A REVISION OF THE GENUS HOPEA ROXBURGH OF THE PHIUPPINES

Hermes G. Gutierrez University 0/ Santo 7'011«18

[ - INTRODUCTION

The genus Hapea, which comprises one of the most important forest trees that we have in the Philippines, is the subject of this botanical investigation. Although quite a number of papers deal­ ing with this local genus have been published, the taxonomic delimi­ tation of each member species have never been fully well estab­ lished.

It is primarily due to the present chaotic status of the mem­ ber species of the genus Hopea in the Philippines, therefore, that after reviewing and evaluating the history and literature, and exam­ ining the extant plants in various herbaria and in the field, a revision of the genus is imperative.

The family DIPTEROCARPACEAE is represented throughout the world by seventeen genera and about four hundred species r37, p. ixJ. The recognized genera comprising the family are: Anisop­ tera., Balanocarpus, Cotylelobium, Dipterocarpus, Doona, Dryoba­ lanops, Hopea, Marquesia, Manotes, Parashorea, Pentacme, Shorea, Stemanoporus, Upuna, Vateria, Vateriopsis and Vaticu.

In the Philippines the family is represented by only eight genera [19, p. 243J: Anisoptera, BalanocaTpus, DipterocaTp1tS, Hopea, Pura­ shorea, Shorea and Vatica. The genus Hopea is the third largest genus in the Philippines, represented by nine species, including two new species. The largest genera are Shorea and Dipterocarpus, re­ presented by fifteen and eleven species, respectively.

The family DIPTEROCARPACEAE is distributed over a large area of tropical Africa and the Indo-Malayan region. It is not, however, ACTA MANILANA represented in tropical America or in tropical Australia. The mem­ bers of the family are found in Africa, Seychelles, India, Ceylon, Andaman Islands, China and Hainan, Burma, Indo-China (Cambo­ dia, Laos and Vietnam), Thailand, Malaya, Sumatra, Borneo, Java, Celebes, Moluccas, New Guinea and the Philippines. The genus Hopea, which comprises about eighty-six species through the world, is found throughout the range of the family, except in Africa, Sey­ chelles, China and Hainan.

In the Philippines there are seven species of Hopea reported by Foxworthy [19, pp. 273-284J Our paper, however, reports on nine species, all of which are endemic, two of which are new to science. They are distributed in almost all parts of the Philippines, from Calayan Island of BABUYANES in the north, to BasOan Island in the south.

The members of our genus Hopea form part of the commercial wood that is known in the trade as the Philippine Mahogany [39, p. 9]. The Philippine Mahogany is represented solely by the mem­ bers of the family DIPTEROCARPACEAE in the Philippines. They are the most important of our forest trC'cs, producing the greatest bulk of timber. They have a wide range of colors, height and hardness, which make them suitable for a varicty of uses. As a whole, thl' members of our family DIPTEROCARPACEAE (Lauan Family as it is known locally) produce one of the finest hardwoods of the world.

Almost all the species of commercial trees imported by Japan, the United States and other countries of the world from the Philip­ pines belong to the family DIPTEROCARPACEAE. Quite a number of papers dealing with the family has becn published, but the delimita­ tion of species from one to the other has never been fully well established, to the extent that disagreement prevails among foresters, botanists, and even betwef:n the two groups, regarding the position and identity of the member species. For this reason the author has chosen as a primer in the study of our family the genus Hopea, which is considered as one of the vaguely treated genera as far as our accumulated knowledge about the group is concerned.

This work endeavors, primarily. to delineate taxonomically the position and identity of the member species of the genus Hopea of the Philippines. It is done purposely with the thought in mind that this work will serve as a guide for foresters, botanists and people engaged in the marketing of lumber, to settle once and for A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. ACTA MANILANA

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AFRICA 2/34

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J ; Fig. i. WORLP DISTRIBUTION OF THE FAMILY DIPTEROCARPACEAE (The I'lgvpes aft"r!he names olferrt'for/esq'v, the number o/genera Bnd s!,eoles represenledJ A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. ACTA MANILANA

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AFRICA

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AUSTRALIA Fig. 2. WORLD DISTRIBUTION OF THE GENUS HOPEA (The figures alter the names oj terr/torles 3/v< I-h& number oj 8j'eoles rel'resenled) A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. all the confusion regarding the position and identity of the member species of our genus Hopea.

II-DISTRIBUTION OF THE FAMILY DIPTEROCARPACEAE

A. World Distribution

The family DIPTEROCARPACEAE comprises about 400 species in seventeen genera, which are distributed over a large area of tropical Africa and the Indo~Malayan region from India to New Guinea [29, pp. 1~32]. Borneo [37, pp. vi~xi1 ':s specifically the richest in dipte~ rocarps, representing nine genera and two hundred seventy-six species. The Malay Peninsula comes next, with one hundred sixty­ eight species in ten genera. The third richest is the Philippines, with eight genera and fifty-three species distributed throughout the country. Java represents a very poor dipterocarp flora, but this may be due to the almost complete destruction, in compara­ tively recent times, of the primary forest throughout the land below an altitude of 1,200 meters. Everywhere in Malaysia, the destruc­ tion of the primary forest means the destruction of the dipterocarps. The Eastern Peninsula, from Assam through Burma, Thailand, Indo­ China (Cambodia, Laos and Vietnam), to the southern part of the Malay Peninsula, presents ten genera and one hundred thirty-five species. No representative reaches Formosa, and only a single species of Shorea is known from the southwestern part of Kwantong Province, China, while a single species of Vatica occurs in Hainan. India presents only fourteen species in five genera. In contrast to this, the small island of Ceylon presents no less than forty-four species in nine genera, approximately the same number of species and genera that occurs in the entire Philippine archipelago.

A striking feature in the distribution of the dipterocarps is their paucity in eastern Malaysia, that is, in the entire region extending from Celebes southward to Lombok, and eastward through the Mollu~ cas to New Guinea, where only fifteen species in four genera are known. Three of these belong in the genus Anisoptera, five in the genus Hopea, two in the genus S.horea and five in the genus Vatica. All of these genera nre of wide geographic distribution, represented in most of the regions in which the family occurs. The genera H~ea, Shorea ~nd Vatica are three of the large genera in the family; Dipterocarpus IS the fourth. 10 ACTA MANILANA

It is perfectly evident that the Eastern Peninsula, and the Sunda IslaT!ds taken as a whole-that is, Sumatra, Java, Borneo and the smaller intervening islands such as Bangka, Billiton, and Lingga­ are essentially the regions in which the family has reached its maxi­ mum development in genera and in species. India may have pos­ sibly supported a much richer dipterocarp flora in the past than today; this is especially probable in view of the fact that the com­ paratively small island of Ceylon, off the southern end of the Indian Peninsula, presents many more genera and species than does the entire Western Peninsula.

That there is a larger distribution of this family in the Sunda Islands, the Eastern Peninsula, and the Philippines than in eastern Malaysia is evident by the number of genera and species found in the regions.

Borneo, which has the largest number of species, is believed to be the original home of the family. From its place of origin the species radiated; but sea, mountain and climatic barriers, and pos­ sibly human influence, have inhibited an evenly graded distribu­ tion.

Although most dipterocarps have winged fruits, they are not definitely adapted for wide dissemination by wind. The fruits are, in general, too heavy for wind distribution, while the wings are adapted to provide a gyratory motion in falling, rather than for horizontal distribution. Except in some species of the genera Vatica and ShoTea, the seeds or fruits do not present the slightest adaptation for dissemination by water. It is said that the fruits of some Vatica species float for an average period of twenty-two days, while those of some species of ShoTea, deprived of its corky sepals, sink within a period of sixty hours. The fact, however, is that it is not possible to regard water distribution as in any way ancestral in the order. In general, the seeds of the dipterocarps are noted for their brief period of viability; they do not withstand drying out, which is perhaps one very potent reason for their prac­ tical non-occurrence in open places. The trees, from seed to seed­ ling stages to full maturity, are clearly adapted to the shade, tem­ perature, moisture and light conditions characteristic of the dense tropical primary forests of India and Malaysia L37, p. 7J. To one, then, familiar with the dipterocarps as they occur in nature, with the habitat complex of most or all their species, with their seed and seedling characters, and especially with their peculiarly short- 13 A REVISION OF T. HE GENUS HOPEA ROXBURGH OF P. I.

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8. Philippine Distribution

As of this writing, the Philippines is credited with 53 species represented in eight genera of the family DXPTEROCARPACEAE. It is likely that this number may be increased as additional data on flowering and fruiting specimens are accumulated. On the other hand, this number may also decrease, depending upon our own interpretation of what constitutes a species, based on whatever additional data we might be able to accumulate.

It seems that most of our species do not flower every year, but only at irregular intervals. And, that these species are not all found in anyone forest area. The dipterocarp forests are tall, characteristically of low-altitude, and usually occupy localities most favorable to tree growth. They occur on all types of topography, but are usually best developed on well-watered and well-drained plains and on the lower gentle slopes of the main mountain masses.

Generally our dipterocarp forests r29, p. 9] reach their maxi­ mum development below an altitude of 700 meters; very few species are found at or above an altitude of 800 meters. Only the species of Vatica manggachapui Blume and Shorea polysperma Merrill are recorded from altitudes of 800 meters in Luzon and the central Philippines. In Mindanao Parashorea warburgii Brandis and Shorea squamata Dyer have been found at 1000 meters altitude, Vatica mindanensis Foxworthy at 1,100 meters altitude.

In a study of the altitudinal range of the DXPTEROCARPACEAE in the Philippines, Brown [7, pp. 76-97] made an interesting investiga­ tion in the mid-mountain forests of Central Luzon, that is, between 600 and 900 meters altitude. At an altitude of 700 meters, a plot was selected with the same slope and exposure as those of the plot surveyed in the dipterocarp forest at lower altitudes. From this investigation it is definitely shown that the DXPTEROCARPACEAE are strictly limited as to their altitudinal range; while representatives of many other families and genera which are associated with the DIPTEROCARPACEAE and form the complex low-altitude forests of the I' ACTA MANILANA Philippines, thrive at altitudes distinctly higher than the DIPTERo­ CARPACEAE themselves. In the transition from the dipterocarp to the mid-mountain forest, the r.hange nom one association to another is usually gra­ dual, and is marked by intermediate conditions. The tall trees, characteristic of the DXPTEROCARPACEAE that form the upper story of the tropical dipterocarp forest, gradually disappear, and the first story of the Quercus (Lithocarpus)-Neolitsea association of the mid-mountain forest is approximately of the S

The most important factor, however, that appears to influence greatly the distribution of DIPTEROCARPACEAE in the country is the amount and distribution of rainfall. The parts of the country [39, p. 15] where there is no dry season, or a very short dry season lasting only from one to three months, have the greatest number of species represented in the forest. These regions include the Babuyan Islands, the Cagayan Valley and the e:lstern part of Luzon, Polillo, Catanduanes, Samar, Leyte, Marinduque, Masbate, eehu, Bohol, eastern part of Mindoro, northeastern part of Panay, the eastern part of Negros, Mindanao and Basilan. Those regions that have distinct dry and wet season have extensive dipterocarp forest, but with relatively small number of species. '{hese regions include the rest of the country, such as tbe western part of Luzon, west of the Cagayan Valley and the Sierra Madre Mountains, the western parts of the islands of Mindoro, Panay, Negros and most of Palawan.

III - REVIEW OF LITERATURE

The generic name Hopea of the family DIPTEROCARPACEAE was described by Roxburgh [32, p. 71. The plant to which this name was first assigned is Hopea odoTata Roxburgh-a plant from Indo­ China (Phu Quoc, Cambodia). Early authors who studied our plants have erroneously placed or assigned member species of this genus to various genera of the family, such as Mocanera [3, p. 447], Dipterocarpus [3, ed. 2, p. 311], Anisoptera [5, p. 52], and Sho,.ea. f13, p. 21]. Mocanera, which is no longer a valid generic name, was at one time a dumping ground of members that were not clear- A REVISION OF THE GENUS HOPEA ROXBURGH OF P. 1. l7 ly understood, which was apparently the case in Blanco's first edi­ tion of the FLORA DE FILIPINAS of 1837. In the second edition, which came out in 1845, Blanco transferred the Mocanera species to Dipterocarpus. The first plant, however, from the Philippines to be assigned correctly to the genus, is Hopea philippinensis Dyer, des­ cribed in 1878 [11, p. 100].

The SINOPSIS ATLAS of Vidal [40, p. 15] in 1883 also includes an illustration of a 15-stamened species which he believed to be a "variety" of Hopea odorata Roxburgh. This, later, was reduced by Foxworthy [16, p. 100] to the synonymy of Hopea plagata (Blanco) VidaL Vidal's [41, p. 62] REVISION PLANTAS VASCULARES DE FILI­ PINAS in 1886 appropriately corrected Blanco's mistakes. He trans­ ferred MocanerQ plagata Blanco (later Dipterocarpus plagatus (Blanco) Blanco) to Hopea plagata (Blanco) Vidal. Aside from these two valid species-Hopea philippinensis Dyer and Hopea pla­ gata (Blanco) Vidal-nothing had been added to the member of our genus prior to 1900. With the coming of the Americans at the start of the twentieth century, extensive study on the flora of the Philippines was under­ taken. The first addition to our genus was described by E.D. Mer­ rill (26, p. 30] as Hopea acu.minata Merr. Foremost among the American botanists, however, on the study of Philippine DIPTERO­ CARPACEAE was Dr. William F. Foxworthy. His first paper concern­ ing the family came out in the PHILIPPINE JOURNAL OF SCIENCE in 1911 [16, pp. 231-285J. He listed eleven local species of the genus, two of which, Hopea basilanica Foxw. and Hopea mindanensis Foxw., were described for the first time. Four species were included at a venture but without specific epithets, while the two other species were found later to have been erroneously attributed to the Philip­ pines. In his LEAFLETS OF PHILIPPINE BOTANY, Elmer l12, pp. 1469-1470] described in 1912 two new species, Hopea foxworthyi Elm. and Hopea glutinosa Elm. While the former is a valid species, the latter was reduced by Foxworthy [18, pp. 163-199]. A subsequent new species described by Foxworthy [17, p. 1953] in 1913 came out in Elmer's LEAFLETS OF PHILlPPWE BOTANY as Hopea malibato Foxw. Another paper of Foxworthy [18, p. 184] concerning the family came out in the PHILIPPINE JOURNAL OF SCIENCE in 1918. This time Hopea maquilingensia Foxw., a supposed new species, was described. On the other hand, he reduced to eight the member species of the genus Hopea. However, Merill [30, vol 3 pp. 88-108] 18 ACTA MANILANA

in his AN ENUMERATION OF PHILIPPINE FLOWERING PLANTS in 1923 reduced the number of species to seven. Finally, the last extensive work, as far as the Philippine family DIPTEROCARPACEAE is concerned, was published by Foxworthy [19, p. 241·331] in the PHILIPPINE JOURNAL OF SCIENCE in 1938. The number of the member species of the genus was maintained at seven. Most of the species names were revalidated, while others were either excluded, or elevated to the rank of that of a species, or reduced to the status of that of a synonymy. To a largf: extent, the author of this paper is in agreement with Foxworthy. but proposes and describes two new species, and reva­ lidates the original Blanco's description of Hopea plagata (Blanco) Vidal.

IV - MATERIALS AND METHODS OF STUDY

A. Materials Examinoo

The materials used in this study are herbarium specimens: these are deposited in the Philippine National Herbarium, National Museum, Manila; in the Instituto Jardin Botanico Herbarium, Madrid; and in six different herbaria of the United States. It is unfortunate to state that all our original collections before the Second World War were completely destroyed during the liberation of Manila in February, 1945. The bulk, therefore, of the specimens studied, which are duplicc>tes of our original collections, is in the United States, Our dipterocarp specimens deposited at present at the Philippine National Herbarium

Aside from the data gathered from field labels attached to herbarium specimens, the field characters of the trees were studied in their natural habitats.

