Kaiser Wilhelm Land (1889) 56. — Ancistrocladus pentagynus WARB. Bot. Jahrb. 13 (1891) 383. — H. pentagyna (WarRB.) K.SCH. in K.Sch. & Laut. Fl. Deut. Schutzgeb. Stidsee (1900) 373; HuB. WINKLER in E. & P. Nat. Pfl. Fam. ed. 2, 19a (1931) 100, f. 48. — Durandea rotundata K.ScuH. in K.Sch. & Laut. Fl. Deut. Schutzgeb. Stidsee, Nachtr. (1905) 278. — Durandea pentagyna (WARB.) K.Scu. /.c. — Durandea jenkinsii (F.v.M.) STAPF in Hook. Ic. Pl. (1906) t. 2822. — H. robinsonii MERR. Philip. J. Sc. 11 (1916) Bot. 277. — Durandea pen- tagyna var. rotundata (K.ScH.) LAuT. Bot. Jahrb. 52 (1915) 117. — Durandea robinsonii (MERR.) HALL./. Beih. Bot. Centralbl. 39, 2 (1923) 41. — Fig. 3. Liana to 40 m, rarely small tree to 3 m, or climbing shrub, up to 10 cm diam. Climbing shoots with first internodium (2.5—)6—18 cm, following internodia very short with some distichously arranged, reduced, linear to orbicular leaves, a few of these shoots with a long-peduncled, reduced inflorescence or a climb- ing hook, or the climbing shoot above the hooks growing out to a leafy and often fertile branch. Hooks distichously or subopposite, woody, (3.5—) 6.5—8.5 cm long, apically often with bract-like or- gans, peduncles, cymes, or reduced leaves. Stipules entire, obtuse to acute, with or without glandular teeth, persistent or sometimes caducous, sometimes only present as a glandular tooth, subulate or trian- gular, 0.5—0.8 by 0.2—0.8 mm. Leaves glabrous (rarely some hairs on midrib), narrowly elliptic to obovate, sometimes broadly elliptic or ovate, some- times asymmetric, (6—)11—27(—36) by (1.6—)3—9.5 (—11) cm; base cuneate, seldom rounded; margin often wavy, glandular-crenulate to crenate, apex acute to acuminate, rarely obtuse, acumen up to 2 cm; nerves 7—16 pairs, slightly arching upwards and almost reaching the margin, meeting in an often somewhat obscure intramarginal vein or in the dense reticulatfon; petiole 3—20 mm. /nflorescence an axil- lary or terminal compound panicle, sometimes a ra- ceme; terminal panicles often leafy, cone-shaped to obovoid, (4—)12—18(—32) by (2.5—)1.5—14 cm with 5—25(—30) more or less closely set subinflorescences, sometimes up to 5(—8) pseudoterminal panicles to- gether; axillary panicles patent, narrowly obovoid to broadly ellipsoid, (2—)6—11(—12) by (1—)2—6.5 cm, with (4—)10—23 subinflorescences; peduncle up to 6.5 cm; cymes up to 15, 0—5 times branched; bracts persistent, obtuse deltoid, (1.5—)2.5—5 by 0.2-1 mm, with 2—6 glandular teeth. Pedicel articulate and often thickened below the flower, 1.5—3 mm. Brac- teoles like the bracts but smaller. Flowers heterodis- tylous, 7—9 by 5—9 mm. Sepals + orbicular to trans- versely broadly elliptic, (1.8—)2—3(—4) by 2—4(—4.5) mm, shortly connate to free, appressed but patent in fruit, outer 2 smaller and thicker. Peta/s recurving during anthesis, narrowly elliptic to -obovate, 6—9.5 by 1.5—3(—4) mm; base often indistinctly 0.8 mm clawed. Stamens shortly persistent, obdiplostemon- ous, in short-styled flowers (4.5—)4.8—6 and (5—)5.5—7.5 mm, in long-styled flowers (2.5—) 3.8—4.5(—4.8) and (3.8—)4.2—5(—5.8) mm; tube 1—1.5(—2) mm, partly persistent in fruit; base often irregularly thickened or marked by darker coloured LINACEAE (van Hooren & Nooteboom) 613 Fig. 3. Hugonia jenkinsii F.v.M. a. Flower, x 4; b. inner sepal, x 5; c. outer sepal, x 5; d. flower, sepals and petals removed, x 4; e. stamens with glandular thickenings at base of tube, x5; f. fruit, x 1.5; g. pyrene, x6; h. seed, 3 (a—h BSIP 10657). semi-ellipsoid, sometimes apically notched, staminal glands at the base of long or short stamens, these later becoming connate with each other; filaments triangular at base, apically filiform. Ovary 5-loculed, ovoid to broadly ellipsoid, 1—2 by 0.8—1.2 mm; styles 5, basally shortly connate and apically (in short- and long-styled forms) hooked or sigmoidly curved or (in long-styled forms) slightly arching to curving downwards, sometimes of different length, in short-styled flowers (1—)1.5—2.8 mm, in long- styled flowers (3—)3.2—3.5(—5.2) mm; stigma 2-lobed. Drupe 15-ribbed, ovoid to subglobose, (8—)10—13 by (7—)8—10.5 mm, splitting into 5 pyrenes; mesocarp fleshy, surrounding the pyrenes, between the pyrenes often disappearing thus giving rise to a false locule; endocarp bony, thin, 3-ribbed, ribs branching, the branches connected by bony sep- ta, forming cavities filled with aerenchyma; septa and aerenchyma dissolving when ripe. Seed | or 0 in each pyrene, ventrally-apically attached by a short funicle, medially flattened, asymmetrically semi-el- lipsoid, c. 8.5 by 3.5 mm; hilum and arillus not ob- vious; testa thin, 2-layered. Embryo c. 8 by 3 mm; radicle flattened, arched, c. 1.5 by 2.5 mm. Endo- sperm fleshy. Distr. W. Pacific (Fiji), Melanesia (Solomons, New Caledonia), Queensland; in Malesia: New Guinea, Moluccas (Ceram, Ambon). Fig. 2. Ecol. Primary and secondary forests, often on well drained places or along rivers, also along estua- ries or mangroves, locally rather scarce to common; up to 2000 m. Fi. Jan.—Nov.; fr. Feb.—Nov. Field notes. Flowers fragrant to very strong sweet-scented with yellowish green to dark green sep- als and bright golden yellow to orange petals. Fila- ments pale olive yellow to orange with deep purple to brown anthers. Styles pale yellow to pale orange with green stigma. Fruits golden yellow to light orange be- coming pinkish red, red to brown or orange brown, Uses. In Queensland the Tully River natives use the climbing hooks as fish hooks. Vern. New Guinea: agref, Vogelkop, abus, Sepik, Waskuk, eamuraka, Sepik, Wagu, quamo, Wikahiri. 614 FLORA MALESIANA [ser. I, vol. 10° 2. PHILBORNEA HALL.f. Arch. Néerl. Sc. Ex. Nat. IIIB, 1 (1912) 110; in Fedde, Rep. 13 (1914) 414; Hus. WINKLER in E. & P. Nat. Pfl. Fam. ed. 2, 19a (1931) 109; van HooreENn & Nootresoom, Blumea 29 (1984) 556. — Fig. 4. Lianas with hooks. Indumentum absent. Stipules simple, entire. Leaves spirally arranged, rarely some leaves opposite. /nflorescences a rather lax flow- ered axillary or terminal raceme, sometimes ramiflorous. Bracts and bracteoles present. Sepals basally shortly connate, subequal. Disk absent; nectary glands not obviously present. Styles 3. Ovary 3-loculed. Drupe indehiscent, 1-seeded with 2 reduced locules. Seed ventrally attached, with arillode. Embryo straight. Endosperm scanty, fleshy. Distr. Monotypic. Malesia: Sumatra, Borneo, Philippines. Ecol. Periodically inundated soil and near rivers in the lowland. Fy ils Fig. 4. Philbornea magnifolia (STAPF) HALL.f. a. Habit, x 0.5; b. bract, bracteoles, and flower bud, x5; c. fruit; d. fruit, exocarp removed; e. seed; f. embryo; all x 3 (a after STAPF, b KOSTERMANS 21142, c—f RAHMAT st BOEEA 5362). 