JOURNAL or tue ARNOLD ARBORETUM HARVARD UNIVERSITY VOLUME 61 NUMBER 1 US ISSN 0004-2625 Journal of the Arnold Arboretum Published quarterly in January, April, July, and October by the Arnold Arboretum, Harvard University. Subscription price $25.00 per year. Subscriptions and remittances should be sent to Ms. E. B. Schmidt, Arnold Arboretum, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. Claims will not be accepted after six months from the date of issue. Volumes 1-51, reprinted, and some back numbers of volumes 52-56 are available from the Kraus Reprint Corporation, Route 100, Millwood, New York 10546, U.S.A EDITORIAL COMMITTEE S. A. Spongberg, Editor — E. B. Schmidt, Managing Editor P. S. Ashton K. S. Bawa P. F. Stevens C. E. Wood, Jr. Printed at Edwards Brothers, Inc., Ann Arbor, Michigan COVER: Th L + = bi + fal 1 : \ 4 1 { ol ibaa | nearly 160 times) of a Fijian Sten Calphtin eucocarpun A. C. Smith form the basis of this year’s stylized cover design. A row of trichomes iS ised in the device on the back cover, while a cluster of eves appears on the offprints. As in recent years, the designs, based on holotype material in the herbarium of the Arnold Arboretum, were drawn by Karen Stoutsenberger. — Calophyllum L. (Guttiferae) is a wide-ranging genus of tropical forest trees in both the New and Old Worlds, and some species are of considerable economic importance as sources of valuable timber. Its use on the cover of this volume of the Journal of the Arnold Arboretum is appropriate since the April and July numbers are devoted to P. F. Stevens’s revision of the Old World ES DESSCHIaHY eS of this genus.—S. A. S. Second-class postage paid at Ann Arbor, Michigan JOURNAL OF THE ARNOLD ARBORETUM VoLumE 61 JANuARY 1980 NuMBER | FRUITS AND SEEDS OF THE BRANDON LIGNITE, V. RUTACEAE Bruce H. TIFFNEY THe Branpon Licnite (lat. 43°50’ N., long. 73°03’ W.) is a small deposit of Early Oligocene brown coal located in the town of Forestdale, Vermont, 27 km. north of Rutland, Vermont, and 230 km. northwest of Boston, Massachusetts. This locality is of particular value both because of the care with which it has been studied, and because it is the only known plant- megafossil-bearing terrestrial deposit of Tertiary age north of New Jersey in eastern North America. Fossil fruits and seeds were first described from the locality by Hitchcock (1853) and by Lesquereux (1861; and in Hitchcock et al., 1861) and were subsequently redescribed by Perkins (1904a, 1904b, 1905, 1906a, 1906b). Barghoorn and his students initiated a modern evaluation of the deposit in the late 1940’s, leading to a series of papers on the wood, pollen, and some of the fruits (Spackman, 1949; Barghoorn & Spackman, 1949; Barghoorn, 1950; Traverse & Barghoorn, 1953; Traverse, 1955; Eyde & Barghoorn, 1963; ae Bartlett, & Barghoorn, 1969). The author is presently investigating the remaining fruits and seeds of the deposit (Tiffney & Barghoorn, 1976, 1979: Tiffney, 1977, 1979). GEOLOGY The lignite and its associated clays, sands, and gravels unconformably overlie the contact of the Lower Cambrian Cheshire Quartzite and Dunham Dolomite, and are in turn unconformably overlain by Quaternary drift (Burt, 1928, 1930; Cady, 1945; Spackman, 1949). Most of the fossiliferous sediment is subsurface, but an area of roughly 82 square meters was exposed at the northern end of the deposit in 1974. The fossiliferous sediment is divisible into two separate units: a fine- to coarse-grained lignite, rich in fruits, seeds, and wood; and a fine-grained, purple-brown silt containing widely scattered fruits, seeds, wood, flowers, broken leaves, and particulate organic matter. © President and Fellows of Harvard College, 1980. Journal of the Arnold Arboretum 61: 1-40. January, 1980. 2 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 These two fossiliferous units are associated with a complex of sediments that appear to be a remnant of an extensive river system that flowed through west-central Vermont in mid Tertiary time. The silt, which is rich in speed lous pollen, 1s interpreted to represent an open body of still water, perhap by vegetated banks and subject to a considerable influx of organic material. The stratigraphic relationship of the silt and lignite has been rendered indefinite by the deformation associated with the passage of the Quaternary ice sheet. However, the present-day proximity of the two sediments, together with the extreme similarity of their contained flora, suggests that the two were deposited within a short period of time. Thus, while the two sediments represent slightly different ecological circumstances, they are here treated as coeval, and their flora considered as a single un MATERIALS AND METHODS The Brandon fruits and seeds are preserved as compactions—original organic matter retaining three dimensions and full morphological and anatomical detail. Larger specimens were recovered by manually breaking the matrix and then picking the fossils off of the freshly exposed face. The resulting matrix fragments were subsequently disaggregated in a solution of either sodium carbonate or hydrogen peroxide and were passed through sieves to recover smaller organic fragments. This material was viewed under a dissecting microscope, and the smaller specimens of interest were removed with a small rush. Adherent mineral matter was removed from the specimens with a toothbrush, and they were treated with 52 percent hydrofluoric acid to remove mbedded mineral matter. All specimens were stored in a 1:1 solution of glycerine and SO percent ethyl alcohol. Photomicrographs were made by means of a Wild M-20 compound microscope with camera attachment; whole specimens were photographed in air by using a Leicaflex camera with extension tubes and a Zeiss Planar or Tessar lens. Scanning electron micrographs were made with a model 1000a AMR microscope. Comparative modern material was obtained from herbarium specimens held by the Arnold Arboretum (a) and Gray Herbarium (Gu) of Harvard University. Although many of these sheets had recently been annotated by specialists working in the Rutaceae, several others had only original label data. occasion- ally over 100 years old. This situation raises the problem of the serious, and often underestimated, potential for utilizing misidentified modern comparative material in the identification of angiosperm fossils. Although the paleobotanist can avoid specimens of particularly dubious provenance, he cannot hope to fulfill his paleontological goals if all modern identifications must also be checked. For this reason, the herbarium sheets examined have been annotated, indicating that they were used in a study of the Brandon flora. Thus, if any of the modern comparative specimens used are subsequently assigned to a different taxon by a neobotanist, it becomes that individual’s responsibility to notify the paleobotanical community of the changes, particu- larly if the specimen is of significance. In addition, fruits and seeds of all 1980] TIFFNEY, BRANDON LIGNITE, V 3 the modern comparative specimens, together with all appropriate label data, have been placed in the fruit and seed collection of the Harvard University Herbaria. All modern material was cleaned and prepared by boiling in 10 percent potassium hydroxide for five to fifteen minutes, then washed, scrubbed with a stiff toothbrush, and dried, thus partially simulating the fossil condition. FOSSIL RUTACEAE Fossil seeds of the Rutaceae are known from Lower Eocene through Holocene sediments and comprise over sixty species in eleven genera, including one extinct genus, Caxtonia (Chandler, 1961b). The other genera reported are Acronychia, Euodia, Fagara, Orixa, Phellodendron, Ruta, Ptelea, Spathe- lia, Toddalia, and Zanthoxylum. The form genus Rutaspermum primarily includes seeds assignable to Zanthoxylum, although some members are rutaceous seeds of uncertain generic affinity. The use of this form genus is discussed further in the description of the Zanthoxylum seeds from Brandon. No extensive summary of the living or fossil seeds of the group is available, although Kirchheimer (1957) discussed the then-known remains of Phelloden- dron, and Gallet (1913) has summarized the development and anatomical structure of the seeds of several genera within the family. Because of this dearth of information, the process of identifying the Brandon fossils com- menced with a survey of the modern fruits and seeds of the family. Approximately 151 species in 55 genera were examined, with emphasis on genera and species presently found in the New World and in the tropical and temperate areas of eastern Asia, although representatives from other areas were included. The number of species examined per genus will be presented in the systematic descriptions. The family Rutaceae is characterized by one, two, or up to a large number of anatropous seeds per carpel, the seeds often being marked by a distinctively elongate hilar scar along their ventral margin. At maturity the carpels can be variously united into berries (Citrus) or drupes (Phellodendron), or can develop into samaras (Prelea, Skimmia). They may also form papery to woody capsules (Euodia, Zanthoxylum). To summarize the seed structure of the family as a whole is not necessary or possible in the present context. In view of the range of variation in fruit and seed morphology of the group, and of the diversity of forms present in the Brandon Lignite flora, this paper will diverge from the pattern set in the earlier portions of the Brandon Lignite investigation (Tiffney, 1977, 1979; Tiffney & Barghoorn, 1976, 1979) and consider the morphology and fossil history of the seeds of each genus independently. In all cases the important morphological characters of the seeds of each genus are summarized in a table and in an illustration accompanying the description of that genus. SYSTEMATIC DESCRIPTIONS Rutaceae A. L. de Jussieu, Gen. Pl. 296. 1789. Lesquereux (1861; and in Hitchcock er al., 1861) described Drupa rhabdo- sperma (later illustrated in Perkins, 1904b, 1905, 1906b) without assigning 4 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 it to a family. This seed is here recognized as belonging to the genus Zanthoxylum. Traverse (1955) reported one species of pollen from the Brandon Lignite, which was provisionally assigned to the tribe Toddalieae, fruiting remains of which are represented at Brandon by seeds of the genus Phellodendron. Euodia J. R. & G. Forster, Char. Gen. Pl. 13. 1776. The genus Euodia consists of some forty-five (Li, 1963; Willis, 1973) to over one hundred (Mai, 1970a) species of trees and shrubs of temperate to tropical forests in the Old World. These range from northern China south to Australia and east to eastern Africa. The majority of these species are evergreen, although many from northern and central China are deciduous (Wang, 1961). Chinese species of Euodia are particularly concentrated in the mixed mesophytic forests of the Yangtze Valley (Wang, 1961). The fruit is a sometimes rather thinly woody, loculicidal capsule, usually four to five lobed (occasionally unilobed), each lobe separating into two valves to reveal one or two seeds loosely enclosed in a bipartite, parchmentlike endocarp. This endocarp may provide a mechanical means of seed dispersal (see section on ecology). In many specimens, the shiny to iridescent seeds are attached to the open carpel or dangle from it on a short funicle. The present investigation is based on a survey of the seeds of 26 species of Euodia, 13 from China, India, and Japan, and 13 from Malaysia, Indonesia, and the Philippines. The recognition of this geographic separation is suggested by a dichotomy in the morphology of the seeds from the two areas. The seeds of the southern species are rugose and conspicuously irregular in shape, while those of the Indian, Japanese, and northern and central Chinese species are smooth to faintly reticulate and vary from nearly spherical to ellipsoid. These characters are all obtained following the removal of the smooth, shiny outer coat that is common to all seeds of the genus. Since the seeds of Euodia from the Brandon locality are comparable to those of the Chinese and Japanese species of the genus, the following discussion of the modern seeds will be limited to this northerly group of species. The seeds of the northern species of Euodia (see Ficure 2 for a generalized example) are small (2.2-5.3 mm. long by 2.0-3.9 mm. in diameter) and vary from nearly spheroidal to ellipsoid. In those species bearing two seeds per carpel the seeds are superimposed, the points of mutual contact appearing as flattened faces on the mature seeds (see PLate |, F). The shiny to iridescent, black or dark brown seed surface is broken only by a light tan ventral hilar scar. This shiny outer surface is formed by a thin, crustaceous layer that is easily removed to reveal the inner layer of the outer integument, thus simulating the fossil condition. The surface of this second, hard layer may be smooth (Euodia daniellii Hemsley ex Forbes & Hemsley, E. hupehensis Dode) or may be marked by gentle longitudinal ridges that are systematically broken by fainter transverse ridges to form a weak but distinct reticulum (E. bodinieri Dode, E. colorata Dunn, E. glauca Miq., E. hirsutifolia Hayata, E. meliifolia Bentham, E. officinalis Dode, and the fossil reported here: see Piate 1, A, D). The distinct hilar scar traverses the ventral face of 1980] TIFFNEY, BRANDON LIGNITE, V 5 the seed from the apex to the base; occasionally it develops a wide, distinctive lip or margin on either side of the hilum. The basal end of the hilum leads to a short raphal canal, which bends around to the large basal chalaza. The apical micropyle is small and inconspicuous. The apex in some species bears a more or less developed knob, which, according to Mai (1970a), results from the adherence of an abortive seed to the apex of the remaining seed. If this is correct, then it is always the upper of the two seeds that aborts in a biovular carpel. The anatomy of the seeds of Euodia was briefly summarized by Gallet (1913). The outer integument consists of a thin external unit and a sclerified internal unit. The outer two or three layers of cells are spongy and have a thicker external face that forms the shiny layer. It is this unit that is removed to reveal the inner layer of spirally thickened, radially elongate sclereids that form the sclerotesta. The inner integument is also bipartite, punctate or spirally thickened walls and an underlying layer, one or two cells thick, of large thin-walled cells. Fossil seeds of the genus have been reported on two previous occasions. Miki and Kokawa (1962) cited seeds of Euodia glauca from Recent deposits of Kyushu, Japan, and Mai (1970a, 1970b) has suggested that a previously unidentified seed of the Oligocene and Miocene of Europe is allied to the genus. This latter report is based on over one hundred specimens of consistent morphology from localities in Germany and western Siberia. Notzold (1963) identified this seed as Aldrovanda praevesiculosa Kirchheimer, which Mai (1970a) correctly argues on morphological grounds it cannot be. Nikitin (1965; and in Dorofeev, 1963) cited it as Carpolithus nitidus Nikitin and tentatively allied it with the Rutaceae. As described by Mai (1970a, 1970b), the seed in question is 1.5-1.8 mm. long and about |.2-1.3 mm. in diameter, pear shaped in ventral (hilar) view and elliptic in lateral view. Its shiny external surface is marked by a broad, light-colored hilum that passes from a terminal perforation identified as the micropyle, to a point approximately halfway down the ventral face of the seed. From the terminus of the hilum, an angular raphe passes over the remainder of the ventral side to the subbasal chalaza (see Ficure 1, A, a schematic drawing). Nikitin (1965) has interpreted the seed differently, placing the micropyle at the end of the hilum near the center of the ventral face of the seed, and the chalaza at the terminus now occupied by the perforation (see Ficure 1, B). The described and illustrated characters, particularly in view of the slight amount of variation in the abundant material (Mai, 1970a, 1970b), make it difficult to accept this as a seed of the genus Euodia. The pearlike shape end (sensu Mai), is distinct t absence of flattened surfaces on any of the fossils suggests that they were consistently borne singly in the carpel. The short hilum that extends about halfway down the ventral face of the fossil contrasts with those of the extant seeds, which run the entire length of the ventral face. The fossils are 6 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 micropyle chalaza raph: hilum igen raphe micropyle chalaza E 1. Two interpretations of Carpolithus nitidus: A, by Mai (1970a, 1970b): B. by Nikitin (1965). characterized by a shiny exterior surface similar to that of Euodia; however, in Euodia this coat is easily abraded away to reveal the dull sclerotesta. The fact that the shiny surface of the fossil is consistently whole and is formed by a layer of radially elongate sclereids (Notzold, 1963; Mai, 1970a) suggests that its structure is quite distinct from that of the membranaceous outer layer of Euodia. No modern species of Euodia exhibit the large ‘‘micropylar’’ perforation of the fossil, although its consistent presence and location eal that it is a true morphological character of the seed and not a chance degradational feature. Since these fossils are thus not really ley with the seeds of Euodia, perhaps it is best to return them to the form genus Carpolithus in the hope that their correct affinities might be elucidated in the future. This reevaluation of Carpolithus (Euodia) nitidus does not signal the demise of the genus from the European Tertiary, however, since seeds originally identified as Phellodendron costatum Chandler from the Eocene of England show a close resemblance to the seeds of modern species of Euodia. The seeds of P. costatum (Chandler, 1925-26, p. 28, pl. 4, figs. 6a-c; 196la, Dp, 125; -1961b,.p. 75, pl. 7, fies. 0; Jd: 1962, p. 73, pl. 10; fie... 19630, p. 92, pl. 14, figs. 24-28) are 3.0-3.5 mm. long, 2.0-2.25 mm. in diameter, and ellipsoid, often with a protruding knob on the micropylar terminus. The hilum and its broad margins extend the length of the ventral face of the seed (see particularly Chandler, 1963b, p/. 14, fig. 28), with the short raphal canal commencing at the basal end of the hilum and leading to the large basal chalaza. The micropyle is situated on the knob at the apical terminus of the hilum. The sclerotesta is marked by a series of strong longitudinal ribs connected by weaker transverse ridges, the whole forming a reticulum. The sclerotesta is approximately 300 pm. thick and is formed of radial rows of equiaxial cells 20-25 ym. in diameter (Chandler, 196la). While modern seeds of both Phellodendron and Euodia have an elongate hilum and a reticulate sclerotestal surface, those of Phellodendron are longer, have a narrower hilum without margins, and are flattened laterally from the mutual pressure of 1980] TIFFNEY, BRANDON LIGNITE, V 7 the five seeds in the drupe. Since the other reported fossil species of Phellodendron conform to these characters (Kirchheimer, 1957), it is best to consider P. costatum not as an aberrant Phellodendron, but as a distinct species of Euodia. Chandler (1925-26) assumed that the morphology of P. costatum was linked to that of the modern species through the Plocene P. elegans C. & E. M. Reid, but Kirchheimer (1957) and Tralau (1963) both expressed doubt as to the assignment of this seed to Phellodendron, and Kirchheimer (1957) suggested its possible affinity with the Toddalieae. However, the rounded shape suggesting derivation from a one-seeded carpel, the elongate hilum with wide margins, the apical knob, and the small size suggest placement of this seed in the genus Euodia. A formal reassignment of this taxon will be proposed and discussed in detail in a pending publication. Euodia lignita Tiffney, sp. nov. Piate 1, A-G. MatTERIALS. Six seeds have been recovered from the lignite at Brandon. The type specimen (PLate |, A, D) is assigned number 51378 of the Paleobotanical Collections of the Botanical Museum, Harvard University. The paratypes are assigned number 51379 in the same collection. Description. The seeds average 4.5 mm. long (range 3.4-4.9 mm.) and 2.8 mm. in diameter (range 2.7-2.9 mm.). With one exception, each is the product of a carpel containing one seed and is ellipsoid. The exception (PLate 1, F) is more rounded and has one face flattened from the pressure of a second seed in the mature carpel. One of the seeds bears a very distinct apical knob (Pate |, G), and two others display it to a lesser degree. All are marked by a hilar scar that extends from the apex to the base of the seed and is bordered by a wide margin (see Figure 2 and Piate |, A, E). At the base of the hilum a short raphe leads to the large, pitlike basal chalaza. The micropyle is at the apex of the seed, just beyond the terminus of the hilum. The dull black external surface of the sclerotesta is marked by many faint longitudinal ridges that are crossed at intervals by weak, short transverse ridges, the whole yielding a very faint reticulum of spaces ranging from 175 to 250 um. on a side. At higher resolution the surface is marked by a faint pattern of pits 20-30 jzm. in diameter in the outermost layer of sclerotestal cells. Two seeds exhibit only the latter pattern and show distinct signs of layer of the outer integument (sclerotesta) is 150-250 ym. thick and is formed of many layers of isodiametric sclereids. The inner integument is thin and hyaline, with an outermost layer of small, longitudinally elongate cells with spiral thickenings, underlain by two (three?) layers of larger isodiametric cells (PLate 1, B, ArrFinities. No single modern species is completely similar to the fossil, and only one that has been examined, Euodia colorata, is as large. Although E. colorata has a reticulate sclerotestal pattern, it lacks hilar margins, any suggestion of an apical knob, and any evidence that more than one seed was ever borne in a carpel. Both E. glauca and E. hirsutifolia, although 8 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Apex Apex Micropyle _— Apical Knob ———ao” g iy ote Bowe AV ates Par al te) Hilar Margin - Dorsal Face Ventral Face . ice eee ep te Meee testes Ventral View Lateral View Ficure 2. Descriptive characters of Euodia seeds. small, have wide hilar margins and a reticulate sclerotestal pattern. Euodia