Practically all Philippine specimens, including Types (duplicates and Isotypes for that matter), studied by former authors, were examined and studied by the author. These materials were gathered from almost all parts of the country. Except for the Types of Hopea plagata (Blanco) Vidal, which is not extant, and Hopea philippinensis Dyer, deposited at the British Museum in London all the Types of the other member species are deposited in the dif~ ferent herbaria of the United States.

Indo-Malaysian specimens were also examined as far as prac­ ticable, primarily for the purposes of authenticating the distribu­ tion, the relationship, the position and the exact identity of our species.

B. Methods of Study

Intensive investigation was undertaken on the morphological characters of herbarium specimens as well as, to some extent, fresh materials from standing trees in their natural habitats, repre­ senting the different member species of the Philippine genus Hopea. Type specimens of our species, including Indo-Malaysian as far as practicable, were also studied carefully. The macroscopic structures of the plants were studied directly without the aid of magnifying equipment. On the other hand, the minute structures of the parts of the flowers, fruits and leaves, were examined under a stereo­ scopic wide-field binocular. The flowers as well as the young fruits were either boiled or soaked in water with detergent for some time, to soften the hardened tissues, before they were dissected. The induments that are microscopic, taken from other parts of the plants, were also studied under th4! same equipment. All measure­ ments mentioned in this paper were taken from dried specimens, with the exception of the parts that were measured in their more or less softened condition, that is, allowed to dry a little. The photo­ graphs were taken by the author, while the illustrations were done by scientific illustrators under his c;upervision. Field data of the member species, when available and gathered in their natural habitats by the author himself, are also incorporated in the description and discussion of the species concerned. 20 ACTA MANILANA

The literature consulted, dealing with this work, is composed mainly of those papers that have something to do with the taxo­ nomy and systematics of the group concerned.

Translation of the descriptions in Latin was done by the author with the assistance of others acknowledged in this paper.

The abbreviations used to indicate the various dialects adopted from Merill [30, Vol. 4, pp. 28-29].

Bik. Bikol Bis. Bisaya P. Bis. Panay Bisaya C. Bis. Cebu Bisaya S.-L. Bis. Samar-Leyte Bisaya Buk. Bukidnon Dum. Dumagat Gad. Gaddang Ibn. lbanag IIk. Iloko Lan. Lanae Mang. Mangyan Mho. Manoba Neg. Negrito Pang. Pangasinan Sbl. Sambali Sml. Samal SuI. Sulu Tag. Tagalog Yak. Yakan

The abbreviations used to indicate the different herbaria visited and materials studied were adopted from Lanjou and Stafleu [20, pp. 205-228].

(A) - The Arnold Arboretum, Harvard University (F) - Chicago Natural History Museum, Chicago (GH) - Gray Herbarium, Harvard University (MA) - Instituto Jardin Botanico, Madrid (NY) - New York Botanical Garden, New York (PNH) - Philippine National Herbarium, Manila (UC) - Herbarium of the University of California, Berkeley (US) - United States National Museum, Smithsonian Institution A REVISION OF THE GENUS HOPEA ROXBURGH OF P. 1. 21

v - RESULTS OF TAXONOMIC FINDINGS

A. Description of the Family DIPTEROCARPACEAE

Trees, usually of large size, with tall stems, often unbranched to a considerable height. Leaves evergreen, simple, alternate, sO pulate, penninerved: stipules often small and early deciduous. Wood, pith, bark, and leaves usually containing resin-canals. Fruits with sepals prolonged into wings, which often exceed the fruit in length and aid in its dispersal. Flowers perfect, numerous, in panicles, usually 5-merous. Receptacle obconical, sometimes coni­ cal. Sepals 5. Stamens: many, 15, 10, or 5. Carpels (3-1) with each of the 2 anatropous ovules with 2 integuments. Stylopodium often present. Style short or long, glabrous. Stamens of various forms; filaments often connate; anthers with the connective often prolonged. Only one of the ovules develops into a perfect seed. Coty­ ledons with long petioles; radicle superior; hypocotyl often as long as the embryo. Cotyledons usually bifid to the base and often divided into numerous lobes. Seeds often albuminous.

B. Key to the Philippine Genera of the Family DIPTEROCARPACEAE

1. Calyx urceolate, enclosing fruit: 2 sepals growing out into long wings. Stamens usually more than 20, anthers iong-awned. Style on a large, fleshy stylopodium.

2. Fruit not grown together with calyx, stamens many, styles filamentous, stipules amplexical . ( 1) Dipterocarpus.

2. Fruit grown together with calyx. Stipules small, fugacious. Stamens 25 to 60, style short. (2) Anisoptera.

1. Calyx imbricate in bud, in the ripe fruit usually 2 or 3 lobes larger than the others, developing into long wings. Stamens mostly 15, sometimes many, rarely 10. Appendage to connective usually long-pointed.

3. Long wings not more than 2.

4. Two long wing~ appendage to connective longpointed. Usually a large stylopodium . (3) Hopea.

4. Lobes of fruiting calyx short, not extending beyond fruit. 22 ACTA MANILANA 5. Calyx lobes not recurved, otherwise as in H opea (4) Balanocorpus.

5. Calyx lobes round and recurved, stamens 30 to 60 (Sect. Isoptera) (6) Shorea.

3. Long wings 3 or more.

6. Fruit with 3 long wings.

7. Anthers with 5 pointed projection (5) Pentacme.

7 Anthers with one projection from the connective, or without projections (6) Shorea.

6. Fruit with long wings; calyx in bud almost valvate; no stylopodium (7) Parashorea.

1. Calyx valvate in bud; fruit usually with 2 long wings._ Stamens 15. Anthers short, oval, smooth; appendage to connective blunt (8) Vatica.

C. Description of the! Genus Hopea Roxburgh

Hopea Roxb., PL. COROM. 3 (1819) 7. Type: H. odorata Roxb. Hopea Roxb., HORT. BENG. (1814) 42, nomen nudum. Neisandra Ratio., SYLVA TELLUR (1838) 163. Hoppea Roxb., fide End!., GEN. PL. (1840) 1014. Petalandra Hassk., HaRT. BOG. DESC. (1838) 104. Balanocarpus Bedd., FOR MAN. BOT. (1873) 236. Doona Burck. ANN. JARD. BOT. BUITENZ. 6 (1887) 231 Hancea Pierre. FL COCH. 16 (1891) sub tab 244. Pierrea Heim, BULL. MENS. Soc. LINN. PARIS'; 2 (1891) 958. Dioticarpus Dunn, KEW BULL. (1920) 337. Pierreocarpus Ridl. ex Sym .. GARD. BULL. S.S. 8 (1934) 958.

Dr. Ashton made a critical study of the synonymy of the genus in his exhaustive studies of Bornean DIPTEROCARPACEAE [1, pp. 229- 284]. He studied apparently all the species recognized by him as Hapea. As I have no access to the entire literature nor to all the herbarium specimens, much less the types, of the species of Hopea in the world, I limit my studies to the Philippine species. A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. :!3

"Receptacle plat ou un peu convexe, portant sur Ie berd 4-5 sepales persistants et en partie tres accrescents, imbri­ ques dans Ie bouton; 15 etamines (parfois 10: sect. Peta­ landTa) sur 3 rangs, les 5 intermediaires epipetales, les autres alternipetales, filets elargis vers Ie bas, antheres a lobes egales, et connectif a long appendice acumine et sou­ vent courbe; ovaire glabre au presque, stylopode soit grand et charnu avec style en general tres court, soit (plus rare­ ment) 0 ou presque, et Ie style long et filiforme, souvent avec une couronne de pails a la base; fruit a 2 ailes formees des sepales exterieurs longuement accrus et lineaires, peri­ carpe mince, tegument de la graine tres delicat, radicule egale a la graine au de moitie plus courte, cotyledons epais charnus, profondement bipartits, en general inegaux, albu­ men present au O.

"Grands arbres; feuilles aiternes, simples, coriaces, glabres, a stipules petites et caduques; infiorecences termi­ nales au axillaires, composees d'epis unilateraux, glabres au velues, fieurs sessiles au pedicellees. "Plus de 55 especes dispersees de Ceylan aux iles Philippines et a la Nouvelle-Guinee.

"Syn.: Hancea Pierre, Hoppea EndJ., NeisandTa Raf., PetalandTa Hassk." [22, p. 640]. Receptacle flat or slightly convex, bearing on the mar­ gin 4-5 persistent, imbricate in bud, partly very accrescent sepals; stamens 15 (sometimes 10: sect. PetalandTa) in 3 rows, 5 episepalous, others alternating with the petals; fila­ ments broadened towards base; anthers uniform, connective of anther lobes with long acuminate, often bent, appendages; ovary glabrous or almost glabrous, stylopodium either large and fleshy, the style then generally short, or rarely absent, the style then long and filiform, often with a crown of hairs at the base; fruit 2-winged, formed from the outer sepals, linear, pericarp thin, integument of seed very delicate, radicle equal to or one-half the length of the seed, cotyle­ dons thickly fleshy, distinctly bipartite, usually unequal, albumen present or nil. Large trees; leaves alternate, simple, coriaceous, glab­ rous, having small caducous stipules; inflorescence terminal or axillary, composed of unilateral heads, glabrous or hairy, pedicellate or sessile flowers. There are more than 55 species distributed in Ceylon. the Philippines and New Guinea. 24 ACTA MANILANA

Syn.: Hancea Pierre, Happea Endl., Neisandra Raf., Petalandra Hassk.·

Medium- to large-sized trees, the branchlets ienticellate, occa­ sionally furrowed, glabrous or pubescent, light brown to dark­ chocolate in color. Hairs simple and stellate, sericeous. Leaves simple, alternate, distichous, the blade chartaceous to coriaceOlls; apex either short or long acuminate, or long and thin caudate-acu­ minate; the base equal, oblique or unequal (the left side then rounded, when viewed from above), the margins entire; venation pinnate, the mid-vein prominent or sunken above, distinct or more or less keeled beneath, the principal lateral veins distinct or less conspicuous, with or without alternating parallel veins, reticulate venules distinct or not distinct; domatia present, pouch like or hol­ low depressions on the lower surface in the axils of the veins, glab­ rous or hairy on the margins. Petioles glabrous, puberulous, or velutinous, wrinkled with age, rarely smooth, generally sulcate, terete or more or less flattened above, light brown to dark-choco­ late in color. Stipules caducous or semi persistent. Inflorescences. racemose panicles of secund flowers, each raCEme subtended by a caducous bract. Flowers pedicellate, subsessile or sessile, complete, pentamerous, each subtended by a caducous bracteole. Receptacle flat or slightly convex, bearing on its margin 5 free, quincuncially disposed, glabrous, pubescent or lepidote sepals. Corolla hypogy­ nous, imbricate, the petals oblong to oblique-oblong, pubescent on the outer half of each petal, exposed in bud. Stamens mostly 15 (10 in H. acuminata, 32-38 in H. plagata), generally in 3 whorls of 5, those of the first and third whorls opposite the petals, the second opposite the sepals; filaments broad~ned towards base, connective prolonged into awnlike terminal appendage; anthers orbicular, basi­ fixed, the lobes of equal length, opening by 2 lateral longitudinal slits. Stigma minute or obscurely 3-parted, obtuse; style short, or long and slender. Stylopodium present or absent; ovary glabrous or hairy, superior, 3-locular, each locule with 2 axile anatropous ovules, the micropyle up. Fruit a nut, invested by the persistent accrescent calyx, the 2 outer sepals becoming winglike, parallel or crossed at the base, oblanceolate, narrowly elliptic to broadly elliptic or suborbicular.

The bark is thin or rather thick, smooth, flaky or fissured. Al­ though our species are usually trees of medium or large size, they

°Tran81 ..ted d""riplion hom Lorn ..... A REVISION OF THE GENUS HOPEA ROXBURGH OF P. 1. 2:3 do not reach the proportion of the large species of Dipterocarpus and Shorea. All of our species have an irregular fruiting period, but they all, or nearly all, fruit at tolerably frequent intervals. The fruiting of a particular species is not uniform throughout the country, but rather dependent upon the climatic condition of the locality. Our species of the genus Hopea are not as abundant as some of the species of the genera Dipterocarpus and Shorea. They are mostly found at low to medium altitude, although certain forms are com· mon on ridges below 800 meters. Some species are characteristically growing on low flat ground or along streams.

The genus is represented by 9 species in the Philippines, all of which are endemic.

D. Key to the Sections and Subsections

1. Lamina nervation dryobalanoid or subdryobalanoid Section I. DRYOBALANOIDES.

2. Nervation dryobalanoid Subsection 1. Dryobalanoides

2. Nervation subdryobalanoid Subsection 2. Sphaerocarpae

1. Lamina nervation scalariform Section II. HOPEA.

2. Lamina base equal or oblique Subsection 1. Hopea.

2. Lamina base unequal Subsection 2. PierTea.

E. Descriptions of Sections and Subsections [1, pp. 258-259J

Section I. DRYOBALANOIDES Miq. Nervation dryobalanoid or subdryobalanoid. Bark smooth, fis­ sured or cracked, not evenly flaky. 26 ACTA MANILANA

Subsection 1. DryobalaDoides Miy.

Nervation dry0balanoid, bracts fugaceous; corolla pale; racemes reguarly branched, branchlets short; flowers many; ovary and stylopodium ovoid to pyriform, rarely truncate.

Distribution. 23 species: Indo-China (1), Thailand (3),

Burma (1) I Malaya (14), Borneo (15), Sumatra (5), Philip­ pines (4).

Subsection 2. ~haerocarpae (Heim) Ashton

Nervation subdryobalanoid; bracts subpersistent; racemes irregularly branched, with long branchlets and few flowers; corolla dark colored; ovary and stylopodium truncate; bark smooth or cracked and with small irregular scales.

Distribution. 9 species: Malaya (6) I Borneo (6) I Sumat­ ra (1), Philippines (None).

Section II. HOPEA Miq.

Nervation scalariform; bark surface smooth or evenly flaky.

Subsection 1. Hopea Miq.

Lamina base usually equal or subequal; raceme generally tomentose, rarely fascicled; ovary and stylopodium more or less ovoid, or if elongate, without median constructions; style evident.

Distribution. 47 species: Ceylon (3), S. India (7), Burma (5), Thailand (8), Indo-China (6), S. China (1), Malaya (7), Sumatra (4), Java (1), Borneo (5), Celebes and Mo­ luccas (3), New Guinea (10), Philippines (3).

Subsection 2. Pierrea (Heim) Ashton

Lamina base unequal; racemes glabrescent, fascicled. Ovary and stylopodium hour-glass (bottle-shaped) shaped, elongate; style short, obscure, bark smooth, rarely shallowly patchily flaked. A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 27

Distribution. 8 species: Malaya (3), Borneo (4) Philippines (2).

F. Key to the Philippine Species of Sections and Subsections

Section I. DRYOBALANOIDES Miq.

Subsection 1. Dryobalanoides Miq.

KEY TO THE SPECIES

1. Leaves characteristically with the midrib sunken above, more or less keeled beneath.

2. Leaves broadly elliptic to s!..lborbicular; ovary hairy. (1) H. quisumbingiana.

2. Leaves narrowly elliptic to ovate-Ianceolate; ovary glabrous (2) H. fOXWOTthyi.

1. Leaves characteristically with the midrib prominent above, not keeled beneath.

2. Leaves broadly elliptic to oblong·elliptic; style stout, shorter than the length of the ovary. (3) H. malibato.

2. Leaves narrowly elliptic to ovate-lanceolate; style slender, as long as or longer than the length of the ovary. (4) H. woodiana.