1988] LINACEAE (van Hooren & Nooteboom) 615 1. Philbornea magnifolia (STAPF) HALL.f. Arch. Néerl. Sc. Ex. Nat. IIIB, 1 (1912) 110; in Fedde, Rep. 13 (1914) 415; van Hooren & NooTEesBoom, Blumea 29 (1984) 556. — Durandea magnifolia STAPF in Hook. Ic. Pl. (1906) t. 2822; in Fedde, Rep. 5 (1908) 268. — P. palawanica Hat..f. Arch. Néerl. Sc. Ex. Nat. IIIB, 1 (1912) 110; in Fedde, Rep. 13 (1914) 415. — Fig. 4. Twigs often roughened by many light coloured lenticels. Climbing shoots patent, first internode 6—12 cm, following internodia very short with cadu- cous, reduced, broadly elliptic leaves. Hooks 2, (sub)opposite, woody, 1.5—5 cm. Stipules appressed to patent, triangular, sometimes gland-like, 0.3—0.5 by 0.2—0.3 mm. Leaves obovate or elliptic, rarely ovate, 7—20(—30) by (2—)4—10(—14.5) cm, some- times asymmetrical; base gradually decurrent to the petiole; margin glandular-crenulate; apex broadly acute to obtuse or acutely acuminate, acumen up to 2.8 cm; nerves 7—14 pairs, slightly arching upwards to and along the margin, often meeting in an intra- marginal vein; reticulation dense; petiole rather stout, often thickened, (3—)6—15(—25) mm. Ra- cemes axillary from (often fallen) leaves or scales, or terminal, sometimes with some cymes, 7—15(—28)- flowered, (2—)3.5—9.5 by (1—)1.5—4 cm. Bracts nar- rowly triangular to rhomboid, 0.5—2 by 0.2-0.5 mm, usually with 2 triangular glandular teeth. Pedi- cels articulate in or above the middle, (2—)4—10(—15) mm. Bracteoles 2, persistent, very small. Sepals basally shortly connate, membranous, + shell- shaped, 3—4 mm diam., the inner 3 sometimes la- cerate up to 2/3. Petals recurving during anthesis, el- liptic to obovate, 7.5—9 by 4—4.5 mm. Stamens ob- diplostemonous, the long ones 5.5—6.2 mm and ba- sally thickened, the short ones 1—1.5 mm shorter, 4.5—4.8 mm; staminal tube 0.5—1.5 mm. Filaments basally flattened. Ovary elliptic to obovoid- pyriform, 1—1.5 by 0.7—1.5 mm; styles free, apically sometimes slightly twisted; stigmas knob-shaped, 2-lobed. Drupe lengthwise wrinkled, often flattened at one side, ovoid, 10—12 by (S—)7—10 mm, exocarp membranous; mesocarp thin-fleshy, + 0.3 mm thick; endocarp leathery in basal and sterile half, tougher and with prominent, subreticulate, firm, + 0.2 mm thick ribs in upper half of fertile part. Seed 1, smooth, flattened ovoid, 10-11 by 5.5—6 mm, ventrally attached by a long raphe, thickened around the raphe and along the whole length by an arillode; testa 0.2 mm thick; tegmen somewhat thinner. Em- bryo straight; cotyledons transversely broadly ovate, + 6 by 6 mm; radicle cylindric, 4.5 mm long. Distr. Malesia: Sumatra, Borneo (Sarawak, E. Kalimantan), Philippines (Palawan). Fig. 5. Fig. 5. Range of Philbornea magnifolia (STAPF) HALL. Ecol. Periodically inundated soil and near rivers, up to 270 m. Field notes. Petals bright yellow. 3. INDOROUCHERA HALL.f. Beih. Bot. Centralbl. 39, 2 (1923) 50; Hus. WINKLER in E. & P. Nat. Pfl. Fam. ed. 2, 19a (1931) 109; Backer & Baku./f. Fl. Java 1 (1963) 241; VAN Hooren & Nootesoom, Blumea 29 (1984) 557. — Fig. 6—8. Glabrous lianas, with hooks. Buds often covered with resin. Stipules simple, entire or dentate, often covered with resin. Leaves distichously arranged. Flow- ers in fascicles in the axils of (often fallen) leaves, rarely solitary, heterodis- tylous. Bracts absent. Pedicel densely beset with often numerous, + spirally ar- ranged, persistent, imbricate bracteoles and often covered with resin. Sepals free, unequal. Petals very thin. Nectary glands absent. Ovary 3(—4)-loculed; 616 FLORA MALESIANA [ser. I, vol. 10° Fig. 6. Indorouchera contestiana (PIERRE) HALL.f. a. Habit; b. twig with climbing hooks provided with re- duced inflorescence at apex; c. climbing hook, thickened after grasping a twig; all x 0.5; d. stipule, x 12; e. flower and bracteoles, x 12; f. flower, sepals and petals removed, x 8; g. fruit; h. fruit in CS; i. fruit, exocarp removed; j. seed; k. embryo; all x 12. — /. griffithiana (PLANCH.) HALL.f. /. Fruit, exocarp removed; m. fruit in CS; n. seed; all x 12 (a, d, g—k HAVILAND 2840, b VAN NIEL 3607, c A 441 RAHM, e, f NBFD 2113, /, m DE WILDE & DE WiLDE-DuyrFfieEs 16540, n Meijer 7335). 1988] LINACEAE (van Hooren & Nooteboom) 617 styles 3—4(—5), basally connate to free. Drupe indehiscent, 1- (rarely 2-)seeded with 2 (or 1) reduced locule(s). Seed ventrally attached, arillate. Embryo straight or slightly curved. Endosperm copious, oily. Distr. 2 spp., SE. Asia (India: Nicobars; ?Burma, Thailand, Cambodia, S. Vietnam); in Malesia: Sumatra, Malay Peninsula, Borneo, Java. Ecol. Rain-forest, also along the seashore, estuaries and on sanddunes, up to 1500 m. KEY TO THE SPEGIES 1. Leaves elliptic to obovate; reticulation densely parallel in the whole leaf. Petals (5—)6.4—8.5 by 1.5—3 mm. Stigma knob-shaped. Drupe 4.5—6 by 3—4.5 mm 1. L. griffithiana 1. Leaves often ovate, rarely elliptic; reticulation predominantly in the basal half laxly and (irregularly) parallel, in the apical half reticulated. Petals 4—5.5 by 1—1.8 mm. Stigma clavate, recurved. Drupe 4—4.5 by 2.5—3 mm 1. Indorouchera griffithiana (PLANCH.) HALL./. Beih. Bot. Centralbl. 39, 2 (1923) 50, incl. var. coria- cea HALL.f., /.c. 52; HuB. WINKLER in E. & P. Nat. Pfl. Fam. ed. 2, 19a (1931) 109; BAcKER & BakH/. Fl. Java 1 (1963) 242; CockBurNn, Tree Fl. Malaya | (1972) 306; Trees of Sabah 1 (1976) 204; vAN HoOREN & NooTtesoom, Blumea 29 (1984) 557. — Roucheria griffithiana PLANCH. Hook. Lond. J. Bot. 6 (1847) . 1. RSE RLS, CUS ee ee 2. I. contestiana 143: ibid. 7 (1848) 527; Warp. Ann. 1 (1849) 97: Hook./. Fl. Br. India 1 (1874) 414; Bogert. Hand. |! (1890) 140; Kina, J. As. Soc. Beng. 62, ii (1893) 190; BoERL. Feestbundel (1894) 91, tab.; Ic. Bog. 1 (1897) 25, t. VII, f. 1-22; Koorp. Nat. Tijd. Ned. Ind. 60 (1901) 384; Backer, Schoolfl. Java (1911) 163; Koorp. Exk. Fl. Java 2 (1912) 415; Rm.ey, FI. Mal. Pen. | (1922) 323; Kew Bull. (1926) 60. — Flacourtia Fig. 7. Indorouchera griffithiana (PLANCH.) HALL./. in blossom. Njarumkop (NW. Kalimantan) (Photogr Father A. E_sener, 9 April 1964) 618 FLORA MALESIANA [ser. I, vol. 10° ?camptoceras Mia. Fl. Ind. Bat., Suppl. (1861) 288; BOERL. Feestbundel (1894) 91; Ic. Bog. 1 (1897) 27; Koorp. Exk. Fl. Java 2 (1912) 415; HALL.f. Beih. Bot. Centralbl. 39, 2 (1923) 50; SLEUMER, FI. Males. I, 5 (1954) 77. — Hugonia sumatrana Mia. Illustr. (1871) 68; BoERL. Feestbundel (1894) 91; Ic. Bog. 1 (1897) 27. — Fig. 61—n, 7, 8. Liana, up to 30 m, rarely shrub or treelet, 4 m. Stem up to 3 cm diam. Climbing shoots with first in- ternodium of 3—10 cm. Hooks | or 2 at the end of a climbing shoot or several (up to 5) distichously ar- ranged in the axils of (often fallen) leaves or inflores- cences, 3—5 cm long. Stipules triangular to ovate, 0.7—0.8 by 1.5—1.8 mm, crenate. Leaves elliptic to obovate, rarely ovate, (4—)7.5—11.5(—19) by (1—) 2.5—4(—6.8) cm; base cuneate; margin glandular to crenate; apex obtusely caudate to acuminate, rarely rounded; acumen up to 2.5 cm; nerves (S—)7—10 pairs, + arching upwards and almost reaching the margin; primary veins often subperpendicular to midrib and many of them + parallel; reticulations Fig. 8. Indorouchera griffithiana (PLANCH.) HALL./. Twig with hook and flowers at Njarumkop (NW. Ka- limantan) (Photogr. Father A. ELSENER, 9 April 1964). densely parallel and transverse to midrib; petiole nar- rowly sulcate above, (0.5—)1—2 cm. Flowers (1—)3—7(—10) together, heterodistylous. Pedicels ar- ticulate, 1-3 mm; bracteoles membranous, ovate, 0.2—1(—2) by 0.2—0.9(—2) mm. Sepals elliptic to ovate or orbicular, 1.8—2.5 by (1.5—)2—2.8 mm. Petals thin, in anthesis straight, elliptic to obovate, (5—)6.5—8.5 by 1.5—3 mm; base obtuse to very short- ly clawed. Stamens in short-styled flowers 3.5—6 and 5—8 mm, in long-styled flowers 2.5—3.8 and 3.5—5.5 mm; staminal tube 0.6—1.3 mm, persistent in fruit. Ovary 3(—4)-loculed, rather smooth to 3—4-lobed when dry, cylindric, obovoid or _ globular, 0.8—1(—1.5) by 0.7—1(—1.5) mm; styles 3—4(—S), sometimes persistent in fruit, straight to sigmoidly curved in the middle, in short-styled flowers 1.2—2 mm, in long-styled flowers 3.2—5.5 mm; stigma flat- tened, knob-shaped, slightly 2-lobed. Drupe + lengthwise ribbed when dry, ovoid to ellipsoid, 4.5—6 by 3—4.5 mm; exocarp membranous, 0.1—0.2 mm thick; mesocarp + ribbed, fibrous-bony, 0.4—1 mm thick; endocarp bony, 0.3—0.4 mm thick. Seed(s) 1 (or 2), ventrally attached, semi-ovoid, 3—4.5 by 2.2—2.3 mm; arillus around the 1-2 mm long hilum, short, slightly thickened, funicular; testa thin. Embryo 3—4.3 by 1—2 mm; cotyledons straight or plicate, elliptic to broadly elliptic, 1.8—2.8 by 1—2 mm; radicle cylindric to flattened, straight to slightly curved, 1.1—1.5 by 0.4—0.5 mm. Distr. India (Nicobar Is.), Thailand; in Malesia: Sumatra, Malay Peninsula, Borneo, Java. Fig. 9. Ecol. Rain-forests, common, but rare in Java, the Nicobar Is. and Thailand; up to 1500 m. Fi. fr. Jan.