belonging to an extinct species. Two of the three modern species discussed (E. colorata and E. hirsutifolia) are members of the evergreen oak communities of China, while the third (£. glauca) is a member of the mixed mesophytic forests of central China (Wang, 1961). ‘*Phellodendron”’ costatum is smaller and has both a more pronounced sclerotestal reticulation and (in many cases) a more prominent apical knob than E£. lignita. Thus, while the two fossil species are similar in their wide hilar margins and their sclerotestal pitting, they definitely represent two distinct entities. The specific epithet /ignita commemorates the source of this fossil. Ecotocy. Although the members of the northerly Evodia group (as distinguished by their seeds) range from deciduous species of northern China (EF. daniellit) to evergreen taxa of the southern Chinese rainforests (E. meliifolia) (Map 1), the strongest concentration of extant species is in the upper Yangtze River valley, where ten occur in the intermixed deciduous and evergreen mesophytic forests of the hills and valleys (Wang, 1961). Both E. colorata and E. hirsutifolia are probably evergreen since they occur in the evergreen oak forests of Yunnan and the mountains of Taiwan, respectively. Euodia 1980] TIFFNEY, BRANDON LIGNITE, V 9 TaBLe 1. Descriptive characters of Euodia seeds. Size Hilum Length Percent length of ventral face Diameter . Hilar margin Shape Present / absent ae Micropylar knob Present / absent Number of seeds/carpel One Seed surface Smooth Predominantly low reticulation Predominantly low parallel ridges Two glauca is a canopy tree of the mixed mesophytic forests, found from 800 to 1400 meters elevation along the Yangtze River. Of interest is its association with species of Alangium, Ilex, Illicium, Magnolia, Nyssa, and Quercus (Lee, 1935; Wang, 1961), all of which are genera found at Brandon. The habitats of the modern species suggest that E. lignita was perhaps a denizen of a mixed mesophytic or evergreen broad-leaved sclerophyllous type of forest. Although possibly deciduous, it could equally well have been evergreen, particularly in light of the presence at Brandon of a species of Magnolia comparable to the extant evergreen M. grandiflora L. The small number of Euodia seeds in the deposit and their abraded condition (two more noticeably so than the others) suggest transport from a distant source. Ridley (1930) has suggested that the dispersal of Euodia seeds is similar to that of Dictamnus, where the seeds are mechanically tossed from the carpel by tensions created within the drying endocarp. The writer has not seen any evidence of mechanical dispersal in Euodia; indeed, the glistening, black seeds hanging free of the carpel on a short funicle seem perfectly adapted for avian dispersal. It is uncertain whether the abrasion of the fossils could have been produced by passage through a bird’s digestive tract, or whether it represents aquatic transport from a distant source. In view of the diversity of Euodia species in the hills of western China, it is tempting to conceive of E. lignita as having grown on the slopes of the Oligocene Green Mountains of Vermont, and then having been transported down to the Brandon deposit. The limited ecological data on the genus (Lee, 1935; Wang, 1961) suggest that it prefers drier, forested slopes, rather than moist riverbottom land, and thus favor a somewhat distant source for the seeds. It is unlikely that the genus was a dominant element in the forests surrounding Brandon in the Oligocene. Phellodendron Ruprecht, Bull. Acad. Imp. St.-Pétersb. 15: 353. 1857. This genus comprises between nine and thirteen species (Kirchheimer, 1957; Tralau, 1963; Willis, 1973) of small and large dioeceous trees. In the absence of a monographic study, the disposition of species is uncertain, and revision may result in a reduction of the total number (Tralau, 1963). Owhi (1965) 10 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 recognizes three of the species considered here (Phellodendron japonicum Maxim., P. lavallei Dode, and P. sachalinense Sarg.) as varieties of P. amurense Rupr. Phellodendron is restricted to the temperate and subtropical regions of eastern Asia. The species with the northernmost distribution is P. amurense, a tree of the coniferous and northern hardwood forests of Siberia, northeastern China, Korea, Japan, and Sakhalin. The southern distribution of the genus centers about the mixed mesophytic forests of the Yangtze Valley, where up to five deciduous species occur as low trees in the understory (Wang, 1961). The fruit of Phellodendron is a resinous, black, five-loculed drupe (Lee, 1935), although it may rarely develop four to six locules (Kirchheimer, 1957). Each locule is lined with a two-valved, thin, membranaceous endocarp that splits on a median plane to release the enclosed seed. The mature carpel usually bears only one seed, but two-seeded carpels are known, with the resulting seeds being small and highly deformed (Kirchheimer, 1957). Dorofeev (1970) noted the absence of herbarium specimens for comparative work, a seemingly not uncommon problem with the genus. The present study is based on seeds from six species (Phellodendron amurense, P. chinense Schneider, P. japonicum, P. lavallei, P. molle Nakai, P. cided. the total number available in the herbaria (a and Gu) at Harvard University. Seed morphology is rather consistent throughout the species of the genus. Excluding abnormal ones, the seeds range from approximately 4 to 6 mm. in length, 2.5 to 3.5 mm. in height (from the ventral to the dorsal margin), and 1.8 to 2.3 mm. in width (perpendicular to the dorsal-ventral plane). The overall shape is that of a laterally compressed hemisphere, the convex dorsal face bending at either end to meet the essentially linear, sharp, raphe-bearing ventral face formed by the junction of the two shallowly arched lateral faces (see Figure 3 and Prate 2, A-E). The flattened lateral faces are shaped by the mutual pressure of the five seeds within the drupe. Occasionally the dorsal face is marked by large, uneven depressions of unknown origin. These may be seen in the present fossil, Phellodendron sachalinense, and in some specimens of P. amurense. The linear hilum extends from one half to the whole length of the ventral face, depending on the species. The raphal canal begins at the basal end of the hilum and leads to the basal chalaza of the seed. The micropyle is situated at the apical end of the hilum, occasionally on a small protrusion or micropylar beak (Ficure 4). The two lateral faces are marked by a reticulate pattern of varying intensity and organization. In some species (e.g., Phelloden- dron lavallei) this consists of a faint pattern of small depressions, while in others more distinct longitudinal ridges parallel the dorsal margin of the seed and are crossed by transverse ridges to form a reticulum. This latter pattern may be strong (P. japonicum) or weak (P. amurense, P. chinense, P. molle, and P. sachalinense), depending on the species. Germination results and passing around to the opposite dorsal face. The outer integument is formed of two layers, an exterior layer of large, open cells, often abraded away in the fossils, underlain by a sclerotesta of many layers of isodiametric \ L's 1980] TIFFNEY, BRANDON LIGNITE, V rey g Fab t oe ra 1 oe ff [ Bs | / e Cs ee — Map 1. Present distribution of Euodia (after Engler (1896), Chiarugi (1933), Lee (1935), and herbarium specimens at a and Gu). 12 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Apex Apex Micropyle Hilum SSN Ventral Face Dorsal Face SONS US SR ys . bd ie kee 7 we eA . Pee as . ae LeeNe:, sata : vee , an eae : iS eee SWS rs See AN : S S oh es P : : bY ¥ Reh oe Se ‘ at 4 Raphal Entrance ss SS SS Raphe Chalaza Base Base Ventral View Lateral View Ficure 3. Descriptive characters of Phellodendron seeds. sclereids. The exterior of these two layers frequently appears to be striate, a pattern impressed upon it by the inner wall of the fibrous endocarp, likewise striate. The inner integument is formed of three cell layers, the central of which consists of flattened, tangentially elongate cells and is bounded by Apex Apex Micropyle Hilar margi n Hilum Raphal pore Dorsal face Raphal crest Chalaza Base Base Ventral View Lateral View Ficure 4. Descriptive characters of Zanthoxylum seeds. 1980] TIFFNEY, BRANDON LIGNITE, V 13 a layer of large, isodiametric, thin-walled cells on either side. The cell walls of these two enclosing layers have cellulosic spiral thickenings (Gallet, 1913). The foregoing suite of characters is presented in TaBLe 2 Unfortunately, even with the aforementioned characters, it is often difficult or impossible to distinguish between the seeds of the modern species, an observation already made by Tralau (1963) and Dorofeev (1970). This might support the view (Tralau, 1963; Owhi, 1965) that a revision of the genus would reduce the number of species. The seeds of Phellodendron japonicum seem to have a stronger sclerotestal pattern than do those of other species, and the seeds of P. chinense are marked by a distinctively angled face (as seen in lateral view), but the stability of these characters in a large sample is not established. Certainly the variation in the length of the hilum in three samples of P. amurense is sufficiently great as to exclude it as a specific character. Reid (1923) presented a table of characters of modern and fossil Phellodendron seeds, but many seeds of the same species examined in the present study do not conform to her descriptions. This suggests that the seed characters cited are not particularly constant. Although some fossil seeds have been determined to an extant species of the genus, the practice is not defensible. In approximately 20 reports, the fossil seeds of the genus have been placed in three extinct and two extant species. This total excludes ‘‘Phellodendron”’ costatum and P. europaeum Menzel (1913), a five-locular drupe that Kirch- heimer (1957) concluded could not be proven to belong to the genus. The remaining species can be divided into those displaying strong sclerotestal sculpturing and those showing the weaker sculpturing equivalent to that of today’s species. The former group includes three species that range from the mid-Oligocene to the Pliocene of Europe. Phellodendron lusaticum Kirchheimer, of the German Oligocene, is represented by a small, thick-walled, strongly sculptured seed first reported by Kirchheimer (1940). More recent fossils from the Miocene of Germany are of fragments (Mai, 1964), and in one case (Notzold, 1963) may be incorrectly assigned in view of their divergent morphology. Phelloden- dron elegans C. Reid (Reid & Reid, 1915), of the Miocene and Pliocene TasLe 2. Descriptive characters of Phellodendron seeds. Size Seed surface Length Smooth Width Rough Thickness Pattern type, if distinctive . Pattern scale Hilum Length Micropylar beak Percent length of ventral face Present /absent Dorsal face Ventral face Smooth or marked by small Notable breaks in straight line depressions f face 14 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 deposits of Europe and western Russia, is a larger seed with sculpturing intermediate between that of P. /usaticum and that of the extant species. Phellodendron ornatum E. M. Reid, of the Pliocene of France (Reid, 1923), is intermediate in size (4.5 mm. long) but very strongly sculptured. The remaining group includes seeds assigned to P. amurense (Miki, 1937, 1938; Szafer, 1946, 1954; Kolakovskii, 1958; Miki & Kokawa, 1962; Kokawa, 1966), P. japonicum (Szafer, 1946, 1954), and Phellodendron sp. (Dorofeev, 1963, 1970), which collectively range from the Miocene through the Recent. Although these assignments are not defensible in view of the variability of the modern seeds, they do correctly imply a greater resemblance to seeds of the present than to the more deeply sculptured and patterned ones of the past. The chronological transition from the older forms with stronger and more orderly sclerotestal patterns, through the increasing dominance of forms having less pronounced patterns, to the present seeds with faint and only marginally ordered patterns, suggests a directional trend. This could conceivably be paralleled by a trend from the thick sclerotestal walls of Phellodendron lusaticum (400 wm., Kirchheimer, 1940) to the thinner walls (average 200 um.) of the present-day species, a tendency not dissimilar to that previously noted for the Vitaceae (Tiffney & Barghoorn, 1976) and the Magnoliaceae (Tiffney, 1977). However, the true significance of these trends in seeds of Phellodendron can only be established in light of an understanding of the whole organism. Phellodendron novae-angliae Tiffney, sp. nov. Piates 1, H; 2, A-E. MarterIAL. Two seeds have been recovered from the lignite at Brandon. Each is broken but is held together by its internal contents. The type specimen (PLate 2, A, C, E) is assigned number 51380 of the Paleobotanical Osa Botanical Museum, Harvard University; the paratype is assigned number 51381 in the same collection. DescripTion. One seed is 5.3 mm. long, 2.9 mm. high (from the dorsal to the ventral margin), and 2.1 mm. wide (perpendicular to the dorsal-ventral plane); the other is 4.8 mm. long, 2.7 mm. high, and 2.0 mm. wide. Their shape is similar to that of the modern seeds: the two gently convex lateral faces join on one margin to form the elliptically curved dorsal surface, which bends upward at either end to join the straight, linelike ventral surface formed by the juncture of the opposite margins of the lateral faces. The narrow hilum extends from the apical end, three quarters of the way toward the base of the seed, where it terminates at the entrance to the raphe, which continues over the remaining portion of the ventral face and leads to the basal chalaza. The micropyle is situated on a beak that protrudes above the ventral margin at the extreme terminus of the hilum (PLate 2, A). A small depression occurs in the ventral margin just below this apical beak. The dorsal and dorsal-lateral surfaces of both seeds appear broadly scalloped (Pate 2, E) and bear approximately three large indentations of indefinite origin on each side. The external layer of sclerotestal wall cells provides a faint pitting to the vaguely vitreous surface of the fossil seed. On the 1980] TIFFNEY, BRANDON LIGNITE, V 1) lateral faces this pitting is dominated by a larger pattern of randomly organized weak ridges and depressions (PLATE 2, A). he outermost layer of the outer integument is missing. The internal portion of the outer integument ranges from 110 to 140 um. in thickness and is composed of eight to ten layers of isodiametric sclereids. The inner integument is rather badly distorted and its cellular arrangement obscured, but its outermost layer consists of irregular cells, 60-75 xm. long by 20-35 um. wide. These have spiral thickenings that are 1-2 ym. wide spaced at intervals of 6-8 wm. The remaining portion of the inner integument includes a thick mass of small (25-45 um. diameter) isodiametric cells with collapsed contents, which could be interpreted as albuminous cells. Arrinities. The laterally compressed shape and long, linear hilum, in conjunc- tion with the spirally thickened cells of the inner integument, mark this as a species of Phellodendron. The somewhat similar seeds of Euodia ae to a lesser degree, Zanthoxylum, can be excluded from compariso e basis of their round shape, which stems from an absence of oan pressure. Similarities can be found with seeds of many of the modern species of Phellodendron. The seeds of an unvouchered specimen of P. amurense in the seed collection at Harvard (a and Gu) are particularly similar to those of P. novae-angliae. No evolutionary link is to be inferred from this similarity, however, since other collections of P. amurense seeds differ distinctly from the fossil, as well as from each other. Thus, the identification can be pursued no further than to the generic level. Phellodendron novae-angliae shares the distinction with P. lusaticum of being the earliest reported seed of the genus, but is not similar to this, or any other, fossil seed. Phellodendron novae-angliae is primarily distinguished by its distinctively subdued sclerotestal pattern, the chronological appearance of which casts doubt on the possible evolutionary trend toward the reduction of the strength of sclerotestal sculpturing suggested earlier. Similarly, its sclerotestal thickness is far less than that expected in view of the aforementioned trend from thicker Paleogene forms toward S. The specific epithet novae-angliae commemorates the geographic source of this paleofloristically important fossil seed. EcoLtocy. The modern genus Phellodendron is restricted to temperate east Asia and is of deciduous habit. Although P. amurense is a northerly species of the Picea-Abies and northern hardwood forests, the majority of species are low trees of the mixed mesophytic forests of the Yangtze River valley (Wang, 1961) growing in conjunction with other species found at Brandon, including Euodia (Map 2). Since no one modern species is particularly similar to the fossil, the assumption that P. novae-angliae is a temperate form rests primarily upon the concentration of modern species in temperate forests, and on its association with other temperate forms, such as Magnolia, Illicium, and Euodia, found at Brandon. Perhaps the greatest significance of Phelloden- dron as an element of the Brandon flora is as a paleobotanical reinforcement of the classic eastern North America-eastern Asia distribution pattern of species, since the genus is restricted today to eastern Asia. 16 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Map 2. Present distribution of Phellodendron (after Engler (1896), Wang (1961), Tralau (1963), and herbarium specimens at a and Gu). No mention is made in the literature of how the seeds of Phellodendron are dispersed, although it is to be hoped that Starshova’s continuing study (1972, 1973) of the genus will ultimately answer the question. One might logically assume that a fleshy drupe would be dispersed by an animal, most likely a bird; however, the drupes of Phellodendron are particularly resinous, even after fifty years on a herbarium sheet. Since strong resins are normally considered repellent to animals, the seeds may be dispersed in another manner. Because only two seeds of Phellodendron were found in the deposit, their source may have been relatively distant. Zanthoxylum L. Sp. Pl. 1: 270. 1753; Gen. Pl. ed. 5. 130. 1754. Zanthoxylum is a large, pantropical genus with outlying species in the Temperate Zone of eastern Asia and North America (Brizicky, 1962a; Maps 1980] TIFFNEY, BRANDON LIGNITE, V 17 3, 4). Zanthoxylum, in the broad sense (including Fagara; Brizicky, 1962b),- consists of approximately 215 species. The genus is varied in habit and encompasses deciduous and evergreen trees and shrubs of both wet and dry habitats. The seeds of 79 species, including 26 species of eastern Asia and 53 of the New World, and representing the available fruiting material in the herbaria of Harvard University (A and Gu), were examined in the present study. The bivalved carpels occur in clusters, each carpel dehiscing along its dorsal margin to expose shiny seeds, which dangle from the carpel by a short funicle. The membranaceous to woody follicular valves are pitted with oil cells and lined by a free or adherent, cartilaginous, bipartite endocarp. Although two ovules are present in each locule, one normally aborts. Thus only one seed is usually found in the mature carpel. Modern seeds of Zanthoxylum range from 2.5 to 6.2 mm. (average 4.1 mm.) in their maximum dimension and have a variety of shapes, reflecting the nature of the hilar scar and raphe. The majority of the seeds are spherical or roundly ellipsoid, but a few are elongate-ellipsoid and some are laterally compressed hemispheres, with one straight (ventral) and one arched (dorsal) margin. This variation in shape often makes it difficult to define an obvious dorsal and ventral face as was done for the seeds of Euodia and Phellodendron. In many of the spherical Zanthoxylum seeds, ihe funicle attaches only at Map 3. Present New World distribution of Zanthoxylum (after Engler (1896) and herbarium specimens at a and GH). 18 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 one point; thus, the ventral face is more properly termed a ventral hemisphere. In the more elongate forms, the long axis of the ellipsoid is parallel to that of the follicle, with the micropyle at the apex and the chalaza at the base of the ellipsoid, thus isolating the ventral face as the one located between these two points and facing into the locule. If an elongate hilum passes along the entire length of this face, the extent of the ventral margin is made clear: if the hilum is shorter and occupies only a portion of the inwardly directed face (Ficure 4), the ventral margin becomes less obvious—in isolated seeds it is best defined by the position of the chalaza and micropyle. This interpretation differs from that of Chandler (1925-26, 1960, 196la, 1961b, 1962, 1963a, 1963b, 1964), who uses the point of funicle attachment (the hilum) to define the extent of the ventral face. In a few seeds (Z. limonella (Dennst.) Alston of the extant flora, and the fossil species Z. bognorense Chandler (Chandler, 196la)) the very long raphe and the hilum oriented perpendicular to the long axis of the seed combine to make Chandler’s interpretation more serviceable, but these cases are rare. The hilum can vary from a circular to an elongate-linear scar, but it 1S most commonly an elongate triangle with the point directed toward the apex of the seed, and with the flat end, bearing the raphal entrance, toward the base. In those cases where the hilum occupies a portion of the ventral face, it often follows the slope of the ellipsoid, thus falling at an angle relative to a line passing from the apex to the base of the seed. Although the presence or absence (in which case the hilum parallels the long axis of the seed) of this angle seems to be a specific character, the magnitude of the angle does not, since it varies widely on seeds of a single species. The area around the hilum is occasionally differentiated from the rest of the sclerotesta by its inflation (see Zanthoxylum rhabdospermum below, and Pvate 3, C) or by the absence of sculpture. The raphe commences at the base of the hilum and passes as a canal around to the hollowed-out basal chalaza. The path of the raphe is often traced externally by a ridge or raphal crest that may occasionally be quite prominent (Z. rhabdospermum, Pate 3, A) and that is frequently more faintly sculptured than is the rest of the seed surface. The micropyle is at the opposite end of the seed, at or just beyond the apical terminus of the hilum (see Ficure 4). The seed surface may be completely smooth (Zanthoxylum americanum Miller, Z. dissitum Hemsley ex Forbes & Hemsley, Z. scandens Blume) or may bear a gross pattern of varying intensity and organization. Many species have a rugose surface of small bumps that in some cases (Z. procerum onn. Sm., Z. piperitum DC., Z. pimpinelloides DC.) are organized into rows paralleling the dorsal margin. In a few species (e.g., Z. microcarpum Griseb.) the bumps coalesce into flanges and ridges that approach spines, while in others (Z. micranthum Hemsley, Z. schinifolium Sieb. & Zucc.) the ridges all join together and form a strong and distinctive reticulum. Occasionally there is a smaller version of this reticulum, which parallels the dorsal margin of the seeds in a series of lunate rows (Z. myriacanthum Wall., Z. panamense P. Wilson). In many species this large-magnitude sculpture is underlain by a finer pattern of very small pits that vary from 10 pm. 1980] TIFFNEY, BRANDON LIGNITE, V and herbarium specimens — Oo > Map 4. Present Old World distribution of Zanthoxylum (after Engler (1896), Hartley (1966) at a and Gu). 