Section II. HOPEA Miq.

Subsection 1. Hopea Miq. 28 ACTA MANILANA

KEY TO THE SPECIES

1. Stamens 10 . (5) H. acuminata.

1. Stamens 15 ...... (6) H. basilanica.

1. Stamens 32 to 38 . (7) H. plagata.

Subsection 2. Pierrea (Heim) Ashton

KEY TO THE SPECIES

1 Leaves large, 39-51 em. long, 13-17 em. wide, base cordate-auri- culate, unequal (8) H. mindanensis.

1. Leaves smaller, not more than 24 em long and 7.5 em wide, base obtuse, unequal. . (9) H. philippinensis

G. Key to the Phili.ppine Species of Genus Hopea

1. Primary lateral veins of lamina distinct; alternating parallel veins absent; reticulations distinct.

2. Stamens 10; inflorescencEs pube~c€'nt; stylopodium absent; ovary globose; mature winglike sepals oblanceolate to nar­ rowly elliptic. 1.5-3 em long, 0.6-1 em wide, with 6-7 princi- pal veins in each. (5 I H. acuminata.

2. Stamens 15; inflorescences pubescent or glabrous; stylopf)­ dJUm present; ovary cylindrical or bottle shaped. mature wmglike sepals narrowly elliptic or distinctly oblanceolate.

3. Inflorescences pubescent; stylopodium and ovary cylindri­ cal; mature winglike sepals narrowly elliptic 3.5-4.5 cm. long, 1.1-14 cm. wide, with 9 principal veins in each. (6) H. basilanica. · A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 29

3. Inflorescences glabrous; stylopodium and ovary bottle shaped; mature winglike sepals distinctly oblanceolate.

4. Leaves large, 30-51 cm. long, 13-17 cm. wide, oblong, base cordate-auriculate, unequal; mature winglike sepals, 11-12 cm. long, 1.4-1.6 cm. wide, with 6-7 prin­ cipal veins in each. ( 8) H. mindanensis.

4. Leaves smaller, (8-) 11.5-19.5 (-24) cm. long, (2.5-) 3.5-6.5 (-7.5) cm. wide, narrowly elliptic to oblong­ elliptic, base obtuse, unequal; mature winglike sepals, 6-10.5 cm. long, 1.2-2.5 cm. wide, with 8-9 principal veins in each (9) H. philippinensis.

2. Stamens 32-38; inflorescences glabrous; stylopodium present, stout, continuous with the ovoid ovary; mature winglike se­ pals broadly elliptic to suborbicular, 2.5-4.5 cm. long, 1.5-3 wide, with 12-14 principal veins in each. ( 7) H. plagata.

1. Primary lateral veins of lamina not distinct; alternating parallel veins present; reticulations not distinct.

2. Leaves characteristically with the midrib sunken above, pro­ minent and more or less keeled beneath.

3. Leaves broadly elliptic to suborbicular, 5-7 cm. long, 2.5- 3.2 cm. wide; ovary distinctly hairy. (1) H. Quisumbingiana sp. nov.

3. Leaves distinctly uniform, narrowly elliptic to ovatelan­ ceolate, (3.5) 4-5 (-5.5) cm. long, 1.3-2 (12.5) cm. wide; ovary glabrous (2) H. forxworthyi.

2. Leaves characteristically with the midrib prominent above, distinct but not prominent beneath.

3. Leaves oblong-elliptic to broadly elliptic, (6-) 8-10 (-12) cm. long, (2.5-) 3.54.5 (-5) cm. wide, the stout caudate tip about one-eight to one-twelfth the length of the ovary .. (3) H. malibato. 30 ACTA MANILANA

FIG. 5 Hopt a quuumbingiano Gutierrez, sp. nov.; Typfl A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 31

3. Leaves narrowly elliptic to ovate-Ianceolate, (4-1 5.5-6.5 (-8) cm. long, 1.5-2.5 (-3) cm. wide, the slender caudate tip about one-third to one-sixth the length of the blade; flowers pedicellate; pedicels 1-2 cm. long; style as long as or longer than the length of the ovary ( 4 ) H. woodiana sp. nov.

VI. DESCRIPTION OF THE SPECIES

1. HOPEA QUISUMBINGIANA Gutierrez, sp. nov. Text fig. 5' Plate I.

Ramuli puberulentes sed maturitate glabri. Folia coria­ cea, glabrata, paene elliptica usque suborbiculata, 5-7 cm. longa, 2.5-3.2 cm. lata, apice caudata-acuminata, apicem ver­ sus obtusa vel retusa usque paene truncata, basi obtusa; costa mediana obverso impressa, dorso prominens, carinata; nervis primariis 9-13 utrinsecus, 1 vel 2 nervis alternantibus, venulibus reticulatis indistinctis; petiolis 0-8 mm. longis, glabratis, valde sulcatis, teretibus. lnflorescentia glabra terminalis et/vel axillaris, singulatim, raro bina, paralieliter constituta, 2-3.5 cm. longa, cum racemis 2 ad flores pedi­ cellatas gerentibus. Pediculi 1-3 mm. longi, glabrati. Calyx haud lepidotus, duabus sepalis extern is suborbiculatis, 2.5-3 mm. longis, 2-2.5 mm. latis, sepalis internis orbiculatis, 2.2-5 mm. longis, 1.8-2.5 mm. latis. Petala ova to-oblongata, obli­ qua, 6.5-8 mm. longa, 2-2.5 mm. lata. Stamina 15, 3 series ex 5, 3 stamina in unoquoque petalo; filamenta variabilis longitudinis; anthem orbicularia, 0.5 mm. longa, cum arista appendice terminali vix filamentibus breviori. Stigma minusculum; stylus filiformis. Stylopodium 0; ovarium oviforme, 1-1.5 mm. latum, pubescens. Fructus ignotus.

Branchlets puberulent, becoming glabrous with age, dark-cho­ colate in color. Leaf-blade coriaceous, glabrous, shiny above, dull beneath, broadly elliptic to suborbicular, 5-7 cm long, 2.5-3.2 cm wide, approximately twice as long as broad, apex caudate-acuminate, the caudate tip about one-fourth to one-fifth the length of the blade, rounded or retuse to more or less truncate, the base obtuse; midvein sunken above, prominent beneath, more or less keeled; the princi­ pal lateral veins 9 to 13 on each side, curving, with 1 or 2 alter­ nating less conspicuous veins, reticulate venules not distinct; do- 32 ACTA MANILANA

matia hollow depressions, confined to the axils of the first pair of veins, occasionally in only one vein, rarely in the axils of succeed· ing veins (UP to 3 domatia), with tufts of hairs on the margins; petioles 6-8 mm long, glabrous, very finely sulcate, terete. Stipules not seen. Inflorescences glabrous, dark-chocolate in color, terminal and/or axillary, borne singly, rarely in pairs, in axils of the leaves or of fallen leaves, those in pairs arranged side by side, 2-3.5 em long, the racemes bearing 2 to 5 pedicellate flowers; bracts and bracteoles not seen. Pedicels 1-3 mm. long, glabrous. Caly:r sparsely lepidote, the 2 outer sepals suborbicular, slightly pointed at the apex, 2.5-3 mm long, 2-2.5 mm wide, the inner sepals orbicular, with hairs on the upper margins, 2.2-5 mm long, 1.8-2.5 mm wide. Petals ovate-oblong, oblique, 6.5-8 mm. long, 2-2.5 wide, approxi­ mately three to four times as long as broad. Stamens 15, in 3 whorls of 5, seemingly 3 stamens to each petal, the first and third opposite the petals, the second opposite the sepals; filaments of dif­ ferent lengths, the first whorl 0.5 mm long, the second 0.8 mm, the third 1 mm.; antheTs orbicular, 05 mm. long, the awnlike ter­ minal appendage as long as the filaments. Stigma minute; style slender, 0.7 mm long, approximately one-half the length of the ovary. StyZopodium absent; ovaTY ovoid, 1.5-2 mm. long 1-1.5 wide hairy. FTuit unknown.

SAMAR: Provo Samar, B.S. 1623 M. Ramos, April, 1914, in flower, (Holotype: CHI, (lsotypes: NY, US).

This species differs from other species of our genus Hopea by its hairy ovary. It may be confused with Hopea foxwoTthyi Elm. in the general appearance of the flowers and the size of the leaves. However, the hairy ovary and the broadly elliptic to suborbicular shape of the leaves distinguishes it readily from the glabrous ovary and the narrowly elliptic to ovate-Ianceolate leaves of Hopea fox­ wOTthyi Elm.

This species is named in honor of Dr. Eduardo Quisumbing who has contributed much to the knowledge of Philippine Botany.

2. HOPEA FOXWORTHYI Elmer Text fig. 6; Plate II.

Hopea foxwoTthyi Elmer, LEAFL. PHILIP. BOT. 6 (1912) 1496.

Hopea pieTTei Foxworthy, pro parte PHILIP. JOUR. SCI. 6 (1911) A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 33

Bot. 265; 13 (1918) Bot. 184; Whitford, PHILIP. BUR. FOR. BULL. 10 (2) (1911) 76; Brown and Matthews, PHILIP. JOUR. SCI. A. 9 (1914) 439,481; Merrill, ENUM. PHILIP. FL. PL. 3 (1923) 94; Reyes, PHILIP. JOUR. SCI. 22 (1923) 339, pl. 31, fig. 1, non Hance.

Hopea foxwoTthyi Foxworthy, pro parte PHILIP. JOUR. SCI. 67 (1938) Bot. 282, non Elmer.

Hopea glutinosa Elmer, LEAFL. PHILIP. BOT. 6 (1912) 1470; Fox­ worthy, PHILIP. JOUR. SCI. 67 (1938) Bot. 282.

ORIGINAL DESCRIPTION:

"A handsome middle sized tree, stem 3 dm thick, terete, 12 m. high, straight; branches chiefly toward the top and forming a spreading crown, the main branches ultimately finely rebranched; wood moderately hard, dingy white, ex­ cept the yellowish brown central portion, odorless, with a distinct bitter taste; bark smooth, brown; young twigs brown, puberulent, finally glabrous. Leaves alternatingly scattered, numerous, all alike in shape and size, subcharta­ ceous, glabrous, folded upon the upper surface, paler green beneath, drying brown, the caudate tips recurved, the en­ tire margins subinvolute toward the apex, ovately lanceo­ late, obtuse at base, gradually tapering into the 1.5 cm. long apical portion which terminates in an obtusely rounded point, the lamina nearly 5 cm. long, 1.5 cm. wide below the middle; mid vein bold beneath, clear to the apical point, smooth and brown, grooved along the upper side, lateral primary veins 9 to 11, parallel, with secondary ones be­ tween, nearly straight, very obscure, reticulations none; pedicels 5 to 8 mm. long, slender, blackish brown in the dry state, ultimately glabrate. Infructescence (rom the leaf axils; stalks 2 to 3 cm. long, similar to the petioles, divari­ cated only very shortly, branched toward the distal end if at all; fruits short pedicelled, 1 to 3 clustered, pendulous; calyx green, ovoid, glabrous, occasionally glutinous at the apex, 6 mm. long, nearly 5 mm. across the base, snugly covering the seed, the 5 obtuse teeth imbricately arranged, the outer 2 segments winged; wings crossed from the base, obtusely rounded at apex, narrowed toward the base, flat, oblanceolately oblong, about 7-veined, reticulations fine, pleasing red and appearing as if glutinous, the normal ones 3 cm. long, 7.5 mm. wide above the middle.

''Type specimen 12071, A.D.E. Elmer, Magallanes (Mt. Giting-giting), Province of Capiz, Island of Sibuyan, March, 1910). " ACTA MANILANA

,~ ...-C " lr

\ I

Fm. 6 Hupeu foxworlklli Elmer; Type A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 3!)

"This lIopea was discovered in red sticky clay of woods at 2000 feet elevation, on the summit ridge along tratl from Magallanes to Espana. Named after Dr. F. W. Foxworthy, who has recently revised our known Philippine DIPTEROCAR­ PACEAE.

"The foliage of this species is practically the same as that of the following (H. glutinosa Elmer) but the fruits are en­ tirely different."

Trees, 30 to 35 m tall, the stem about 60 to 65 cm diameter at breast height. Branchlets puberulent, becoming glabrous with age, brown to dark-chocolate in color. Leaf-blade subcoriaceous to coriaceous, glabrous, shiny above, dull beneath, narrowly elliptic to ovate-Ianceolate, (3.5-) 4-5 (-5.5) cm. long, 1.3-2 (-2.5) cm. wide, approximately 2 to 3 times as long as broad, apex caudate-acumi­ nate, the thin caudate tip about one-third to one-fourth the length of the blade, rounded or retuse to more or less truncate, the base cuneate or obtuse; midvein sunken above, prominent beneath, more or less keeled, the principal lateral veins 9 to 12 on each side, curv­ ing, with 1 or 2 alternating less conspicuous veins, reticulate ve­ nules not distinct; domatia hollow depressions, confined to the axils of the first distinct pair of veins, rarely in axils of succeeding veins (up to 4 domatia), with tufts of hairs on the margin, mostly on the side of the midrib; petioles 6 to 8 mm long, glabrous, finely sulcate, more or less terete. Stipules not seen. Flowers unknown. Infructes­ cences borne singly in axils of the leaves, 1 to 1.5 em long, with few scattered minute hairs, bearing 1 to 2 short pedicelled, pendu­ lous frUlts. Fruit conically elongated, smooth, shiny, glutinous, 1.5 cm long, 0.5 cm wide near the base, apiculate, the winglike sepals, when dry, dark brown, crossed at base, narrowly elliptic to oblan­ ceolate and narrowed toward the base, 3-3.5 cm long, 0.7-0.8 cm wide, with 5-6, rarely 7, principal veins in each, the inner s(!pals appressed to and enclosing about one-third to one-half the length of the fruit.

SIBUYAN ISLAND: Provo Capiz (Romblon), Magallanes, Mt. Giting-giting (Mt. Guiting-guiting), ElmeT 12071, March 1910, in fruit, (Holotype: PNH, destroyed in 1945), (Lectotype: GH), (Iso­ type: A, NY, US, F. DC); ElmeT 12289, April 1910, in fruit, (Type for H. glutinosa) , (Al, (GH), (NY), (US), (F); Espana, Mt. Guiting-guiting, PNH 97268, 97269 H. Gutierrez & E. Reynoso, April 28, 1966, in fruit buds, (PNH). 36 ACTA MANILANA LOCAL NAMES: malungai (Visayan of North Sibuyan), manggachapui (Sibuyan), manggachapui-tagokan (Sibuyan).

Hopea foxworthyi is a species described by Elmer (rom a single collection. He found the tree growing in red sticky clay of woods on the summit ridge along the trail from Magallanes to Espana in Sibuyan Island, Province of Capiz (Ramblon), at an altitude of 2000 feet. He also described Hopen glutinosa from a single collec­ tion, gathered a month later, from the same area. He found the tree growing in a clay soil on wet gravelly subsoil of wooded banks along Pauala River at 750 feet altitude. He distinguished these two species primarily on fruit differences.

Merrill, later, reduced Hopea foxworthyi Elm. and Hopea glu­ tinosa Elm. to the synonymy of Hopea pierrei Hance, and states: "I fail to distinguish Hopea foxworthyi Elm. and H. glutinosa Elm. from each other or from H. pierrei Hance. The fruit differences in­ dicated in the original descriptions are apparently differences of development." [30 vol. 3, p. 94J. From his second statement, it is likely that Merrill reduced, based on the original descriptions alone, both species without having examined critically any of the Type specimens, and for that matter, neither Hopea pierrei Hance.