—Dec. g B ¢ 199° * gs ae Fig. 9. Range of Jndorouchera griffithiana (PLANCH.) HALL./f. 1988] Field notes. Petals white to bright yellow, pale orange or reddish brown. Stamens yellow, anthers bright yellow, pistil light yellow. Drupe yellow to red, 8 mm diameter, with edible pericarp and tomato-like flavour. Uses. According to ScHotz (in Engler, Syllabus 2, 1964, 243) the plant contains saponin-like constit- uents. They are used in mixtures for arrow poisons. See also BuRKILL, Dict. 2 (1935) 1949. In Sarawak used for parang handles. Vern. Sumatra: andor pijom, Tapanuli, akar tandu, takkolan, East Coast, olor nanas, olor sila- yur, surango etem, Simalur. Malay Peninsula: akar garam garam, a. tenggadin, Selangor. Borneo: wa bakar, Iban, akar katjap, Brunei, dingkai, Sabah, akar tuai, Kenyah. 2. Indorouchera contestiana (PIERRE) HALL./. Beih. Bot. Centralbl. 39, 2 (1923) 52; Hus. WINKLER in E. & P. Nat. Pfl. Fam. ed. 2, 19a (1931) 110; vAN Hoo- REN & NooTesoom, Blumea 29 (1984) 558. — Rou- cheria contestiana PrERRE, Fl. For. Coch. (1893) t. 281; GumLLauMIN, FI. Gén. I.-C. 1 (1911) 586, f. 61-5 & 61-6. — I. rhamnifolia HA...f. Beih. Bot. Cen- tralbl. 39, 2 (1923) 52; HuB. WINKLER in E. & P. Nat. Pfl. Fam. ed. 2, 19a (1931) 110. — Fig. 6a—k. Liana, up to 10 m, sometimes a shrub up to 4 m. Stem + 1 cm diam. Climbing shoots patent, c. 45°, first internodium 3.5—7 cm, hooks | or 2 at the end of a climbing shoot or up to 3 in the axils of leaves or inflorescences, at the apex sometimes with re- duced inflorescences, (0.5—)1.5—4.5 cm long. Stip- ules liguliform to shallowly triangular, 0.8—1.5 by 1—1.5 mm, entire to crenate. Leaves elliptic to ovate, 3—11.5 by 1.5—4.7 cm; base obtuse, rarely cuneate; margin glandular-crenulate; apex obtusely acumin- ate, rarely obtuse, acumen up to 1.5 cm; nerves (4—) 5—8 pairs; veins + perpendicular to midrib and many parallel; reticulation rather lax, irregularly pa- rallel, predominantly in basal half and along midrib, densely reticulate in apical part of leaf; petiole 0.5—1 cm. Flowers in (1—)4—7(—14)-flowered fascicles. Pedicel articulate, 0.5—1.5 mm. Bracteoles membra- nous, ovate, 0.3—1(—1.5) by 0.3—1(—1.5) mm. Sepals elliptic to ovate, 1.7—2.5 by 0.7—1.6 mm. Sta- mens 1.8—2.2 and 2.5—3.5 mm; staminal tube 0.7—0.8 mm high. Ovary 3-loculed, rather smooth, cylindric to ovoid, 0.8-1 by 0.6—0.7 mm; styles 2-3.5 mm; stigma clavate, recurved, slightly 2-lobed, c. 0.25 mm diam. Drupe ovoid, 4—4.5 by 2.5-3 mm; exocarp thin-fleshy, c. 0.2 mm thick; LINACEAE (van Hooren & Nooteboom) 619 mesocarp chartaceous, + lengthwise grooved, c. 0.2 mm thick; endocarp woody, crescent-shaped in CS, + 0.2—0.4 mm thick. Seed(s) 1 (or 2), ventrally at- tached, semi-ovoid, 2.5—2.7 by 1.7—1.9 mm, often asymmetric; hilum narrowly elliptic, + 1.5 mm long; aril indistinct, short, + thickened below the hilum; testa thin. Embryo 2.8—3 by 1.3—1.5 mm; cotyle- dons straight, broadly elliptic to + orbicular, 1.5—1.7 by 1.3—1.5 mm; radicle cylindric to slightly flattened, 0.8—1 by 0.2—0.3 mm. Endosperm gran- ular. Distr. S. Vietnam, Cambodia; Borneo. Fig. 10. in Malesia: Fig. 10. Range of /ndorouchera contestiana (PIERRE) HALL. Ecol. Edge of young secondary forest and of swampy forest, rare; up to 10 m altitude. Fi. fr. April—June. Field notes. Petals yellow, fruit red. Uses. According to Scnouz (in Engler, Syllabus 2, 1964, 243) the plant contains saponin-like constit- uents. They are used in mixtures for arrow poisons. See also BuRKILL, Dict. (1935) 1949, For details on chemical contents see HEGNAUER, Chemotaxonomie der Pflanzen 4 (1966) 397. On the label of Cutn 2829 is mentioned: ‘a ‘‘T’’-point can be made into a pa- rang handle. The wood is tough, not excessively hard, and will not split easily.’ Vern. Borneo: kait kait puteh, Brunei. IXONANTHACEAE (R. Kool, Leiden) This small family of 2 or 3 Old World rain-forest genera was already recognized as a separate suprageneric taxon by PLANCHON (1847) and PLANCHON & KLOTZSCH (1856), who relegated it to the affinity of Ochnaceae, later correctly referred to Linaceae as a subfamily /xonanthoideae by Hus. WINKLER (1931) and finally recognized as a family of its own by EXELL & MENDONGA (1951). As to the number of genera contained in the family, there is no unanimity of opinion. FORMAN (1965: 523) referred 8 genera to the family, but NooTEBOoM (1967) argued that several belong to Simaroubaceae. After careful consideration 3 genera are admitted here in Jxonanthaceae sensu stricto: Cyrillopsis KuHLtM. from South America, Ochthocosmus BENTH. (incl. Phyllocosmus KLotzscuH) from tropical America and Africa, and /xonanthes Jack from Indo-Malesia. The three genera form a close-knit group. They are all small and the wide distribution in the tropics points to a high age of the group. We must mention that there is a, doubtful, fourth genus, A//antospermum FoRMAN, which the author reckons to the /xonanthaceae. It occurs with one species in Borneo and another one in Ma- dagascar. NooTEBOOM (1967, 1972) included this genus in Simaroubaceae. The morphology, chemotaxonomy, and palynology corroborate this affinity, but the anatomy of wood and leaf (VAN WELZEN & BAAS, 1984) is just in favour of affinity with /xonanthaceae. We refrain from a long discussion of the merits of HALLIER’s attempt (1923) to have Linaceae as a huge complex centre of affinities and confine ourselves to what is usually accepted nowadays in recognizing a few families grouped around Linaceae sensu Stricto. In the treatment of the family Linaceae (page 607, see there) the families Linaceae, Ixonan- thaceae, and Ctenolophonaceae are opposed by concise diagnoses. Leaf anatomy has clarified many points but a fair number remains unsolved, which should be the subject of further multidisciplinary studies (VAN WELZEN & BAAS, /.c.). References: ExELL & MENDONGA, Bol. Soc. Brot. ser. 2a, 25 (1951) 105; FoRMAN, Kew Bull. 19 (1965) 521—526; Hauer f. Beih. Bot. Centralbl. 39, 2 (1923) 1-178; NoorEBoom, Adansonia Il, 7 (1967) 161—168; Fl. Males. I, 6 (1972) 970, f. 1; PLANCHON in Hook. Lond. J. Bot. 6 (1847) 588—603; PLANCHON & KiotzscH, Abh. Kon. Ak. Wiss. Berlin 1856, Physik. Abh. (1857) 235; VAN WELZEN & Baas, Blumea 29 (1984) 453—479; Hus. WINKLER in E. & P. Nat. Pfl. Fam. ed. 2, 19a (1931) 123, f. 55 & 56. — C.G.G.J. VAN STEENIS (1985). Vegetative anatomy. The leaves of /xonanthes have paracytic stomata, a simple vascular strand in petiole and midrib with sclerenchyma forming a complete cylinder enclosing paren- chymatous ground tissue adaxially, and rhomboidal crystals and druses, as more or less constant characters. Variation has been recorded in presence or absence of anticlinal division walls of the epidermal cells, mucilage cells and extent of the vascular bundle sheaths of the minor veins. This diversity partly coincides with the boundaries between the sections Brewstera and Ixonanthes. The wood of Lxonanthes is characterized by solitary vessels with simple perforations, fibres with numerous distinctly bordered pits, largely apotracheal axial parenchyma bands of 2—S cells wide, and narrow 1—3(—4)-seriate, weakly heterogeneous rays. The