20 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 to approximately 70 pm. in diameter, depending on the species, and occur over the entire seed. They do not seem to be derived from the cells of the sclerotesta and may, as implied by Corner (1976), represent imprints of cells of the external layer of the outer integument upon the surface of the internal layer of the outer integument. The anatomy of these seeds is similar to that of the other rutaceous genera discussed. The outer integument is subdivided into an easily removed papery exterior layer and an underlying thick sclerotesta having the described sculpture. The exterior layer is thin and invariably shiny, while the inner sclerotesta 1s dull black and averages 300-400 ym. in thickness, ranging up to | mm. in extreme cases (Zanthoxylum dissitoides Huang, Z. nitidum DC.). Within a single seed the wall is often 100-200 um. thicker beneath the raphe mbranaceous layer in the mature seed (Gallet, 1913; see also Pare 4, The majority of the modern seeds seen are rounded or slightly elliptic and have a randomly organized, rough external sculpture. With the exception of certain particularly distinctive seeds (Zanthoxylum melanostictum Schlecht. & Cham., for example), it is often difficult to discern clear specific differences between seeds of separate species. In the species examined, smooth-walled forms occurred with equal frequency in both the Old and the New World (four species in each), but seeds with an organized sculptural pattern paralleling the dorsal margin were more common in the New World. Circular hila, representing the attachment of the funicle at a single point, are restricted to New World seeds in the sample examined, while seeds of the Old World species most commonly have hila that extend one-quarter to one-half the length of the ventral face. A summary of these characters is presented in TABLE 3. Twenty-three species of fossil seeds ostensibly related to Zanthoxylum (Chandler, 1925-26, 1957, 1960, 196la, 1961b, 1962, 1963a, 1963b, 1964; Palamarev, 1973; Mai, 1976) or Plio-Pleistocene in age (Miki, 1937, 1938; Miki & Kokawa, 1962; Kokawa, 1966), although a few Miocene species have been reported (Palamarev, 1968; Gregor, 1975, 1978a, 1978b). All of the Plio-Pleistocene reports are from Japan and are of modern species. Although Berry (1929) did cite an Eocene Zanthoxylum seed (as Fagara) from western Peru, the majority of the Paleogene reports are from Europe and include seeds of the form genus Rufaspermum. This latter genus was erected by Chandler (1957) for rutaceous seeds of uncertain affinity, although many of the seeds included are quite similar to those of Zanthoxylum. Chandler (196la), in transferring Zanthoxylum ornatum Chandler to Rutaspermum, stated that ‘‘it is now clear that no living representative of Zanthoxylum has such clearly defined regular superficial ornamentation.’’ However, Pala- marev (1973) transferred two species of Rutaspermum(R. bognorense Chandler and R. rugosum Chandler) to Zanthoxylum. Since seeds of Rutaspermum excavatum Chandler (Chandler, 1962, 1963b), R. glabrum Chandler (Chandler, 1980] TIFFNEY, BRANDON LIGNITE, V 21 Tas_e 3. Descriptive characters of Zanthoxylum seeds. Size Raphal crest Length Present /absent Diameter Prominent / faint Smooth /rough aeolian Percent length of seed Shape y Surface Spherical ae Smooth Ellipsoid Compressed-hemispheric mouer Random pattern Hilum Spines Shape Bumps Circular i Triangular Reticulation Slitlike Organized pattern Hilar angle in degrees Pattern strength Percent length of seed Pattern scale Hilar margin ’ Small pits Present / absent Present /absent Smooth/rough Di lameter Inflated 1962), R. magnificum Chandler (Chandler, 1962), and R. striatum Chandler (Chandler, 1962) are likewise all similar to those of Zanthoxylum, they too are perhaps best considered to be representatives of this modern genus. A formal proposal of this transfer must await examination of all the fossils involved. However, it would seem wise to avoid placing a specimen in a form genus when its characters, although not entirely comparable to those of any single modern species, conform closely to those of only one modern genus. Rutaspermum sp. (Chandler, 1960, p. 225) is rather similar to the previously reported R. ornatum (Chandler) Chandler (Chandler, 1925-26, 196la), while R. bognorense (Chandler, 1961b; Palamarev, 1973) may be conspecific with Zanthoxylum compressum Chandler (Chandler, 1925-26, 1961a) on the basis of morphological similarity. Of the remaining Rutaspermum seeds, R. minimum Chandler (Chandler, 1961b), Rutaspermum sp. (Chandler, 1960, p. 226) and Rutaspermum sp. (Chandler, 1964) are distinctly different from Zanthoxylum and are appropriately assigned to Rutaspermum in its function as a form genus. Mai(1976) proposed the transferral of seeds identified as R. ornatumto the extant genus Acronychia. The differing seed morphologies of these two forms argue against this proposal. These Paleogene seeds range from 2.0 mm. (Rutaspermum excavatum) to 8.0 mm. (Rutaspermum sp., Chandler, 1960, p. 226) in length, the latter fossil exceeding the size of the seeds of all of the modern species of Zanthoxylum examined, A few seeds show patches of the external papery layer of the outer integument, but most exhibit only the inner sclerotestal layer. In three cases (assuming R. bognorense and Z. compressum Chandler a2 JOURNAL OF THE ARNOLD ARBORETUM [voL. 6] to be conspecific) this is smooth, while in nine others the sclerotesta is marked by an organized coarse sculpture of ridges paralleling the dorsal margin of the seed. Excluding R. minimum (not included here in Zanthoxylum), all of the Paleogene seeds have an elongate-triangular hilar scar, which varies from 0° to 30° inclination relative to the micropyle-chalazal axis. The sclerotestal thickness of these fossils ranges from approximately 100 to 200 pm. up to 1000 um. (Zanthoxylum cf. planispinum Sieb. & Zucc.; Palamarev, 1968) but may vary within a single seed. The Miocene species described by Gregor (1978a, 1978b) include Z. ailanthiforme Gregor, Z. tiffneyi Gregor, and Z. giganteum Gregor. The last-named is particularly distinctive for its large size (5.0-8.0 mm. long), which approaches that of Z. hawaiiense (Hillebr.) Engler and Z. kauaiense A. Gray. The seeds all appear to be the products of one-seeded carpels. These various European Zanthoxylum and Zanthoxylum-like Rutaspermum seeds indicate the distinctive nature of the genus Zanthoxylum by Eocene time, while the contemporaneous existence of a Peruvian species suggests its widespread distribution. Although the earliest forms are comparable in size, general morphology, and sclerotestal thickness to today’s species, the sclerotestal patterning of the Eocene seeds tends to be stronger and more organized than that of modern ones, an observation also made by Chandler (196la, 1962, 1963b). The constancy of this distinction between Eocene and modern seed sculpture mirrors a decrease in the randomness of its organization through time. This may represent an evolutionary trend, in view of its predominance, although some modern seeds do have weak but regular sclerotestal patterns and a few Eocene seeds are smooth surfaced. Zanthoxylum rhabdospermum (Lesquereux) Tiffney, comb. nov. PLates 3, A-I; 4, A, B. Unnamed seed, E. Hitchcock, Am. Jour. Sci. II. 15: 100. fig. 19. 1853; Geol. Vermont 1: 231. fig. 150. 1861. Drupa rhabdosperma Lesquereux, Am. Jour. Sci. II. 32: 360. 1861, in Hitchcock et al., Geol. Vermont 2: 716. 1861; Perkins, Rep. Vermont State Geol. 4: 210. pl. 81, figs. 168-170. 1904, Bull. Geol. Soc. Am. 16: 514. pl. 87, fig. 25. 1905, Rep. Vermont State Geol. 5: pl. 53, fig. 25. 1906 MatTeRIAL. Approximately 370 seeds have been recovered from the lignite at Brandon. The type specimen described by Lesquereux is in the collection of the Division of Paleobotany of the U.S. National Museum and bears the number USNM 222845. Ten selected specimens in the Paleobotanical Collections of the Botanical Museum, Harvard University, have been assigned number 51382 and are stored there with the remainder of the material. DescripTIoN. The seeds average 4.9 mm. long (range 4.0 mm.—5.7 mm., standard deviation 0.64 mm.) and 3.3 mm. in diameter (range 2.9 mm.—4.4 mm., standard deviation 0.29 mm.). They range from ovoid to ellipsoid and have a rounded basal terminus and a somewhat pointed apical terminus, the latter formed by the intersection of the curving dorsal face and the flattened ventral hilar 1980] TIFFNEY, BRANDON LIGNITE, V 23 scar. In some cases this point is a ventrally directed beak or hook. Although the laterally compressed shape of one specimen (PLateE 3, E) indicates the presence of two seeds in one carpel, the remaining seeds lack flattened surfaces and are probably from single-seeded carpels. The elongate-triangular to rectangular hilar scar is inclined at an angle of 20-25° relative to the long axis of the seed; in side view it appears as an inclined plane cutting off the apical one third to one half of the ventral face of the seed (PLATE 3, B, H). This hilum averages 2 mm. in length (range 1.5 mm.-2.6 mm.) by 0.7 mm. in width (range 0.6 mm.—0.9 mm.) and is surrounded by a smooth, inflated margin. Occasionally the remains of a vascular strand are seen within this scar, leading to the raphal entrance at the basal end of the hilum. The path of the raphe is marked by a slight ridge or crest with reduced external sculpture, which passes from the base of the hilum over the remainder of the gently curved ventral face to the large basal chalaza (Plate 3, G). The micropyle is located at the extreme apex of the hilar scar. The vitreous black surface of the seed is marked by 30 to 32 regular longitudinal ridges that occasionally anastomose and that parallel the curved dorsal margin. These range from 60 to 120 ym. in width and are spaced 180-320 wm. apart. They are absent from the hilar margin and are considerably reduced on the raphal crest. Underlying this pattern is an all-pervasive one of small pits in the seed surface (PLate 3, D). These pits average 50 wm. in diameter (30 ym. on the hilar margin and portions of the raphal crest). Occasionally they are absent from the crests of some of the longitudinal ridges of the seed coat due to abrasion. In no case were any remains of the external layer of the outer integument seen. The thickness of the inner sclerotestal portion of the outer integument is 200-300 wm. (to a maximum of 400 um.) on the lateral and dorsal walls, ranging up to 600 pm. underlying the ventral raphal canal. The inner integument consists of three to five layers of rectangular cells ranging from 40 to 170 wm. in length and from 30 to 45 wm. in width. All the inner integumental cells have spiral thickenings 1-2 «wm. wide that are spaced every 3-4 pm along the cell wall (Plate 4, A, B). Arrinities. Although the elongate hilar scar, rounded shape, surficial sculpture, and spirally thickened cells of the inner integument clearly identify this as a member of the genus Zanthoxylum, no single modern species is comparable. As with the seeds described by Chandler, this is particularly true of the regular sclerotestal structure of the fossil, which is much more pronounced than that of any modern seed of the genus examined in this study. However, other characters of Z. rhabdospermum may be found in modern species, and although they do not occur in the same combinations as those of the fossil seed, they confirm the placement of the fossil in the genus Zanthoxylum. Of the modern seeds examined, the most similar were those from Central America and the Caribbean, including Zanthoxylum acuminatum caribaeum Lam., Z. pimpinelloides, and Z. pringlei S. Watson. All are reece ly smaller and more spherical than the fossil, but each has a few characters in common with Z. rhabdospermum. Zanthoxylum acuminatum has an inflated 24 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 hilar margin, Z. caribaeum a surficial pitting and sclerotestal sculpture similar to that of the fossil, Z. pimpinelloides an inflated hilar margin, and Z. pringlei a similar surficial pitting. Zanthoxylum rhabdospermum is most similar to previously reported fossil species in its strong and regular sculptural pattern, which is held in common with a number of English Eocene forms including Rutaspermum magnificum, R. striatum, and R. ornatum. Of these, R. striatum has the pattern most similar to that of the Brandon form, although more weakly displayed. In size, shape, hilar length and angle, fine sclerotestal pitting, and general appearance R. magnificum is the most similar to Z. rhabdospermum, but it lacks the inflated hilar margin and the parallel sclerotestal ridges of the Brandon species. Although R. ornatum is of the correct size and has a similar sclerotestal pattern, the hilum is too long, and while the hilar margin is free of sculpture, it is not inflated. With reference to previously reported fossils, perhaps the most significant factor is that Z. rhabdospermum conforms to the previously noted tendency for Paleogene Zanthoxylum seeds to have a stronger and more regular sclerotestal pattern than the modern seeds of the genus. Two other fossil seeds have been ascribed to Drupa rhabdosperma since Lesquereux’s publication of the species. That figured by Hartz (1909, pp. 18, 19, 275, pl. 2, fig. 5) as D. rhabdospermus exhibits a similar pattern of sclerotestal ridges, but the greatly elongate ‘‘beak’’ and the clear hilumlike area on one face are quite unlike the Brandon form. The seed illustrated exhibits little relation to Zanthoxylum and may not belong to the Rutaceae. On the other hand, the specimen of D. rhabdosperma described by Hofmann from the Eocene of Gaumnitz (1930, pp. 49, 50, pl. 5, figs. 28, 29) is definitely a seed of Zanthoxylum. Although these seeds are quite similar to those of Z. rhabdospermum from Brandon, particularly with respect to their size and to the strength of seed coat sculpturing, some distinct differences do separate the two. The German form has strong longitudinal ridges joined by short transverse walls; its long, narrow hilum extends for most of the length of the ventral face and is parallel to the long axis of the seed. This is in contrast to the Brandon form, which lacks conspicuous transverse sculptural elements and has a short hilum inclined at an angle of 20-25° relative to the long axis of the seed. For these reasons, Hofmann’s material cannot be considered conspecific with that of Z. rhabdospermum from Brandon. The characters of Hofmann’s specimens are quite similar to those of Rutaspermum ornatum, and Mai (1976) has united the two under Acronychia ornata (Chandler) Mai in his study of the Eocene Geiseltal flora. Ecotocy. The wide ecological range of the species of Zanthoxylum (Maps 3, 4), coupled with the lack of a single modern species comparable to the fossil, forestalis any worthwhile ecological conjectures. Even among the four most similar modern species, habitat preferences range from wet (Z. caribaeum, Little et al., 1974; Z. acuminatum, Fawcett & Rendle, 1920) to quite dry (Z. pringlei, Standley, 1923). The presence of a shiny, slightly fleshy, exterior layer around the seeds, and the manner in which they dangle from the carpel 1980] TIFFNEY, BRANDON LIGNITE, V ps) by a short funicle suggests that they are bird dispersed (van der Pijl, 1969). They have been observed to be eaten by a species of thrush in Java (Ridley, 1930) and occasionally by white-winged doves (Zenaisa asiatica) and American pipits (Anthus spinoletta) in the southern United States (Martin er al., 1961). Although some Zanthoxylum seeds may have arrived at Brandon through bird dispersal, the large number of seeds in the lignite indicates that the plants, either trees or shrubs, grew quite close to the site of deposition and thus presumably preferred wet ground. The slightly abraded nature of a few of the seeds suggests that they were transported to the site of deposition from a distant upstream source, and thus that Z. rhabdospermum was a widespread plant—at least along the river margins of the area—in Brandon time. Also of ecological note is the presence of a circular hole, 1.0-1.2 mm. in diameter, in the sclerotesta of two of the seeds examined (see PLaTE 3, I). These closely resemble insect holes observed in some modern species (Zanthoxylum foetidum Rose of Mexico, Z. limoncello Planchon & Oersted ex Triana & Planchon of Costa Rica, and Z. microcarpum of Mexico) and in one other fossil (Chandler, 1925-26). This similarity between modern and fossil holes suggests that perhaps an herbivore-host relationship was established in early Tertiary time and has since persisted. According to Prof. Horace Burke of Texas A & M University (pers. comm.), the holes in the modern seeds of the genus are most likely made by weevils when they mature and emerge from within the seed. The Rutaceae are well known for their distinctive plant chemistry, an example of which may be evident in these fossils. A single seed of Zanthoxylum rhabdospermum, removed from the glycerine-alcohol preservative mixture (to which phenol was finally added), washed, and placed in water, developed a mass of fungal hyphae in four days. This distinction is seemingly specific, since the seeds of other Brandon species were not similarly afflicted; it may provide the basis for future investigation. Zanthoxylum echinospermum Tiffney, sp. nov. PLaTeE 4, E-G. MATERIAL. One whole seed and several fragments were recovered from the silt at Brandon in 1949; the whole seed was carefully photographed but was subsequently broken. Of necessity, the type now consists of a photograph and a number of fragments, which are collectively assigned number 51384 of the Paleobotanical Collections of the Botanical Museum, Harvard Universi- ty. Description. The whole seed is 6.0 mm. long, 3.8 mm. high (from the dorsal to the ventral margin), and 3.4 mm. wide (from lateral face to lateral face). It is essentially ellipsoid (PLate 4, F, G), with the apical half of the ventral margin appearing flat in lateral view due to the hilum, which, in ventral view, protrudes to form an apical beak. The elongate- triangular hilum, 4.0 mm. long by 0.9 mm. wide, is inclined at an angle of 25° relative to the long axis of the seed and is surrounded by a smooth, inflated margin. The raphe commences at the widened base of the hilum and passes along the 26 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 remainder of the ventral margin to the basal chalaza. The external path of the raphe is marked by a low raphal crest, which bears a surficial pattern similar to that of the rest of the seed. The micropyle is located at the apical terminus of the hilum. The slightly vitreous black surface is traversed by rows of conical to faintly laminar spines that parallel the curved dorsal margin of the seed. These spines project approximately 230 um. from the surface of the seed and are about 200 ym. in diameter. The laminar forms are usually 200 pm. wide and up to 400 ym. long, with the long axis always parallel to the long axis of the seed. The spines may be from 130 to 250 um. apart. This pattern of spines is absent from the hilar margin and is slightly suppressed on the top of the raphal crest, but otherwise covers the seed. A pattern of finer pits, 35-50 jm. in diameter, covers the entire surface of the seed including the faces of the spines. This pattern may be derived in a manner similar to that of Zanthoxylum rhabdospermum. Presumably a crustaceous or fleshy external portion of the outer integument was present in life, but no evidence of it is found in the fossil. The patterned sclerotesta, assumedly the inner portion of the outer integument, ranges in thickness from 175 wm. on the lateral walls to 300 um. beneath the ventral raphe. The inner integument is composed of three or four layers of oblong cells ranging from 60 to 80 um. in length and from 40 to 60 um. in width. All of these inner integumental cells have anastomosing spiral thickenings on their walls, the bands being 1-2 ym. wide and occurring at intervals of up to 4 wm. (PLate 4, E). Arrinities. The rounded shape, indicative of development in a one-seeded carpel, the elongate-triangular hilum, the nature of the sclerotestal pattern, and the presence of spiral thickenings in the walls of the cells of the inner integument all conform with the seeds of Zanthoxylum. However, no modern species of Zanthoxylum seen has such simultaneously large and regularly placed spines, or combines a strongly patterned sclerotesta with such a large and distinctive hilum. Zanthoxylum melanostictum Schlecht. & Cham., of Central America, bears rows of small spines 300 ym. in diameter and 60-80 um. high; however, the size and shape of the seed and its hilum, the absence of a hilar margin, and the lack of surficial pitting clearly separate it from Z. echinospermum. Other modern species (e.g., Z. coco Engler, Z. monophyllum P. Wilson, and Z. tomentellum Hooker f.) have spines, but theirs are really elongate flanges and are not regularly spaced. From the present evidence it may be concluded that Zanthoxylum echinospermum represents a highly distinctive and extinct species of the genus; its characters do not match those of any one modern species but do fall well within the range of variation of eae of Zanthoxylum. the age reported Zanthoxylum fossil seeds, much the same eanrie obtains. Rutaspermum excavatum (Chandler, 1962, 1963b) has an lms ee pattern of protrusions intermediate between bumps and spines, but its small size, different shape, flat hilum, and lack of a hilar aren disqualify it from comparison with Z. echinospermum. Rutaspermum 1980] TIFFNEY, BRANDON LIGNITE, V rag, magnificum (Chandler, 1962) is similar in size and shape to Z. echinospermum, but lacks the spinose exterior and the inflated hilar margin. Although the overall configuration of R. striatum (Chandler, 1962) is also similar, this species may be dismissed for the same reasons; it is also too small. The common denominator between Z. echinospermum and the previously reported Paleogene forms is in the strength and regularity of their collective sclerotestal patterns. Although Z. echinospermum is represented by only a single specimen, itis rather unlikely that it is a morphological variant of the previously described Z. rhabdospermum. In addition to the distinctly different sclerotestal patterns, Z. echinospermum has a stronger pattern on the raphal crest and a less vitreous sclerotesta that is decidedly more brittle than that of Z. rhabdosper- mum. Additionally, the large sample of Z. rhabdospermum seeds has presented a fairly clear knowledge of the range of variation within that species. The specific epithet echinospermum is in recognition of the echinate nature of the testa of this form. Ecotocy. The restriction of this form to the Brandon silt, coupled with its limited representation, suggests that the parent plant was a minor component of the surrounding vegetation. Alternatively, in view of the avian dispersal of at least some modern species (as discussed under Zanthoxylum rhabdosper- mum), these seeds may have been carried from a distant source by birds. Zanthoxylum cf. tiffneyi Gregor, Acta Paleobot. 