Hopea foxwortyi Elm. together with Hopea quisumbingiana, sp. nov. and Hopea woodiana sp. nov., have, at one time or another, been identified as Hopea pierrei Hance. Hopea pierrei Hance is a species that does not occur in the Philippines. It was collected in Indo-China (Phu Quoc, Cambodia) and was identified by Hance as Hopea micrantha Hook. f., but later described and named as Hopea pierrei Hance. It has an impressed midrib, a stylopodium and a dis­ tinctly bottle-shaped ovary. Symington L36, p. 3231 states, "that the species has also been recorded from the Philippine Islands from 1909 onwards, as a result of mistaken identification." This mistake was corrected accordingly by Foxworthy [19, pp. 282-2831. Conse­ quently, since Hopea pierrei Hance does not occur in the Philippines and none of the Philippine flowering material shows a stylopodium (which is a distinct character of the species), he elevated Hopea foxworthyi Elm. to the rank of a species and reduced Hopea gluti­ nasa Elm. to a synonymy, the former name having page priority.

The author is in agreement with Symington, and for that mat­ ter with Foxworthy, in considering Hopea pierrei Hance as a species that does not occur in the Philippines. Nevertheless, while it is A REVISION OF THE GENUS HOPEA ROXBUlWH OF P. I. 37 true that the stylopodium is absent in Hopea quisumbingiana sp. nov., and in Hopea woodiana sp. nov., and is present in Hopea pier­ rei Hance (I have had the opportunity of examining the lsotype de­ posited at the Arnold Arboretum), it is rather doubtful at the present to consider the stylopodium as a distinguishing charactE'r between Hopea pierrei Hance and Hopea foxworthyi Elm. No col­ lection of Hopea foxworthyi Elm. bearing flowers has ever been gathered yet from the Type-locality; only specimens bearing fruit buds and matured fruits have been gathered. The species with which Foxworthy presumably compared Ho­ pea foxworthyi Elm. is here treated as Hopea woodiana sp. nov. It could be likewise presumed that the material used by Reyes (31, pp. 339-340) in studying the wood of the supposedly Hopea pierTei Hance of the Philippines came from Hopea woodiana sp. nov. Hopea foxworthyi Elm. differs from the true Hopea pieTTei Hance, a plant of Indo-China (Cambodia), in having uniformly smaller coraceous, narrowly elliptic to ovate-Ianceolate leaves. The status of Hopea foxworthyi Elm. as a valid species, and the reduction of Hopea giutinosa Elm. to a synonymy, is maintained here, following Foxworthy's view. However, given the present condition of our herbarium specimens, the author believes that the status of Hopea foxworthyi Elm. and Hopea glutinosa Elm. may be better understood when more complete materials will have been examined. In Sibuyan Island where the author had the opportunity to gather specimens in the Type-locality, one can readily observe the differences of the individual members of the species by simply look­ ing at the external features of the bark alone. These features, as a matter of fact, are used by woodcutters to distinguish the pro­ fusely sappy ("tagokan" = bearing sap) individuals from the re­ latively non-sappy. Doubtless, Elmer described them as two sepa­ rate species: Hopea foxworthyi Elm. and Hopea glutinosa Elm. This gives us, therefore, a reason to doubt; these field characters may be only within the range of variation of the species. This species is found only in the island of Sibuyan. The author not only did not find the trees in flower, except for fruit buds, but he broke his wrist while in search for them. The wood of Hopea foxworthyi Elm. is used locally for genera] house construction, for posts, bridge timber and other uses requir­ ing strength and durability. 3' ACTA MANILANA

\

FIG. 7 Hopea malibato Foxworthy A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 39

3. HOPEA MALIBATO Foxworthy Text fig. 7; Plate HI.

Hopea malibato Foxworthy, Elmer LEAFL. PHILIP. BOT. 6 (1913) 1953; PHILIP. JOUR. SCI. 13 (1918) Bot. 184; 67 (1938) Bot. 281; Merrill, ENUM. PHILIP. FL. PL. 3 (1923) 93.

ORIGINAL DESCRIPTION:

"Ramuli, petioli et Jolia materia resinosa incano-obtecti, glabri. Folia firmiter chartacea, modice petiolata, a basi cuneata vel acuta, elliptica vel ovato-elliptica, longiter cau­ dato-acuminata, 8-12 cm. longa, 3-4.5 cm. lata, costa media supra prominens. Petioli 10-12 mm. longi. FloTes ignoti Racemi fructiferi breves, calycis fructiferi laciniae majores 4-5.5 cm. longae, 12-15 mm. latae, chartaceae, rubrae, glab­ rae, 8-9 nerviae. FTuctus breviter pedunculati, resinoso­ nitentes, 8-9 mm. longi. "This species is exceedingly close to Hopea beccaTiana Burck., differing from that species only in the greater size of leaves and fruit, and in the greater number of veins on the larger fruit wings. "No. 13526, Cabadbaran (Mt. Urdaneta), Province of Agusan, Mindanao, August, 1912. The field notes with this material are: "Rather small tree on a forested ridge at 2000 ft. altitude, in well drained soil among some rocks; stem 1 foot thick, terete, slightly crooked, a trifle buttressed at base, 30 feet high, mostly branched toward the top; wood hard, heavy, the outer 0.33 sappy white, the heartwood light ixabellinus, odorless and tasteless; bark mottled, smoot ish, testaceous except the epidermis; main branches spreading, repeatedly rebranched, the lax twigs brown; leaves horizontally spreading, subchar­ taceous, dull green, yet lighter so beneath, flat with recurved tips; infrutescence axillary, few inches long at most, the green stalks horizontal; fruits pendant, the nut somewhat glutinous, the 2 twistingly recurved wings paler green or very light green. - Malibato in Manobo, also called dangin­ ginan by the same natives.

"Hopea beccaTiana Burck, produces a wood of the same grade as that produced by several of the Philippine species of H opea, and known here by the name dalingdingan. The wood of H opea beccariana is known in Sarawak by the name of mahan-besi and mang-besi." ACTA MANILANA

A tree up to 36 m tall. the stem about 50 cm diameter at breast height. Branch lets glabrous. dark-chocolate in color. Leaf-blade chartaceous, glabrous. broadly elliptic to oblong-elliptic, (6-) 8-10 (-12) cm long, (2.5-) 3.5-4.5(-5) cm wide, approximately 2 to 3 times as long as broad, apex caudate-acuminate, the stout caudate tip about one-eight to one twelfth the length of the blade, obtuse, the base cuneate to more or less obtuse; midvein distinct on both sides. the principal lateral veins 8 to 13 on each side, curving, with 1 to 3 alternating less conspicuous veins, reticulate venules not dis­ tinct; domatia hollow depressions, confined to the axils of the first distinct pairs of veins, occasionally in succeeding veins (rarely up to 12). with tufts of hairs on the margin. Petioles 0.7-1.2 em long, glabrous, finely sulcate, more or less terete. Stipules not seen. In­ florescences glabrous. occasionally with few scattered minute hairs, dark-chocolate in color, terminal and/or borne singly or in pairs in axits of the leaves or of fallen leaves, those in pairs arranged side by side, 2.5-4 cm long, the racemes bearing 1 to 7 pedicellate flow­ ers; bracts and bracteoles not seen. Pedicels 1 mm long or less, glabrous. Calyx lepidote, the 2 outer sepals 9vate to ovate-elliptic, inner and outer surfaces of the blunt apices densely lepidote, 0.5-1 mm long, 0.4-0.6 mm wide, the inner sepals suborbicular, slightly pointed at the apex, 0.5-1 mm long, 0.5-0.8 mm wide, with minute hairs on the margin. Petals light red, 5-6 mm long, 1.5-2 mm wide, approximately 3 to 4 times as long as broad. Stamens 15, in 3 whorls of 5, seemingly 3 stamens to each petal, the first and third opposite the petals, the second opposite the sepals; filaments of different length, first and second whorls about 1 mm long, the third 1.2 mm.; anthers orbicular, 0.3 mm. long, the awnlike terminal appendage as long as the innermost filaments. Stigma minute: style slender, 0.8 mm long, occasionally bearing hairs at the base. Stylo­ podium absent; ovary ovoidal, I mm long, 0.7-1 mm wide, glabrous. Fruit ovoid, 1-1.2 cm. long, 0.6-0.8 cm. wide, apiculate, the winglike sepals green, dark brown when dry, crossed at the base, oblanceolate, generally rounded. at the apex and narrowed toward the base, 5.5-6 cm long, 1-1.5 cm wide, 7-9 distinct principal veins in each, the inner sepals appressed to and enclosing about two-thirds the length of the fruit.

LUZON: Provo Tayabas (Quezon), Mt. Binuang, B.S. 28486, M. Ramos & G. Edana. May 1917, in fruit, (A), (NY), (US); Provo Camarines, R. Rosenblu.th F.B. 12810, in fruit, (NY). A REVISION OF THE GENUS HOPEA ROXBURGH OF P. 1. 41

MINDANAO: Provo Agustan, Cabadbaran, Mt. Urdaneta, Elmer 13526, August 1912, in fruit, (Holotype: PNH, destroyed in 1945), (Lectotype: A) (]sotypes: GH, NY, US, F); Butuan, San Mateo, Tungao, PNH 42503 D.R. Mendoza, May 30 -June 26, 1961, in fruit, (PNH), (A); Basilan Island, Maluso, Canas, PNH 97155 Gutierrez, Reynoso & Fraga, April 18, 1965, in flower, (PNH); PNH 97149 Gu­ tierrez, Reynoso & Fraga, May 10, 1965, in flower, (PNH).

LOCAL NAMES: malibato (Mbo.), danginginan (Mbo.), da­ lindingan (Tag.), manggachapui (Yak.), yakat-kaliot (Official Com­ mon Name).

Hopea malibato is distinct from other species of Section DRYO­ BALANOIDES for having characteristically large, broadly elliptic to oblong elliptic leaves which have short, stout caudate tips about one-eight to one-twelfth the length of the blade. The fruit, especially when young, is very similar to that of Hopea foxworthyi and Hopen woodiana, but can be readily differentiated by the number of dis­ tinct veins in the expanded sepals. This species has 7 to 9 veins in each expanded sepal, while the latter two species have 5 to 6, rarely 7 veins. The shape and size of the leaves are distinct features of Hopea malibato Foxw.

The position of PNH 97149 collected in the same site as PNH 97155, although very similar to this species, is rather doubtful. It deviates in the general form and texture of the leaves and the dark red color of the flowers. In Basilan Island, where it was col­ lected, it is known by loggers as the female manggachapui. Hopea malibato Foxw .. \ PNH 971551. is simply called mangachapui, while PNH 97150 (see Hopea woodiana sp. nov.) is known as male mang­ gachapui. Whether this is only a variant or a different species. it is included here temporarily, until more complete materials are available to delineate its proper position.

The Holotype of Hopea malibato Foxw. was destroyed in the Philippine National Herbarium in 1945. The Isotype deposited at the Arnold Arboretum, Harvard University, has been chosen as the Lectotype of this species.

Hopea malibato Foxw. belongs to the Yakal Group. [39, p. 84]. It is a small to medium size tree, attaining a height of 36 meters, and a diameter at breast height of 50 centimeters. The bole is straight, attaining 20 to 26 meters in length. The bark is smooth, " ACTA MANILANA grayish to almost dark-chocolate in color. The outer bark is thin, while the inner bark is ahout 6 millimeters thick. The wood is brownish white, turning brown on exposure. It is medium hard to hard. The features of the wood are very similar to those of Hopea basilanica Foxw.

It is locally used for structural timber, bridge construction, railroad ties and other permanent construction. The supply is very limited. This species is rather rare.

4. HOPEA WOODIANA Gutierrez, sp. nov. Text fig. 8; Plate IV.

Hapea pieTTei Foxworthy, pro parte PHILIP. JOUR. SCI. 6 (1911) Bot. 265; 13 (1918) Bot. 183; Merrill, ENUM, PHILIP FL. PL. 3 (1923) 94; Reyes, PHILIP. JOUR. Sc. 22 (19'23, pI. 31, jig. 1, non Hance.

Hapea foxworthyi Foxworthy, pro parte PHILIP. JOUR. SCI. 67 (1938) Bot. 282, non Elmer.

. Arbor 1?-45 m: alta. Ramuli puberulentes, sed adult.e­ rlO.re~ glabn. Foha ehartaeea usque subcoriaeea, glabra, elhptlca ad ovato-Ianceolata, (4-) 5.5-6.5 (-8) cm. longa, 1.5- 2.5 (-3) em. lata, apice caudato-acuminata, summa caudato-- ~~t~!~~r~~is~u~u~~f~~rfi~s~~l~e~ti~~~e~~;,inc~~ ui~~~s~~~d' ;1~; venis alternantibus, venulibus reticulatis indistinctis; petiolis 0.5-1 cm. longis, puberulibus, ultimatim glabris, valde sulca­ tis, evidenter planis. Infloreseentiae puberulae, ultimatim gla­ brae, terminales et/vel axillares, singulatim vel binae, strata­ formae ordinatae, 1-1.5 em. longae; racemi 1-4, vix 6, nores pedicellatos gerentes; braeteolae fugaces, ovato-lanceolatae, 1 mm. (vel breviores) longae, pubescentes extus, glabrae intus. Pediculi 1-2 mm. longi, haud lepidoti. Calyx lepidotus, cum 2 sepalis externis ova tis, 1.5-2 mm. latus. Stamina 15, 3 series ex 5, evidenter 3 stamina unicuique petalo; filamenta variabilis longitudinis; anthem orbiculata, 0.5 mm. long.a, cum arista appendice terminali vix filamentibus brevio:ri. Stigma minuseulum; stylu.s filiformis, 1-15 mm. longus. Sty­ lopodium 0; ovarium oviforme, 1-1.5 mm. longum, 1 mm. latum, glabrum. Fructus oviformis, cum sepahs alae-formis, rubidis, basi cruciatis, oblanceolatis, basim versus acuminatis, 2.5 em. longis, 0.6 em. latis, cum 5-6, raro 7, venis primariis in unaquaquae, cum sepalis interioribus fructus longitudinem totam adhaerentibus atque circumtenentibus. A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 43

Trees 10 to 45 m tall, the stem about 15 to 60 em in diameter at breast height. Branchlets puberulent, becoming glabrous with age, dark-chocolate in color. Leaf-blade chartaceous to subcoriaceous, glabrous, narrowly elliptic to ovate-Iancolate, (4-) 3.5-6.5 (-8) cm long, 1.5-2.5 (-3) cm wide; when dry, grayish brown above, light brown beneath, approximately two to three times as long as broad, apex caudate-acuminate, the caudate tip about one-third, rarely one­ half, to one-sixth the length of the blade, obtuse, the base cuneate, midvein distinct on both sides, glabrous, the principal lateral veins 8 to 13 on each side, curving, with 1 to 2, rarely more, alternating less conspicuous veins, reticulate venules not distinct; domatia hol­ low depressions, confined to the axils of succeeding veins (up to 6 domatia, rarely 11), with tufts of hairs on the margins; petioles 0.5-1 cm long, puberulous, ultimately glabrous, finely sulcate, some­ what flattened. Stipulps not seen. Inflorescences puberulous, ul­ timately glabrous, dark-chocolate in color, terminal and/or axilla­ ry, borne singly or in pairs in the axils of the leaves or of fallen leaves, those in pairs arranged one above the other, 1-1.5 cm long, the racemes bearing 1 to 5, rarely 6, pedicellate flowers; bracts not seen; bracteoles fugacious, ovate-Ianceolate, 1 mm long or less, pubescent outside, glabrous within. Pedicels 1-2 mm long, sparsely lepidote. Calyx lepidote, the two outer sepals ovate, densely lepidote on the outer and inner surfaces of the blunt apices, 1-1.5 mm. wide; the inner sepals suborbicular, slightly pointed at the apex, with hairs on the upper margin, 2-2.5 mm long, 1.8-2.5 mm wide. Petals pink, 6-8 mm long, 1.5-2 mm wide, approximately three to four times as long as broad. Stamens 15, in 3 whorls of 5, s~emingly 3 stamens to each petal, the first and third opposite the sepals; filaments of different lenghts, the first whorl 0.5 mm long, the second 0.8 mm, the third 1 mm; anthers orbicular, 0.5 mm long, the awn like ter­ minal appendage as long as the filaments. Stigma minute; style slender, 1-1.5 mm long, as long as or longer than the length of the ovary, occasionally bearing hairs at the base. Stylopodium absent; ovary ovoid, 1-1.5 mm long, 1 mm wide, glabrous. Fruit ovoid, the winglike sepals brick red, crossed at the base, oblanceolate and nar­ rowed toward the base, 2.5 cm long, 0.6 cm wide, with 5-6, rarely 7, principal veins in each, the inner sepals appressed to and en­ closing the whole length of the fruit.