vessel-ray pits are large and simple. The above characters are in fairly good agreement with the treatment of /xonanthes, together with Allantospermum, Cyrillopsis, Ochthocosmus, and Phyllocosmus in one family of the Linaceae alliance. Allantospermum would fit here better than in the /rvingiaceae, although its wood stands out in the /xonanthaceae on account of its minute, half-bordered vessel-ray pits. References: Buraess, Sabah For. Rec. 6 (1966) 247—248; Descu, Mal. For. Rec. 15 (1957) 156—157; MeTCALFE & CHALK, Anatomy of the Dicotyledons | (1950) 268-273; METCALFE, Lescor & Lopreau, Adansonia sér. II, 8 (1968) 337-351; Roso, Adansonia sér. II, 8 (1968) 73—83; vAN WeLzeN & Baas, Blumea 29 (1984) 453-479. — P. Baas. Palynology. Pollen of xonanthaceae, which is described by LoBREAU (1969: 526, Cyrillop- sis) and OLTMANN (1971), is subprolate to prolate (Cyrillopsis, Ochthospermum) or subspherical (Ixonanthes), and measures from 27 to 52 4m. The apertural system is always tricolporate. The colpi are long and the endoapertures have costae on their polar sides. Exine stratification can easi- ly be observed. Tectum and nexine are thin, while the infratectal layer consists of long columellae, (621) 622 FLORA MALESIANA [ser. I, vol. 103 The thickness of this columellate layer clearly exceeds that of tectum and nexine together. Sculp- ture is mostly somewhat scabrate. Cyrillopsis has a rugulate or striate exine. Pollen of Ixonanthes is characterized by the presence of distinct supratectal spines. A/lantospermum, which genus was at first included in /xonanthaceae, has tricolporate pollen with endoapertural costae, long col- umellae, and a rugulate to striate sculpture. As for its pollen morphology, the genus A//antosper- mum is considered to be related to Ixonanthaceae by BORTENSCHLAGER C.S. (1966), METCALFE C.S. (1968) and OLTMANN (/.c.) and to Simaroubaceae (especially to Jrvingia) by MULLER (1972), which latter opinion is accepted here. METCALFE c.s. (/.c.) consider pollen of Ixonanthaceae, Allan- tospermum, and Irvingiaceae together more similar to Simaroubaceae pollen than to that of Linaceae. However, OLTMANN (/.c.) concluded on pollen morphological evidence that [xonan- thaceae (Allantospermum excluded) are nearest to Erythroxylaceae and also, but to a lesser ex- tent, related to Linaceae. References: BORTENSCHLAGER, ERDTMAN & PRAGLOWSKI, Bot. Notis. 119 (1966) 160—168; LoBREAU, Pollen et Spores 11 (1969) 499-555; METCALFE, LEscot & LoBREAvU, Adansonia sér. II, 8 (1968) 337—351; MULLER, Fl. Males. I, 6 (1972) 972; OLTMANN, Pollenmorphologisch- systematische Untersuchungen innerhalb der Geraniales. Diss. Bot. 11 (1971). — R.W.J.M. vAN DER HAM. 1. IXONANTHES Jack, Mal. Misc. 2, 7 (1822) 51; ENpi. Gen. Pl. (1840) 1055, ‘Ixionanthes’; HaLL.f. Beih. Bot. Centralbl. 39, 2 (1923) 6; HUB. WINKLER in E. & P. Nat. Pfl. Fam. ed. 2, 19a (1931) 124, f. 55, 56A—E; Kooi, Blumea 26 (1980) 195. — Em- menanthes Hook.f. & ARN. Bot. Beech. Voy. (1836) 217. — Brewstera M.J. RoEMER, Syn. Monogr. | (1846) 141. — Pierotia BLume, Mus. Bot. Lugd. Bat. 1 (1850) 179; ibid. 1 (1851) 396. — Discogyne SCHLTR, Bot. Jahrb. 52 (1915) 123. — Fig. 1-3. Evergreen, buttressed trees or treelets growing monopodially with flushes, glabrous. Branches ascending. Bark brownish, finely fissured, rich in tannin; younger parts smooth. Stipules caducous, free, scale-like, about obliquely tri- angular, entire, acutish, glabrous. Leaves simple, spirally arranged, with slight- ly incrassate, entire or glandular-serrate margin, pinnately nerved, mostly obovate, tapering towards the base into a short petiole; midrib sulcate above, prominent beneath; venation obscure or slightly prominent on both surfaces; reticulations fine, irregular. Inflorescences axillary, dichasially corymbose. Peduncle smooth, glabrous; primary axes paired or sub-4-whorled, secondary axis paired, otherwise as the primary ones but usually much shorter; bracts scale-like, triangular, persistent, acutish, entire, smooth; bracteoles absent. Flowers bisexual, 5-merous, actinomorphic, perigynous. Young buds sticky. Calyx and corolla indurated and persistent in fruit, increasing in size. Sepals connate for up to 0.2 of their length, quincuncial, glabrous, fleshy to coriaceous in fruit. Petals quincuncial, almost free, glabrous, distinctly veined. Stamens 10 or (15—)20, in one whorl; filaments inserted outside and against the disk, ir- regularly coiled in bud, glabrous, subpersistent; anthers (basi-)dorso-versatile with a large peltate connective, introrse with 2 longitudinal slits, finely verru- 1988] IXONANTHACEAE (Kool) 623 Fig. 1. View on /xonanthes forest, seen from the resthouse at Malili, Central Celebes (Photogr. J. vAN ZULL DE JONG, 1933). cose. Disk well-developed, bowl-shaped, with free margin, erect, entire or slightly lobed. Ovary superior, 5-celled, glabrous. Ovules 2 per locule, axile, one on each margin of the carpel, collateral, pendulous, epitropous. Sty/e 1, ir- regularly coiled in bud, either elongating to 25 mm or remaining very short (up to 2.5 mm in /. petiolaris), glabrous, subpersistent in fruit; stigma mushroom- shaped, margin slightly lobed, fleshy. Capsule septicidal and septifragal, 5-celled, 5-valvate, without a central column, ovoid-conical or ellipsoid, acute or obtuse, glabrous, smooth; valves sometimes ultimately apically bifid, in transverse section W-shaped; exocarp sometimes fibrous and subtended by a thin membrane (hypoderm) folding along the septs and apparently into the 5 epicarp clefts; mesocarp dark, spongy; endocarp glossy. Seeds 1 or 2 per cell, either with a basal wing or with a suprahilar arillode; testa tenacious, brown; endosperm spongy, white, containing oil; embryo straight, + half as long as the dorso-ventrally appressed, + asymmetrically elliptic cotyledons. Germination epigeal. Distr. In continental Southeast Asia and Malesia 3 spp. (absent in Java, the Lesser Sunda Islands, and the Moluccas). Ecol. Primary forest on various soils below 1000 m 624 FLORA MALESIANA [ser. I, vol. 103 ; &. i illode, x 15: h, stipules, x2 (KEP FR] 31211) . 1988] IXONANTHACEAE (Kool) 625 Field notes. According to CORNER (Wayside Trees, 1940) the withering leaves are characteristic ochre- brown. Uses. Though they may be sizeable trees, the timber is of little value and has no features for special pur- poses. KEY TO THE SECTIONS AND THE SPECIES 1. Petiole 2-3 mm. Leaf margins slightly glandular-serrate. Primary branches of the inflorescence sub-4-whorled. Stamens (15—)20. Ovules and seeds with a suprahilar arillode, without a basal wing. — PPP EEAIUSION TL ns oe ele i Soc sk we eee CRT COE Oe EIR Stet. Sees 1. I. icosandra 1. Petioles 10-25 mm. Leaf margins entire, eglandular. Primary branches of the inflorescence paired. Stamens 10. Seeds with a basal wing, suprahilar arillode absent. Sect. Ixonanthes. 2. Inflorescence dense. Flowers at anthesis 1.5—3 by 1—2.5 mm diam. Style 1.5—2.5 mm. Fruit at most 1.5(—2) cm long. Valves ultimately apically 1-2 mm deep bifid. Seeds 10—13 by 3—4 mm 2. I. petiolaris 2. Inflorescence lax. Flowers at anthesis 3—5 by 2—7 mm. Style 15—20 mm. Fruit (2—)3—4(—4.5) cm long. Boaes mot bifid. seed, 18—20 by: 4=9.mm. 2. ates 209 baer Was bles ous Reet ele 3. I. reticulata 1. Section Brewstera (M.J.ROEMER) HALL.f. Beih. Bot. Centralbl. 