19: 33. 1978. PLATE 2, F-I. MarteRIAL. One whole seed and four fragments have been recovered from the lignite of the deposit. The entire seed is assigned number 51385 in the Paleobotanical Collections of the Botanical Museum, Harvard University. The fragments are collectively assigned number 51386 in the same collection. Description. The whole seed is 6.25 mm. long, 3.9 mm. high (from the dorsal to the ventral margin), and 3.7 mm. wide (from lateral face to lateral face). It is essentially ellipsoid, the dorsal portion being well rounded, while the ventral part is slightly compressed laterally to form a ridgelike ventral margin. The hilum is 4.4 mm. long; it lacks a true hilar margin but is surrounded by a raised lip. It follows a slightly curved path along most of the ventral face, widening from 0.5 mm. at the apical end to 1.0 mm. at the basal end (Pate 2, F). The short raphe passes from the hilar terminus around to the large basal chalaza and is marked by a very faint raphal crest. The vitreous black surface of the seed is completely covered with small depressions that range from isodiametric pits 40-50 pm. in diameter, to elongate pits 40 ym. wide by 70-90 um. long, which parallel the long axis of the seed. This pattern is overlain by a very faint reticulation formed by weak transverse and cross ridges; this occurs over the entire surface of the seed, including the raphal crest. No evidence of the external portion of the outer integument was seen. The hard inner portion of the outer integument (the sclerotesta) is 130-150 wm. thick, except beneath the raphe, where it may exceed 280 pm. The inner integument consists of two or three layers of isodiametric to rectangular cells that average 35 by 80 um. in size, all of which have spiral thickenings 28 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 61 on their walls. These thickenings, which range from | to 3 wm. in width and are separated by 4-10 ym; often anastomose and create a particularly spidery pattern (PLate 2, H). Arrinities. Although this seed is superficially similar to those of Euodia (notably with respect to the faintly reticulate sclerotesta and the length of the hilum), its large size, broad but marginless hilum, and, in particular, its inner integument of entirely spirally thickened cells, place it in the genus Zanthoxylum. However, no single modern seed has all the characteristics of the fossil. The seeds of many species of Zanthoxylum have a long, straplike hilum, but often in conjunction with a rough sclerotestal surface. Only Z. williamsii Standley, of Honduras, has a large, smooth seed with an elongate hilum comparable to that of the fossil, but it lacks the appropriate faint surficial pitting and reticulation and is more ovoid than ellipsoid. Among other modern species, a surficial pattern similar to that of the fossil can be found in Z. jamaicense P. Wilson (Jamaica) and Z. obtriangulare (Urban) Jiménez (Dominican Repub- lic), but neither has the fine pitting of the fossil form. Thus, although parallels can be drawn with a number of modern seeds, the fossil is distinct and has no modern counterparts among the species examined. The fossil is very similar in several respects to Zanthoxylum tiffneyi, described by Gregor (1978a, 1978b) on the basis of three fossils from Middle Miocene (floral zone 6) sediments near Wackersdorf, West Germany. The parallels include large size, thick sclerotestal walls, and, in particular, an elongate hilum and a smooth sclerotestal surface marked by faint pitting. Certain of the reported Paleogene forms (Rutaspermum bognorense (Chandler, 1961b); Z. compressum Chandler (Chandler, 1925-26, 1961a)—here considered probably conspecific with R. bognorense) do resemble the Brandon fossil in overall shape but deviate from it in significant respects (hilar shape, testal surface). While the Brandon form resembles Zanthoxylum tiffneyi more closely than any other known fossil, the limited amount of fossil material presently available, together with the absence of completely distinctive seed characters below the generic level in modern Zanthoxylum, renders the present identification somewhat tentative. In addition, certain fragments, which are assumed on the basis of sclerotestal characters to be of the same species, suggest that the morphology of the Brandon form varies beyond that seen in the single whole seed. For these reasons, the identification is cited as ‘‘conforming to” Z. tiffneyi pending the collection of further material. EcoLocy. The limited material is not sufficient to permit any ecological inferences. The rarity of this form may reflect its importation from a distant source, either by birds, or—less likely in view of its unabraded condition—by flowing water. DISCUSSION Wolfe (1975) has proposed that a rather homogeneous flora existed in the Northern Hemisphere in early Tertiary time. This view is based on 1980] TIFFNEY, BRANDON LIGNITE, V 29 considerable evidence, which includes the similarity of the Paleogene fruit and seed floras of Clarno, Oregon (Scott, 1954; Manchester, 1976), and the London Clay (Reid & Chandler, 1933; Chandler, 1961b, 1962, 1963b, 1964) of southern England. The presence of Euodia and Phellodendron in the Paleogene of eastern North America tends to support this hypothesis; both genera are known from the European Tertiary and are presently restricted to eastern Asia. On the other hand, Zanthoxylum does not fit this pattern quite as clearly. While it follows a boreotropical distribution in the Tertiary of Europe and eastern North America, the modern distribution of the genus is much larger and includes the Southern Hemisphere (Maps 3, 4). It is possible that the genus could have attained its present range by post-early Tertiary dispersal, particularly if it was primarily bird dispersed. However, the presence of an apparently correctly identified seed of Zanthoxylum (Fagara) from the Eocene of Peru (Berry, 1929) indicates that, at least in the New World, the genus had spread beyond the Northern Hemisphere boundary of the boreotropical forest by Eocene time aven and Axelrod (1974) consider the Rutaceae to have a pre-Tertiary, presumably Late Cretaceous, origin and to have been widely dispersed throughout both hemispheres by the early Tertiary. Since floral exchange between North and South America is assumed to have been minimal prior to the Early Miocene (Raven & Axelrod, 1974), the Paleogene Zanthoxylum species of Europe and North America were probably more closely related to each other than either was to the group of South American species represented by Z. piurianum Berry (Berry, 1929). This possibility is supported by the strong differences in morphology between Z. piurianum and the other known Paleogene rutaceous seeds, particularly with respect to the strong rectilinear pattern of the square pits on the testa of the Peruvian specimen. he postulated early Tertiary division of the range of Zanthoxylum would explain the dominance of strong, regular sclerotestal patterns among the Paleogene species of the Northern Hemisphere, since they would all presuma- bly belong to one lineage or a group of related lineages. This postulate does not explain why, of the modern forms examined, all of those most similar to the Paleogene Z. rhabdospermum are restricted to the New World. Based on the strong similarity of the European Tertiary flora to the extant flora of southeastern Asia, together with the close affinities of the North American and European Paleogene seeds of Zanthoxylum, it is logical to expect to find the modern relatives of the Brandon form in southeastern Asia. A complete resolution of the relationships and distributions of the various lineages of Zanthoxylum can result only from the collection of fossil material of the Rutaceae from the Southern Hemisphere. The taxonomic similarity of the Early Eocene London Clay flora and the Mid Eocene Clarno flora occasionally extends to the specific level. However, those known elements of the presumably Oligocene Brandon flora that are also common to contemporaneous European floras are similar only at the generic level (e.g., Phellodendron, Euodia, Illicium, Turpinia). This also holds in the cases of Magnolia waltonii Tiffney (Tiffney, 1977) and Zanthoxylum cf. tiffneyi (described here), which, although similar to previously described 30 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 European species, are not sufficiently so to warrant being considered con- specific. This taxonomic distinctiveness of the Brandon flora relative to contemporary European floras may be explained in one of two ways. The Brandon flora may represent the initial stages in the evolutionary divergence between the Old and New World elements of the boreotropical flora. Such a divergence would presumably be a direct result of the disappearance of the Early Eocene land bridge postulated to have linked eastern North America and Europe (Lehmann, 1973; McKenna, 1975), thus cutting off biotic exchange between the two areas. On the other hand, it is also possible that the boreotropical flora was not a completely homogeneous unit, but contained distinct geographic and ecological subunits. Indeed, the proposed geographic extent of the boreotropical flora (Wolfe, 1975) is such that one would expect some degree of local differentiation. In this light, the London Clay and Clarno floras might represent an ecologically specialized facies within the larger flora that occurred together with other facies restricted to different habitats. The western European floras of the late Paleogene could then be descendents of one such facies, while the Brandon flora could be derived from a different Of these two possibilities, the former is presently felt to be the more acceptable. The concept of the boreotropical forest provides an adequate explanation of the patterns seen in both the past and present flora and vegetation of the Northern Hemisphere. Additionally, it does so within a framework that acknowledges the individualistic nature of the geographic migration of plant species through time. e presence in the Brandon flora of several genera restricted in modern time to eastern Asia demonstrates that the classic floristic affinities between eastern Asia and eastern North America, which are established on modern distributions, become stronger with increasing age through the Tertiary. The question remains, however, whether both areas commenced in the early Tertiary with an entirely similar flora. Certainly the extant flora of eastern Asia 1s considerably more diverse than that of eastern North America, even if one includes related Central American floras in the latter catagory. Wolfe (1977) has suggested four historical factors that could account for this modern disparity in diversity, one of which is ‘‘isolation.’? Such isolation could be enforced from the west by the development of continental climates following Paleogene eastern North America. Since the Brandon flora is presumed to 1980] TIFFNEY, BRANDON LIGNITE, V 31 predate the Paleogene climatic deterioration (Wolfe, 1978), it might provide the best sample of this eastern North American boreotropical forest to date, but one that had undergone some 20 million years of evolution since its last exchange with the boreotropical forest at large. ACKNOWLEDGMENTS I wish to thank the owner of the Brandon Lignite site, Mr. Welland Horn, of Forestdale, Vermont, for his continuing friendship, interest, and support of the Brandon Lignite project; Professor E. S. Barghoorn, of the Botanical Museum, Harvard University, for his support, friendship, and guidance; the staff members of the Arnold and Gray herbaria of Harvard University for their assistance in many stages of this research, and particularly Dr. Shiu-Ying Hu, of the Arnold Arboretum, for her advice on Chinese botany; Dr. Richard H. Eyde, of the Smithsonian Institution, for his assistance in obtaining Russian literature; and Dr. Thomas G. Hartley, of the Herbarium Australiense, for advice on the distributions of living species of Rutaceae. The research was supported by grants from the Milton Fund of Harvard University, the Bache Fund of the National Academy of Sciences, and Sigma Xi. Publication is supported by National Science Foundation grant DEB79-05082. Scanning electron microscopy was conducted on a machine established under NSF grant BMS-741-2494. LITERATURE CITED Barcuoorn, E. S. 1950. The Brandon Lignite. Joint Bull. Vermont Bot. Bird Clubs 18: 21-36. & W. SpackMan, Jr. 1949. A eee study of the flora of the Brandon Lignite. Am. Jour. Sci. 247 Berry, E. W. 1929. Early Tertiary fruits me ae from Belén, Peru. Johns Hopkins Univ. Studies Geol. 10: 137-179. Brizicky, G. K. 1962a. The genera of Rutaceae in the southeastern United States. Jour. Arnold Arb. 43: 1-22. 2b. Taxonomic and nomenclatural notes on Zanthoxylum and Glycosmis (Rutaceae). Ibid. 80-93. Burt, F. A. 1928. The origin of the Bennington kaolins. Rep. Vermont State Geol. 16: 65-84. . 1930, The geology of the Vermont ocher deposits. /bid. 17: 107-136. Capy, W. M. 1945. Stratigraphy and structure of west-central Vermont. Geol. Soc. Am. Bull. 56: 515-588. CHANDLER, M. E. J. 1925-26. The Upper Eocene flora of Hordle, Hants. 2 pp. Palaeontological Society, London. 1957. The Oligocene flora of the mee Tracey lake basin, Devonshire. Bull Brit. Mus. (Nat. Hist.) Geol. 3: 123. 1960. Plant remains of the sae and Barton beds. I[bid. 4: 191- 238. 196la. Flora of the Lower Headon beds of Hampshire and the Isle of Wight. Ibid. 5: 93-157 32 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 . 1961b. The lower Tertiary floras of southern England. I. Palaeocene floras. London aes flora (suppl.). 354 pp. British Museum (Natural History), Londo ——. 1962. The eae Tertiary floras of southern England. II. Flora of the Pipe-Clay Series of Dorset (Lower Bagshot). 176 pp. British Museum (Natural History), London. ——— .. 1963a. Revision of the Oligocene floras of the Isle of Wight. Bull. ae Mus. (Nat. Hist.) Geol. 6: 323-383. 1963b. The lower Tertiary floras of southern England. III. Flora of the Bournemouth Beds; the Boscombe, and the Highcliff Sands. 169 pp. British Museum (Natural History), London 1964. The lower Tertiary floras of southern England. IV. A summary and survey of findings in the light of recent ee observations. 151 pp. British Museum (Natural History), Lond Cuiaruc!, A. 1933. Legni fossili della Somalia ia (Palaeontologia della Somalia; Fossili del Pliocene e del Pleistocene). Palaeontogr. Ital. 32(suppl. 167. Corner, E. J. H. 1976. The seeds of dicotyledons. Vol. I. 311 pp. Vol. II. 552 pp. Cambridge University Press, Cambridge. DororeEy, P. I. 1963. Tretichnye flory zapadnoi Sibiri. 346 pp. Izdat. Akademia .5.S.R., Botanicheskii Institut V. L. Komarov, Leningrad. (In Russian.) 1970. Tretichnye flory Urala. 66 pp. Izdat. Nauka, Botanicheskii rasta V.L. Komaroy, Leningrad. (In Russian. Enc_Ler, A. 1896. Uber die geographische Verbreitung der Rutaceen im Notas ures zu ihrer systematischen Gliederung. Abh. Akad. Wiss. Berlin 1896: Eype, R. a | E. S. Barcuoorn. 1963. Morphological and paleobotanical studies of the Nyssaceae, II. The fossil record. Jour. Arnold Arb. 44: 328-370. , A. Bartcett, & E. S. BarGHoorn. 1969. Fossil record of Alangium. Bull. Torrey Bot. Club 96: 288-314. Fawcett, W., A. B. Renpce. 1920. Flora of Jamaica. Vol. 4. Dicots. Leguminosae to Callitrichaceae. 369 pp. British Museum (Natural His- tory), London. Ga. et, F. 1913. Développement et structure anatomique du tégment séminale des Rutacées. 66 pp. These, Univ. Paris, Paris. Grecor, H.-J. 1975. Die mittelmiozane Mastixioideen-Flora aus dem Braun- kohlen-Tagebau Oder II bei Wackersdorf (Oberpfalz). 249 + xli pp. Unpubl. Inaugural-Dissertation, Ludwig-Maximilians Univ., Munchen. 78a. Die miozanen Frucht- und Samen-Floren der Oberpfalzer Braunkohle. I. Funde aus den cree Zwischenmitteln. Palaeonto- graphica (Abt. B, Palaophytol.) 167: 8-103. 1978b. Subtropische Elemente im europaischen Tertiar III. Rutaceae. Die Gattungen Toddalia und Zanthoxylum. Acta Palaeobot. 19: 21-40. Harter, T. G. 1966. A revision of the Malesian species of Zanthoxylum (Rutaceae). Jour. Arnold Arb. 47: 171-221. Hartz, N. 1909. Bidrag til Danmarks tertiaere og diluviale Flora. 292 pp. anmarks Geologiske Undersogelse I]. Raekke, Copenhagen. Hitrcucock, E. 1853. Description of a brown coal deposit in Brandon, Vermont, with an attempt to determine the age of the principal hematite ore beds in the United States. Am. Jour. Sci. Il. 15: 95-105. 1980] TIFFNEY, BRANDON LIGNITE, V 33 —., E. Hitcucock, Jr., A. D. Hacer, & C. H. Hitcucock. 1861. Report on the geology of Vermont: descriptive, theoretical, economical and scenographical. 988 pp. Claremont Manufacturing Co., Claremont, N. H. Hormann, E. 1930. Palaobotanische Untersuchungen von Braunkohlen aus dem Geiseltal und von Gaumnitz. Jahrb. Halleschen Verb. n. s. 9: 43-54. KirCHHEIMER, F. 1940. Ein neuer Beitrag zur Kenntnis der Frucht- und Samenfossilien aus den Braunkohlenschichten Sachsens und Thiringens. Bot. Arch. 41: 276-294, 5 . Die Laubgewachse der Braunkohlenzeit. 783 pp. Wilhelm Knapp, Halle (Saale). Koxawa, S. 1966. 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Die Mastixioideen-Floren im Tertiar der Oberlausitz. Palaontol. Abh. (Abt. B, Palaobot.) 2: 1-192. . 1970a. Subtropische Elemente im europaischen Tertiar I. Ibid. 3: 441- 503. 1970b. Neue Arten aus tertiaren Lorbeerwaldern in Mitteleuropa. Feddes Repert. 81: 347-370. 976. Fossile Fruchte und Samen aus an Mitteleozan des Geiseltales. Abh. Zentr. Geol. Inst. Palaontol. 26: 93-149. MancuesTer, 8. R. 1976. Description and Se of fossil woods, leaves and fruits from the Clarno Nut Beds, Wheeler County, Oregon. Oregon Mus. Sci. Industr. Tech. Rep. 4: ee 102. Martin, A. C., H. S. Zimm, & A. L. NeEtson. 1961. American wildlife and piants: a guide to wildlife food eats 500 pp. Dover Publ. Co., New York. or eee P. 1913. Beitrag zur Flora der Niederrheinischen Braunkohlenforma- Jahrb. Konigl. Preuss. Geol. Landesanst. Berlin 34: 1-98. Winer ‘S. 1937. Plant fossils from the Stegodon beds and the Elephas beds near Akashi. Jap. Jour. Bot. 8: 303-341. On the change of flora of Japan since the Upper Pliocene. Ibid. 9; 213 252. . 1941. On the change of flora in eastern Asia since Tertiary period (1). The clay or lignite beds flora in Japan with special reference to the Pinus trifolia beds in central Hondo. Jbid. 11: 237-303. . Koxawa. 1962. Late Cenozoic floras of Kyushu, Japan. Jour. Biol. Osaka City Univ. 13: 65-85. 34 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Nikitin, P. A. 1965. Akvitanskaya Semennaya Flora Lagernogo Sada (Tomsk). l p. Tomskogo Universiteta, Tomsk. (In Russian.) Norzo_p, T. 1963. Einige karpologische Pflanzenfossilien aus dem mitteldeut- schen Tertiar. Monatsber. Deutsch. Akad. ey 5: 439-445. Ounwi, J. 1965. Flora of Japan. (English ed.; F. G. Meyer & E. H. Wacker, eds.) 1067 pp. Smithsonian Institution, Fe ey bm © PALaMAREV, E. 1968. Karpologische Reste aus dem Miozan Nordbulgariens. Palaeontographica (Abt. B, Palaophytol.) 123: 200-212. 1973. Fosilnata flora na V’glenosniya Eotsen v Burgasko (Die eozane Flora des Burgas-Beckens). Izv. Bot. Inst. Sofia 24: 75-124. (In Bulgarian, erman summar Perkins, G. H. 1904a. On the lignite or brown coal of Brandon and its fossils. Rep. Vermont State Geol. 4: 153-162 . 1904b. eee of species call in the Tertiary lignite of Brandon, Vermont. Ibid. 12. 1905. are Tonite of Brandon, Vermont, and its fossils. Geol. Soc. Am. Bull. 16: 499-516. 1906a. The lignite or brown coal of Brandon. Rep. Vermont State Geol. 5: 188-194. . 1906b. Fossils of the lignite. Jbid. 202-230. Pu, L. van per. 1969. Principles of dispersal in higher plants. 154 pp. Springer Verlag, oe Raven, P. H., & D. I. Axerop. 1974. Angiosperm biogeography and past continental movements. Ann. Missouri Bot. Gard. 61: 539-673. Rew, C., & E. M. Rep. 1915. The Pliocene floras of the Dutch-Prussian border. 178 pp. Mededeelingen van de Rijksopsporing van Delfstoffen, The Hague. Rew, E. M. 1923. Nouvelles recherches sur les graines du Plieeene Infeérieur du Pont- a Gail (Cantal). Soc. Geol. France Bull. IV. 23: 308-355 & M. E. J. CHanpter. 1933. The London Clay flora. 561 pp. British Museum ol History), London Ripcey, H. N. 1930. The dispersal of plants throughout the world. 744 pp. L. Reeve & Compa ent. Scott, R. A. 1954. Fossil fruits and seeds from the Eocene a formation of Oregon. Palaeontographica (Abt. B, Palaophytol.) 