LUZON: Provo Laguna, F.B. 17638 H.M. Curran, Feb. 1910, in fruit, (NY), (US-2 sheets); F.B. 24665 F.L. Amarillas, Nov. 1915, in flower, (NY), (US); San Antonio, M. Ramos 397, Aug. 1910, in .. ACTA MANILANA

FIG. 8 Hope4 WOOdia11Q, Gutierrez, Sf!. noY. A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 45 fruit, (US); Paete, M.D. Sulit 27, Feb. 1933, in fruit, (NY); Sulit & Canicosa 13, March 1933, in fruit, (A); PNH 3137 Sulit & Canico­ sa, April, 1933, in fruit, (A); Famy, Minayotan, Sigsigon, PNH 83707 M. Lagrimas, Feb. 18, 1958, in fruit, (PNH); PNH 83577 Sulit & Canicosa, May 15, 1968, sterile, (PNH); PNH 83732 Sulit & Canicosa, May 23, 1958, in fruit, (PNH); Mt. Makiling, PNH 33392 T.B. Banaga, March 8, 1955, in flower, (PNH), (A), (US-2 sheets); PNH 33378 D. Olegario, Jr., Jan. 30, 1955, in flower, (PNH), (US); PNH 33440 Severino Raneses, Jan. 16, 1955, in flower, (PNH-. sheets); (US); PNH 33407 P.P. Lazo, Feb. 22, 1955, in flower, (PNH), (US); PNH 37310 R. Qll.itoles, Jan. 30,1955, in flower, (PNH-2 sheets); PNH 35360 H. P. Amis, Feb. 5, 1955, in flower, (PNH); PNH 35514 R. Galutera, Jan. 22, 1955, in flower, (PNH); PNH 37535 Melchor Tolentino, Jan. 27, 1955, in flower, (PNH-2 sheets), (Al; PNH 34349 J. Gerardo, Feb. 26, 1955, in flower, (PNH); Makiling National Park, PNH 35802 M.D. Sufit, Feb. 16, 1956, in fruit, (PNH-3 sheets); PNH 22873 E. Canicosa, Feb. 26, 1955, in fruit, (PNH); Provo Tayabas (Quezon); F.B. 30971 M. Oro, June­ July, 1929, in fruit, (NY); Tayabas, F.B. 24753 F. Manuel, Oct.-Nov. 1915, in flower, (A), (US); Infanta, Kalumpit, PNH 37270 M. Lag­ rimas, March 24, 1956, in fruit, (PNH-2 sheets); Infanta, Subayat, Nonok, PNH 83334 M.Q. Lagrimas, March 27, 1956, sterile, (PNH); Infanta, Llavac, Kalauang, PNH 83578 M. Lagrimas, Oct. 30 1955, in flower, (PNH); Llavac, Real, PNH 74057 M. Lagrimas, May 30, 1962, in fruit, (PNH); Provo Albay; F.B. 10579 H.M. Curran, June, 1908, in fruit, (NY); Polillo Island, F.B. 26249 Valderama & Soriano, Oct. 1916, in flower, (A), (US); PNH 6501 A.P. Castro & P. Ano­ nuevo, Dec. 2, 1948, in flower & fruit, (PNH), (Al, (UC).

MINDANAO: Distr. Zamboanga, F.B. 13900 Foxworthy, De Mesa & Villamil, May-June, 1912, in fruit, (US); F.B. 21883,21991 & A. Villamil, May-June, 1914, in flower, (US); Basilan Island, Maluso, Canas, PNH 97150 Gutierrez, Reynoso & Fraga, May 10, 1965, in flower, (Holotype: PNH), (lsotypes: to be distributed).

LOCAL NAMES: barakbakan (Mba.), dagingdingan (Tag., Bik.), dala (Neg.), dalingdingan (Bik., Tag., Dum.), dalingdingan­ isak (Tag., Bik.l, isak (Tag.), kaliot (Pang.), lito (Ibn.), malata­ gum (Bik. ), manggachapoi (Bik., Tag., Mba.), manggachapui (Pang. ), mullllogum (Bik. l, pilak (Bik. l, pisak (Ibn.), sarabsaban " ACTA MANILANA (Mang.). siyau. (S. L. Bis.), sugkad (S. L. Bis.): the above common names which were referred to Hopea pieTTei by Merrill [30, Vol. 3, p. 94], presumably belong to this species; dalingdingan (Official Common Name): this name is adopted by Salvosa [32, pp. 36, 106] for Hopea foxworthyi, whose members for the most part belong to Hopea woodiana; manggachapui male (Basilan loggers).

This species has a distinct midrib, an ovoid ovary, and no styla­ podium. It has occasional small leaves which may be confused with those of Hopea foxworthyi Elm., but the presence of a distinct mid­ rib readily distinguishes it from the impressed character of the latter.

The author believes that the wood specimens collected and studied by Reyes [31, p. 3391 for the supposedly Hopea pieTTei Hance were gathered from this species. It is rather unfortunate that his work is not documented by herbarium specimens which we could at least verify. On the other hand, we can at least presume from his statement that the species is best known in Laguna and Tayabas (Quezon) Provinces. Our findings show that, while Hopea woodia­ na sp. nov. is widely distributed in the non seasonal part of the Philippines, Hopea fOXWOTthyi Elm. is found and confined only in the seasonal island of Sibuyan.

The species of Hopea fOXWOTthyi Elm. which belongs to the Manggachapui Group, [39, p. 70J is, for the most part, Hopea woodia­ na sp. nov. The bole of the tree is regular and straight; mature trees, often, strongly buttressed. The crown is rather heavy, dark­ green, with few heavy branches. The bark is grayish brown, rather smooth when young, fissured and flaking irregularly with occasional hardened resins. The sapwood is thin and lighter-colored than the heartwood, which is brown and turning to dark brown upon ex­ posure. The wood is fine to medium texture, fairly straight grained, moderately hard to hard, moderately heavy to heavy (Basilan mem­ bers are used as sinkers in log ponds), specific gravity is 0.793. [31, p.339].

It is locally used for bridges and other heavy constructions for which woods of great strength are required. It is a favorite wood for doors, window frames, flooring boards, furniture and agricultu­ ral implements.

This species is named in honor of Dr. Carrol E. Wood, Jr., of the Arnold Arboretum, Harvard University. A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 47

5. HOPEA ACUMINATA Merrill Text fig. 9, Plate V.

Hopea actlminata Merrill, BUR. GOVT. LAB. PUBL. (Philip.) 29 (1905) 30; PHILIP JOUR. SCI. SUPPL. 1 (1906) 98; ENUM. PHILIP. FL. PL. 3 (1923) 93; Whitford, PHILIP. JOUR. SCI. 4 (1910 I Bot. 703; PHILIP. BUR. FOR. BULL. 10 (2) (1911) 75, pI. 80; Foxworthy, PHILIP. JOUR. SCI. 2 (1907) Bot. 389; 4 (1909) Bot. 514, pI. 27, fig. 68; 6 (1911) Bot. 264; 13 183; 67 (1938) Bot. 280; Reyes. PHILIP. JOUR. SCI. 22 (1923) 339, pl. 30, fig. 1; Symington, CARD. BULL. S.S. 8 (1934) 22.

Hopea maqtlilingensis Foxworthy, PHILIP. JOUR. SCI. 13 (1918) Bot. 184.

ORIGINAL DESCRIPTION:

"A tree reaching a height of 40 m., with broadly lan­ ceolate, narrowly acuminate, glabrous leaves, 8 em. long or less, small flowers in unilateral racemes, which are arranged in terminal and axillary panicles. Branches nearly black when dry, submembranous, shining above, the base inequilateral, rounded at least on one side of the la­ mina, the apex long slender, acuminate; nerves 8 to 10 pairs. rather prominent beneath; petioles 5 to 8 mm. long, rugose, glabrous. Inflorescence garyish-stellate-pubescent, the panicles 5 to 6 cm. long or less. Calyx rusty-pubescent, the sepals 5, imbricate, the outer two slightly larger t~an the inner. Petals slightly pubescent on the outside, falcate, 4 mm. long, 1.8 mm. wide, obtuse. Stamens 10; filaments thick; anthers 0.4 mm. long, the single appendage slender, equaling the anther in length. Ovary glabrous, 3-cell~d, each cell 2-ovuled; style short; stylopodium O. Fruit gla­ brous, the two wings oblong, about 2 cm. long, 5 mm. wide, the apex rounded.

"Specimens examined all from Lamao River, Mount Mariveles, Province of Bataan, Luzon: No. 786 (Forestry Bureau), collected by Borden, May, 1904 (flower); No. 335 (Whitford), "May, 1904 (flower); No. 3864 (Merrill), and No. 1592 (Forestry Bureau), Borden, August, 1904 (fruit); Nos. 825, 1175, and 1245 (Forestry Bureau), Borden, May­ June, 1904 (sterile).

"A tall tree, with a slender, straight trunk, growing in the hill forests from an altitude of 100 to about 800 m. above the sea. T. Dalindingan." .8 ACTA MANILANA

FIG. 9 Hopea acuminata M~rrill A REVISION OF THE GF.NUS HOPEA ROXflURGH OJ.' 1'. 1.

A tree 30 m tall, the stem about 30 cm in diameter at breast height. Branchlets pubescent, becoming glabrous with age, dark~ chocolate in color. Leaf~blade chartaceous, with minute waxy scales, occasionally with few scattered hairs on the veins, oblong to ob~ long-elliptic, (4-) 6.5-9 (-12) cm long, (2-) 3-4 (-5.5) cm wide, approximately 2 to 3 times as long as broad, apex long acuminate. base obtuse, oblique, rounded on the side above the midrib; mid­ vein prominent on both sides, sparsely hairy, the principal lateral veins (8-) 9-12 pairs, prominent beneath, curving, alternating pa­ rallel veins absent, reticulate venules distinct beneath, more or less parallel; domatia pouch like, confined to the axils of one to three pairs of veins, absent in some leaves, hairs absent on the margins; petioles 0.7-1 cm long, finely sulcate, more or less terete. Stipules not seen. Inflorescences velutinous, light brown, terminal and/or in the axils of leaves or of fallen leaves, 1 to 3 panicles-when more than one, arranged side by side, (1.3-) 2-5 (17) cm long, the racemes bearing 3 to 9 small, pedicellate flowers; bracts not seen; bracteoles pubescent, ovate-lanceolate, about 1 mm long. Pedicels 1 mm long or less, pubescent. Calyx densely pubescent, the 2 outer sepals ovate to ovate-elliptic, blunt at the apex, 1-1.3 mm long, 1 mm wide, the inner sepals orbicular, slightly pointed at the apex. 1-1.2 mm long, 0.8-1 mm wide. Petals 3-3.5 mm long, 1.5-2 mm wide, appro­ ximately twice as long as broad. Stamens 10, in 2 rows of 5, the first opposite the petals, the second opposite the sepals; filaments of one length, 0.5 mm long; anthers orbicular, 0.4 mm long, the awnlike terminal appendage as long as the anthers. Stigma minute; style slender, short, 0.2 mm long, about one-fourth to one-fifth the length of the ovary. Stylopodium absent; ovary globose, 1 mm. long, glabrous. Fruit ovoid, 7 mm. long, 6 mm. wide, apiculate, the winglike sepals yellow, glabrous, occasionally with minute hairs, parallel at the base, oblong-elliptic to oblanceolate and narrowed toward the base, 1.5-5 cm long, 0.6-1 cm wide, with 6-7 distinct principal veins in each, the inner sepals tightly appressed to and enclosing almost the whole length of the fruit.

LUZON: Provo Babuyanes, Calayan Island, F.B. 26706 N. Pe­ nas, May, 1917, in flower, (A), (US); Provo Cagayan, Penablanca, B.S. 263 M. Adduru, May 1 - June 18, 1917, in flower, (A), (US), IF); Provo Tayabas (Quezon), F.B. 10390 H.M. Curran, May, 1908, sterile, (NY); Provo Bataan, F.B. 27147 A. Udasco, May, 1918, in !lower & fruit, (A), (US); F.B. 22044 J. Medina, July, 1914, in fruit, (Topotype: Hopea maquilingensis Foxw - US); Lamao Ri­ ver, Mt. Mariveles, F.B. 786 T.E. Borden, May, 1904, in flower, 50 ACTA MA:-':ILA:-\A

Holotype: PNH, destroyed in 1945), (Lectotype: NY" Ilsotype: US); F.B. 717 T.E. Borden, May, 1904, sterile. (NY" IFI; F.B. 825. 1175 & 1245 T.E. Borden, May·June, 1904, sterile, I NY I, I US); T.E. Borden s. n., 1903-4, sterile, (A I, I UC-Nos. 216221. 238465 238468 & 2385631; F,B. 7504 H.M. Curran, Sept. 1907, in fruit. IUS), NY-sterile); E.D. Merrill 3864, August, 1904, in fruit, (Note: de­ signated by Foxworthy to represent the Type of the species in fruin, (NY), (US); H.N. Whitford 335, May, 1904, in flower, (NYI, IUSI. (F); Provo Laguna, F.B. 24670 F.L. Amarillas, Nov,1915, sterile, (US); Mt. Makiling, F.B. 22966 F. Canicosa, Oct. 1914, in fruit, I Paratype: Hopea maquilingensis Foxw.J, (NY), (US), I A-sterile I; PNH 3138 M. Sulit, April, 1933, in fruit, I A); F.B. 21988 A. Villamil, August 1914, in fruit, (US I; Makiling National Park. Sulit & Canicosa 9. April. 1932, in flower, (NYI; Sulit & Canicosa 12, May. 1933. in flower, (A); Provo Camarines, Mt. Isarog, F.B. 10459 H.M. Curran. May, 1908, sterile, (NY), (US).

MINDANAO: Provo Zamboanga, F.B. 22758 E. Nave, June, 1914, in flower, (US).

LOCAL NAMES: baniakau (Ibn.), barosingsing (Ilk.), daling­ ding (Tag.), dalingdtngan (Tag., S.L. Bis.), kalot (Ilk.)., mangga­ chaput (Tag., Bik., Bis., Sul., Sml.) manggachinoro (Tag.), mang­ gasinoTo (Tag. 1, siayu (S.L. Bis. J, 1jakal (Tag.), manggachapui (Of­ ficial Common Name).