39, 2 (1923) 7. — Brewstera M.J.ROEMER, Syn. Monogr. | (1846) 141. Young inflorescence axes, petioles efc. not glaucous. Lenticels inconspicuous. Petiole pul- vinate. Leaf margin slightly glandular-serrate. Peduncle slightly flattened and distally faintly grooved with four ridges; primary branches sub-4-whorled around the usually developed terminal flower of the peduncle. Disk entire. Stamens (15—)20. Ovules 2 per cell, usually only one fertile. Capsule ovoid-conical, acute. Seeds with a suprahilar arillode, without a basal wing; arillode as long as the seed, fleshy, cream-coloured. 1. Ixonanthes icosandra Jack, Mal. Misc. 2, 7 (1822) 53; Mra. Fl. Ind. Bat. 1, 2 (1859) 494; Illust. (1870) 68, incl. var. cuneata Miq.; Hook.f. Fl. Br. India 1 (1874) 416; Kina, J. As. Soc. Beng. 62, ii (1893) 191; Hocure. Pl. Bogor. Exsic. (1912) 40; GuILLAUMIN, Fl. Gén. I.-C. 1 (1911) 584; Rrptey, Fl. Mal. Pen. 1 (1922) 325, f. 31, incl. var. obovata RipLey; HALL./. Beih. Bot. Centralbl. 39, 2 (1923) 7; CoRNER, Wayside Trees (1940) 221, f. 60; GumLAumin, FI. Gén. I.-C. ed. 2, 2, 1 (1945) 501; Merr. J. Arn. Arb. 33 (1952) 227; CockBuRN, Tree Fl. Mal. | (1972) 307; Koo, Blumea 26 (1980) 197, f. 2. — Brewstera cre- nata M.J.Roemer, Syn. Monogr. | (1846) 141, nom. illeg. — Pierotia lucida Biume, Mus. Bot. Lugd. Bat. 1 (1850) 180. — /. lucida Biume, ibid. | (1851) 396. — I. dodecandra Grirr. J. As. Soc. Beng. 23 (1854) 632, t. 1 (‘subdodecandra’). — Macharisia isosandra PLancu. [in Herb. Hook.] ex Cuoisy, Mém. Soc. Phys. & Hist. Nat. 14 (1855) 168, nom. inval. — I. cuneata Mig. Suppl. (1896) 484. — J. obovata Hook.f. Fl. Br. India 1 (1874) 417. — Fig. 2. Trees or treelets up to 30 m, bole up to 1.3 m diam. Leaf scars + orbicular to triangular. Stipules up to 0.6 mm long. Leaves oblong to slightly obovate ob- long, 6-19 by 3—6.5 cm, pergamentaceous, base tap- ering; apex obtuse, often retuse, apiculate; petiole 2—3 mm, pulvinate. Peduncle 6—14 cm; pedicels 3—15 mm; bracts up to 5 mm long. Flowers at an- thesis 2—3 by 1-2 mm diam. Sepals elliptic, 1—1.5 by 0.8—1 mm (in fruit enlarging to 1.5—2.5 by 1—1.5 mm), c. 0.3 mm thick at the base, margin + hyaline, subcoriaceous in fruit. Petals orbicular, 2—2.5 by 2—2.5 mm (in fruit enlarging to 3—4 by 3—4 mm), subcoriaceous in fruit. Fi/aments ultimately up to 15 mm long; anthers basi-dorso-versatile. Style up to 10 mm. Ovary top-shaped, + S-angular, c. 0.5 by 0.7 mm. Capsule 15(—20) by 5-6 mm. Seeds ellipsoid, 10 by 2 mm; arillode adaxillary, tripartite. Distr. Thailand; in Malesia: Sumatra and Malay Peninsula, Ecol. Primary and secondary forests on slopes and ridges, 0-600(—900) m. Field notes. Bole straight, sometimes with short stiltroots. Bark smooth or slightly shallowly fissured, red, fawn, brown, or grey, soft. Inner bark red, purplish red, reddish brown, or brown, Wood white, orange, yellow, brown, cream-brown, or brownish red, hard. Crown conical, dense. Flowers whitish, yellow, green, sticky. Fruits green to dark brown, sticky. 626 FLORA MALESIANA [ser. I, vol. 10° beluks, Banka, buah tui, pagar anak, sankau merah, Malaya, injau belukar, Pahang, jenjulang, men- julong, punggong kijang, Kedah. Uses. Sometimes used for house-building. Vern. Kayu leja-leja, Sum. E. Coast, kassi branah, kayu ratuh, pempaaga, Palembang, kayu 2. Section [xonanthes Emmenanthes Hook.f. & ARN. Bot. Beech. Voy. (1836) 217. — Ixonanthes sect. Emmenanthes HAL..f. Beih. Bot. Centralbl. 39, 2 (1923) 8. Young inflorescence axes, petioles, efc. glaucous. Lenticels punctiform or slit-like. Petiole flat- tened, upper side more or less deeply sulcate, sometimes very narrowly winged. Leaf margin slightly incrassate, entire, eglandular. Peduncle angular to flattened, not grooved; primary branches of inflorescence paired, terminal flower of the peduncle usually developed. Disk entire or sometimes slightly 10-lobed. Stamens 10. Ovules 2 per cell, usually both fertile. Capsule short- or long-ellipsoid, obtuse. Seeds with a basal wing, without a suprahilar arillode; wing oblong, fairly stiff, concolorous, with a distinct dark-coloured raphe. 2. Ixonanthes petiolaris BLUME, Mus. Bot. Lugd. Bat. 1 (1851) 396; Mia. Fl. Ind. Bat. 1, 2 (1859) 494; Illust. (1870) 69; KooL, Blumea 26 (1980) 199, f. 1d, 3b. — Pierotia reticulata BLUME, Mus. Bot. Lugd. Bat. 1 (1850) 180; ibid. 1 (1851) 396, non I. reticulata Jack. — J. multiflora StapF ex RIDLEY, Kew Bull. (1930) 75. — I. philippinensis ELMER, Leafl. Philip. Bot. 10 (1939) 3758, descr. angl. — Fig. 3. Trees or treelets up to 30m, bole up to 50 cm diam. Young bark with punctiform lenticels, older parts striped with numerous lanceolate lenticels. Leaf scars orbicular with 3—5, sometimes distinct vascular scars. Stipules up to 0.5 mm. Leaves elliptic-oblong, 6—15 by 3—7.5 cm, pergamentaceous to subcoria- ceous; base acute; apex slightly obtuse; petiole 1.5—2 cm. Inflorescences densely flowered. Peduncle 3.5—7(—9.5) cm; pedicels c. 5 mm; bracts up to 1 mm. Flowers at anthesis 1.5—3 by 1—2.5 mm. Sepals elliptic to orbicular, 1—1.5 by 1—1.5 mm (in fruit en- larging to 1.5—2 by 1.5—2 mm), thickened at base, laterally with a c. 1.5 mm wide hyaline band; fleshy in fruit. Petals orbicular to elliptic, 2—2.5 by 1—1.5 mm (in fruit enlarging to 2.5—3 by 1.5—2 mm), thick- ened at base, margin narrowly hyalinous, charta- ceous in fruit. Filaments ultimately up to 15 mm; an- thers dorso-versatile. Sty/e up to 2.5 mm. Ovary flat- tened globose, c. 1.5 by 0.5 mm. Capsule short- ellipsoid, 1.5(—2) by 0.8(—1.2) cm, valves ultimately apically 1-2 mm deep bifid; septa after dehiscence (long-)persistent, adaxially connate with the adjacent ones. Seeds 1—1.3 by 0.3—0.4 cm. Distr. Thailand; in Malesia: Sumatra, Malay Peninsula, Borneo, Philippines (Luzon, Sulu Is.), and Central Celebes. Perhaps also in New Guinea; see the note under J. reticulata. Ecol. In primary and secondary forests on gran- itic sand and on slopes and ridges, 0—800 m. Field notes. Bole deeply fluted, buttresses grad- ually merging into the bole. Outer bark smooth, green, red, pale brown, yellowish brown, or black, flaking in small pieces, minutely ridged. Inner bark orange-whitish, yellow, pink, red, or redbrown, granular, sticky, soft. Wood white or reddish brown. Sapwood white, pinkish white, honey-coloured, yellow, or brown, with distinct lamination. Crown large, spreading, medium dense. Flowers cream to green; calyx green; stamens white. Fruits green to brown. Vern. Mara jening, meribikang, Sumatra; tinjau laut, Sum. W. Coast; kayurdori bunga, Sum. E. Coast; gerungang, jurung, Malaya; inyang burong, N. Sembilan, Selangor; inggi burong, pinang pinang, Borneo. 3. Ixonanthes reticulata JAcK, Mal. Misc. 2, 7 (1822) 51; Mia. Fl. Ind. Bat. 1, 2 (1859) 494; Illust. (1870) 69; Hance, J. Bot. 14 (1876) 243; Kina, J. As. Soc. Beng. 62, ii (1893) 192; RrpLey, Fl. Mal. Pen. 1 (1922) 326; HALL.f. Beih. Bot. Centralbl. 39, 2 (1923) 9; CORNER, Wayside Trees (1940) 222, f. 60; MERR. J. Arn. Arb. 33 (1952) 228; BRowneE, For. Trees Sa- rawak Brunei (1955) 175; CocKBuURN, Tree Fl. Mal. 1 (1972) 307; Koo, Blumea 26 (1980) 200, f. 3A. — Hypericinea macrocarpa WALL. Cat. (1831) 4833, nomen. — Gordonia decandra Roxs. FI. Ind. ed. Carey 2 (1832) 573. — Emmenanthus chinensis Hook.f. & ARN. Bot. Beech. Voy. (1836) 217. — J. chinensis (Hook.f. & ARN.) CHAMP. in R.Br. Proc. Linn. Soc. 2 (1850) 100, and many later authors. — I. khasiana Hook.f. Fl. Br. India 1 (1874) 416. — /. hancei PIERRE in Laness. Pl. Util. Col. Fr. (1886) 306. — I. cochinchinensis PreRRE, Fl. For. Cochin. 4 (1893) t. 284A; GuiLLauMIn, FI. Gén. I.-C. 1 (1911) 584; ed. 2, 1 (1945) 501. — J. grandiflora Hocnr. Pl. 1988] IXONANTHACEAE (Kool) 627 Ixonanthes Fig. 3. florescence, x2; b. flower, x 6 (SAN 40370). petiolaris BLUME. a. In- Bog. Exsicc. (1904) 39; Merr. Enum. Born. (1921) 313; Heyne, Nutt. Pl. Ned. Ind. (1927) 854. — Dis- cogyne papuana ScuiTr, Bot. Jahrb. 