96: 66-97. SpackKMAN, W., Jr. 1949. The flora of the Brandon Lignite: pee aspects and a comparison of the flora with its marae equivalents. 187 pp. Unpubl. h.D. dissertation, Harvard Univ., Cambr STANDLEY, P. C. 1923. Trees and shru bs of ene, (Oxalidaceae to Turnera- ceae). Contr. U. S. Natl. Herb. 23(3): 517-848. Starsuova, N. P. 1972. K antekologii Phellodendron amurense Rupr. v srednem olzh’e. I. Ekologiyatsveteniya. Bot. Zhur. 57: 1402-1412. (In Russian.) . 1973. K antekologii Phellodendron amurense Rupr. v srednem volzh’e. I. Ekologiya opyleniya. [bid. 58: 43-52. (In Russian.) Szarer, W. 1946. The Pliocene flora of Kroscienko in Poland, I-II. Polska Akad. Umiejetn. Rozpr. Wydz. Mat.-Przyr., DziaY B, Nauki Biol. 72: 1-162, 1-213. (In English and Polish.) ———. 1954. Pliocene flora from the vicinity of Czorsztyn (West Carpathians) and its relationship to the Pleistocene. Prace Inst. Geol. 11: 1-238. Tirrney, B. H. 1977. Fruits and seeds of the Brandon Lignite: Magnoliaceae. ot. Jour. Linn. Soc. 75: 299-323 1980] TIFFNEY, BRANDON LIGNITE, V 35 1979. Fruits and seeds a the Brandon Lignite III. Turpinia (Staphy- leaceae). Brittonia 31: 39- E. S. BarGHoorn. oe Fruits and seeds of the Brandon Lignite. I. Vitaceae. Rev. Palaeobot. Palynol. 22: 169-191. & . 1979. Flora of the Brandon Lignite. IV. Illiciaceae. Am. Jour. Bot. 66: 321-329. Tracau, H. 1963. Asiatic dicotyledonous aa in the Cainozoic flora of Europe. Sv. Vet.-akad. Handl. 9(3): | TRAVERSE, A. 1955. Pollen analysis of the Sas Lignite of Vermont. 107 pp. U.S. Dept. Interior Bur. Mines Rep. Invest. 5151. & E BarGcHoorn. 1953. Micropaleontology of the Brandon a an early Tertiary coal in central Vermont: preliminary note. Jour Paleontol. 27: 289-293. Wana, C.-W. 1961. The forests of China, with a survey of grassland and desert eee 313 pp. Maria Moors Cabot Foundation Publ. 5. Cambridge, Mass. Wits, J. C. 1973. A dictionary of the flowering plants and ferns. ed. 8. (Revised by H. K. Airy Suaw.) 1245 + Ixvi pp. Cambridge University Press, Cambridge, England. Wotre, J. A. 1975. Some aspects of plant geography of the northern hemisphere during the Late Cretaceous and Tertiary. Ann. Missouri Bot. Gard. 62: 264-279. 1977. Paleogene floras from the Gulf of Alaska region. U. S. Geol. Survey Prof. Paper 997: 1-108. . 1978. A paleobotanical interpretation of Tertiary climates in the northern hemisphere. Am. Sci. 66: 694-703. DEPARTMENT OF BIOLOGY New Haven, Connecticut 06520 EXPLANATION OF PLATES PLATE I Figures A-G. Seeds of Euodia lignita. A, #51378, SEM of ventral face, apex directed forward, x 12. B-G, #51379: B, SEM of cross section of sclerotestal wall (outer integument of small rounded cells at bottom of picture is succeeded in a vertical direction by one layer of spirally thickened cells of exterior coat of inner integument, followed by a layer of large, isodiametric parenchymatous cells of interior coat of inner integument), x 211; C, SEM of spirally thickened cells of exterior layer of inner integument, < 291; dorsal view, apex directed downward, x 8.5; E, ventral view, apex directed downward, x 8.5; F, ventral view of ae of two-seeded carpel (note flattened area to lower left), x 8.5; G, dorsal view (note apical knob on lower end of seed), x 8.5. Ficure H. Seed of Phellodendron novae- ws oe spirally thickened cells of outer portion of inner integument, x 339 36 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 PLATE I Figures A-E. Seeds of Phellodendron: A, P. novae-angliae (#51380), lateral view, ventral margin on right; B, P. amurense (C. S. Sargent, August 23, 1903 (a)), lateral view, ventral margin on right; C, P. novae- ae (#51380), ventral view, break in seed disrupting path of hilum; D, P. amurense (C S. Sargent, August 23, 1903 (a)), ventral view; E, P. novae-angliae (#51380), dorsal view (note scalloped nature of dorsal face); all x 10. Ficures F-I. Seeds of Zanthoxylum cf. tiffneyi: F, #51385, ventral view, x 6.5; G, #51385, dorsal view, x 6.5; H, #51386, spirally thickened cells of inner integument, x 166; I, #51385, lateral view, ventral margin (hilum) facing upward, x 6.5. PLATE III Figures A-I. Seeds of Zanthoxylum rhabdospermum (#51383): A, SEM of ventral face, apex to left (note central raphal crest), x 9.5; B, SEM of lateral face, apex and hilar area to right, x 9.5; C, SEM of apex, looking basally past hilum and inflated hilar margin, x 11; D, SEM of surficial pits of sclerotesta, Lindera cercidifolia Hemsley, Cheng & Hwa 1108. Lindera communis Hemsley, Hwa 34, 165, 386 (carpellate plant in flower; he isotype in all ora characters except that leaves are glabrous beneath), 5/5. Lindera communis Hemsley var. gee (Levl.) 8S. Y. ra comb. nov. ibe esquirolit Lévl. in Repert. Sp. Nov. 9: 459. Benzoin commune (Hemsley) Rehder i in Jour. oa ee 1: 144. 1919, pro ae Rehder in 1919 considered Lindera communis Hemsley and Litsea midrib, and primary lateral nerves. I have examined eight sheets of the syntypes of Lindera communis (Henry 1204, 1207, 1523, 2662, 3413, 3413B, 4551), all from western Hupeh. All these specimens have a glaucous lower leaf surface and are sparingly and very finely pubescent on the midrib and lateral nerves. The taxon with a rufous lower leaf surface and densely pubescent veins and twigs has a more southerly distribution than the typical variety. Additional specimens of var. esquirolii examined are Cheng & Hwa 743, 858, 1050, and Hwa 418, 516. Lindera fruticosa Hemsley, Cheng & Hwa 576, 636; Gressitt 2572. Lindera glauca Sieb. & Zucc. Cheng & Hwa 575; Gressitt 2487, 2549; Hwa , 100. Lindera hemsleyana (Diels) Allen, Hwa 307, 408, 414, 507. Lindera ichangensis Gamble, Djou 110; Gressitt 2571. Lindera megaphylla Hemsley, Cheng & Hwa 857; Hwa 103. Lindera stewardiana Allen, Cheng & Hwa 779. Lindera subcaudata (Merr.) Merr. Cheng & Hwa 619, 963; Hwa 35, 235 Lindera umbellata Thunb. Cheng & Hwa 952 (too immature to be identified with slag oe Lindera sp., Hw Litsea chunii chee. Hwa 246. Litsea cubeba (Lour.) Pers. Cheng & Hwa 797, 885, 1011; Djou 125; Hwa De 9; Litsea elongata (Nees) Hooker f. Cheng & Hwa 672, 895, 1047, 1106; Hwa 09, 266 Litsea glutinosa (Lour.) C. B. Robinson, Cheng & Hwa 815. Litsea pungens Hemsley, Hwa 259, 264 Litsea verticillata Hance, Hwa 37] (sterile; identification not certain). Machilus chinensis (Champ. ex Bentham) Hemsley, Hwa 141, 468. Machilus ichangensis Rehder & Wilson, Cheng & Hwa 886. Machilus lichuanensis Cheng ex Li, Cheng & Hwa 1017 (syntype); Hwa 476 (syntype; det. W. C. Che Neolitsea sieboldii (O. Kuntze) Nakai, Cheng & Hwa 668. Neolitsea viridis Cheng & S. Y. Hu, sp. nov Arbor sempervirens, ramulis glabris; foliis subcoriaceis, ellipticis, 4.5-7 cm. longis, 1.2-2.5 cm. latis, acuminatis, ad basin acutis, supra prominente 1980] HU, METASEQUOIA FLORA Vd ee glabris, viridis, subtus foveolatis, sub lente pilosis, penninervis, ervis 9- vel 10-jugatis, subalternatis; petiolis 10-14 mm. longis, glabris eee inflorescentiis fasciculatis, axillaribus, subsessilibus; floribus 4-lobatis, bees utrinque ferruginoso-hirsutis, lobis ovatis, ciliatis, 3 mm longis, 1.5 mm. latis, extus pubescentibus, intus glabris; staminibus 6, exsertis, ARS 4mm. longis, glabris, 2 raro | interioribus bi-glandulosis, glandulis breviter stipitatis, antheris magnis; ovariis neem 1-2 mm. longis, glabris; floribus femineis et fructibus ignot Hurex: Li-chuan District, Tuan-pao-hsiang, C. rT Hwa 125 (holo- type, A). This collection was distributed as a new species of Litsea. After having checked all the collections of the genera Litsea and Neolitsea in the herbarium of the Arnold Arboretum and having dissected the flowers, I decided to place it in Neolitsea. The species is closely related to N. wushanica (Chun) Merr., which differs in having glaucous lower leaf-surfaces, and ‘ ‘pedicellis sericeo-hirsutis.’” From N. pinninervis es & Huang, it differs in having glabrous filaments and rudimentary ovari Sassafras tzumu (Hemsley) Hemsley, ao & Hwa 764; Djou 105; Gressitt 2440; Hwa 208. Papaveraceae Macleaya cordata (Willd.) R. Br. Cheng & Hwa 1021. Saxifragaceae Astilbe chinensis Franchet & Sav. Gressitt ee sac ae febrifuga Lour. var. glabra S. Y. Hu, var. nov. typo differt in ramis glabris; foltis mena ak us, 5-11 cm. longis, 2-4 cm. latis, utrinque glabris; inflorescentiis, hypanthiis, calycibus, petalis et antheris omnino glabris. Cheng & Hwa 587 (holotype, a). Other specimens examined. Hupeu: En-shih, H. C. Chow 1881; Pa-tung, H. C. Chow 697 (a better specimen than the holotype; in flower), 1025. SzecHwan: Chung Hsien, W. P. Fang 492; Nan-chuan, W. P. Fang 1246, 5720 Dichroa febrifuga Lour. was described on the basis of Loureiro’s collection from the vicinity of Canton, now deposited in the British Museum. Many specimens from this area, as well as from many other provinces in the warmer region of ae were examined. A wide range of variation of trichomes on the leaves and inflorescences is constant. A distinctive species with strigose-villose indumentum, D. longipila Merr., was described from the mountains on the border of Kwangtung and Kwangsi. Here a glabrous variety of D. febrifuga is reported from the mountains on the border of Hupeh and Szechwan provinces Hydrangea chinensis Maxim. Cheng & Hwa 884. Hydrangea paniculata Sieb. Cheng & Hwa 1065. Hydrangea strigosa Rehder, Cheng & Hwa 812; Gressitt 2437, 2464. Hydrangea umbellata Rehder, Cheng & Hwa 1082. Parnassia wrightiana Wall. var. flavida Franchet, oe & Hwa 778. Philadelphus sericanthus Koehne, Cheng & Hwa 78 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Pittosporaceae Pittosporum adaphniphylloides Hu & Cheng, Hwa oe Pittosporum glabratum Lindley var. glabratum, Hwa 133. Pittosporum glabratum Lindley var. oo Rehder & Wilson, Hwa 340. Pittosporum truncatum Pritzel, Hwa 405 Hamamelidaceae Corylopsis henryi Hemsley, Hwa 245. Corylopsis platypetala Rehder & Wilson, Hwa 281, 283. Corylopsis sinensis Hemsley, Cheng & Hwa 1057; Hwa 49. Liquidambar formosana Hance, Cheng & Hwa 708; Gressitt 2415. Loropetalum chinense Oliver, Cheng & Hwa 1024. Sycopsis sinensis Oliver, Cheng & Hwa 747, 1179; Hwa 260, 271, 300, 542. Rosaceae Agrimonia pilosa Ledeb. Cheng & Hwa 658. Chaenomeles cathayensis Hemsley, Hwa 292, 469. Cotoneaster salicifolia Franchet, Cheng & Hwa 940; Hwa 123. Eriobotrya cavaleriei (Lévl.) Rehder, Hwa 146, 448, 588. Eriobotrya deflexa Hemsley, Hwa Eriobotrya japonica (Thunb.) Lindley, Cheng & Hwa 1159; Hwa 526. Kerria japonica DC. Cheng & Hwa 595; Hwa 251. Malus halliana Koehne, Cheng & Hwa 1025; Hwa 279. Malus prunifolia (Willd.) Borkhausen, Hwa 401. Photinia amphidox (Schneider) Rehder & Wilson, Cheng & Hwa 1151; Hwa 9. Photinia beauverdiana Schneider, Cheng & Hwa 999; Hwa 94, 110, 114, 306, 482. Photinia davidsoniae Rehder & Wilson, Cheng & Hwa 1118; Hwa 106. Photinia franchetiana Diels, Cheng & Hwa 1001; Hwa 409, 523. Photinia parvifolia Schneider, Cheng & Hwa 563, 831, 1094; Hwa 470. Photinia sp. Cheng & Hwa 608, 799. Prunus conradinae Koehne, Djou 126; Hwa 297, 358. Prunus glyptocarpa Koehne, Hwa 60, 247, 263, 331, 349, 552. Prunus grayana Maxim. Cheng & Hwa 703; Gressitt 2552; Hwa 285, 347, aac, Prunus microbotrys Koehne, Cheng & Hwa 669, 737, 1149; Hwa 170, 338. m Nepal extending eastward via China to Japan, there exists a group of species of evergreen Prunus characterized by subcoriaceous leaves that are eglandular beneath and by simple axillary racemose inflorescences. The names proposed for the species are Prunus wallichii Steudel (1841) for the Nepalese species, P. spinulosa Sieb. & Zucc. (1843) for the Japanese species, and P. microbotrys Koehne (1911) for the species from Central China. Kalkman (1965) renamed some of the Chinese specimens P. wallichii and others P. spinulosa (including P. microbotrys). However, it is thought that there are three species in this complex. Typical P. wallichii has large, thin leaves 12-13 cm. long and 5-7 cm. wide. The racemes are usually 1980] HU, METASEQUOIA FLORA 719 fasciculate, and the rachis and pedicels are glabrous. The basal portion of the ovary is pilose. Typical P. spinulosa from Japan has leaves that racemes are always solitary, and the rachis and pedicels are strigose. The ovary is glabrous. Typical P. microbotrys from Central China has oblong or ovate-oblong leaves 6-9 cm. long and 2.5-3.5 cm. wide. The racemes are solitary, and the rachis and pedicels are pilose. The ovary is hairy at the base. = acl from Central and West China are variable, especially morphological cee that do not exist in the specimens from Japan, Nepal, and adjacent northern India. The name P. microbotrys, originally published for the taxon of Central ae West China, a be maintained. Prunus microbotrys Koehne var. obovat OV Arbor, ramulis glabris; foltis anes oblongo- ieee 10-11 cm. longis, nee 5 cm. latis, basi acutis, apice caudato-acuminatis, acuminibus 1.2-1.4 cm. longis; racemis axillaribus, solitariis, 4-7 cm. longis, rachibus et pedicellis pilosis, calycibus glabris, petalis 2 mm. diametro Ch Hwa 735 (holotype, a). Prunus mume (Sieb.) Sieb. & Zucc. Gressitt 2524 (sterile). Prunus padus L. Gressitt 5248; Hwa Prunus persica (L.) Batsch, Djou 135. Prunus pseudocerasus Lindley, Gressitt 2534. Prunus salicina Lindley, Djou 109; Gressitt 2439; Hwa 256. Prunus szechuanica Batalin, Hwa 248, 343. Prunus venosa Koehne, Cheng & Hwa 816; Hwa 7. Prunus wilsonii (Diels) Koehne, Gressitt 2570; Hwa 369. Prunus wilsonii (Diels) Koehne var. leiobotrys Koehne, Cheng & Hwa 592. Pyracantha crenato-serrata (Hance) Rehder, Cheng & Hwa 977; Djou 127; Gressitt 2426. Pyrus calleryana Decaisne, Djou 106. Pyrus pyrifolia (Burman f.) Nakai, Hwa 319. Rosa helenae Rehder & Wilson, Djou 116 Rosa roxburghii Tratt. Cheng & Hwa 956. Rubus adenophorus Rolfe, Cheng & Hwa 637, 742. Rubus ampelinus Focke, Cheng & Hwa 702. Rubus cavaleriei Lévl. Cheng & Hwa 826; Gressitt 2432, 2519 oy nes allied to R. setchuensis Bur. & Franchet, but with leaf apex trun Rubus ichangensis Hemsley & Kuntze, Cheng & Hwa 905. Rubus malifolius Focke, Ge 2459. Rubus pectinellus Maxim. Cheng & Hwa 591. Sorbus aronioides Rehder, Cheng & Hwa 901, 1101. Sorbus caloneura (Stapf) Rehder, Cheng & Hwa 671; Hwa 52, 311. Sorbus folgneri (Schneider) Rehder, Cheng & Hwa 674, 1103; Hwa 58, 501, I53: Sorbus wilsoniana Schneider, Cheng & Hwa 899, 1070. Spiraea chinensis Maxim. Cheng & Hwa Spiraea japonica L. f. var. fortunei Rehder, Gressitt 2436. Spiraea prunifolia Sieb. & Zucc. Hwa Stephanandra chinensis Hance, Cheng & Hwa 878. Stranvaesia davidiana Decaisne var. undulata (Decaisne) Rehder & Wilson, heng & Hwa 900, 1034; Hwa 159, 325, 380. 80 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Leguminosae Albizia kalkora Prain, Cheng & Hwa 1054; Gressitt 2472; Hwa 37. Amphicarpaea edgeworthii Bentham, Cheng & Hwa 832 (det. C. E. Wood, Jr.), 1144. Cercis chinensis Bunge, Hwa 69 (sterile). Cladrastis sinensis Hemsley, Gressitt 2461; Hwa 417. Dalbergia dyeriana Prain, Cheng & Hwa §&22. Dalbergia mimosoides Franchet, Cheng & Hwa 804 (det. Y. T. Lee). Desmodium podocarpum DC. Cheng & Hwa 626, 820; Gressitt 2497. Gleditsia macracantha Desf. Gressitt 2443; Hwa 45, 477. cans chinensis Baillon, Cheng & Hwa 1038; Hwa 478 (det. Y. T. ee). ey ae amblyantha Craib, Gressitt 2514. Indigofera pseudotinctoria Matsumura, Cheng & Hwa 1116 ae ¥. T, Lee), Lespedeza cuneata G. Don, Cheng & Hwa 659, 727; Djou 12 Lespedeza thunbergii (DC.) Nakai, Cheng & Hwa 895; Ae ee Bg Maackia chinensis Takeda, Cheng & Hwa 1146. Ormosia puberula Cheng, Hwa 485 (isotype; det. W. C. Cheng). Pueraria lobata (Willd.) Ohwi, Cheng & Hwa 731; Gressitt 2562. Robinia pseudo-acacia L. Djou 139 (sterile; introduced). Oxalidaceae Oxalis griffithii Edgew. & Hooker f. Cheng & Hwa 787 (sterile; Dr. A. Lourteig consulted). Rutaceae Boenninghausenia pa ale amas Meissner, Cheng & Hwa 870. Euodia fargesii Dode, Euodia meliaefolia ener vie a7, Euodia officinalis Dode, Cheng & Hwa 706, 1013; Gressitt 2421, 2442, 2475, 2540; Hwa 83. Orixa japonica Thunb. Hwa 346. Phellodendron chinense Schneider, Gressitt 2451; Hwa 13. Toddalia asiatica (Lam.) Kurz, Cheng & Hwa 927, 942; Hwa 423. Zanthoxylum dissitum Hemsley, Cheng & Hwa 601. Zanthoxylum nitidum (Roxb.) DC. Hwa 337 Zanthoxylum simulans Hance, Hwa 152. Simaroubaceae Ailanthus altissima (Miller) Swingle, Cheng & Hwa 720, 818, 1156; Djou 133; Gressitt 2554. Meliaceae Toona sinensis (A. Juss.) Roemer, Cheng & Hwa 1152; Hwa 151. Polygalaceae Polygala wattersii Hance, Hwa 5, 416. Euphorbiaceae Aleurites fordii Hemsley, Djou 136. Croton tiglium L. Cheng & Hwa 1182. 1980] HU, METASEQUOIA FLORA 81 Glochidion pubescens (L.) Hutch. Cheng & Hwa 687; Djou 115, 124; Gressitt 2538 Phyllanthus flexuosus (Sieb. & Zucc.) Mueller-Arg. Cheng & Hwa 648 (det. G. L. Webster). Ricinus communis L. Cheng & Hwa 1183. Sapium sebiferum (L.) Roxb. Cheng & Hwa 1181. Daphniphyllaceae Daphniphyllum glaucescens Bl. Cheng & Hwa 1030; Hwa 454. Daphniphyllum longistylum Chien, Cheng & Hwa 660, 1072; Hwa 533, 534. Coriariaceae Coriaria sinica Maxim. Gressitt 2442, 2563; Hwa 238, 249, 250. Buxaceae Buxus microphylla Sieb. & Zucc. Cheng & Hwa 806. Anacardiaceae Choerospondias axillaris (Roxb.) Burtt & Hill, Cheng & Hwa 920; Hwa 447, 481, 498, 525, 555. Rhus chinensis Miller, Djou 123; Gressitt 2470, 2533. Rhus potaninii Maxim. Hwa 71. Rhus trichocarpa Miq. Cheng & Hwa 744, 1104. Rhus verniciflua Stokes, Cheng & Hwa 722; Gressitt 2473; Hwa 10, 149. Aquifoliaceae Ilex dasyphylla Merr. var. lichuanensis S. Y. Hu A typo differt in ramulis petiolisque sparse ane ‘foliis subglabris, costa supra elevata; fructibus 6 mm. diametro, sepalis persistentibus glabris. Cheng & Hwa 1055 (holotype, A). In shape, size, and texture, the leaves of this variety are similar to those of Ilex dasyphylla Metr. var. dasyphylla, a variety of the mountainous area of northern Kwangtung and eastern Kwangsi, characterized by dense, ferrugineous, long hairs on the branchlets, leaves, and inflorescences. The fruit of var. lichuanensis is small, 4-5 mm. in diameter, and the persistent sepals are hirsute. Ilex elmerrilliana 8. Y. Hu, Cheng & Hwa 767; Hwa 487. Ilex ficoidea Hemsley, Hwa 134 (sterile). Ilex intermedia Loes. Hwa 132, 518, 521. Ilex intermedia Loes. var. fangii (Rehder) S. Y. Hu, Cheng & Hwa 937. Ilex macrocarpa Oliver, Hwa 97, 409, 520. Ilex pernyi Franchet, Cheng & Hwa 829; Hwa 4. Ilex suaveolens (Lévl.) Loes. Cheng & Hwa 729, 1035; Hwa 41. Ilex wilsonii Loes. Cheng & Hwa 775, 1139; Hwa 541. Celastraceae Celastrus gemmatus Loes. Djou Celastrus hypoleucus (Oliver) ae Cheng & Hwa 1062. 82 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Euonymus acanthocarpa Franchet, Cheng & Hwa 1037. Euonymus acanthocarpa Franchet var. sutchuenensis Franchet ex Loes. Cheng & Hwa 1004, Euonymus alata Regel, Hwa 112, 524. ne hae Thunb. var. acuta Rehder, Cheng & Hwa 1121; Gressitt 2518; Hwa 102. Euonymus ene Hemsley, Cheng & Hwa 949. Euonymus sanguinea Loes. var. camptoneura Loes. Djou 122; Gressitt 2413; Hwa 82. Aceraceae Acer amplum Rehder, Cheng & Hwa 740; Hwa &, 360. Acer cordatum Pax, Hwa 428, 446, 540. Acer davidii Franchet, Cheng & Hwa 714, 902, 1069; Hwa 288, 345. Acer erianthum Schwerin, Cheng & Hwa 704; Gressitt 2565. Acer faberi Hance, Cheng & Hwa 991; Hwa 428, 489. Acer henryi Pax, Hwa 335 (a very es shoot), 472. Acer longipes Franchet ex Rehder, Hwa 55. Acer mono Maxim. Hwa 28. Acer oliverianum Pax, Cheng & Hwa 862. Acer sinense Pax, Cheng & Hwa 783; Hwa 139, 339. Acer wilsonii Rehder, Cheng & Hwa 569; Hwa 348, 465. Hippocastanaceae Aesculus wilsonii Rehder, Cheng & Hwa 1134; Hwa 96 (sterile). Sabiaceae Meliosma beaniana Rehder & Wilson, Hwa 59, 67, 131, 545 (59 and 131 both sterile). Meliosma pendens Rehder & Wilson, Cheng & Hwa 577; Gressitt 2460. Meliosma sinensis Nakai, Cheng & Hwa 772. Balsaminaceae Impatiens lucorum Hooker f. Cheng & Hwa 642, 796, 838 (very young). Staphyleaceae Euscaphis japonica (Thunb.) Kanitz, Cheng & Hwa 768; Gressitt 2476; 2531; Hwa 565. Tapiscia sinensis Oliver, Cheng & Hwa 564; Hwa 50, 538. Rhamnaceae ee (Wall.) Brongn. Cheng & Hwa 621; Djou 119; Gressitt 2406, 2 ices Sees aii Wall. ex Lawson, Cheng & Hwa 1046. Hovenia trichocarpa Chun & Tsiang, Cheng & Hwa 561, 734, 1039; Hwa 168. Rhamnus esquirolii Lévl. Cheng & Hwa 810. Rhamnus paniculiflorus Schneider, Cheng & Hwa 1115. Rhamnus rugulosus Hemsley, Cheng & Hwa 613, 1045. Rhamnus utilis Decaisne, Cheng & Hwa 607, 629, 833, 909, 970, 971; Djou 131; Gressitt 2448, 2523; Hwa 68, 98, 512. Sageretia henryi J. R. Drumm. Cheng & Hwa 864. 1980] HU, METASEQUOIA FLORA 83 Vitaceae Ampelopsis chaffanjoni (Lévl.) Rehder, Cheng & Hwa 766, 827. Parthenocissus heterophylla (Bl.) Merr. Cheng & Hwa 917. Tetrastigma serrulatum Planchon, Cheng & Hwa 943. Elaeocarpaceae Elaeocarpus chinensis (Gardner & Champion) Hooker f. Cheng & Hwa 680; Hwa 434 wa Sloanea hemsleyana (Ito) Rehder & Wilson, Cheng & Hwa 1097; Hwa 293. Tiliaceae Tilia tuan Szysz. Cheng & Hwa 741, 1059, 1114. Tilia sp., Cheng & Hwa 1095. Malvaceae Hibiscus syriacus L. Cheng & Hwa 721; Hwa 99. Actinidiaceae Actinidia arguta Planchon ex Miq. ae & Hwa 1109. Actinidia callosa Lindley, Cheng & H Actinidia chinensis Planchon, Gressitt 2419, 2495. Actinidia coriacea (Finet & Gagnep.) Dunn, Cheng & Hwa 567. Theaceae Camellia caudata Wall. Cheng & Hwa 620, 931. oe eee cuspidata (Kochs) Veitch, Cheng & Hwa 609, 686; Gressitt 2471; Hwa 270. Camellia japonica L. Hwa 255. Camellia oleifera Abel, Cheng & Hwa 568, 1044; Gressitt 2450. Camellia sinensis (L.) O. Kuntze, Cheng & Hwa 689, 1140; Gressitt 2490; Hwa 320. Eurya nitida Korthals var. aurescens (Rehder & Wilson) Kobuski, Hwa 136, 273. eth sinensis (Hemsley & Wilson) Airy Shaw, Cheng & Hwa 890; Gressitt 2480. Schima superba Gardner & Champion, Cheng & Hwa 1023. Schima parviflora Cheng & H. T. Chang, Cheng & Hwa 953 (isotype; det. n Ternstroemia nitida Merr. Hwa 455 (flower buds very young). Guttiferae Hypericum ascyron L. Gressitt 2431. Hypericum attenuatum Choisy, Cheng & Hwa 819. Hypericum petiolulatum Hooker f. & Thomson, Cheng & Hwa 736, 1092. Violaceae Viola cf. confusa Champion ex Bentham, Cheng & Hwa 1087 (sterile). Viola inconspicua Bl. Cheng & Hwa 965, 1091. Viola triangulifolia W. Beck, Cheng & Hwa 1086. 84 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Flacourtiaceae Carrierea calycina Franchet, Cheng & Hwa 928. Idesia polycarpa Maxim. Cheng & Hwa 723, 979; Djou 118; Hwa 539. Idesia polycarpa Maxim. var. vestita Diels, Cheng & Hwa 1060 Poliothyrsis sinensis Oliver, Cheng & Hwa 1132; Hwa 511. Stachyuraceae Stachyurus himalaicus Hooker f. & Thomson, Cheng & Hwa 604, 988, 994. Begoniaceae Begonia evansiana Andr. Cheng & Hwa 590. Begonia pedatifida Levl. Cheng & Hwa 840. Elaeagnaceae Elaeagnus bockii Diels, Cheng & Hwa 800; Gressitt 2462. Elaeagnus henryi Warb. Hwa 158. Elaeagnus schnabeliana Hand.