Hopea acuminata Merr. is the only Philippine species that has 10 stamens. It may be confused with Hopen plagata (Blanco) Vid. by the oblong to broadly elliptic shape and the size of the leaves. However, the velutinous inflorescences and the oblanceolate wing­ like sepaJ.s of the mature fruit of the former are readily distin­ guished from the glabrous inflorescences and the orbicular, mature winglike sepals of the latter. The number of the stamens is a dis­ tinct character of the species, not found in other species of Philip­ pine 11opea.

Merrill [26, p 30] described Hopea acuminata Merr. from se­ veral collections gathered in Mt. Mariveles, Bataan, but he did not designate the Type specimen. Foxworthy [16, p. 364] designated F.B. 786 T. E. Borden, in flower, as the Type chosen from Syntypes, when he revised the genus for the Philippines in 191 L In his sub­ sequent revision of the genus in 1938, [19, p. 280] he designated E. D. Merrill 3864, as the other specimen to represent the Type of the A REVISION OF THE GENUS HOPEA ROXBL"RGH OF P. I. 51 species in fruit. Unfortunately these specimens were destroyed in the Philippine National Herbarium during the Second World War. Thus the author is designating here a new set of Lecto-syn­ types from the extant Iso-syntypes deposited at the Herbarium of the New York Botanical Garden, to represent the species.

The reduction of Hopea maquilingensis Foxworthy to the sy­ nonymy of Hopea acuminata Merr. was done by Merrill (26, p. 93]. Merril states that the Type of the former is a specimen with ma­ ture fruits, beautifully matched by a Topotype, Medina F. B. 22044. of the latter. This reduction was accepted by Foxworthy, stating that the features mentioned may be within the natural range of variation of the species. The descriptions of these two species show that they are very alike, differing only in the size of the fruits. Fox­ worthy [18, p. 184] states, "Arbor, H. acuminatae similis sed fructi­ bus majoribus" The author, having studied the materials cited by Merrill. confirms the reduction valid.

Hopea acuminata Merr. belongs to the Manggachapui Group [39, pp. 69-70]. The tree reaches a height of about 35 meters and a diameter of from 32 to 40 inches. It has a straight regular bole that goes up to 18 meters in length. The buttress is not prominent.

The sapwood [31, p. 339] is thin, pale white in color and sus­ ceptible to dark brown sap stain. The heartwood is pale grayish yeJIow, becoming golden brown upon exposure. The wood is fine to medium textured and fairly straight grained. It is moderately hard and heavy, with a specific gravity of 0.783.

The wood of (Anisoptera thurifera [Blanco] Blume) palosapis, which is an inferior grade of timber, is frequently substituted for this species. However, palosapis can be distinguished from mang­ gachapui by its coarser texture and larger pores, which are plai'nly visible to the naked eye. The resin cysts of manggachapui are ar­ ranged in concentric rings, while those of palosapis are diffused.

It is generally used for bridges and rough construction requir­ ing wood of medium strength and durability. It is a favorite wood for doors, window frames, sills and flooring boards. It is one of the best-known woods for shipbuilding, although less durable than the yakal. FIG. 10 Hopea basilanka Foxworthy A REVIS lOX OF THE GENUS HOPEA ROXBURGH OF P. 1. 53

6. HOPEA BASILANICA Foxworthy Text fig. 10, Plate VL

Hopea basilanica Foxworthy PHILIP. JOUR. SCI. 6 (1911) Bot. 260. pI. 42; 13 11918! Bot. 183; 67 (1938) Bot. 275; Merrill ENUM. PHILIP. FL. PL. 3 t 19231 93.

ORIGI~AL DESCRIPTION:

"Abor magna: folds chartaceis. anguste oblongis vel ('llip­ tieis. glabris. supra nitidis. 10 ad 15 em long is. 2.2 ad 4.8 em latis. brevissime caudato-aeuminatis. basi paulo obliquis. obtusis. utnnque glaberrimis. costa media utrinque nervis lateralibus subtus prominentibus; petiolo brcvissim{l; paniculis brevibus ter­ minalibus vel laterulibus: floribus. : fructu majusculo. calyce acereto cinch): calvcis fru(:tifcri lobi aucl! daubus alis 3.5 ad -1:.3 em longis. latitudl'ne fere 10 ad 15 mm.

"Large tree 18 m tall, with thick. blaci;:. ful'ro ... ·C'ci bark. like that of H. aClimimlto. Lcaves chartaceous. narrowly oblanq or elliptic, glabrou.:' <;hining above. 10 to 1:') {'m l{lll.~. 2.Z to !.P em wide. Apex acuminate. tapering and u'lsymm{'tric~tl at bils('. the part above the midrib being the larger. Pctiok' dark-colore.d. and wrinkled when dry. Midrib slightly pubescent above; se­ condary nerves 10 to 12 pairs. ascending and b{'1lI upward :\t the margin of t:1e leaf. Domatia in the axils of at least the lowest pairs of veins. Tertiary veins parallel and retieulatc Inflorescences axillary in few-flowered raeemose J-lnnidcs Oc­ casionally two inflorescences from one axil. Rachis Jf inflores· eences pubescent. Fruit 3 to 4 mm in diameter. -l to 5 mm. lOll.!.:. resin-cavities in lower part of fruit. Fruiting calyx with 2 Ion>..: wings 3.5 to 4.3 cm long and 10 to 15 mm wide. with about 9 principal veins. Wings yellow whcn dry. the vcin<; b('in!:{ or darker color and the fruit and base of wings very dark-brown The bases of these two larger wings are expanded so a;, prac­ tically to conceal the small wings.

"Differs from H. odorata in shape of leaves and in its thick short petioles, in size of fruit and in expanded bases of the two enlarged ealyx wings; from H. plagata in its narrower leaves. shorter petioles, smaller fruit, with expanded bases to calyx wings; from H. philippinensis in fruit and the absenee of the lon~ stipules. "Basilan. For. BUT. 152208 Pray. Klemme, Aug. 1910 For. BUT. 15408 "Common name: dalingdingan' .

A large tree up to 64 m tall. the stem about 62 em in diameter at breast height. Branchlets pubescent, becoming glabrous with age 54 ACTA MANIl.ANA

Leaf-blade chartaeeous, glabrous, ,>hiny above, dull beneath, oblong to oblong-elliptic, (8-) 10-5 em long. (3-1 3,5-5.7 em wide. appro· ximately three to four times as long as broad, apex long acuminate base obtuse, oblique, slightly larger on the side above the midrlb: midvein prominent on both sides. with minute hairs above. glabrou~ beneath. the principal lateral veins prominent beneath. 10 or 11 pairs curving. alternating parallel veins absent. reticulate venules faintl~' visible. more or less parallel; domatia pouehlike. confined to the axils of succeeding veins (up to 6 domatia I. hairs absent on the margin: petioles 7-8 mm. long, puberulous. ultimately glabrous. more or less terete. Stipules not seen. Inflorescences densely pubescent. grayish in color, borne in the axils of leaves or of fallen leaves. 1 to 3 pa­ nicles; when more than one, arranged side by side; 2.5-9 cm. long. the- racemes bearing 2 to 6 subsessile, yellow flowers; bracts and bractEOles not seen. Pedicels 0.5 mm or less long. velutinous. Calyx densEly pubescent. the 2 outer sepals ovate. apex acute. 1-1.2 mm lon~, 0.8-1 mm wide, the inner sepals orbicular. slightly pointed at the apex. 1.2-1.4 mm long, 1-1.2 mm wide. Petals 3.5-4 mm long. 1.8-2 mm. wide. approximately two times as long as bread. Stamem 15. in 3 whorls of 5, the first and third opposite the petals. the second opposite the sepals; filaments of different lengths. the first and second whorls 0.3 to 0.4 mm. long, the third 0.2 mm.; anthers orbicular. 0.3 mm long, the awn like terminal appendage as long as the filaments. Stigma minute; style slender. short. 0.3 mm long. approximately one-fifth the length of the ovary. Stylopodium and ovary cylindrical, glabrous, slightly constricted above the middle. 1.3-1.5 mm long. 0.8-1 mm wide. Fruit ovoid, 0.7-1 cm long. 0.5-0.8 cm. wide. the winglike sepals, when dry, yellow to light brown. crossed at the base, narrowly elliptic, with expanded bases. about 3.5-4.5 cm. long, 1.1-1.4 cm. wide, with 9 distinct principal veins in each, the inner sepals tightly appressed to and enclosing almost the whole length of the fruit.

MINDANAO: Basilan Island. F.B. 15220 W. Klemme. August. 1910, in fruit, I Holotype: PNH, destroyed in 1945). I Lectotype: US); F.B. 31232 G.S. Corbett, June. 1930, in flower. rA). IUS); habela. Fabiawan. F.B. 27248 Deogracias Mendoza. June. 1918, in flower. IAr, IUS); Lamitan. Yawas. Jose Vera Santos 4131. May 7.1948, in flower, IUS).

LOCAL NAMES: dalingdingan (SuI. I. yakal I Sui.. Yak .. Chab.J. Brw/an yakal (Official Common Namel. A REVISION OF THE GENUS HOPEA ROXBURGH OF P. t. 55

Hopea basilanica Foxw. has densely pubescent. grayish inflo­ rescences. small flowers and yellow winglike mature fruiting calyces which are almost similar to Hopea aCliminata Merr. The latter may also have occasional large leaves that approach very much the shape and size of the former. However. Hopea basi/anica Foxw. has 15 stamens. in contrast to the 10 stamens of Hopea aCZlminata Merr. It is unlikely to be confused with other species of Philippine Hopea in having a large stylopodium \vhich is continuous with the ovary. forming a broadly cylindrical structure.

The original description of the species was based entirely on fruiting material. Howeyer. Foxworthy (19. p. 2751 amended the description and included a diagnosis of the flower. The flower description was based on the collection of Deogracias Mefldoza: F.B. 27248. made in the same locality.

Since nothing has been left of the original materials used by Foxworthy in describing the species. except an Isotype deposited at United States National Herbarium. it is hereby designated as the Lectotype of the species.

Hopea basilanica Foxw. belongs to the Yakal Group [39. p. 831- It is a large tree that grows up to 64 meters tall. It has a thin and open crown. The bark is furrowed. and the color is gray-brown to brown. The inner bark is yellowish when fresh, turning to brown when dry. The sapwood is narrow and the color is lighter than the heartwood. which is yellowish. The heartwood turns dark brown with age. The texture of the wood is fine and cross grained. It is very hard and very heavy, having an specific gravity of 0.917.

It is used locally for purposes requiring strong and durable wood. It is excellent for structural timber. railroad ties, bridge construction and other permanent constructions. The wood may be confused with manggachapui (Hopea acuminata Merr.), except for its harder and heavier characteristics.

This species is known only from Basilan Island, and the supply is rather limited.

7. HOPEA PLAGATA (Blanco) Vidal Text fig. 11, Plate VII.

Hopea plagata (Blanco) Vidal. REV. PL. VASCo FILIP. (1886) 62; 56 ACTA MANILANA

FIG. 11 HopeD, poogata (Blanco) Vidal A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 57

Brandis, JOUR. LINN. Soc. BOT. 31 (1895) 64; Merrill, GOVT. LAB. PUBL. (Philip.) 27 (1905) 22; SP. BLANCOANAE (1918) 269; E~UM. PHILIP. FL. PL. 3 (1923) 94; Merrill, PHLIP. BUR. FoR. BULL. 8 (19081 48; Whitford. PHILIP. JOUR. SCI. 4 (19101 715; PHILIP. BCR. FoR. BULL. 10 121 (1911173. pIs. 76, 77; Fox~ worthy. PHILIP. JOUR. SCI. 2 119071 Bot. 396; 4 (19091 Bot. 515; 6119111 Bot. 262; 13 (19181 Bot. 183; 67 (19381 Bot. 276.

Mocanera plagata Blanco. FL. FILIP. (1837 I 447

Dipterocarplls plagallls Blanco op cit. ed. 2 118451 311; ed. 3. 2 118781 212.

Anisoptera plagata Blume. Mus. BOT. LUGD.-BAT. 2 (19521 42; A. DC. PRDR. 16 121 118681 616,

Shorea reticltlata F,~Villar. NOVISS. App (18801 21, non Dyer.

Hopea sp. odorata Foxworthy, PHILIP. JOUR. SCI. 13 (1918) Bot. 183, non Roxb.

Hopeo plagota Foxworthy, pro parte PHILIP JOUR. Sq. 67 (19381 Bot. 278, non Blanco.

Hopea sp. "Gyam" Foxworthy, PHILIP. JOUR. SCI. 6 (1911) Bot. 263: paino Foxworthy. PHILIP. JOUR. Sc. 6 (1911) Bot. 285.

ORIGINAL DESCRIPTION:

"Rami nigri. Folia alterna, lanceolata. integra, glabra. rigida, longiuscula glandula longitudinaliter dehlscente in basi uniuscujusque venae lateralis paginae inferae. brcvis­ simeque petiolata. Flores axilla res, cernui, globosi, racemo­ compositi. Cal. maturus adhaerens, monophyllus, altissime 5~partitus; partibus circinatus carnosis, extern is internas tegentibus. Cor. fere usque ad basim 5-partita; partibus rotundis. superpositis. Stam. pluri-30, ovarii basi inserta. Filam. brevia. Anth. 4-angulatae, compressae, in aristam desinentes. Germen immaturum intra florem sessile, pyra­ midatum, 4~angulare. StyL O. Stigm. quasi 2 cuspides. Nux a 5 calycis partibus superata, alas simulantibus: 2 majori­ bus (sesquiuncialibus) erectis, 33 alteris parvis, Nuclei testa tenUIS et coriacea, monosperma. Semen 5-partitum. "T. Yacal - Sapolongan". .,R ACTA MANJLANA

"Vid. ApPEN. GENER.: DIPTEROr'ARPACEAE arbor magna, ob lignum valde utile notissima, resi!:am non fundens. Fruc­ tus alae in hac specie et sequenti viridulae. Nomen speci­ ficum ex foliorum glandulis, plagas simulantibus - Flor. mens. Mart .. et non alia tempore".

ORIGI:\"AL DESCRIPTlO~: Mocanera plagata.

··.1Jocanera llagada Ramas negras. Hojas alternas. lanceoladas. enteras. lampinas y tiesas. con una glandula largulta que se rompe a 10 largo. en carla vena lateral de la pagina inferior PeclOlos cortlslmus. Flores axiJares. cabiLbaJ8s. globosas. en racimos compuestos. Cal. en la madurez adherente. monoftlo. en cmco partes muy profun­ das rcdondeadas carnosas. y las de afuera cubren a las in­ tel"lOl"CS Cor. en CIllCO partes que Began casi hast a la base. redondas y sobrepuestas unas a otras. Estam. pasan de trclnta. fijos en la base del germen. Fllam. cortos. Anteras de cuatro angulus. comprimldas. terminadas con una arista Germen asentado dentm de la flor antes de la madurez. piramidal de cuatro lados Estilo ninguno. Estigm

A tree up to 55 m tall, the stem about 180 em in diameter al breast height. Branchlets glabrous. Leafblade subcoriaceous to coriaeeous, glabrous, elliptic-Ianeeolate to broadly elliptic. (4.5-) 6- 12.51-151 em long, (1.5-)2.5-6(-8.51 cm wide. approximately tw(. to three tImes as long as broad, the apex shortly acuminate. baSE: cuneate to obtuse, equal or cblique; when oblique, the base raunded and slightly larger on the side above the midrib; mid vein prominent .)n both sides. glabrous, the principal lateral veins 8 to 12 pairs on each side. curving, alternating parallel veins absent. reticulate ve­ nules distinct beneath, more or less parallel; domatia pouch like. confined to the axils of one to seven pairs of veins. absent in some leaves. hairs wanting on the margins; petioles 0.8-1 cm long, gla- A REVISION OF THE GENUS HOPEA ROXBIJRGH OF P. I. 59 brous, occasionally finely sulcate, smooth, terete. Stipules not seen. Inflorescences glabrous, dark-chocolate in color, terminal and/or borne singly or in pairs in axils of leaves or of fallen leaves. those in pairs arranged one above the other, 1.5-3 cm long. the racemes­ bearing 3 to 6 pedicellate flowers; bracts and bracteoles not seen Pedicels about 1 mm long, glabrous. Calyx glabrous. the 2 out('J sepals ovate to suborbicular, blunt at the apex. 2 mm long. 1.8 mm wide, the inner sepals orbicular, rounded at the apex. 2.3 mm long 2.6 mm wide. Petals 4.2 mm long, 2.7 mm wide. approximately two times as long as bread. Stamens 32 to 38, arranged secmmgly in 3 whorls around the ovary; filaments of several different length~ the innermost whorl about 1 mm long, outer whorls shorter; anther!; orbicular, 0.2 mm long, the awnlike terminal appendage ~horter than the length of the filaments. Stigma obtuse, obscurely 3-parted style stout, short, 0.2 mm long, continuous with the stout. fairl) distinct stylopodium, approximately one-fifth the length of the ovary Sty]opodium and ovary conical, glabrous. 2-2.5 mm L 11«. 1.3 mm wide. Fruit conical. 0.6-1.2 em. long, 0.5-1 cm. wide, apiculate; the winglike sepals, in dry state, brown, somewhat parallel at the base, broadly elliptic to almost erbicular. 2.5-4.5 cm long, 1.5-3 wide. with 12 to 14 faintly visible principal vc.ins in each, the inner sepals appressed to when young, rather lax when matured, and enclosing about one-half to one-third the length of the fruit.