52 (1915) 123. — I. longipedunculata Merk. Philip. J. Sc. 17 (1921) Bot. 264. — J. crassifolia Ha... Beih. Bot. Cen- tralbl. 39, 2 (1923) 10. — /. beccarii HALLS. Lc. — I, petiolaris (non BLuMe) HALL. Lc. 8, pro syn, — I. grandifolia Riptey, Kew Bull. (1930) 74. — /. pa- puana (SCHLTR) Hus. WINKLER in E. & P. Nat. Pf. Fam. ed. 2, 19a (1931) 126, t. 55, 56 A—E. Treelet or tree up to 40 m, bole up to 1 m diam. Lenticels slit-like or punctiform. Leaf scars round to ovate. Stipules up to 1 mm. Leaves elliptic, elliptic oblong, or slightly ovate, entire, 5.5—14 by 3—10 cm, pergamentaceous to coriaceous; base acute, apex slightly obtuse to emarginate; petiole 1—2.5 cm. Jn- Jlorescences lax. Peduncle (3—)5—8(—14) cm; pedi- cels c. S mm, bracts up to 1 mm. Flowers at anthesis 3—5.5 by 2—7 mm. Sepals elliptic, 4—5 by 3—4 mm (in fruit enlarging to c. 8 by 4mm), within from top to bottom with a slightly thickened and paler band, fleshy to coriaceous in fruit. Perals orbicular to ellip- tic, 4—5.5 by 3—4 mm (in fruit enlarging to c. 10 by 6 mm), within from the base upwards thickened by a bundle of nerves, margin broadly hyaline, sub- coriaceous in fruit. Filaments ultimately up to 2 cm. Style up to 2 cm. Ovary globose, c. 3 by 2 mm. Cap- sule long-ellipsoid, (2—)3—4(—4.5) cm long, valves not apically bifid; septa after dehiscence (long-)per- sistent, connate with the adjacent ones. Seeds 1.8—2 by 0.4—0.9 cm. Distr. From Assam through Indochina to S. China (Kwangsi, Kwantung, Yunnan, Kweichow, Hainan); throughout Malesia (but not in Java and Lesser Sunda Islands, and not yet collected in the Moluccas). Ecol. Primary (rarely secondary) forests, fre- quently on hillsides and ridges, often on sand and granite, sandstone and kerangas, in swamp and heath forest, largely below 500 m, but occasionally found as high as 1000 m. Field notes. Bole straight and fluted. But- tresses narrow, small. Bark smooth, lengthwise fis- sured, scaly or cracked, pink, yellowish, brown, fawn, or grey to blackish. Inner bark red, light brown, or dark orange brown, granular, soft. Wood white, dirty white, or dirty yellowish, hard, heavy. Sapwood ochre, with white or yellow lamination, medium hard. Flowers white or greenish. Stamens yellowish-brown. Ovary brown, style green. Vern. Obah, Malaya, jinjagong, sansak china, Penang, ingeran or inggi burong, nyiran burong, pagar anak, sakit hudang, Selangor, angaran buron, Trengganu, djurung, Palembang, sentulu, N. Borneo, /angudai, perepat rimba, Sabah, kanju Junong, Brunei, redin, S. Sarawak, /ura, Celebes, keseruok, New Guinea, Tehid lang. Note. This species is the only one with certainty found in New Guinea. As identification of sterile col- lections is not possible, such collections from New Guinea are tentatively included in /. reticulata, but might belong in part to /. petiolaris. mit Bau ath al aw) wv Veo My saiie, roviav enol itp (¢.b— dh 7, | '29Q(-S0p bande Bed tetin Gaeta yi fad ~ ‘Bah Roce pete hei a Wal) aa iid. (a TH) ; i 0 ay dai tiled shawty) ree noes | iu MeN AipT eee hie emt) Bid geal at fog iif 208. Wogan uly oor ec tt rsa ete wietildsh id end, Way) 2798, Azer. ary! PeADYil 7 venwun th, Sheard).: + ee Dan ah t wggilnrne re Se eg a a y anne nm eg + pyres ee a ‘ie ay ye ; CTENOLOPHONACEAE (A.M.N. van Hooren & H.P. Nooteboom, Leiden) The systematic place of the tropical lowland rain-forest tree Ctenolophon OLIVER has a chequered history. Originally it was referred to affinity with Olacaceae (OLIVER, 1873; MASTERS, 1875; ENGLER, 1889; BaILLon, 1892) or Jcacinaceae (BECCARI, 1877). HALLIER f. (1912, 1918) held another view and arranged the genus in the Celastrales, deriving this group from Linaceae. HUTCHINSON (1959, 1973) referred the genus to the Malvales. In a meticulous exposé of the anatomy, flower and fruit structure, PIERRE (1893) concluded that the affinity of Crenolophon is with the Linaceae and he was followed by ENGLER (1907), EXELL (1927), and Hus. WINKLER (1931). At present Ctenolophon is almost unanimously recognized as belonging to the Linaceous af- finity, together with /xonanthaceae. Within Linaceae HuB.WINKLER (1931) had raised the genus to the rank of a monogeneric subfamily. Later ExELL & MENDONCA (1951) recognized it as repre- senting a family of its own, a view now almost unanimously accepted, as fully discussed by VAN HOoOREN & NooTEBOOM (1984). In the treatment of the family Linaceae (page 607, see there) the families Linaceae, Ixonan- thaceae, and Ctenolophonaceae are opposed by concise diagnoses. On account of studies of special features some other affinities have been put forward. HEIMSCH (1942) found a remarkable resemblance in xylem structure with Humiriaceae. CRONQUIST (1981) referred to Cteno/ophon as an aberrant member of Hugoniaceae, a view with which we cannot agree. VAN WELZEN & Baas (1984) compared the leaf anatomy of Ctenolophon with that of Hu- miria and some Malpighiaceae and found that it is very different from both Humiriaceae and Malpighiaceae, but also from other Linaceae, and they supported the status of a family of its own, adding that close affinities are still unknown. References: BAILLON, Hist. Pl. 11 (1892) 445; Beccart, Malesia 1 (1877) 119; CRongutstT, Inte- grated system efc. (1981) 759; ENGLER, Nat. Pfl. Fam. 3, 1 (1889) 237; ibid., Nachtr. 3 (1907) 204; ExeE.L, J. Bot. 65 (1927) Suppl. 1: 50; Exett & MENDoNGA, Conspectus flora Angolensis 1, 2 (1951) 248, 392; HALLER f. Arch. Néerl. III B, 1 (1912) 109; Meded. Rijksherb. 35 (1918) 28; Hermscu, Lilloa 8 (1942) 83; van HoorEN & NootTesoom, Blumea 29 (1984) 547; HUTCHINSON, Fam. Fl. Pl. 1 (1959) 265; ed. 3 (1973) 324; Masters, Fl. Br. India 1 (1875) 577; Outver, Trans. Linn. Soc. 28 (1873) 516; Prerre, Fl. For. Coch. (1893) t. 281; VAN WELZEN & Baas, Blumea 29 (1984) 477; Hus. WINKLER in E. & P. Nat. Pfl. Fam. ed. 2, 19a (1931) 122. 1. CTENOLOPHON OLIVER, Trans. Linn. Soc. 28 (1873) 516; HUB.WINKLER in E. & P. Nat. Pf. Fam. ed. 2, 19a (1931) 122; BULLock, Kew Bull. 14 (1960) 41; VAN HoorEN & Nootesoom, Blumea 29 (1984) 547. — Fig. 1, 2. Trees. Hairs stellately tufted and simple. Stipules interpetiolar, caducous. Leaves entire, opposite, petiolate, simple, pinnately nerved. /nflorescence an axillary or terminal cymoid panicle. Bracts present, bracteoles absent. Flowers bisexual, 5-merous, actinomorphic, hypogynous. Sepals basally shortly con- nate, quincuncially imbricate, subequal, indurate, swollen and persistent in fruit, with stellate hair tufts. Petals free, contorted, caducous, often shortly clawed. Disk extrastaminal. Stamens 10, free, inserted halfway the disk, alternately longer and shorter, the longer epipetalous and the shorter episepalous; anthers dorso-versatile, 2-celled, introrse; connective protruding, acute-triangular. Extrastaminal nectary glands absent. Ovary superior, 2-celled; style 1, apically (629) 630 FLORA MALESIANA [ser. I, vol. 10° forked with 2 capitate stigmas; ovules 2 per cell, axile, collateral, pendent, epi- tropous. Fruit a 1-celled capsule, the woody pericarp finally lengthwise splitting into 2 valves. Seed 1, persisting after falling of pericarp and pendulous from the top of a filiform columella; arilloid papillose, surrounding lower half of the seed. Endosperm copious. Embryo straight. Distr. Two species, one (C. engleriana Mi.ppr.) in West Africa (Angola, Zaire, Nigeria, Gabon) and one throughout Malesia (but not in Java and Lesser Sunda Islands and not yet reported from Celebes and Moluc- cas, where it is expected to occur). For Malesia see fig. 3. Ecol. Mixed lowland rain-forest. Vegetative