-Mazz. Cheng & Hwa 608. Elaeagnus viridis Servettaz, Cheng & Hwa 809. Lythraceae Lagerstroemia indica L. Cheng & Hwa 765. Alangiaceae Alangium chinense (Lour.) Rehder, Cheng & Hwa 678; Gressitt 2492. Alangium handelii Schnarf, Hwa 105 (closely related to A. chinense; recognized by hairy fruits and strigose branchlets and inflorescences). Nyssaceae Nyssa sinensis Oliver, Cheng & Hwa 611; Hwa 61, 432, 433. Melastomataceae Sarcopyramis nepalensis Wall. Cheng & Hwa 572, 730 (very small and young; more material needed for positive determination). Onagraceae Circaea erubescens Franchet & Sav. Cheng & Hwa 842. Epilobium angustifolium L. Cheng & Hwa 712 Araliaceae Aralia decaisneana Hance, Cheng & Hwa 888. Dendropanax chevalieri (Vig.) Merr. Cheng & Hwa 684; Hwa 156, 174 (both sterile). Evodiopanax evodiifolius (Franchet) Nakai, Cheng & Hwa 900, 903, 980; Hwa 464. Hedera nepalensis K. Koch var. sinensis (Tobler) Rehder, Cheng & Hwa 780, 1129; Hwa 129. Kalopanax septemlobus rane oo (Kalopanax pictus (Thunb.) Nakai), Cheng & Hwa 711; Hw Nothopanax bodinieri (Lévl.) S. Y. “a comb. no Aralia bodinieri Lévl. In Bull. Geogr. Bot. 4: 143 saa alii delavayi sensu Rehder in Jour. ica ee 15: 115. 1934, pro par 1980] HU, METASEQUOIA FLORA 85 Rehder in 1934 interpreted Heptapleurum esquirolii Lev. (Cavalerie 871, holotype), and Aralia bodinieri Lévl. (Cavalerie 2696, syntype) as synonyms of Nothopanax delavayi (Franchet) Harms. He was correct with the first epithet. Fragments and a photograph of the syntype of Aralia bodinieri show distinctive characteristics in leaves and inflofescences that do not appear in over 40 sheets of Yunnan specimens of Nothopanax delavayi examined. Aralia bodinieri (Cavalerie 2696) is a species of Nothopanax and is transferred above. Nothopanax bodinieri has unifoliolate, trifoliolate, and pentafoliolate leaves with the leaflets 2-4 times wider than those of N. delavayi. The fruiting peduncles of the umbels are half as long as those of N. delavayi. It is a distinctive species of northeastern Kweichow and the adj ae areas of Szechwan and Hupeh, while N. delavayi is endemic to western Y Nothopanax davidii (Franchet) Harms, Cheng & Hwa 783; Gressitt 2525; Hwa 43. Nothopanax rosthornii Harms, Cheng & Hwa 773; ene 2556. Schefflera delavayi (Franchet) Harms, Cheng & Hwa 978. Schefflera venulosa (Wight & Arnott) Harms, Hwa da. Cornaceae Aucuba chinensis Bentham f. angustifolia Rehder, Hwa 315, Cornus controversa Hemsley, Cheng & Hwa 634; Hwa 458. Cornus kousa Buerger var. chinensis Bean, Cheng & Hwa 570, 664, 908; Hwa 31, 107. Cornus macrophylla Wall. Cheng & Hwa 632, 919, 1014; Djou 137; Gressitt 2428; Hwa 29, 91, 130. Helwingia japonica (Thunb.) Dietrich, Cheng & Hwa 640, 930, 1093; Gressitt 2429 Torricellia intermedia Harms, Cheng & Hwa 1145; Hwa 12, 421. Umbelliferae Daucus carota L. Djou 130; Gressitt 2520. Clethraceae Clethra fargesii Franchet, Cheng & Hwa 821, 1068, 1110. Ericaceae Enkianthus chinensis Franchet, Cheng & Hwa 1100. Enkianthus serrulatus Schneider, Cheng & Hwa 616; Hwa 171, 275. Lyonia ovalifolia (Wall.) Drude, Cheng & Hwa 746, 1053; Hwa 73. Pieris formosa D. Don, Cheng & Hwa 802, 1041; Hwa 44, 243. Rhododendron argyrophyllum Franchet, Hwa 66, 274 Rhododendron caeruleum Lévl. Cheng & Hwa 891. Rhododendron ciliicalyx Franchet, Hwa 277. Rhododendron farrerae Tate ex Sweet, Cheng & Hwa 897, 1027; Hwa 377. Rhododendron fortunei Lindley, oF & Hwa 896; Hwa 353. Rhododendron lutescens Franchet, Hw Rhododendron ovatum (Lindley) Peon “Cheng & Hwa 1029; Hwa 298, 445, Rhododendron oxyphyllum Franchet, Cheng & Hwa 682; Hwa §4. Rhododendron simsii Planchon, Cheng & Hwa 879; Gressitt 247. Hwa 453. 86 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Vaccinium bracteatum Thunb. Cheng & Hwa 990. Vaccinium carlesii Dunn, Hwa 299. Vaccinium hirtum Thunb. Cheng & Hwa ss 675, 1135. Vaccinium iteophyllum Hance, Cheng a se 610 Vaccinium japonicum Miq. Cheng & H 685. Vaccinium mandarinorum Diels, cee & Hwa 673, 898, 1056. Myrsinaceae Ardisia chinensis Bentham, Cheng & Hwa 1154. Ardisia crenata (Thunb.) DC. Cheng & Hwa 1031. Maesa hupehensis Rehder, Cheng & Hwa 922 Myrsine cf. semiserrata Wall. Cheng & Hwa 1148 (leaves and inflorescences match well, but uniformly punctate lower leaf surface differs). Primulaceae Lysimachia fortunei Maxim. Cheng & Hwa 1085. Ebenaceae Diospyros lotus L. Cheng & Hwa 710; Hwa 30. Styracaceae Alniphyllum megaphyllum Hemsley, Cheng & Hwa 984, 1008; Hwa 490. Pterostyrax hispidus Sieb. & Zucc. Cheng & Hwa 1107; Hwa 491, 566. Styrax bodinieri Lévl. Cheng & Hwa 559, 662, 975; Gressitt 241, 2573; Hwa 16, 427 (young inflorescences of some specimens infested by insect galls). Styrax suberifolius Hooker f. & Arn. Cheng & Hwa 677, 918; Hwa 121, Symplocaceae Symplocos anomala Brand, Cheng & Hwa 770, 1064; Hwa 147. Symplocos botryantha Franchet, Cheng & Hwa 690, 868, 904; Hwa 74, 140, 316, 373, 382. Symplocos caudata Wall. Cheng & Hwa 603, 1061; Hwa 244, 310, 314. Symplocos discolor Brand, Hwa , 544, sa lancifolia Sieb. & Zucc. ‘Cheng & Hwa 691, 984; Djou 107; Hwa 309. lca stellaris Brand, Cheng & Hwa 1112; Hwa 425. Oleaceae Fraxinus chinensis Roxb. Cheng . Hwa 807; Djou 101 (sterile); Hwa 402. Fraxinus Saori 5. ¥. Huy sp, vel frutex, glabra, | al robustis, hornotinis 4-5 mm. diametro, fénticellis albidis numerosis notatis; foliis trifoliolatis, petiolis 3.5-5 cm. longis, canaliculatis, basi incrassatis, foliolis petiolulis medianis 2-3 cm. longis, lateralibus 6-8 mm. longis, laminis aon ovatis, medianis 10-13 cm. longis, 6-7 cm. latis, lateralibus 8.5-12 cm. longis, 4-5 cm. latis, basi acutis, apice acuminatis, acumine 1-1.5 cm. oe subtus glabris et punctatis, margine crenulato- serratis, costa supra impressa, subtus elevata spathulatis, 2.5-3 cm. longis, 5 mm. latis, apice retusis; calycibus persis- 1980] HU, METASEQUOIA FLORA 87 tentibus patelliformibus, lobis deltoideis, apice subulato-acuminatis vel is. Cheng & Hwa 946 (holotype, 4). The trifoliolate, subcoriaceous leaves and the linear-spathulate samaras of this species suggest a close relationship to Fraxinus trifoliolata W. W and the persistent calyx of the fruit is subcampanulate with very shallow erose lobes. The ovary of this species is covered by peltate glandular scales that are persistent in fruit and become rather conspicuous on the basal one-third of the samara. In contrast, the lower leaf surface of the species in the metasequoia area is glabrous and punctate, and its persistent calyx is patelliform and has well-defined lobes that are subulate-acuminate or acute at the apex. The glandular scales on the fruit are very sparse and obscure Jasminum lanceolarium Roxb. Cheng & Hwa 989. Ligustrum lucidum Aiton, Cheng & Hwa 1157; Hwa 68. Ligustrum quihoui Carr. Cheng & Hwa 713. Ligustrum sinense Lour. Cheng — oe 1077; Hwa 115. Osmanthus cooperi Hemsley, Hwa 430. Osmanthus fragrans Lour. Cheng “& Hwa 724; Hwa 272, 303. Loganiaceae Buddleia davidii Franchet, Cheng & Hwa 811; Gressitt 2501; 2527. Gentianaceae Swertia bimaculata Clarke, Cheng & Hwa 955. Apocynaceae Trachelospermum jasminoides (Lindley) Lemaire, Hwa 261. Asclepiadaceae Cynanchum auriculatum Royle, Cheng & Hwa 638A; Djou 114, 132. Convolvulaceae Cuscuta chinensis Lam. Cheng & Hwa 860. Verbenaceae Callicarpa bodinieri Lévl. var. giraldii (Rehder) Rehder, Cheng & Hwa 1124; Gressitt 2526. Callicarpa rubella Lindley var. hemsleyana Diels, Cheng & Hwa 1042. Clerodendron bungei Steudel, Cheng & Hwa 631, 914; Gressitt 2454, 2553. Clerodendron mandarinorum Diels, Cheng & ae 769, 894; Hwa 40. Clerodendron trichotomum Thunb. Cheng & Hwa 913. Premna puberula Pampan. Cheng & Hwa 560; ee 2427, 2483, 2558. Labiatae Mosla chinensis Maxim. Cheng & Hwa 788, 1096. Mosla dianthera (Buch.-Ham.) Maxim. Cheng & Hwa 789. Perilla frutescens (L.) Britton, Gressitt 2567. Pogostemon glaber Bentham, Cheng & Hwa 835. 88 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Prunella vulgaris L. Cheng & Hwa 793. Teucrium bidentatum Hemsley, Cheng & Hwa 1032. Solanaceae Solanum lyratum Thunb. Cheng & Hwa 1177. Scrophulariaceae Botryopleuron nervosum (Hemsley) Hemsley, Cheng & Hwa 1015. Paulownia fargesii Franchet, Hwa 95, 462. Phtheirospermum chinense Bunge ex Fischer & Meyer, Cheng & Hwa 1088. Bignoniaceae Catalpa fargesii Bur. Cheng & Hwa 855, 935; Gressitt 2418: Hwa 510. Gesneriaceae Didissandra cf. sesquifolia C. B. Clarke, Cheng & Hwa 718, 841 (specimens resemble D. sesquifolia of northern India in having pubescent leaves and short-pedicellate flowers, but differ in having leaf margins subentire, not subduplo-serrate). Didymocarpus hwaianus S. Y. Hu, Vv. Herbae succulentae, perennes, aire internodis caulium 4~7 cm. longis, 3 mm. crassis; ars oppositis, sessilibus, integris, ellipticis, 12-15 cm. longis, 4.5-5 cm. latis, supra glabris vel leviter hirsutis, subtus glabris, fere ad basin libris, obovato- -oblongis, 8-9 m ene 5-6 mm. glabris, apice rotundatis; corollis et staminibus rene capsulis juvenilibus compresso-cornutis, 2.5 cm. longis, 3 mm. latis; annulis interruptis; stylis 1 cm. longis, stigmatibus inaequaliter bilobatis. Cheng & Hwa 923 (holotype, a). The foliaceous sepals free almost to the base, the short axillary inflores- cences and the laterally compressed, curved, cornute young capsules of this Ra suggest a close relationship with Didymocarpus bicornutus (Hayata) S. Y. Hu,’ comb. nov., from Taiwan, which has petiolate leaves. Another closely related species is D. tibeticus Franchet, which differs in having villose, lanceolate sepals and petiolate leaves. Lysionotus pauciflora Maxim. Cheng & Hwa 1003. Acanthaceae Hemigraphis cuneata S. Y. Hu, sp. erba prostrata, caulibus. aad: 4 late: glabrescentibus; foliis parvis, siembeldels 1-2.5 cm. longis, 0.7-2 cm. latis, basi cuneatis, apice acutis, margine remote serratis, utrinque glabrescentibus, petiolis 5-15 mm. longis, glabris; floribus solitariis, in axillis foliorum sessilibus, bracteis foliaceis, obovato-cuneatis, 3 mm. longis, 2 mm. latis, apice obtusis; calycibus glabris, fere ad basim partitis, laciniis linearibus, 1.5 cm. longis, 1.5 mm. latis, *Didymocarpus bicornutus (Hayata) S. Y. Hu, comb. nov. Chirita bicornuta Hayata, Ic. Pl. Formos. 3: 154. 1913. 1980] HU, METASEQUOIA FLORA 89 eee obtusis; capsulis oblongis, ad basim constrictis, 12-13 mm. longis, . diametro, glabris; seminibus Pee -compressis, suborbicularibus, ee mm. diametro, atro-fuscis, papillosis Cheng & Hwa 887 (holotype, a This species is related to Hemigraphis procumbens (Lour.) Merr., from which it is readily distinguished by its glabrous leaves, sepals, and capsules. The leaves of H. pro he eae are hirsute and the sepals hispid-strigose; the capsules are hairy at the apex. Peristrophe roxburghiana (Schultes) Bremek. Cheng & Hwa 715. Plantaginaceae Plantago major L. Cheng & Hwa 717. Rubiaceae Emmenopteryx henryi Oliver, Cheng & Hwa 1173; Hwa 166. Galium bungei Steudel, Cheng & Hwa 784, 1143. Hedyotis lindleyana Hooker var. glabra Hara, Cheng & Hwa 790. Lasianthus hartii Franchet, Cheng & Hwa 622. Paederia chinensis Hance, Cheng & Hwa 926, 973, 996. Paederia scandens (Lour.) Merr. Cheng & Hwa 628, 824. Serissa foetida Commerson, Cheng & Hwa 983. Caprifoliaceae Abelia myrtilloides fgeree Cheng & Hwa 861, 1133. A belia uniflora . Hwa 452. Lonicera henryi Hemsley, ‘Cheng & Hwa 676, 1036, 1083. Lonicera macranthoides Hand. ae Cheng & Hwa 882. Sambucus racemosa L. Hwa 11, 372. Viburnum ichangense ae Rehder, Cheng & Hwa 814, 1105; Gressitt o59: Viburnum ovatifolium Rehder, Cheng & Hwa 649, 992; Hwa = Viburnum propinquum Hemsley, Cheng & Hwa 867; Hwa 265, 5. Viburnum propinquum Hemsley var. mairei W. W. Sm. Cheng i Hwa 860, Li22: Viburnum rhytidophyllum Hemsley, Hwa 359. Viburnum rosthornii Graebner var. xerocarpa Loes. Cheng & Hwa 892; Hwa Viburnum setigerum Hance, Chang & Hwa 803, 993, 1071; Djou 117; Gressitt 2434, 2521. Viburnum sympodiale Graebner, Hwa 341 or 355 (number not clear). Viburnum ternatum Rehder, Cheng & Hwa 589, 732, 1019; Gressitt 2474; Hwa 56. Viburnum theiferum Rehder, Hwa 473. Viburnum tomentosum Thunb. Hwa 282, 426. Viburnum tsangii Rehder, Cheng & Hwa 883, 1161. Weigela japonica Thunb. var. sinica (Rehder) Bailey, Cheng & Hwa 573; wa 47]. Valerianaceae Patrinia villosa (Thunb.) Juss. ex DC. Cheng & Hwa 964. 90 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Cucurbitaceae Gymnostemma pentaphyllum Makino, Cheng & Hwa 925, 1014. Campanulaceae Campanumoea javanica Bl. Cheng & Hwa 625; Gressitt 2424. Lobelia davidii Franchet, Cheng & Hwa 643 (very similar to L. fangiana (E. Wimmer) S. Y. Hu, comb. nov.,° from which it may be distinguished by the subglabrous ovary and hairy anthers). Lobelia radicans Thunb. Cheng & Hwa 578 Pratia nummularia (Lam.) A. Br. & Ascherson (Lobelia angulata Forster), Cheng & Hwa 599 Compositae Adenocaulon adhaerescens Maxim. Cheng & Hwa 719. Ainsliaea fulvipes Jeffrey, Cheng & Hwa 786 Anaphalis margaritacea (L.) Bentham & Hooker f. Cheng & Hwa 762. Carpesium abrotanoides L. Cheng & Hwa 679. eo ales chinense L. Cheng & Hwa 967. Eupatorium japonicum Thunb. Cheng & Hwa 837. Ixeris Pace (DC.) Stebbins, Cheng & Hwa 1084. Leontopodium sinense Hemsley, Gressitt 2515. Ligularia duciformis (C. Winkler) Hand.-Mazz. Cheng & Hwa 850. Monocotyledoneae Alismataceae Sagittaria pygmaea Miq. Cheng & Hw Sagittaria sagittifolia L. subsp. oe. ee ) Rataj. Cheng & Hwa 760. Hydrocharitaceae Ottelia alismoides (L.) Pers. Cheng & Hwa 847 (material too young to show species characteristics). Gramineae Chimonobambus sp., Hwa ae 496 (sterile; det. T. R. Soderstrom). Miscanthus sinensis Andersson, Gressitt ee Phyllostachys sp., Gressitt 2410 (sterile; det. T. R. Soderstrom). Sinarudinaria sp., Gressitt 2494 (det. T. R. ene Hwa 494. “Lobelia fangiana (E. Wimmer) S. Y. Hu, comb. Pratia fangiana E. Wimmer, Repert. io Nov. 38. e. Lobelia omiensis E. Wimmer, Ann . Mus. vee wn 1948. SZECHWAN: Mt. Omei, W. P. Fang oe renee NY; isotype, GH), E. H. Wilson 5034 (isotype of L. omiensis, GH), H. C. Chow 7927, 8106 (a), S. C. Sum & K. < 5 oO 5 a oo Ss n ctions from the same area give a fuller picture of morphological variation of the pe and the gaps caused by age differences in immer’s s samples are bridged. The species is characterized by hirsute inflorescences, pubescent ovaries, and glabrous anthers. A transfer of the earlier specific epithet from Pratia to Lobelia is necessitated. 1980] HU, METASEQUOIA FLORA 9] Cyperaceae Bulbostylis teeta (L.) Clarke, a & Hwa 1089. Carex brunnea Thunb. Cheng & Hwa 791 (det. T. Koyama). Cyperus pence Franchet & Sav. Cheng & Hwa 753 (det. T. Koyama). Eleocharis lea a D. Don subsp. japonica (Miq.) Koyama, Cheng & Hwa 758 (det. T. Koyama). Kyllinga brevifolia Rate Cheng & Hwa 596, 757. Pycreus flavescens (Poiret) Koyama, Cheng & Hwa 754 (det. T. Koyama). Pycreus sanguinolentus Vahl, Cheng & Hwa 755 (det. T. Koyama). Scirpus wallichii Roxb. Cheng & Hwa 756 (det. T. Koyama). Eriocaulaceae Eriocaulon buergerianum Koern. Cheng & Hwa 759. Commelinaceae Commelina communis L. Cheng & Hwa 785, 957. Pontederiaceae Monochoria vaginalis (Burman f.) Presl, Cheng & Hwa 763, 848. Juncaceae Juncus setchuensis Buchenau, Cheng & Hwa 644. Liliaceae Cardiocrinum giganteum (Wall.) Makino, Cheng & Hwa 849. Disporum cantoniense (Lour.) Merr. Cheng & Hwa 941. Lilium leucanthum Baker var. — eae Wilson, Gressitt 2513. Paris chinensis Franchet, Cheng & Hw a Protolirion sinii Krause, Cheng & Hwa ea Reineckea carnea Kunth, Cheng & Hwa 597. Smilax lanciifolia Roxb. Cheng & Hwa 982. Smilax megalantha A. DC. Cheng & Hwa 633; Hwa 350. Smilax myrtillus DC. Cheng & Hwa 929. Smilax riparia DC. Cheng & Hwa 972, 974, 1155. Iridaceae Belamcanda chinensis (L.) DC. Gressitt 2416. Iris japonica Thunb. Cheng & Hwa 652. Dioscoreaceae Dioscorea cirrhosa Lour. Cheng & Hwa 1018 Dioscorea kamaonensis Kunth var. fargesii (Franchet) Prain & Burkill, Cheng & Hwa 638, 656, 836, 1016 (det. B. G. Schubert). Orchidaceae Cymbidium ensifolium Sw. Cheng & Hwa 1006. Cypripedium japonicum Thunb. Cheng & Hwa 1162. Pleione henryi Rolfe, Cheng & Hwa 911. 92 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 ACKNOWLEDGMENTS Grateful acknowledgment is due all my friends who have helped with the identification of specimens and the preparation of the manuscript. I should like to take this opportunity to express my special appreciation to Drs. L. M. Perry, B. G. Schubert, S. A. Spongberg, P. F. Stevens, R. M. Tryon, and C. E. Wood, Jr., for numerous helpful suggestions. Particular thanks are also extended to R. A. Howard for the opportunity to undertake this project. I should also like to acknowledge the careful work of Karen Stoutsenberger, who prepared the final versions of the maps that illustrate this article. REFERENCES Anprews, H. N., Jr. 1961. Studies in paleobotany. xi + 487 pp. Wiley, New York. Anonymous. 1948. Living fossil. Jour. N. Y. Bot. Gard. 49: 949. Rand McNally cosmopolitan world atlas. xvi . a pp. Rand McNally & Co., New York. 1972-1976. Iconographia cormophytorum Sinicorum. Vol. Bryophyta, Pteridophyta, Gymnospermae, Angiospermae (Casuarina- ceae-Hermanniaceae). vi + 1157 pp. 1972. Vol. Il. Papaveraceae—Corna- ceae. iv + 1312 pp. 1972. Vol. III. Diapensiaceae-Solanaceae. ii + 1083 pp. 1974. Vol. IV. 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H. 1976. Metasequoia—fossil and living. Bot. Rev. 42: 215-315. [Additions. [bid. 43: 281-284. 1977.] Goop, R. 1974. Geography of flowering plants. ed. 4. 557 pp. Longman, London GressITr, J. L. 1953. The California Academy-—Lingnan dawn-redwood expedi- Proc. Cal. Acad. Sci. 28: 25-58. Hara, H. 1966. Taxonomic comparison between corresponding taxa of 1980] HU, METASEQUOIA FLORA 93 Spermatophyta in eastern Himalaya and Japan. Pp. 627-657 in H. Hara, comp., The flora of eastern Himalaya. x + 744 pp. University of Tokyo Press, Tokyo. Hu, H. H. How Metasequoia, the “‘living fossil,’ was discovered in China. Jour. N. Y. Bot. Gard. 49: 201-207. & W. C. Cuenca. 1948. On the new family Metasequoiaceae and on Metasequoia glyptostroboides, a living species of the genus Metasequoia found in Szechuan and Hupeh. Bull. Fan Mem. Inst. Biol. II. 1: 153-166. Kackman, C. 1965. The Old World species of Prunus subg. Laurocerasus including those formerly referred to Pygeum. Blumea 13: 1-115. Lee, J. S. 1939. The geology of China. 527 pp. Thomas Murby & Co., London. Li, H. L. 1971. Floristic relationship between eastern Asia and eastern North America. Trans. Am. Philos. Soc. 42: 371-429. Matuews, R. H. 1931. A Chinese- English dictionary. xxiv + 1226 pp. Presbyterian Mission Press, Shanghai. MerriLL, E. D. 1948. Metasequoia, another ‘‘living fossil.’? Arnoldia 8: 1-8. Miki, S. 1941. On the change of flora in eastern Asia since Tertiary Period. . The clay or lignite bed flora in Japan with special reference to the Pinus ee beds in Central Hondo. Jap. Jour. Bot. 11: 237-303. . On es al fossil and living. 141 pp. Nippon Kobutsu eet -no, Kyoto . 1954. The occurrence of the remains of Taiwania and Palaeotsuga (n. subg.) from Pliocene beds in Japan. Proc. Japan Acad. 30: 796-981. Nairn, A. E. M., ed. 1961. Descriptive palaeoclimatology. xi + 380 pp. Interscience Publishers, Inc., New : Onwi, J. Flora of Japan. (English edition; F. G. Meyer & E. H. Wacker, eds.) ix + 1067 pp. Smithsonian Institution, Washington, D. C. Orr, M. Y. 1933. A new Chinese silver fir. Notes Bot. Gard. Edinburgh 18: 1-5. . 1933a. Plantae Chinenses forrestianae: Coniferae. [bid. 119-158. REHDER, A. 1940. Manual of cultivated trees and shrubs. ed. 2. xxx + 996 fe) ———.. 1949. Bibliography of cultivated wee a shrubs. xl + 825 pp. Arnold Agborenitn. Jamaica Plain, Massachus & E. H. Wirson. 1914. Taxaceae a Pinaceae. In: C. S. SARGENT, Sma, J. K. 1913. Flora of the southeastern United States. ed. 2. 1394 pp. ‘‘Published by the author,’’ New York. Wana, Z., & P. Y. Fu. 1974. Plantae novae Salicacearum Tibeticarum. Acta Phytotax. Sinica 12: 191-213. Witson, E. H. 1926. The taxads and conifers of Yunnan. Jour. Arnold Arb. 7: 37-68. Wo re, J. A. 1969. Neogene floristic and vegetational history of the Pacific Northwest. Madrono 20: 83-110. 5 17a. A a ania meg one of Tertiary climates in the Northern Hemisphere. . 66: 694-703. Woop, C. E., Jr. 1971. ae rales relationships between the Southern Appalachians and western North America. Pp. 331-404 in P. C. Hott, ed., The distributional history of the biota of the Southern Appalachians. 94 JOURNAL OF THE ARNOLD ARBORETUM [voc. 61 Part Il. Flora. Research Div. Monogr. 2. Virginia Polytech. Inst. State Morphology and phytogeography: the classical approach to the study of disjunctions. Ann. Missouri Bot. Gard. 59: 107-124. ARNOLD ARBORET 22 DIvINity cee CAMBRIDGE, ee aa 02138 1980] DARWIN, AIROSPERMA 95 NOTES ON AIROSPERMA (RUBIACEAE), WITH A NEW SPECIES FROM FIJI STEVEN P. DARWIN Tue Genus Airosperma Lauterbach & K. Schumann has not been widely discussed by taxonomists concerned with Rubiaceae of the western Pacific, although it has recently been studied by botanists revising the Rubiaceae for Flora Malesiana (Ridsdale, Bakhuizen van den Brink, & Koek-Noorman, 1972). Although a formal revision of the genus would be premature, notes on the distribution, ee. oe pollen, and nomenclature of Airosperma species are presented Future workers will have to cer an the problematic tribal affinity of irosperma. | previously (Darwin, 1979) referred the genus to the tribe Alberteae, for the most part following earlier opinion. Interested readers should consult that discussion for a historical outline as well as for an enumeration of the morphological characters that define Airosperma and suggest a relationship with Boholia Merr. from the Philippines As interpreted here, Airosperma comprises six species, four in New Guinea and two in Fiji. Schumann and Lauterbach (1900) established the genus with two species, A. psychotrioides and A. ramuense, both from New Guinea. I have already designated A. psychotrioides the lectotype species (Darwin, 1979); of the two original species, it is by far the better collected. In addition, it was provided by its authors with the more expanded description and was the only species illustrated. Valeton added a third species in 1912, and S. Moore a fourth in 1927. Smith (1945) later transferred Gillespie’s monotypic Fijian genus A bramsia to Airosperma. A sixth described species of Airosperma, also from Fiji, is presented here as new. It has been pointed out (Darwin, 1979) that nearly all genera of Pacific Rubiaceae center in Malesia, or at least their closest allies can usually be found there. To varying degrees, many of those genera have probably expanded their geographic ranges into the Pacific by migrating southward over an island chain now represented by the Solomon Islands, the New Hebrides, Fiji, and associated smaller islands. Smith (1979, p. 25 et seq.) has outlined in a general way the probable geologic events that made such a migration route possible, and pointed out that such a route may have been only intermittently present. If archipelagoes have come and gone along the eastern edge of the Australian continental plate, it would not seem unusual to find two closely related species of Airosperma in Fiji, well isolated from the New Guinean members of the genus. It may well be that the southernmost species of Airosperma trace their ancestry to one or a few diaspores reaching Fiji by © President and Fellows of Harvard College, 1980. Journal of the Arnold Arboretum 61: 95-105. January, 1980. 