LUZON: Provo Isabela, San Mariano, Sierra Madre Mountains PNH 78186 H.G. GutierTez, April-May, 1961, in flower, (PNH I, I A I: Provo Nueva Vizcaya, Bagabag, Bungol, F.B. 29530, 29531 P. Afalla July 20, 1923, in fruit, (UC); F.B. 29532 P. Atalla, in fruit, (AI, (Ue); Provo Nueva Ecija, Magat, PNH 41720 Off. in Charge, Refor. Adm., Oct. 3, 1961, fruit only, (PNH); Provo Bataan, F.B. 25889 26133 F. Alambra & H. Borromeo, July 19, 1916, in fruit, I US I: Provo Bulacan, Sibul, PNH 83806 Felipe M. Salvosa, March 1, 1962, sterile, IPNH); Sibul, Binilangoan, PNH 41835 M. Lagrimas & J.P. Raja Dec, 19, 1962, steTile, (PNH); Provo Rizal, Bosoboso, B.S. 1513 Ma­ ximo Ramos, Oct. 1906, in fruit, (US), (NY-sterile); B.S. 21692 Maximo Ramos, Sept. 1913, in fruit, (US); Langkaya, M. Ramos 190 (SPECIES BLANCOANAE, Illustrative Specimen), Nov. 3, 1914, in fruit, (NY), (US), (F), (A-sterile), (GH-sterileJ; Oriud, A. Loher 14901 & 14892, July, 1913, in fruit, (A), (UC); A. Loher 12914, March 1912, in flower, (UC); Provo Camarines, F.B. 22645 R.J. Alvarez. Nov. 1911, in fruit, (US); F.B. 21250 R, J. Alvarez, March-April. 1914, sterile, (US); Provo Albay-Sorsogon, F.B. 10623 H.M. Curran. June. 1908, sterile, (US). 60 ACTA MANILANA

MINDANAO: Provo Zamboanga, F.B. 22783 D.P. Miranda, July, 1914, in flower, (US).

LOCAL NAMES: banutan (Gad.) batik (Ilk.), gisok-gisok (Bik.), hams (P. Bis.), nutik (Buk.), paina (Tag.), panggaian (Ilk.), saplungan (Tag.), siakol (Tag.), siggai (Ilk.), siggai-na-nalaba.ga (Ilk.), siggai-mulato (Ilk.), siggai-apurau (Ilk.), taggai (Ilk., neg.), yakal (Ilk., Pang., Sbl., Tag., SuI.) I yakal-saplungan (Official Com­ mon Name).

Hopea plagata (Blanco) Vid. is the only Philippine species of Hopea that has more than 30 stamens. The expanded bases of the broadly elliptic to almost orbicular winglike calyx-lobes of the ma­ ture fruit are another distinct feature of the species. It may be confused with Hopea acuminata Men. as regards the near similari­ ties in the size and the oblong to broadly elliptic shape of their leaves. However, the number of the stamens and the size and shape uf the expanded fruiting calyces are distinctive features that dis­ tinguish Hopea plagata (Blanco) Vid. from all the other species of Philippine Hopea.

This species was originally described by Blanco, but without designating any Type specimen. Merrill [28, p. 269] states that no botanical materials representing Blanco's species were preserved by him (Blanco), or if preserved, are no longer extant.

Foxworthy's [19, pp. 278, 279] description of the species having :mly 15 stamens is wrong. Blanco's "more than 30-stamens" is the true Hopea plagata (Blanco) Vid. The author has examined flow· ering materials that fit very well Blanco's description of the more than 30-stamen condition of the species. The collection (Loher Nos. 12914 & 14901) from which Foxworthy based his description of the flower, which has 15 stamens, and the fruit, which is tomentose, is certainly not a Hopea plagata (Blanco) Vid. species. These speci­ mens are in the herbarium of the University of California at Berke­ ley It is rather confusing, however, because the features of the leaves are very much similar to Hopea plagata (Blanco) Vid. But the tomentose character of the fruit is definitely distinct from the glabrous fruit of Blanco's species. Merrill's illustrative material of Hopea plagata (Blanco) Vid. in SPECIES BLANCOANAE has the glabrous character of the fruit. It is not unlikely, therefore. that the mater­ ials cited by Foxworthy might even be another species, the identity of which, however, we shall defer until more materials have been examined. On the other hand, the author, having examined flower- A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 61 ing materials, is definitely convinced that Hopea plagata (Blanco) Vid. has more than 30 stamens.

Since it has not yet been fully established that Blanco's speci­ mens are no longer extant, the designation of the Type to represent the species shall be deferred until we have more valid reasons to do so.

Hopea plagata (Blanco) Vid. belongs to the Yakal Group [39, p. 82). ¥akal as a common name was formerly referred to this species. It is stated that, since over 95 per cent of the wood known as yakal in the lumber trade is produced by Shorea gisok Foxw., the name yakal be adopted for the latter. Saplungan is one of the common names of Hopea plagata (Blanco) Vid., and since this is the only species that carries the name saplungan, the name was there­ fore, assigned to it. Salvosa [32, pp. 81, 1061 later coined yakal­ saplungan as the official common name of the species. According to Merrill [30, Vol. 3, p. 94], "the tree is a valuable timber, widely distributed in the Philippines. It is one of the characteristic species of the primeval forest at low and medium altitudes. It is commer­ cially known as yakal, and its timber enters the Manila market in large quantities".

The tree is large, having a regular and straight bole reaching up to 15 meters in length, and reaches to about 72 inches in dia­ meter. It is strongly buttressed. The crown is semi-open, broad, and with few heavy branches. The sapwood [31, p. 3351 is thin and light-colored, gradually merging into the light-yellow heartwood which becomes dark known on exposure. The wood is finely textured with crossed grain. It is very hard and very heavy, and one of the strongest wood:; of the group. The wood is stronger and harder than the other types of yakal. Its specific gravity is 1.14. A very durable wood, closer in texture and general appearance to narig (Vatica manggachapui

Foxw. I than the other species of the genus. Yakal-saplungan can be distinguished for its resin dUcts, which are arranged in tangen­ tial rows, while those of narig are diffused.

It is used locally for general house construction, especially for posts, shipbuilding, bridge timber, wharf, railroad ties and other uses requiring strength and durability. ACTA MANILANA

FIG. 12 Hopea mindammsis Foxworthy A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 63

8. HOPEA MINDANENSIS Foxworthy Text fig. 12, Plate VHI.

Hopea mindanensis Foxworthy, PHILlP. JOUR. SCI. 6 (1911) Bot. 261, pl. 43; 13 (1918) Bot. 183; 67 (1938) Bot. 277; Merrill, ENUM. PHILlP. FL. PL. 3 (1923) 93; Reyes, PHILlP. JOUR. SCI. 22 (1923) 334, pI. 29, fig. 2; Symington, GARD. BULL. S.S. 8 (1934) 32.

ORIGINAL DESCRIPTION:

"Arbor magna, 12 ad 15 m. alta, 25 cm. diam.; foHis anguste oblongis, brevissime acuminatis, basi irregulariter corda tis, auriculatis, costa media utrinque nervis laterali­ bus subtus prominentibus 18 ad 24; petiolo brevissimo (5 ad 15 mm.), pubescente. Fructu majusculo calyce accreto cinc­ to; calycis lobis majoribus basi ovato tumido. limbo late spathulato, apice rotundato, basim versus valde attenuato, nervis 7 percurso.

"Mindanao, District of Zamboanga, For BUT. 9029 Whit­ fOTd & Hutchinson (Type). Also represented by FOT. BUT. 9376 WhitfoTd & Hutchinson, Jan. 1908, FOT. BUT. 9436, Feb. 1908, all three numbers from the same locality.

"This species differs from H. philippinensis in the larger size of leaves and fruit, the pubescent petioles and the auriculate leaf-base.

"Common name: magasusu."

A tree up to 34 m tall, the stem 45 cm in diameter at breast height. Branchlets robust, velutinuous, becoming glabrous with age. Leaf-blade chartaceous to subcoriaceous, glabrous, narrowly oblong, 39-51 cm long, 13-17 cm wide, approximately three to four times as long as broad, the apex shortly acuminate, base cordate, auricu­ late, unequal, larger on the side above the midrib; midvein distinct above, prominent beneath, the principal lateral veins prominent, 24-26 curving, alternating parallel veins absent, reticulate venules distinct beneath, more or less parallel; domatia hollow depressions, present in the axils of almost all the veins, provided with tufts of hairs on the margin; petioles 1.5-2.5 cm long, velutinous, finely sul­ cate, more or less terete. Stipules not seen. Inflorescences borne singly or in pairs in axils of the leaves or of fallen leaves, or on the nodes appearing as though adventitious, those in pairs arranged side by side, 1-10 em long, glabrous, dark-chocolate in color, the racemes bearing 4 to 7 subsessile flowers; bracts and bracteoles not seen. Pedicels 0.5 mm or less long, glabrous. Calyx glabrous, " ACTA MANILANA the 2 Quter sepals ovate, blunt at the apex, 3.5 mm long, 2.5 mm wide, the inner sepals orbicular, rounded at the apex, 3.5 mm long, 3 mm wide. Petals 8-9 mm long, 2.5-3 mm wide, approximately 3 to 4 times as long as broad. Stamens 15, in 3 whorls of 5, the first and third opposite the petals, the second opposite the sepals; fila­ ments of different lengths, first whorl 0.5 mm long, second 0.9 mm, the third 1.2 mm; anthers orbicular, 0.3 mm long, the awnlike ter­ minal appendage as long as the filaments. Stigma minute, style slender, short, 0.2 mm long, approximately one-seventh the length of the ovary. Stylopodium and ovary bottleshaped, glabrous, 2.5 mm long, 1.2 mm. wide. Fruit large, conical, 1.5 em. long, 1.2 em. wide, apiculate, the winglike sepals crossed at the base, oblanceolate, more or less rounded at the apex and narrowed toward the base, 11-12 cm. long, l.4-l.6 cm. wide, with 6 to 7 principal veins in each, the inner sepals appressed to and enclosing more than half the length of the fruit.

MINDANAO: Distr. Zamboanga, F.B. 13782 Foxworthy, De Mesa & Hutchinscn, December 2, 1907, in fruit, (Holotype: PNH, destroyed in 1945), (Lectotype: NY), (Isotype: US) Mt. Tubuan, B.S. 36652 M. Ramos & G. Edana, October, 1919, in fruit, (US); Ma­ iangas, B.S. 37069 M. Ramos & G. Edano, October-November, 1919, in flower, (A), (US); Naga-naga, F.B. 27402 S. Babaran, December, 1918, in flower (A), in fruit (US).

LOCAL NAMES: bagasusu (Zamb.); magasusu (Sul.); ganon (Sub.); yakal-magasusu (Official Common Name).

Hopea mindanensis Foxw. is a very distinct species which can easily be differentiated from all the other species of the Philippine genus Hopea. It has very large leaves which have cordate-auricu­ late bases, with velutinolls petioles. The fruits and the flowers are comparatively larger. Aside from the possible exception of Hopea philippinensis Dyer, which can have occasional large leaves (but never having more than 22 pairs of veins), it is not likely to be confused with any species of Hopea known in the Philippines.

Since the Holotype was destroyed in the Philippine National Herbarium in 1945, the Isotype deposited at the herbarium of the New York Botanical Gardens has been chosen as the Lectotype of the species.

Hopea mindanensis Foxw. belongs to the Yakal Group [39, p. 84] The bole is generally twisted and attains a length of from 8 A REVISION OF THE GENUS HOPEA ROXBURGH OF P. 1. 65 to 10 meters. The bark is distinctly longitudinally fissured, and nearly black with reddish tinge. It has a thin, grayish white sap­ wood, subject to sap stain. The heartwood is reddish brown in color, with concentric greenish zones which turn nearly black with age. The wood is fine to medium textured and fairly straight grained. It is very hard and very heavy, with a specific gravity of 0.849. [31, p. 334].

The supply is fairly abundant and the wood is used locally for structural timber, railroad ties, bridge construction, and other purposes where strong and durable wood is required. According to Reyes, [31, p. 334], it is similar to narek (Balanocarpus cagayanensis Foxw.), but it can be distinguished from the latter by the reddish tinge of the wood. It is more durable than the member species of the Manggachapui Group, but less durable than the other mem­ ber species of the Yakal Group belonging to the genus Hopea, such as Hopea ba.silanica Foxw., Hopea plagata (Blanco) Vid., and Hopea malibato Foxw. It may be confused with the wood of Hopea basila­ nica Foxw., except for the presence of greenish longitudinal streaks.

9. HOPEA PHILIPPINENSIS Dyer Text fig. 13, Plate IX.

Hopea philippinensis Dyer, JOUR. BOT. 16 (l878) 100; Vidal, REV. PL. VASCo FILIP. (1886) 62; Ceron, Cat. PI. Herb. Ma­ nila (1892) 26; Brandis, JOUR. LINN. Soc. BOT. 31 (1895) 64; Everett & Whitford, PHILIP. BUR. FOR. BULL. 5 (1906) 16,28,53; Whitford, PHILIP. BUR. FOR. BULL. 10 (2) (1911) 75, pI. 79; Foxworthy, PHILIP. JOUR. SCI. 4 (1909) Bot. 515; 6 (1911) Bot. 261; 13 (1918) Bot. 183; 67 (1938) Bot. 276; Merrill, ENUM. PHILIP. FL. PL. 3 (1923) 94; Reyes, PHILIP. JOUR. SCI. 22 (1923) 388, pl. 30, fig. 2; Symington, GARD. BULL. 5.5. 8 (1934) 32; (1935) 279.

Hopea odorata Vidal, SINOPSIS ATLAS (1883) 15, t. 15 fig. a, excl. fruit, non Roxb.