anatomy. Cfenolophon has glabrous leaves, but the young shoots and floral parts have tufted, stellate hairs. Stomata are anisocytic to anomocytic. Crystals are mainly solitary and rhomboidal, more rarely clusters intergrading with druses. Crystalliferous bundle sheath cells have unilateral sclerified thickenings (cristarque cells). The petiole and midrib have a simple collateral vascular strand. The wood of Ctenolophon is characterized by solitary vessels with scalariform perforations, fibres with dis- tinctly bordered pits, parenchyma which is scanty paratracheal and diffuse-in-aggregates, and heterocellular 1—3-seriate rays. The above attributes are not very helpful in determining the phylogenetic affinity of this monogeneric fami- ly Ctenolophonaceae. Its stomatal type removes it from the Linaceae complex, but other leaf anatomical char- acters are of common occurrence throughout the dicotyledons, including the Linaceae. The wood anatomy of Ctenolophon is very plesiomorphic and cannot therefore be used to support or reject various suggestions of natural affinity of the genus, although similarities with Humiriaceae have been pointed out by HEIMSCH & TSCHABOLD (1972) and METCALFE & CHALK (1950). References: HEIMSCH & TSCHABOLD, Bot. Gaz. 133 (1972) 242—253; METCALFE & CHALK, Anatomy of the Dicotyledons 1 (1950) 268—273; Saab, Bot. Notis. 115 (1962) 44—57; vAN WELZEN & Baas, Blumea 29 (1984) 453-479. — P. BAAS. Palynology. Pollen of Ctenolophonaceae shows two types: the Crenolophon englerianus type and the C. parvifolius type. Pollen grains of both types measure 31—66 um. The apertural system is zonocolporate (‘stephanocolporate’), the number of apertures ranging from 3-8 in the parvifolius type and 5—9 in the engle- rianus type. Contrary to the rounded subspherical parvifolius type, the englerianus type shows distinct thick- enings on its mesocolpia and polar areas, making this type characteristically angular and barrel-shaped. Exine thickness ranges from 3—6 um. Exine stratification is rather obscure in the light microscope and dif- ficult to determine, especially in the englerianus type. A thick nexine can mostly be observed. The infratactal layer appears labyrinthine, sometimes exhibiting granulate-columellate structures. The tectum is psilate or finely perforate in the englerianus type, perforate to foveolate in the parvifolius type (THANIKAIMONI C.S., 1984). SAAD (1962) noted striking resemblance with pollen of some Malpighiaceae, which led him to the conclusion that Ctenolophon may be considered as related to Malpighiaceae. Pollen morphological affinity to Linaceae and Humiriaceae is regarded as respectively being absent and not close. Here, it is put forward that pollen of Ctenolophon is neither close to that of Malpighiaceae, as the bipartite nature of wall stratification in both Ctenolophon and some Malpighiaceae noted by Saab (/.c.) actually represents only superficial similarity. Pollen of Ctenolophon cannot be convincingly connected with that of other families, which supports attribu- tion of family rank to the taxon. Fossils. The distinctness of Ctenolophon pollen is illustrated by its extensive fossil record. Both types can be easily recognized (MULLER, 1981; THANIKAIMONI C.S., /.c.). The englerianus type (Ctenolophonidites) oc- curred in Africa nearly uninterruptedly from the Upper Cretaceous onwards, and can be linked with the recent area of distribution of C. englerianus. The type was also present in South America (Paleocene, Eocene) and India (Eocene, Miocene), but disappeared from those areas in the course of the Tertiary. In recent time the englerianus type also exists in the Philippines on Samar, in C. parvifolius (THANIKAIMONI Cc.s., l.c., as C. philippinensis, PNH 6385). This Samar material is distinct from that of Africa by having mostly less apertures (5 to 7 versus 5 to 9) and by showing a perforate rather than a psilate or finely perforate tectum. Another collection from Samar (PNH 6146) shows pollen which seems to be intermediate between PNH 6385 and the parvifolius type (original observation). The parvifolius type (Retistephanocolpites) was first recorded from the Paleocene of Africa. In India it oc- curred in the Paleocene and the Eocene, while it is known from Borneo starting from the Upper Eocene con- tinuing up to the present (Mor.LEy, 1977; manuscript comm. MorRLEY). 1988] CTENOLOPHONACEAE (van Hooren & Nooteboom) 631 The fossil record of both types clearly demonstrates the relic nature of the present area of distribution of Ctenolophonaceae. It points to separation of the two types in the Upper Cretaceous in Africa (GERMERAAD c.S., 1968: 276). However, the Samar collections may reflect another, much more recent as well as easterly contact between both types. Possibly, the Indian eng/erianus stock did not become extinct in the Miocene be- fore branching off into the Malesian area. Probably the Indian parvifolius stock branched off into Malesia as early as the Eocene, becoming extinct in India at the Eocene-Oligocene transition. Macromorphological difference between recent C. englerianus and C. parvifolius is very small. The occurrence of intermediate pollen between the englerianus and the parvifolius type is suggestive of introgression between the two species, although independent origin of the englerianus type on Samar cannot be excluded. References : GERMERAAD, HoppinG & MULLER, Rev. Palaeobot. Palynol. 6 (1968) 189-348; Mor ey, Proc. Indon. Petr. Ass., 6th Ann. Conv. (1977) 255—276; MULLER, Bot. Review 47 (1981) 73—74; Saab, Bot. Notis. 115 (1962) 49—57; THANIKAIMONI, CARATINI, VENKATACHALA, RAMANUJAM & Kar, Trav. Sect. Sci. et Techn. Inst. Frang. Pondichéry 19 (1984) 23, 80. — R.W.J.M. VAN DER Ham. Notes. 1. The African species C. englerianus Mitpprr. is hardly different from the Malesian species, C. parvifolius. The only differences observed are the simple cymose panicle and the stamens being up to 15 mm long in C. englerianus, versus the compound cymose panicle and the stamens up to 10 mm in C. parvifolius. 2. The fruit and seed structure was wrongly described by HUTCHSINSON and by Hus. WINKLER, who said that the seed dangles from a long funicle. The so-called funicle, however, is a filiform columella on which the seed is attached apically. 1. Ctenolophon parvifolius OLtver, Trans. Linn. Soc. 28 (1873) 516, t. 43, f. 1—7; Mast. Fl. Br. India 1 (1875) 577; BEcc. Malesia 1 (1877) 120; Kina, J. As. Soc. Beng. 62, ii (1895) 106; RipLey, FI. Mal. Pen. 1 (1922) 423; Burk. Dict. (1935) 695; ed. 2 (1966) 705; Descu, Man. Mal. Timbers 2 (1954) 15; BuLLock, Kew Bull. 14 (1960) 41; Mewer, Bull. Herb. For. Dep. Sandakan 10 (1968) 151, fig.; Cock- BURN, Gard. Bull. Sing. 24 (1969) 6; Tree Fl. Malaya 1 (1972) 306; Sabah For. Rec. 10 (1976) 205; ANDER- son, Checklist Trees Sarawak (1983) 239; VAN HooreEN & NooTesoom, Blumea 29 (1984) 547. — C. grandifolius OutveR, Trans. Linn. Soc. 28 (1873) 517, t. 43, f. 8—10; Mast. FI. Br. India 1 (1875) 577; Becc. Malesia | (1877) 119; Kina, J. As. Soc. Beng. 62, ii (1895) 106; Rrptey, Fl. Mal. Pen. 1 (1922) 423. — C. philippinensis Hat..f. ex SCHNEIDER, Philip. Bur. For. Bull. 14 (1916) 127, nomen; MerRR. Enum. Philip. 