96 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 means of long-distance dispersal, but there is no apparent adaptation for such dispersal, except, possibly, floating fruits. The more likely hypothesis, it seems to me, is that other geographically intermediate populations have become extinct, leaving survivors only at the extreme ends of a once more continuous range. A similar distributional pattern is exhibited by the genus Mastixiodendron Melchior, in which three Fijian species are quite isolated from four other species in the Solomon Islands, New Guinea, and the Moluccas; the genus is absent only from the New Hebrides, although it may once have occurred there or on an earlier island group in the same area. Airosperma Lauterb. & K. Schum. in K. Schum. & Lauterb. Fl. Deutschen Schutzgeb. Stidsee, 565. 1900; Krause in Engler & Prantl, Nat. Pflanzenfam. Nachtr. 3: 328. 1908; A. C. Sm. Jour. Arnold Arb. 26: 107. 1945, ibid. 36: 288. 1955. Abramsia Gillespie, Bishop Mus. Bull. 91: 27. 1932. Key To SPECIES OF AIROSPERMA 1. Branchlets usually more than 5 mm. broad toward apex; leaves with at least 15 secondary nerves and petioles more than 3 mm. broad near middle: raearanes ee frequently with as many as 200 flowers, the pedicels more than 5 mm. long; New Guinea. ............, 1. A. grandifolium. l. Brancliets usually less than 4 mm. broad toward apex; leaves with fewer than 15 secondary nerves and petioles less than 3 mm. broad near middle: inflorescences usually with fewer than 200 flowers, the pedicels less than 5 mm. long; New Guinea and Fiji. 2. Stipules frequently more than 4 mm. long; pedicels not conspicuously elongating in fruit; corolla limb less than 4mm. broad; anthers apiculate by the produced connectives; mature fruits less than 10 mm. long and 6 mm. broad; New Guinea. 3. Calyx lobes at least 2.5 mm. long, frequently much longer. 4. Leaf blades lanceolate to broadly elliptic; inflorescences usually with fewer than 50 flowers. ........, A. psychotrioides. 4. Leaf blades elliptic to Pacis obovate; inflorescences usually with more than 50 flowers. ............, . A. fuscum. 3. Calyx lobes less than 2 mm. coe ee ee ee 4. A, ramuense. to ‘ Stipules usually less than 3 mm. ae pedicels conspicuously elongating in fruit; corolla limb usually more than 4 mm. broad; anthers emarginate to obtuse or acute at apex, not ee mature fruits at least 10 mm. long, usually more than 8 mm. bro ijl. 5. Corolla tube less than 3 mm. long, Te throat care See ee i oe Ha. eee ME ee ar Su ae ae gs ee 5. A. trichotomum. 5. Corolla tube more than 3 mm. long, the throat Pome ie ae eee Se Ae Gh os ee Fe os Se Seg ae Ss we a ee es . A. vanuense. 1. Airosperma grandifolium Valeton, Nova Guinea Bot. 8: 760. 1912 (as grandifolia); Bot. Jahrb. 61: 32. 1927 (as grandifolia). Ficure |. 1980] DARWIN, AIROSPERMA 97 Ficures 1-6. Scanning electron micrographs of pollen grains from species of Airosperma: 1, A. grandifolium (from NGF 48274), x 1000; 2, A. psy chotrioides (from Brass 23535), x 1000; 3, A. ramuense (from NGF 39326), x 1000; 4, A. aff. ramuense (from Brass 3884), x 640; 5, A. trichotomum es Degener 14549), x 1000; 6, A. vanuense (from Smith 1849), x 1200. 98 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Probably the most distinct species of Airosperma, readily distinguished by its large (20-40 cm. long x 9-18 cm. wide), elliptic to obovate leaf blades, which taper to a short, stout petiole. The inflorescences are much congested, often with as many as 300 relatively long-pedicellate flowers. Like flowers of other Airosperma species in New Guinea, those of A. grandifolium have the same relative calyx and corolla size, a dense ring of white hairs at the corolla throat, and short-apiculate anthers. As discussed below, the pollen of this and all other Airosperma species from New Guinea is 3- to 4-porate and tectate-perforate to semitectate. Collections of Airosperma grandifolium are very few. Valeton (1912, cited ene founded the species on a single collection, Gjellerup 239, collected at ‘‘Salzquelle am Beguwri’’ (Beguwri River, Djajapura, West New Guinea aiien Barat)). Valeton evidently studied the material at eee (now Bogor, see Valeton, op. cit., p. 755); I have seen neither the holotype (at Bo?) nor any isotypes. I have not examined two other collections referred to this species by Valeton (1927, cited above), Schlechter 20175 and Ledermann 8351, both from the Sepik River drainage of Papua New Guinea. More than a dozen other collections have since been annotated as A. grandifolium, but most of the specimens previously attributed to this species are better referred to other genera on the basis of the presence of raphid crystals, basally attached ovules, or valvate corolla lobes. Many such excluded collections may be allies of Psychotria, some species of which bear a strong superficial resemblance to A. grandifolium. Only one specimen seen, NGF 48274 (cited below), conforms to Valeton’s precise description in nearly every detail, except that the stipules of A. grandifolium were described as being strongly caducous, whereas in the collection examined they are relatively persistent. NGF 48274 was collected in the West Sepik District of Papua New Guinea, reasonably close to the locality given by Valeton for the type, and agrees also in its label information—the specimen taken from a herbaceous shrub about | meter tall with white flowers. Valeton reported the type collection as having only flower buds, these much damaged by insects. This is also true of the single collection I have seen. Dr. C. E. Ridsdale (pers. comm.) suggests that recent additional collections of Airosperma grandifolium may have been sent to Leiden, but that they have not as yet been processed or distributed. Collections with mature, undamaged flowers and fruits would be very welcome. PECIMEN EXAMINED. Papua New Guinea. West Sepik: sago swamp, N. of Kilifas Village, 305 m. alt., NGF 48274 (Foreman & Kumul) (Lae). 2. Airosperma psychotrioides Lauterb. & K. Schum. in K. Schum. & Lauterb. Fl. Deutschen Schutzgeb. Sudsee, 565. ¢. XXJ. 1900; Valeton, Bot. Jahrb. 61: 32. 1927 (as psychotroides); Merr. & Perry, Jour. Arnold Arb. 26: 14. 1945. Ficure 2. This species is recognizable by its relatively few-flowered inflorescences (I have counted 15 to 40 flowers in each), and its calyces, which are 2.5— mm. long, the subulate to narrowly oblanceolate lobes considerably longer 1980] DARWIN, AIROSPERMA 99 than the cupular base. This and the two following species are distinct from Airosperma grandifolium in their smaller leaves and relatively slender petioles, among other characters. Lauterbach and Schumann based Airosperma psychotrioides on four collec- tions made by Lauterbach, all from Kaiser Wilhelmsland, all presumably deposited in the Berlin Herbarium and now destroyed. I have located duplicates of three of those collections at the Wrocklaw Herbarium (wrsL) and here designate Lauterbach 510 as lectotype, cited below. The lectotype is noted (on a Berlin Museum label) as having been taken from Sattelberg, Kekagalla, at an altitude of 970 meters on 22/26 June, 1890; a Herb. Lauterbach label with greenish flowers. Lauterbach 510 represents flowering material only, flowers being much more abundant on the lectotype than on the isolectotype. Information from herbarium labels of other collections describes this species as a ‘‘woody herb’’ or erect shrub to 2 meters tall growing in rich rain forests from 20 to more than 1800 meters altitude. The bark is variable, smooth or flaking, the wood straw colored and sometimes with white latex; the flowers are usually greenish white when young, at maturity cream-white and often with brownish or pale purple centers. The young fruits are greenish white, eventually becoming bright blue. Flowering and fruiting collections have been gathered throughout the year. Further investigation into the Airosperma psychotrioides ‘‘complex’’ must also take into account floral dimorphism, possibly associated with out-breeding mechanisms. In the herbarium material studied, there are two flower types. In the first, growth of the style keeps pace with the elongation of the corolla tube; in late bud the stigma is at the corolla throat and is to varying degree exserted beyond the limb after separation of the corolla lobes. The mature stigma lobes are about | mm. long and widely divergent. In the same flowers, the anthers are much reduced (to | mm. long) and apparently sterile. Collections with flowers of this type include NGF 37976, Schodde & Craven 4635 and 4328, and the type of A. fuscum (see below). In the second flower type, the anthers are at least 1.5 mm. long and shed abundant pollen before the corolla bud opens; in such flowers the style was never observed to be more than about 2 mm. long. The flowers of A. psychotrioides thus appear to be either functionally staminate or pistillate. As far as I am able to determine, these two floral morphologies are confined to separate plants and suggest dioecism in this species. SPECIMENS EXAMINED. Papua New Guinea. Mapanc: Amaiaba River, NGF 45871 (Foreman et al.) (LAE), NGF 45890 (Foreman et al.) (CANB, LAE); Ramu River, tributary 9, Lauterbach 3112 (wrsL); upper course of Gogol River, Lauterbach 1124 (wrst). Sea vicinity of Kaiapit eae Clemens 10756 bis (A); Umi River, Markham Valley, Brass 32539 (a, canB, K); Boana, Clemens 41578 (a); Atzera Range, NGF 11923 (Henty) (a); Busom River, NGF 37976 (Katik) (a, Lae); vicinity of Butibum River, Hartley 11634 (a, LAE); Masba Creek, ca. 3 miles S. of Pindiu, Hoogland 8873 (a, CANB, K, LAE); Sattelberg, Weinland 312 (BRI, WRSL); Sattelberg, Kekagalla, Lauterbach 510 (wrst, lecto- type & isolectotype); Matap, Clemens 41091 (a). Gur: ca. 1 mile E. of 100 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 junction of Vailala and Lohiki rivers, Schodde & Craven 4328 (a, CANB); hill on southwest margin of junction of Kapau and Tauri rivers, Schodde & Craven 4635 (a, CANB, LAE). CENTRAL: Dieni, Ononge Road, Brass 3879 (A, BRI, NY); Mafulu, White 572 (pm). NorTHERN: Pongani Valley, vicinity of Dareki Village, Pullen 5721 (cans); between Budiand Anara barracks, Hoogland 4622 (A, CANB, on MixneE Bay: junction Ugat and Mayu rivers, NGF 28925 (Streimann) (A); Mayu River, ca. 15 km. WNW. of Biniguni, Pullen 8363 (canB); north slopes, Mt. Dayman, Brass 23535 (A, CANB, LAE, US), 23657 (a, CANB, LAE, US). 3. Airosperma fuscum S. Moore, Jour. Bot. 65: 266. 1927 (as fusca); in C T. White, Jour. Arnold Arb. 10: 268. 1929 (as fusca). The type of Airosperma fuscum is Brass 1050, the only collection seen by Moore, collected at Hohoro, Vailala River, Gulf District, Papua, from a weak shrub | meter tall growing on ridges in open rain forest at an altitude of 300 feet. The label of the holotype describes the flowers as white with pink centers, the fruits as yellow. In the sense of its type, 4. fuscum appears to be reasonably distinct from A. psychotrioides, especially in its leaf blades, which may be more than 2 cm. wider and more strongly obovate, and in its inflorescences with as many as 100 flowers. Moore further differentiated A. fuscum on the basis of its leaves drying darker and its flowers with judge from the Papuan collections at hand, it is probable that additional populations of the A. psychotrioides alliance will be found in southeastern New Guinea. No collections as yet approach the type of A. fuscum in every regard, but additional material from that type locality would greatly help in determining the proper taxonomic status of A. fuscum, very possibly supporting its recognition at the rank of species. Some aspects of the floral morphology of the present species are discussed under A. psychotrioides. SPECIMEN EXAMINED. Papua New Guinea. Gutr: Hohoro, Vailala River, Brass 1050 (sm, holotype; A, BRI, K, isotypes). 4. Airosperma ramuense Lauterb. & K. Schum. in K. Schum. & Lauterb. 1. Deutschen Schutzgeb. Stidsee, 566. 1900 (as ramuensis); Valeton, Bot. Jahrb. 61: 32. 1927 (as ramuensis); Merr. & Perry, Jour. Arnold Arb. 26: 14. 1945 Ficures 3, 4. This species, the second described by Lauterbach and Schumann, was based ona single collection, Rodatz & Klink 206, from the Bismarck Mountains (Western and Eastern Highlands of Papua New Guinea). I have not examined the holotype, which presumably was deposited in the Berlin Herbarium and is now destroyed; if no isotypes exist, then designation of a neotype will be required. However, based on the excellent original description, I have been able confidently to refer a number of more modern collections to Airosperma ramuense and to distinguish that taxon from the two preceding species. Inflorescences of A. ramuense frequently have more than 50 flowers (usually about 100), and calyces hardly 2 mm. long. In A. psychotrioides 1980] DARWIN, AIROSPERMA 101 the inflorescences have up to 50 flowers; in A. psychotrioides and A. fuscum the calyces are longer than 2 mm., frequently as long as 6 mm. Those same differences were emphasized by Lauterbach and Schumann and by Valeton (1927, cited above). Attention has also been drawn to the generally larger leaves of the present species, which often dry nearly black; however, I have found too much variation in leaf size, shape, and drying quality to regard such characters as being very useful in drawing up an analytical key to A. ramuense and A. psychotrioides. From label data, the species may be described as a shrub to nearly 2 meters tall, growing in lowland or hillside rain forests at altitudes from 30 to 610 meters. The few collections available suggest that the species may be found in flower or fruit throughout the year. The fruits are described as greenish, becoming purple at maturity, and thus contrasting with those of Airosperma psychotrioides, which are frequently described as bright blue. Only specimens from the Sepik area of Papua New Guinea can confidently be referred to Airosperma ramuense. One collection from the Central District, Brass 3884, was included in the present species by Merrill and Perry (1945, cited above), but it is well outside its otherwise recorded range. It is composed of flowering material resembling Airosperma ramuense but differs from the other specimens examined in its somewhat longer (to 30 cm.) leaves, which are oblong in outline, and in its calyces, the lobes of which may be proportionately longer. Furthermore, the inflorescences are noted as bein axillary, on branchlets, or ‘‘low down on the stem.’’ These lateral inflorescenses have peduncles (modified axillary branchlets?) to 2 cm. long, a feature not observed in any other collection here assigned to the genus Airosperma. The flowers of Brass 3884 are 5-merous, while those of specimens from the Sepik area are usually 4-merous. A new species may be represented here, but I cannot describe it formally without more material. It should be noted that the pollen from Brass 3884 is essentially identical with that of the one other specimen of A. ramuense examined, NGF 39326 (Ficures 3, SPECIMENS EXAMINED. Papua New Guinea. West Sepik: Krisa~Vanimo Road, NGF 39326 (Streimann & Kairo) (a, LAE). East Sepik: Prince Alexander Range, SE. of Mt. Turu, vicinity of Ambakanja Village, Pullen 1524 (cans); head of Giligama Creek, between Kuminim and Lawon villages, W. of Wewak, Pullen 1357 (cans). CENTRAL: Dieni, Ononge Road, Brass 3884 (a, ny). 5. Airosperma trichotomum (Gillespie) A. C. Sm. Jour. Arnold Arb. 26: 108. 1945, ibid. 36: 288. 1955; J. W. Parham, Pl. Fiji Is. 187. 1964, ed. 2. 264. 1972. Ficure 5. Psychotria Seem. Bonplandia 9: 257. 1861. Psychotria insularum sensu A. Gray, Proc. Am. Acad. 5: 319. 1862, Bonplandia 10: 36. 1862; Seem. Viti, 437. 1862; ee = Gray, 185 Abramsia trichotoma Gillespie, Bishop Mus. Bull. 29. fig. e 1932: Fosberg, Bull. Torrey Bot. Club 67: 422. 1940, el Te 125, 1942. This species was well described by Gillespie under Abramsia and was 102 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 accompanied by a lengthy discussion concerning the possible tribal relation- ships of that genus. Similar, although much expanded, commentary was offered by Smith in 1945, at which time he transferred Gillespie’s genus to Airosperma, where it belongs. As interpreted here, Airosperma trichotomum is positively referable only to the island of Viti Levu, Fiji, where it has been collected many times. The type of Abramsia trichotoma is Gillespie 3388, cited below. In his published account, Gillespie gave the locality of the type collection as ‘‘Viti Levu, Naitasiri Province, vicinity of Nasinu, 14 kilometers (9 miles) from Suva.’” However, labels (in Gillespie’s hand) accompanying the holotype and isotypes give the locality as the slopes of Mt. Lomalangi (*‘Loma Langa’’), vicinity of Nandarivatu, November 12, 1927; notes with one isotype (uc) give the altitude as 1000 meters. Since collecting numbers adjacent to Gillespie 3388 (and gathered on the same day) are also from the Nandarivatu area (A. C. Smith, pers. comm.), the label data must be taken as correct. Notes accompanying other collections portray this species as a shrub or small tree, occasionally to a height of [5 meters, growing in dense or secondary forests on wooded ridges or along streambanks at altitudes from 15 to 1250 meters. The flowers are described as white or yellow, the fruits as green, becoming white and sometimes tinged with yellow. Fruiting specimens have been gathered throughout the year, but I have seen no flowering material taken between the months of May and October. Native names recorded are ‘‘maskarawa’”’ (Smith 4426), ‘‘malinimbia’’ (Gillespie 2652), and ‘‘silasila’’ (St. John 18285), notes with the last collection further state that the plant is used locally as a cathartic. This and the following species differ from the New Guinean representatives of Airosperma in their smaller stipules, broader corolla limbs, anthers without conspicuously apiculate apices, and larger fruits borne on pedicels that elongate as the fruits mature. The present species is very closely allied to A. vanuense, differing in its shorter corollas, which are essentially glabrous within. A close relationship between the Fijian taxa is also suggested by their intectate pollen; the pollen of the Papuasian species is variously tectate (Ficures |~—6). In this and another study (Darwin, 1977), pollen was found to be exceedingly helpful in suggesting that Fijian species of some Malesian Rubiaceae may be traceable to one ancestral population. Such pollen characters will undoubt- edly prove useful in similar studies of Pacific Rubiaceae. SPECIMENS EXAMINED. Fiji. Viti Levu. Mpa: Mt. Koroyanitu (Mt. Evans), Fiji Dept. Agr. 14148 (BisH); slopes of Mt. Nairosa, eastern flank of Mt. Evans Range, Smith 4074 (a, BISH, BRI, K, NY, oT 4426 (A, BISH, BRI, K, NY, us); Nauwanga, Degener 14549 (a, BISH, NY, UC, US), 14740 (A, BISH, NY, US); vicinity of Nandarivatu, Degener 14807 (a, ny, us), Degener & Ordonez 15542 (A, BISH, BRI, NY, UC, US), Parks 20577 (uc); slopes of Mt. Lomalangi, Gillespie 3388 (BISH, holotyp e of Abramsia trichotoma; BISH, GH, UC, isotypes), 3686 (BISH, UC, US); Mt. Lomalangi, Gillespie 4065 (BIsH, GH, UC), 4364 (BISH, UC), Gresnaod 872 (a, K, UC), Tothill 266 (kK), 267 (Kk); hills E. of Nandala Creek, ca.3 mi. S. of Nandarivatu, Smith 5935 (a, BISH, BRI, K, NY, US); Navai Ranges, Fiji Dept. Agr. 2319 (B. E. Parham) (a); west and south slopes of Mt. Tomanivi 1980] DARWIN, AIROSPERMA 103 (Mt. Victoria), Smith 5743 (a, BISH, BRI, K, US); Mt. Tomanivi (Mt. Victoria), Degener et al. 32080 (BisH), 32086 (BISH). NANDRONGA AND Navosa: northern portion Rairaimatuku Plateau between Nandrau and Nanga, Smith 5517 (a, BISH, BRI, K, NY, US). SERUA: hills W. of Waivunu Creek between Ngaloa and Korovou, Smith 9473 (Bisu, US); Ngaloa Nature Reserve, Fiji Dept. Agr. 16592 (BisH). NaAmosi: near summit of Mt. Naitarandamu, Gillespie 3315 (BISH, NY, Creek, Smith 8634 (BISH, GH, NY, UC, US); vicinity of Namosi, Gillespie 2652 (BisH), Parks 20275 (pis, uc, us), Seemann 250 (cu, kK); Wui Voma, Fiji Dept. Agr. 11680 (sisH, BRI). Naitasiri: Wainimala Valley, Rarandawai to Nairairai- kinasavu, Wainisavulevu Creek, St. John 18285 (a, BISH); Waimanu region, southeastern Nasle, Fiji Dept. Agr. 15427 (Bri, Mass); track to Mendrausuthu Range, Fiji Dept. Agr. 15025 (a, Bri, uc). Taitevu: Ndakuivuna, Fiji Dept Agr. 11016 (BisH). Viti Levu, without further locality, Tothill 268 (k). Fu, without further locality, Fiji Dept. Agr. L. 13370 (Berry) (pri). 