ORIGINAL DESCRIPTION: "Arbor, ramulis ad angulum 45° fere divergentibus, teretibus, siccitate rugosis, pallide fuscis, novellis fusco­ canescentibus; foHis anguste oblongis, brevissime caudato­ acuminatis, basi paulo obliquis, obtusis, utrinque glaberri- 66 ACTA MAN1LANA

FIG. 13 Hopea philippine1l8i. Dyer A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 6i

mis, costa media utrinque nervis lateralibus subtus promi­ nentibus; petiolo brevissimo; paniculis brevibus terminali­ bus vel lateralibus; floribus ... ; fructu majusculo calyce accreto cincto; calycis lobis majoribus basi ovato tumido, limbo late spathulata, apice rotundato, basim versus valde attenuato, nervis 8 percurso.

"Philippine Island, H. Cuming, 879.

"Folia 4-6 poll, longa, 1-1/4 - 2 poll. lata; petiole 1/4 poll. longo. Calycis fructiferi lobi aucti 2-1/2 - 3 poll. longi, latitudine fere pollicares. Capsula semi-pollicaris.

"The dried fruits assume the same rich chocolate-brown colour which is observable in the New Guinea species."

A tree 7-20 m tall, 25-40 cm in diameter at breast height. Branch­ lets pubescent, becoming glabrous with age, brown to dark-chocolate in color. Leaf-blade chartaceous to subcoriaceous, glabrous, narrowly elliptic to oblong-elliptic, (8-) 11.5-19.5 (-241 cm long, (2.5-) 3.5- 6.5 (-75) cm wide, appromately 3 to 4 times as long as broad, apex long acuminate, base obtuse, unequal, rounded on the side above the midrib; mid vein prominent on both sides. the principal lateral veins prominent beneath, 14-22 pairs, curving, alternating paraUel veins absent, reticulate venules not distinct above, fairly visible be­ neath, more or less parallel; domatia hollow depressions, confined to the axils of ten to nineteen pairs of veins, with tufts of hairs on the margin; petioles 5-8 mm long, velutinous. finely sulcate, more or less terete. Stipules semi-persistent. ianceolate, pubescent, 2-3.5 mm long, 1 mm wide. Inflorescences glabrous, dark-chocolate in color, tErminal and/or borne in the axils of leaves or of fallen leaves, or on the nodes, appearing as though adventitious; 1 to 4 panicles; when more than one, arranged one above the other; 2.5-7 cm long, the racemes bearing 2 to 7 subsessile and sessile flowers; bracts and bracteoles not seen. Pedicels of subsessile flowers 0.5 mm long. Calyx glabrous, the 2 outer sepals ovate, blunt or acute at the apex, 2.5-3 mm long, 2-2.3 mm wide, the inner sepals orbicular, rounded to slightly pointed at the apex, with minute hairs on the upper margin, 2-2.2 mm long, 1.8-2 mm wide. Petals 5-6 mm long, 2-2.5 mm wide, approximately 3 times as long as broad. Stamens 15, in 3 whorls of 5, the first and third opposite the petals, the second opposite the sepals; filaments of different lengths, first whorl 0.5 mm long, second 0.8 mm, the third 1 mm; anthers orbicular, 0.3 mm long, the awnlike terminal appendage as long as the innermost fila- f;g ACTA MANILANA

ments. Stigma minute; style short, 0.2 mm long, approximately one-tenth the length of the ovary. Stylopodium and ovary bottIe­ shaped, glabrous, 2 mm long, 1 mm wide. Fruit conical, 6-7 mm long, 5-6 mm, apiculate, the winglike sepals red when fresh, light to dark brown when dry, crossed at the base, linear-oblong to ob­ lanceolate and narrowed toward the base, 6-10.5 cm long, 1.2-2.5 cm wide, with 8-9 principal veins in each, the inner sepals ap­ pressed to and enclosing the whole length of the fruit.

LUZON: Provo Nueva Ecija, Bongabong, Vidal 84, (M); Provo Laguna, F.B. 17650 H.M. Curran, Feb. 1910, in flower, (NY), (US); Makiling National Park, PNH 83515 M.Q. Lagrimas & J.P. Rojo, Apnl 26,1961, in fruit, (PNH); PNH 74059 Justo P. Raja, Sept. 21, 1962, sterile, (PNH); Mt. Makiling, PNH 09559 B.T. Tesoro, Feb. 27, 1949, in flower, (PNHI, (A); Provo Tayabas (Quezon), PNH 9790 Canicosa. May 9, 1949, in fruit, (PNH) (A); F. B. 25323 F. Manuel, March, 1914 in flower, (A), (US); F. B. 25410 G.J. Labitag. April, 1916, in fruit, (A), (US); F.B. 10389 H.M. Curran, May, 1908, in fruit. (NY), (US); Tagkauayan, Mapalot, PNH 39420 M. Lagrimas, June 4, 1959, in fruit, (PNH); Kinakutan, F.B. 30798 M. Oro, Jan. 1929, in flower, (NY); Provo Camarines, F.B. 21711 Penas, Soriano & Abellanosa, April, 1914, in fruit, (US); F.B. 28068 R. V. Alambra, July, 1920, sterile, (UCl; Mt. Labo, PNH 4933 M.D. Sulit, May. 1939, in fruit, (A); Lagonoy, F.B. 30332 M.A. Tamano, April, 1926, in fruit, (A), (US), (UC); Provo Albay, F.B. 10578 H.M. Curran. June, 1908, sterile, (NY); Matnog, Vidal 2164, (M); Provo Catanduanes, F.B. 75105 M. Ramos & G. Edano, July-Sept. 1928, sterile, (NY), (UC).

SAMAR. B.S. 1612 M. Ramos, April, 1914, in flower & fruit, (A), (NY), (US), (UC); F.B. 21066 Sherfesse, Cenabre & Cortes, April, 1914, in flower, (US); Wright, Mt. Malingon. PNH M.D. Sulit, April­ May, 1949, in fruit, (PNH), (A).

LEYTE: Provo Leyte. F.B. 12738 R. Rosenbluth, March, 1909, in flower, IUS); C.A. Wenzel 1706, April 1, 1916, in flower, (A), (GH), (F); C. A. Wenzel 1799, May 1, 1916, in fruit, (A), (GH), (F); F.B. 28411 J.,G. Pacis, April, 1921, in fruit, (A); Baba­ tugon, F.B. 27213 P. Tomeldan, June, 1918, in fruit, (A), (US); Biliran Island, B.S. 18666 R.C. McGregor, June, 1914, in fruit, (PNH), (A), (US).

NEGl )5: Provo Negros, F.B. 1153,22667 H.M. Curran, April, 1912, in fru t, (US); Occidental Negros, F.B. 30180 Masias, Sareno A REVISION OF THE GENUS HOPEA ROXBURGH OF P. J. fi9

& Torrible, May-Sept. 1925, sterile (UC); Faraon F.B. 13757 F. W. Foxworthy, Sept. 1909, sterile, (NY).

MINDANAO: Provo Surigao, Mt. Kabatuan, PNH 10314 D. Men­ doza & P. Convocar, March, 1914, in flower, (PNH), (A); Provo Agu­ san, Butuan, San Mateo, PNH 41886, 41890 Demetrio R. Mendoza, June 12, 1961, in fruit, (PNH); Provo Zamboanga, F.B.13280 De Mesa & Villamil, May-June, 1912, in flower, (US); Provo Lanao, F.B. 24049 R. A. Acuna, May 1915, in fruit, (A), (US); Kulasihan, F.B. 31193 H. Orilloza, April-May, 1930, in flower, (NY); Basilan Is­ land, Isabela, Malique, Jose Vera Santos 4324, May 25, 1948, in fruit, (US); Manluso, Canas, PNH 97148 Gutierrez, Reynaldo & Fraga, May 15, 1965, in fruit (PNH).

LOCAL NAMES: bagitarim (Tag.), baguatsan (Bik), bagupsan (Bik.), bantaya (S.L. Bis.), gisok (P. Bis., Mbo., Lan., C. Bis.), gisok-gisok (P. Bis., SuL, S.L. Bis.), gisok nga-salngan (S.L. Bis.), kulilisiau (Tag.), makatayring I Tag. ), makitarem (Tag.), malatam­ ban (Bik.), malibato (Mba.), manglaum (Tag.). rnalaLamba (Bik. I. maLatagum (Bik.), pagakson (Bik.), paina 1 Tag. I, pongo I S.L. Bis. I, subigan (S.L. Bis.l. gisok-gisok (Official Common Namel.

Hopea phHippitlE'nsis Dyer. though it may have occasional large leaves almost similar to Hopea mindanensis Fox ...... is most likely to be confused with Hopea basilanica Foxw in the cblong or ellip­ tic to oblong-elliptic shape and in the sIze of the leaves. However. the leaf-base of the former is distinctly inequilateral. while that of the latter is oblique. The glabrous. almost black. when dry. inflo­ rescences; the larger flowers and the enlarged. long. linear-oblan­ ceolate, dark brown. calvx lobes on the mature fruit of Hopea philippinensis Dyer. are ~'ery distinct from the densely pubescent. grayish inflorescences. from the small flowers and the short. oblan­ ceolate, yellow calyx lobes on the mature fruit of Hopeo basi/onica Fox\\,.

This species has the most number of representative specimens studied and examined. However. the author has not seen any of the original materials used by Dyer in describing the species. It is likely that the Type specimen is extant and deposited at the British Museum in England. The designation of the Type, therefore. shall be deferred until such time that ~he status of Dyer's onginal ma­ terials shall have been fully ascertained. According to Foxworthy, [19, p. 276], an lsotype I Luzon. Albay Prov .. H. Cuming 879) was ACTA MANILANA deposited in the Philippine National He!'barium. Unfortunately this specimen was destroyed in the Second Ylorld War. Dyer's original description is based on fruiting materials alone. But Foxworthy amended it and added the description of the flowers based on the collection made by Ramos & Edano, B.S. 31296, in Panay, Capiz Province, Jamindan, April 15, 1918. This specimen was also des­ troyed in the Philippine National Herbarium in 1945.

Hopea philippinensis Dyer belongs to the Manggachapui Group [39, p. 71]. The bole is straight, and may reach up to 8 meters, but the girth is rather small. It has a short but distinct buttress. The bark is rather smooth and dark-chocolate in color. The sapwood 131, p. 3381 is thick and white. It is not very distinct from the heartwood, which is light brown and becoming darker brown with age. The wood is fine to medium textured and straight grained. It is moderately hard to hard, and moderately heavy to heavy. A good grade of Hopea philippinensis Dyer which is denser and finer­ textured approaches the wood of Hopea basilanica Foxw. or Hopea mindanensis Foxw. in hardness and texture. The heartwood r42, p. 741 is hard and probably as durable as yakal I Hopea plagata (BIan­ co I Vid.J and could be substituted by it. It is locally used for posts and piling, but rarely sawn into lumber, owing to the small size of the tree.

VII - SUMMARY AND CONCLUSION

1. The result of this botanical investigation has added two new species to the genus Hopea of the Philippines.

2. The species of Hopea quisumbingiana and Hopea woodiana are described here for the first time as new to science.

3. The revalidation of the original Blanco's description of Ho­ pea plagata I Blanco I Vidal has been proven.

4. The interpretation of the works of other botanists on the genL:S Hopea of the Philippines has been based on the major prin­ ciples and practices specified in the International Code of Botanical Nomenclature [2, pp. 341-3871.

5. It is likely that the nine species, including two new species, we have found comprising the genus Hopea of the Philippines, may A REVISION OF THE GENUS HOPEA IWXHCflGH OF P. I 'l1 increase, as additional data are obtained on future collections from other areas in the Philippines not yet explored. On the other hand, this number may also decrease, depending upon our own in­ terpretation as to what constitutes a species, on the basis of additional data we might be able to accumulate.

ACKNOWLEDGMENT

Acknowledgment is given to the United States Educational Foundation in the Philippines, the Philippine Government and the Arnold Arboretum of Harvard University for giving me the oppor­ tunity to study, as Special Student, at the Harvard University, Cambridge, Massachusetts, under a Partial Fulbright-Hays Grant and a Fellowship Grant from the Arnold Arboretum.

To my Advisor-Professors, Dr. Richard Allen Howard, Director of the Arnold Arboretum, Dr. Carroll E. Woed, Jr., and Dr. Eduardo Quisumbing, former Director of the National Museum, Manila, who guided and taught me the modern approach, procedures and techni­ ques of research in plant taxonomy and systematics, I am greatly indebted.

My sincere thanks to the Director-Curators of the Arnold Ar­ boretum and the Gray Herbarium of Harvard University, the United States National Museum Herbarium Garden Herbarium, the Natural History Museum Herbarium at Berkeley for extending to me the use of the facilities of their institutions while studying the materials for my research. The same goes to the Institute of International Education for the kind assistance extended to me while in the United States.

To the Executive Secretary and Staff of the Foundation, U.S. Embassy, Manila, to Dr. Galo B. Ocampo, Director of the National Museum, to Mr. Demetrio R. Mendoza, Curator for Botany and Staff of the Philippine National Herbarium, to Mr. Ricardo C. Aguilar, Cu­ rator for Art and History, for the fine illustrations, to Mr. Eric Ca­ sino, for the description in Latin, Mr. Daniel W. Tantoco, Jr .. Libra­ rian, and to Mr. Rodolfo V. Mallari, Information Editor of the National Mus€um and also to all of those who hav~ helped me made this undertaking a success, my deepest appreciation and gratitude. 72 ACTA MANILANA BIBLIOGRAPHY

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EXPLANATION OF PLATES

PLATE I. H opea quisumbingiana Sp.

Fig. 1. Habit sketch showing a portion of a flowering branch. Xl Fig. 2. Leaf. X 2 Fig. 3. Petal showing the position of stamens. XIO Fig. 4. Ovary. X 14

PLATE II. Hopea joxwoTthyi Elmer

Fig. l. Habit sketch showing a portion of a branch. X 1 Fig. 2. Leaf. X 2 Fig. 3. Fruit. X 2

PLATE III. Hopea malibato Foxworthy

Fig. l. Habit sketch showing a portion of a branch. X 1 Fig. 2. Basal portion of a leaf. X 2 Fig. 3. Fruit. X 2

PLATE IV. Hopea woodiana sp. nov.

Fig. 1. Habit sketch showing a portion of a flowering branch. Xl Fig. 2. Leaf. X 2 Fig. 3. Petal showing the position of stamens. X 18 Fig. 4. Ovary. X 30 Fig. 5. Fruit. X 2

PLATE V. H opea acuminata Merrill

Fig. l. Habit sketch showing a portion of a branch. X 1 Fig. 2. Fruit. X 2 A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 77

PLATE VI. Hopea oasilanica Foxworthy

Fig. 1. Habit sketch showing a portion of a fruiting branch. X 1 Fig. 2. Fruit. X 2/3

PLATE VII. Hopea plagata (Blanco) Vidal

Fig. 1. Habit sketch showing a portion of a flowering branch. X 2/3 Fig. 2. Flower. X 27 Fig. 3. Petal showing abaxial side. X 53 Fig. 4. Petal showing the position of stamens. X 53 Fig. 5. Ovary. X 27 Fig. 6. Fruit. X 2/3

PLATE VIII. Hopea mindanensis Foxworthy

Fig. 1. Habit sketch showing a portion of a branch. X 1/3 Fig. 2. Fruit X 1

PLATE IX. H opea philippinensis Dyer

Fig. 1. Habit sketch showing a portion of a branch. X 1 Fig. 2. Fruit. X 1 78 ACTA MANILANA

PLATE I A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. '19

PLATE II 80 ACTA MANILANA

PLATE III A REVI510N OF THE GENUS HOPEA ROXBURGH OF P.l. 81

PLATE IV

A REVISION OF THE GENUS HOPEA ROXBURGH OF P. I. 83

PLATE VI " ACTA MANILANA

PLATE VII 2

PLATE: Vnl A C TA MANILANA 86

PLATE IX