2 (1923) 326. — Fig. 1, 2. Tree, 12-40 m, 15—120 cm diam., buttressed. Twigs with distinct leaf-scars and thickened at the in- sertion of the buds, glabrous. Innovations marginal- ly and apically tufted-hairy, hairs to 1 mm long. Stip- ules boat-shaped, triangular to ovate, 1.5—2.5(—3) by 1—1.5 mm; entire, apex rounded to emarginate. Leaves glabrous, narrowly ovate to elliptic, some- times obovate, often with + parallel margins, 5—15 by 2—6 cm; base broadly cuneate, sometimes round- ed; margin often + thickened; apex broadly acute to obtusely acuminate, sometimes rounded, acumen to 2.5 cm; nerves (5—)7—10(—12) pairs, slightly curving upwards to near the margin, often meeting in | (or 2) looped intramarginal veins; reticulation rather lax to dense; petiole sulcate above, 4—10(—15) mm. Pan- icles rather densely flowered, terminal and up to 7 to- gether, or 2—3 in the axil of a leaf near the end of the twigs, sometimes ramiflorous, up to 3. times branched, broadly ovoid to depressed obovoid, 3—9(—16) cm long. Axes patently tufted hairy, especially above the nodes. Bracts caducous just below the articulation, densely or sparsely appressed- ly to patently tufted-hairy without, especially on margin and midrib, more or less boat-shaped, nar- rowly ovate-liguliform to triangular, 1—3.5 by 0.5—1.7 mm; base with a row of bristles within. Pedi- cel articulate, grooved, swollen in fruit, 1—1.5(—2.5) mm. Hypanthium thickened. Sepals appressed, densely appressedly to patently tufted-hairy to (the inner 3 marginally) glabrous, glabrous within but in- ner base with a row of up to 0.8 mm long bristles, boat-shaped, obovate to depressed ovate, ciliate, 1—2.5 by 1.5—4.7 mm, the outer two smallest. Petals recurved and more or less twisted in anthesis, stout, thin-leathery, densely appressedly tufted-hairy with- out except the overlapping margin and the base, gla- brous within, slightly asymmetric, narrowly elliptic to narrowly ovate, 7-12 by 2—2.5 mm; claw absent or indistinct, at most 1 by 1 mm; margin sometimes with simple hairs towards its base. Disk membranous or slightly fleshy, cup-shaped, 1—1.5 mm high; mar- gin often more or less dentate to undulate. Stamens inserted halfway up to just below the margin of the disk, short filaments 3.7—6.5 mm, long filaments 4.5—7.5 mm. Pistil up to halfway the style covered with straight, up to 2 mm long simple hairs; style straight, rather stout, slightly flattened, equalling to up to 3 mm exceeding the long stamens, sometimes as long as the short stamens, 5~10 mm fong. Ovary + constricted at base, ellipsoid to ovoid, 1.5—3 by 1-2 mm. Fruit 1|-celled, densely appressedly tufted- hairy, ellipsoid to obovoid, 13-24 by 6-13 mm, sometimes basally curved; pericarp woody, ribbed, smooth inside, consisting of 4 layers, 1.5—2 mm diam. Seed ellipsoid to obovoid, 12—20 by 5-9 mm, 632 FLORA MALESIANA [ser. I, vol. 10° Fig. 1. Ctenolophon parvifolius Outver. a. Habit, x 0.5; b. innovations, with budscales, stipules and plicate leaves; c. flower, x 3; d. sepal from inside, x 12; e. flower (sepals and petals removed), x3; f. flower in LS, x5; g. fruit, x2; h. seed, x2 (a from OLIVER (1873), b Dumas 1513, c—f SAN 4595, g, h LAMBACH 1251). 1988] CTENOLOPHONACEAE (van Hooren & Nooteboom) 633 Fig. 2. Ctenolophon parvifolius O.tver. A large mature tree at Dolok Puhutan Lajan, Tapanuli, E. Central Sumatra (Photo G.A.L. DE HAAN, 1939; coll. no. 697). WE . are PY ; / ‘ oe 40. , hy c 4 ’ ~ 7 3 f . ’ Se %) f Ae ‘ J é ‘, i! Pe ? tre e % f o f° be: s é P ae ‘ , P . vs ; ‘_ Ge ) nh Tegks. on ys. | j oe ev — * el dangling from a filiform columella (10—)15—20 mm long; hilum apical, slightly protruding from testa, obtriangular, 4—7 by 2—2.5 mm; arilloid from slight- ly above the base up to around the hilum, oblong tri- angular, up to § by 5 mm, with hair-like papillae which are reddish when dry, surrounded by a gelatin- ous transparent layer. Testa 0.1—0.2 mm thick, outer layer smooth, crustaceous, dark olive-brown to pur- ple black when dry, sometimes fissured, finally cov- ered by a thin membrane; second and third layer red- brown, free from the outer wall. Endosperm fleshy, spongy. Embryo stout; cotyledons elliptic to obo- vate, 8—10.5 by 4.7—5 mm; radicle (1—)2—2.5 mm long. Distr. Malesia: Sumatra, Malay Peninsula, Bor- neo, Philippines (Samar, Leyte, Mindanao), and New Guinea (incl. Japen I.). Fig. 3. Ecol. Mixed primary forest, also in heath and swamp forest, indifferent to soils (latosols, humic podsols, loam, ultrabasic) and also frequent on sand (Banka, Borneo), locally common to sometimes very common in lowland and on hills, generally below 850 m, but in Borneo rarely up to 1650 m. Fi. /r. Feb.—Dec. Fig. 2. Field notes. Bole up to 35 m; buttresses, if present, up to 4 m high and wide up to 12.5 cm thick. Bark reddish brown to black, often greyish, very var- iable in appearance, scaly or cracked, often peeling off, hard. Inner bark redbrown to pink, granular or fibrous. Exudate colourless to iodine-coloured, clear, slowly appearing, thick, or absent. Heartwood often reddish or brown, rather hard, heavy, sinks in water. Corolla from white to yellow, orange or 130° mo". 150° to” M( be Sa _. ant & 1 ny SA 4 10" , s } ‘ ’ ‘ W Sc [ =p o4 ~ ’ -? _ v . - bd ro" ‘whe J f ts ae: 2 i We . ’ vm * - > *. , IV dé. 7 . iL '0' r ) P * i ‘ i ; 7 Syuct #° “ on, ? . ; 2 4 bert A . be 4 ~ - sey oh , se S| red “4 ee ro iS A Fig. 3. Localities of Crenolophon parvifolius OLiveRr, 634 FLORA MALESIANA [ser. I, vol. 10° bright red. Style pinkish. Fruit yellow to pink, later brown. Seed brown or glossy black; arilloid white to orange or brilliant vermilion. Uses. The hard and very durable timber is used for house construction and as general purpose timber (SCHNEIDER, 1916; RipLEy, 1922; Descu, 1954; BURKILL, 1966). Vern. Sumatra: babi kurus, batu, djambu ayer, jinkinai, kaju batu, k. bawang, kalek bung cung, klawar, medang tembatu, petjah pingan; Malay Peninsula: bunga tanjong hutan, kumus beruang, mata ulat, membatu hitam, perichat, Malaya; ban(g)kal, bankal paya, kas, Mersing; karu karu, kelabau, Perak; kelat, k. bukit, pompong paya, Pahang, kelat hitam, mertas, Kuala Lumpur, mertas kuning, Kuala Trengganu; jambu paya, Johore; Bor- neo: belama’a, tamana’a, Sarawak: litoh, Iban; besi, obah, Sabah; merandi, W. Kalimantan; jarmgin, kayu batu, latak manuk, E. & SE. Kalimantan. New Guinea: gun, Ilwur I.; Kodawu, kodowu, kotawu, In- gembit I.; mamdai, tora, Ormu dial., dnok, nook, Tehid I.; samber, sosopi, Japen dial. Volume Volume Volume Volume Volume Volume Volume Volume 1. Taxonomical Revision. 1959-1982. pp. (20) + xxiv + 600 L a Subscription orders and requests for All requests for further information conce specimen copies should be sent to: the Flora Malesiana should be sent to: Kluwer Academic Publishers Group i Kluwer Academic Publishers Distribution Centre P.O. Box 17 P.O. Box 322 3300 AA Dordrecht 3300 AH Dordrecht The Netherlands The Netherlands Printed in the Netherlands FLORA MALESIANA under the Auspices of Lembaga Biologi Nasional (Botanic Gardens of Indonesia), Bogor, Indonesia, and the Rijksherbarium, Leiden, Holland, executed by Foundation Flora Malesiana Scientific Communications concerning the Flora Malesiana should be addressed to the General Editor, Dr. W.J.J.O. de Wilde c/o Rijksherbarium, P.O. Box 9514 2300 RA Leiden, Netherlands Series I— Spermatophyta (Flowering Plants) 1. Cyclopaedia of collectors & collections. 1950. pp. clii +639 (microfiche edition) 4. General chapters and revisions. 1948—1954. pp. ccix + 631 (microfiche edition) 5. Bibliography, specific delimitation & revisions. 1955—1958. pp. cccxlii+ 596 are 6. Systematic revisions. 1960—1972. pp. 20+ 1023 (microfiche edition) PEG 7. Systematic revisions. 1971—1976. pp. 18 +876 p 8. Cyclopaedia of Collectors, Suppl. 2. Systematic revisions. 1974-1978. pp. 19+577 9. lesion revisions. 1979—1983. pp. 5 en Si Series II — Pteridophyta (Ferns & Fern Allies) KLUWER ACADEMIC PUBLISHERS DORDRECHT/BOSTON/LONDON