6. Airosperma vanuense S. Darwin, sp. nov. Ficure 6. Frutex (vel arbor gracilis?) praeter inflorescentiam et partes juveniles plus minusve glaber; ramulis subteretibus apicem versus 1-2 mm. diametro fuscis; stipulis maturis persistentibus membranaceis ovatis vel late deltoideis ad 1.5 mm. longis apiculatis sed saepissime demum bifidis vel erosis, extus puberulis pilis minutis et dispersis intus pubescentiis densioribus et ad basem cum aliquot glandibus gracilibus circiter 0.2 x 0.1 mm.; petiolis gracilibus semi- teretibus vel aliquantum canaliculatis 5-40 mm. longis ad medium circiter | mm. latis glabris; foliorum laminis membranaceis ellipticis vel oblongis vel aliquantum oblanceolatis 5-15 x 2-8 cm. apice acutis saepe obtuse vel acutissime acuminatis basi anguste cuneatis et in petiolum decurrentibus, integris, supra glabris vel pilis minutis appressis et dispersis subtus pilis frequentioribus in costa et nervis, costa conspicua supra prominula et canali- culata subtus elevata, nervis secundarilis utrinsecus 8-9 late patentibus et Ubnue prominulis, coe eras et rete venularum utrinque subplanis; puberulis pilis dispersis appressis albis vel stramineis ad 0.1 mm. tone cymosis plerumque basi trichotomis ramificatione opposita inordinatescenti, sub anthesi 1.5-3.5 x 2-6 cm. et 30-50-floribus, bracteis subulatis vel linearibus ad | mm. ee apice acutis, pedicellis 1-3 mm. longis (sed sub fructu multo longioribus?); calycis limbo ad 1.5 mm. lato, calycis parte discreta ad 1 mm. longa cupulata et minute 5-denticulata, extus minute puberula intus glabra; corolla hypocraterimorpha vel anguste infundibulari, limbo 4.5-8 mm. lato, tubo 5-8 mm. longo ad medium 0.8-1.2 mm. lato extus glabro vel pilis dispersis intus glabro sed sub fauce hirsuto-villoso pilis albis laxis ad 1 mm. longis, corollae lobis 5 in aestivatione contortis ovatis 2-3.5 x 1-2 mm. apice acutis basi subcordatis intus minute et dense puberulis vel glabris; staminibus 5 sub corollae fauce insertis, filamentis circiter 0.5 mm. longis, antheris plus minusve dorsifixis clavatis circiter 2 x 0.8 mm. apice rotundatis vel emarginatis basi loculis in filamentum decurrentibus; ovario (hypanthio) subgloboso vel ellipsoideo 0.5-1 x 0.5 mm. minute puberulo biloculari, ovulis in quoque loculo solitariis et prope verticem loculi affixis, juxta dissepimentum aliquantum complanatis; 104 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 disco circiter 0.5 mm. alto in centro depresso; stylo ad 4 mm. longo filiformi vel subclavato glabro, stigmate bifido in alabastro ad 0.5 mm. longo glabro, ultra corollae tubum vix exserto. Fructus jam ignoti vel fortasse ad speciem praesentem non referentes. Airosperma vanuense is thus far known with certainty from only two collections from the island of Vanua Levu, Fiji. The type collection, cited below, was gathered in the province of Mathuata, near the Wainunu-Ndreketi Divide, May 17, 1934, from a shrub 3 meters tall growing in a dense forest at an altitude of 200-300 meters. The flowers are nearly mature in the holotype. I am able to separate Airosperma vanuense from A. trichotomum on the basis of two floral characters. In the present species te corolla tubes are 5-8 mm. long, vs. only |-2 mm. in A. trichotomum. Corolla tube length associated with various pollination adaptations in some species. However, in this instance the tube length is correlated with differences in pubescence: the corolla throats in A. vanuense are densely villose-hirsute with white hairs to 1 mm. long, while those of A. trichotomum are glabrous within. Pubescent corolla throats, as far as I am able to determine, are found in all New Guinean species of Airosperma, but none of those species approaches A. vanuense in the length of the corolla tube. Since the diagnostic features of Airosperma vanuense are entirely floral characters, ut 1S not possible for me to assign certain fruiting collections material of Airosperma from Viti Levu is all readily assignable to 4A. trichotomum, and the fruiting specimens from the same island probably represent the same species. However, only two flowering collections are currently known from Vania Levu; as a result, it is possible that some fruiting collections of Airosperma from that island may represent A. trichoto- mum rather than A. vanuense. Such fruiting collections, for the present probably best regarded as ‘“‘aff. vanuense,’’ include the following: Mt. Vatunivuamonde, Savu Savu Bay region, Degener & Ordonez 13964 (a, BISH, NY, UC, US); vicinity of Drayton Peak, Bierhorst F154 (Mass); Taveuni, Mt. Manuka, Smith 796 (BIsH, GH, US), 52/4 (BISH, GH, NY, Uc, us). More field work and additional collections (especially from Vanua Levu) are required before the ranges of A. trichotomum and A. vanuense can be determined with accuracy. SPECIMENS EXAMINED. Fiji. Vanua Levu. Matuuata: Wainunu-Ndreketi Divide, Smith 1849 (Gu, holotype; BisH, Ny, UC, US, isotypes). THAKAUNDROVE: Navavau, Fiji Dept. Agr. 16046 (Bis). ACKNOWLEDGMENTS Special thanks are due to Dr. Albert C. Smith, whose detailed notes on the Fijian species were an invaluable aid. Dr. Edith Raadts kindly provided information about specimens at the Botanischer Garten und Botanisches 1980] DARWIN, AIROSPERMA 105 Museum, Berlin- ease ZS OTUUSHALOUS of ne following herbaria have kindly made herbar fter Holmgren & Keuken, 1974): A, BISH, BM, BRI, CANB, GH, _ LAE, MASS, NY, UC, US, WRSL. The cost of scanning electron micrographs was generously underwritten by the Gray Herbarium of Harvard University. REFERENCES Darwin, S. P. 1977. The genus Mastixiodendron (Rubiaceae). Jour. Arnold Arb. 58: 349-381. 979. A yee of the indigenous genera of Pacific Rubiaceae. Allertonia 2: 1-4 Hocmcren, P. K. a Keuken. 1974, Index herbariorum. Part I. The herbaria of the world. ed. 6. Reg. Veg. 92: 1-397 Moore, S. 1927. New Rubiaceae collected ‘by Mr. L. J. Brass in British New Guinea. Jour. Bot. London 65: 241-247. Ripspace, C. E., R. C. BAKHUIZEN VAN DEN Brink, Jr., & J. KoEK-Noorman. 1972. Notes on New Guinea Rubiaceae. Versteegia and Maschalodesme. Blumea 20: 339-348. ScHuMANN, K. M., & C. A. G. Lautersacu. 1900 [‘'1901’’]. Die Flora der eau Schutzgebiete in der Sudsee. xvi + 613 pp. Gebriider Born- raeger, Leipzig. SMITH, i C. 1945. Studies of Pacific ee plants, IV. Notes on Fijian flowering plants. Jour. Arnold Arb. 26: 97-110. 1979. Flora Vitiensis Nova. Vol. 1. xv + 495 pp. Pacific Tropical Botanical Garden, Lawai, Hawaii. VaLeTon, T. 1912. Rubiaceae. Jn: H. A. Lorentz, ed., Nova Guinea 8(4): TIS- 119s DEPARTMENT OF aa TULANE UNIVER New ORLEANS, es 70118 HU, ELEUTHEROCOCCUS VS. ACANTHOPANAX 107 ELEUTHEROCOCCUS VS. ACANTHOPANAX SHIU YING Hu IN RESPONSE TO a growing interest in the use of ginseng, chemists and physicians have investigated the chemistry and the physiological activities of crude drugs prepared from the roots and bark of Panax ginseng C. A. Meyer and other araliaceous plants. Included in these studies have been the Chinese drugs known as Wu-chia-p’i, which are prepared from the bark of different species of Eleutherococcus. The investigations have resulted in the isolation of sitosterol glycosides from Eleutherococcus senticosus (Rupr. & Maxim.) Maxim., and in the discovery of similar physiological a rag induced by these glycosides and the ginsenosides of P. ginseng (Brekham, 1969; Anonymous, 1976). Because of these similarities, extracts from ee of Eleutherococcus have been used as substitutes for ginseng. Communicating the results of the chemical and physiological studies has often proved difficult and confusing since species of Eleutherococcus have sometimes been included in the genus Acanthopanax, and the nomenclature has been unstable. A summary of the taxonomy of these two genera and necessary nomenclatural changes are presented here in order to clarify this situation and to provide a more stable and less confusing nomenclature. A survey of the pertinent literature indicates that Eleutherococcus Maxim. (typified by E. senticosus) is the earliest validly published name for the genus that some authors have called Acanthopanax (Decaisne & Planchon) Miquel (typified by A. spinosus (L. f.) Miquel). Decaisne and Planchon (1854) used Acanthopanax as a subgenus of Panax L., and later Miquel (1863) raised the subgenus to generic level. Harms (1894) combined A canthopanax and Eleutherococcus, but he used Acanthopanax as the generic name and recognized Eleutherococcus at sectional rank. Harms’s treatment has been followed by Rehder (1940, 1949), Li (1942), Bere satey authors, and floristic botanists in China and Japan. However, in 1924 Nakai recognized both Acanthopanax and Eleutherococcus as distinct genera, and his treatment has been adopted by Poyarkova (1973 The characters utilized by Poyarkova to separate the two genera are ‘‘ovary 2-locular, styles 2; fruit with 2 stones; petioles glabrous or scarcely pubescent, articulate with rachis’? for Acanthopanax, and ‘‘ovary 5-locular, styles 5, fruit with 5S stones; petioles not articulate, densely pubescent’’ for Eleuthero- coccus. These characters may hold for distinguishing the two species (one placed in each genus) that occur in the relatively limited area of the Soviet Far East, which constitutes the periphery of the generic range, but they are of no generic value when species from over the entire geographic range are considered. Even for the small number of Japanese species, Nakai (1924) © President and Fellows of Harvard rome e, 1980. Journal of the Arnold Arboretum 61: -I11. January, 1980. 108 JOURNAL OF THE ARNOLD ARBORETUM [voc. 61 remarked that the ‘‘characters distinguishing Eleutherococcus from Acanthopanax are neither the cohesiveness of the styles nor the presence of articulation in the flowers. . . . The seeming articulation eal in the dried specimens by the contraction of tissue... [but is alse articulation.’’ Nakai separated the genera by utilizing the cece of ‘‘styles and the number of cells in the ovary as well as the shape of the pyrenes.”’ He attributed 2 styles, 2 or 3 ovary cells, and flat pyrenes with roundish ventral sides to Acanthopanax, and 5 styles, 3 to 6 ovary cells, and pyrenes with an acute or somewhat acute ventral side to Eleutherococcus. Like the characters used by Poyarkova, Nakai’s characters do not hold when all of the taxa attributed to the two genera are examined. In all species the endocarps mature separately into hard shells, each covering an individual seed to form a pyrene. The shape of the ventral (adaxial) side of the pyrene depends upon the number of ovary cells. The variation in the number of cells in the ovaries (and, in turn, in the number of pyrenes) is correlated with the number of styles—observations of one flowering branch of Eleuthero- coccus trifoliatus (L.) S. Y. Hu has shown some flowers with 2 or 3 styles (these united up to the middle) and others with 6 (these free to the base). hen all of the species attributed to the two genera are examined, there is not one constant character that can be used to separate Eleutherococcus from Acanthopanax. The genus is a natural one consisting of spinose shrubs with compound leaves, sessile or pedicellate flowers arranged in simple, solitary or racemose umbels or heads, and flowers with 2- to 6-celled ovaries. Eleutherococcus (from the Greek eleuthero, free, and kokkos, seed, in allusion to the separated pyrenes) is the correct generic name. Rehder’s treatment (1940, 1949) of the genus and its sections is followed here with two exceptions. The unarmed, arboreus species with trifoliolate leaves, corymbose-paniculate umbels, and drupes consisting of 2 dorsiventrally flattened pyrenes are placed in Evodiopanax Nakai (see Ohwi, 1965). Another unarmed, arboreus species with (3-, 4-, or) 5-foliolate leaves and loosely arranged corymbose-paniculate umbels is left in Kalopanax (K. sciadophyl- loides (Franchet & Savatier) Harms). A total of fifteen species and seven varieties from China, Korea, and Japan are transferred below from Acantho- panax to Eleutherococcus, and one variety from Acanthopanax to Evodiopan- ax. Important literature is cited to provide access to descriptions of the taxa as well as more complete references. sia aoe Maxim. Mém. Acad. Sci. St.-Pétersb. Sav. Etr. 9: 132. 1859; m & Hooker f. Gen. Pl. 1: 941. 1867; Nakai, Jour. Arnold re 5: ‘9. 1924, Fl. Sylv. Kor. 16: 26. 1927. Panax subgen. Acanthopanax Decaisne & Planchon, Revue Hort. 1854: 105. 1854. Ht es (Decaisne & Planchon) Miquel, Ann. Mus. Bot. Lugd.-Bat. 1: ae 676. 1940, Bibliogr. 492. 1949; Li, Sargentia 2: 69. 1942; Ohwi, Fl. Japan (Eng. ed.) 664. 1965; Anonymous, Ic. Corm. Sin. 2: 1035. 1972. 1980] HU, ELEUTHEROCOCCUS VS. ACANTHOPANAX 109 Acanthopanax sect. Eleutherococcus (Maxim.) Harms in Engler, Nat. Pflanzenfam. III. 8: 49. 1894. Eleutherococcus divaricatus (Sieb. & Zucc.) S. Y. Hu, comb. nov. Panax divaricatus Sieb. & Zucc. Abh. Bayer. Akad. Math.-Phys. Kl. 4(2): 200. 1845 (as divaricatum). Acanthopanax divaricatus (Sieb. & rie Seemann, Jour. Bot. London 5: 239. 1867; Li, Sargentia 2: 76. 1842 Eleutherococcus giraldii (Harms) Nakai var. pilosulus (Rehder) S. Y. Hu, comb. nov. Acanthopanax giraldii Harms var. pilosulus Rehder, Jour. Arnold Arb. 9: 99. 1928; Li, Sargentia 2: 81. 1942. Eleutherococcus gracilistylus (W. W. Sm.) S. Y. Hu, comb. nov. Acanthopanax gracilistylus W. W. Sm. Notes Bot. Gard. Edinburgh 10: 6. 1917. Eleutherococcus gracilistylus var. pubescens (Pampanini) S. Y. Hu, comb. nov. Acanthopanax spinosus (L. f.) var. pubescens Pampanini, Nuovo Giorn. Bot. Ital. II. 17: 678. 1911. Acanthopanax gracilistylus W. W. Sm. var. pubescens (Pampanini) Li, Sargentia 2: 85. 1942. Eleutherococcus henryi Oliver var. faberi (Harms) S. Y. Hu, comb. nov. Acanthopanax henryi (Oliver) Harms var. faberi Harms, Mitt. Deutsch. Dendrol. Ges. 27: 12. 1918; Li, Sargentia 2: 75. 1942. Eleutherococcus lasiogyne (Harms) S. Y. Hu, comb. nov. Acanthopanax lasiogyne Harms in Sargent, Pl. Wils. 2: 563. 1916; Li, Sargentia 2: 78. 1942. Eleutherococcus nodiflorus (Dunn) S. Y. Hu, comb. no Acanthopanax nodiflorus Dunn, Jour. Bot. London en 199. 1909. Acanthopanax gracilistylus W. W. Sm. var. nodiflorus ae Li, Sargentia 2: 86. 1942. Eleutherococcus phanerophlebius (Merr.) S. Y. Hu, comb. nov. Acanthopanax phanerophlebius Merr. Sunyatsenia 2: 12. pl. 6. 1934. Eleutherococcus rufinervis (Nakai) S. Y. Hu, comb. Acanthopanax rufinervis Nakai, Fl. Sylv. Kor. ms eH 1927 (as rufinerve). Eleutherococcus seoulensis (Nakai) S. Y. Hu, comb. nov. Acanthopanax seoulensis Nakai, Fl. Sylv. Kor. 16: 24. fig. 4. 1927 (as seoulense Eleutherococcus sessiliflorus (Rupr. & Maxim.) S. Y. Hu, comb. nov. Panax sessiliflorus Rupr. & Maxim. Bull. Phys.-Math. Acad. Sci. St.- Pétersb. 15: 133. 1857. 110 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Acanthopanax sessiliflorus (Rupr. & Maxim.) Seemann, Jour. Bot. London 5: 239. 1867; Li, Sargentia 2: 77. 1942. Eleutherococcus sessiliflorus var. parviceps (Rehder) S. Y. Hu, comb. nov. Acanthopanax sessiliflorus (Rupr. & Maxim.) Seemann var. Parviceps Rehder, Mitt. Deutsch. Dendrol. Ges. 21: 129. 1912. Eleutherococcus setulosus (Franchet) S. Y. Hu, comb. nov. Acanthopanax setulosus Franchet, Nouv. Arch. Mus. Hist. Nat. Paris II. 8: 249. 1885; Li, Sargentia 2: 82. 1942 Eleutherococcus sieboldianus (Makino) Koidzumi f. variegatus (Rehder) S. Y. u b. nov Acanthopanax neni Marchal var. variegatum Hort. ex Rehder in Bailey, Cycl. Am. Hort. 1: 11. 1900. Pee mere sieboldi lanus Makino f. variegatus Rehder, Jour. Arnold Arb. 7: 243. 1926; Bibliogr. 493. 1949, Eleutherococcus spinifolia (Merr.) S. Y. Hu, comb. nov. Acanthopanax spinifolia Merr. Philip. Jour. Sci. 15: 249. 1919, Eleutherococcus spinosus (L. f.) S. Y. Hu, comb. nov. Panax spinosum L. f. Suppl. Pl. 441. 1781. Acanthopanax spinosus (L. f.) Miquel in Ann. Mus. Bot. Ludg.-Bat. 1: 10. 1863 (as spinosum), p. p., quoad basionym, exclud. descript.; Rehder, Man. 678. 1940, Bibliogr. 493. 1949. Eleutherococcus ee (Harms) Nakai f. angustissimus (Rehder) S. Y. , comb. nov Pere ee eeuapilis Harms f. angustissimus Rehder, Jour. Arnold Arb. 9: 99. 1928; Li, Sargentia 2: 82. 1942. a eeu (Harms) Nakai f. dilatatus (Rehder) S. Y. Hu, comb. n Mesa anne senoeivlis Harms . ‘aa Rehder, Jour. Arnold Arb. 13: 339. 1932; Li, Sargentia 2: 82. Eleutherococcus trifoliatus (L.) S. Y. Hu, comb. nov. Zanthoxylum trifoliatum L. Sp. Pl. 270. 1753. Acanthopanax trifoliatus Voss, Vilmor. Blumengartn. 1: 406. 1894; Merr. Philip. Jour. Sci. (Suppl): 217. 1906; Li, Sargentia 2: 86. 1942: Rehder Bibliogr. 493. 1949 5 Eleutherococcus villosulus (Harms) S. Y. Hu, comb. nov. Acanthopanax villosulus Harms in Sargent, Pl. Wils. 2: 562. 1916. A canthopanax gracilistylus W. W. Sm. var. villosulus (Harms) Li, Sargentia 2: 85. 1942. Eleutherococcus wardii (W. W. Sm.) S. Y. Hu, comb. nov. Acanthopanax wardii W. W. Sm. Notes . Gard. 7 eras 10: 7. 1917; Li, Sargentia 2: 88. 1942; Rehder, are 493. 1949. 1980] HU, ELEUTHEROCOCCUS VS. ACANTHOPANAX 111 Acanthopanax ternatus Rehder, Jour. Arnold Arb. 2: 124. 1924; Man. ed. 2. 678. 1940. Eleutherococcus yui (Li) S. Y. Hu, comb. nov. Acanthopanax yui Li, Sargentia 2: 79. 1942. Evodiopanax evodiifolius (Franchet) Nakai var. gracilis (W. W. Sm.) S. x b nov. Acanthopanax evodiifolius var. gracilis W. W. Sm. in Notes Bot. Gard. Edinburgh 10: 6. 1917 (as evodiaefolius). LITERATURE CITED Anonymous. 1972. Iconographia cormophytorum Sinicorum. Vol. II. + 312 pp. Institutum Botanicum, Academia Sinica, Peking. eralovanas 1035-1038. . 1976. Chung ts’ao yao hsiieh. [Chinese pharmacognosy.] Vol. II. 2 = 897 pp. College of Pharmacology, Nankin BrekuaM, I. I., & I. V. DarpyMov. 1969. Pharmacological Lala of glycosides from ginseng and Eleutherococcus. Lloydia 32: 46-51. DEcAISNE, J., J. E. Prancuon. 1854. Esquisse d’une Beale des Araliacées. Revue Hort. IV. 3: 104-109. Harms, H. 1894. Araliaceae. Nat. ‘see III. 8: 1-62. [Acanthopanax, 49-5 1918. Ubersicht iiber die Arten der Gattung Acanthopanax. Mitt. Deutsch. Dendrol. Ges. 27: 1-39. pls. 1-8. Li, H. L. 1942. The Araliaceae of China. Sargentia 2: 1-134. Maximowicz, C. J. 1859. Primitiae florae Amurensis. Mém. Acad. Sci. St.-Pe- rsb. Sav. Etr. 9: 1-504. pls. 1-10; 1 map. [Eleutherococcus, 132.] Naka, T. 1924. Araliaceae imperii Japonici. Jour. Arnold Arb. 5: 1-36. —___—., 1927. Flora sylvatica Koreana 16: 1-92. OHwI, i 1965. Flora of Japan. (English edition; F. G. Mever & E. H. WaLke eds.) 1067 pp. Smithsonian Institution, Washington, D. Gc. erracene 661-667. ] Povarkova, A. I. 1973. Araliaceae. (Translated by R. Lavoort.) Fl. USSR 16: 6. 3-3 Renper, A. 1940. Manual of cultivated trees and shrubs. ed. 2. xxx + 996 p. Macmillan, New Yor . 1949. Bibliography of cultivated trees and shrubs. xl + 825 pp. Arnold Arbo retum, Jamaica Plain, Massachusetts. Ruprecut, F. J. 1956. Die ersten botanischen Nachrichten tiber das Amurland. Erste Abtheilung. Bull. Phys.-Math. Acad. Sci. St.-Pétersb. 15: 120-144. SEEMANN, B. 1864-68. Revision of the natural order Hederaceae. Jour. Bot. London 2: 289-309. 9 figs. 1864, 3: 173-181. 1865; 5: 236- 239. 1867; 6: 52-58, 129-142, 161-165. pls. 79, 80. 1868 ARNOLD ARBORETUM HarvarD UNIVERSITY CAMBRIDGE, MassacHusetTts 02138 HUNZIKER, CESTRUM 113 A NEW SPECIES OF CESTRUM (SOLANACEAE) FROM SOUTHERN COLOMBIA ARMANDO T. HUNZIKER DuRING A BRIEF VisiT to the National Herbarium of Colombia, Bogota, m attention was drawn by the curious morphology of an unnamed collection of Cestrum. After a thorough analysis of its peculiarities, I have come to the conclusion that it represents an undescribed species. Its essential charac- teristics are noted in the following description. Cestrum morae' A. Hunziker, sp. nov. Ficure |. Arbor cauliflorus 5 m. altus. Folia magna, glaberrima, manifeste petiolata; laminae ellipticae vel oblongo-ellipticae ad apicem acuminatae vel breviter cuspidatae, basi + obtusiusculae, 30-34 cm. longae et 19.3-19.8 cm. latae, nervo mediano supra tenuiimy subtus valde prominente, nervis lateralibus utrinque ca. 20 adscendentibus supra impressis subtus valde eminentibus et nervis secundarlis numerosis, utrinque bene conspicuis dense reticulatis; petioli suberosi 44.7 cm. longi. Inflorescentiae laxae 20-28 florae; axes glabri 14.5—26 cm. longi, fertiles in tertia vel octava parte distali; bracteae lineares, angustae, (6.5—)11—15(-18) mm. longae, (0.25—)0.5—0.6(-1) mm. latae. Calyx 4.5-5.8 mm. longus et ca. 2 mm. latus, extus pilis minutissimis argentatis praeditus. Corolla extus glabra, alba, 37-39 mm. longa; tubus 31-32 mm. longus; lobi 6-7 mm. longi. Stamina aequalia ca. 30 mm. longae; filamenta ad ca. 4/5 longitudine tubo adnata (24-25 mm.), 4-7 mm. longe libera, ad insertionem non dentata et non gibbosa, valde pilosa; pars adnata superne dense pilosa, ceterum parcissime puberula; antherae parvae, orbiculatae, ca. 0.6-0.7 mm. longae. Ovarium diminutum, oe glabrum, ca. 0.5 mm. longum et ca. 1 mm. latum, stipite (ca. 0.8 mm. altus) brevior; stylus filiformis ca. 28-29 mm. ongus, apice minutissime See stamina superans; stigma exsertum. Baccae non suppetunt. Type COLLECTION. Colombia, Dpto. Narino, Municipio de Altaquer, entre Junin y Buenavista, 700 m. alt., L. E. Mora 4278, 21 Nov. 1967 (cot 113604, two leaves and one inflorescence; cot 113605, two inflorescences). OsseRVATIONS. A distinctive species, unique in its cauliflory, with inflores- cence axes 14.5-26 cm. long. Its large leaves and long corollas (to almost 4 cm.) are two additional features not frequently encountered in the genus. 'The specific epithet honors the collector of the only specimen known up to this me, Professor Luis Eduardo Mora, presently Director of the Botany ee Tennuie de Ciencias Naturales, Universidad Nacional, Bogota, Colombia © President and Fellows of Harvard ae 1980. Journal of the Arnold Arboretum 61: 113-115. January, 1980. 114 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Ficure 1, Cestrum morae: A, inflorescence, x 0.73; B, leaf, upper surface, x 0.4; C, flower, longitudinal section, x 2.7; D, gynoecium, basal portion, x 21; E, calyx, articulation with axis, x 7.5; F, flower, upper part, x 4.3. 1980] HUNZIKER, CESTRUM ba) The most closely related species appears to be Cestrum diversifolium Francey, from Colombia (Antioquia) and Ecuador;’ in addition to lacking cauliflory and having comparatively small leaves (less than 5 cm. long), this particular species differs in its somewhat smaller flowers with ovoid ovaries, and in its gibbous filaments that are slightly indented at the point of insertion on the corolla tube. ACKNOWLEDGMENTS The author wishes to thank the authorities of the Instituto Colombiano de Ciencias Naturales (Bogota) for the facilities made available to him, and for the loan of an important set of specimens; furthermore, he gratefully acknowledges the unfailing assistance of Dr. Gustavo Lozano during his stay at the institution. Thanks are due also to Dr. Bernice G. Schubert for reviewing the manuscript, and to Mrs. N. M. de Flury for her work on the illustration. BIBLIOGRAPHY D’Arcy, W. G. 1974. Solanaceae. Jn: R. E. Woopson, Jr., & R. W. ScHerRy, eds., Flora of Panama. Ann. Missouri Bot. Gard. 60: 573-780. Francey, P. 1935. Monographie du genre Cestrum L. Candollea 6: 46-398. 1936. Monographie du genre Cestrum L. Partie II. [bid. 7: 1-132. Chon. J., & P. C. Stanpbiey. 1974. “ane In: Flora of Guatemala. Fieldiana Bot. eee BOS. ba 2)21 Macpsripb_e, J. F. 1962. Solanaceae. Jn: ae of Peru. Publ. Field Mus. Bot. 13(V-B, no. ‘* fe 267. Museo Botanico; CoNnIcET CASILLA DE Correo 495 5000 CorpoBa, ARGENTINA *Francey, 1935, p. 321. SIAL SERVICE STATEMENT OF OWNERSHIP Beste een AND CIRCULATION utred by 39 U.S. ' 2, OATE OF FILING Journat of phe Arnold Arboretun 0) 6) > /13/79 FREQUENCY OF ISSUE Ta. NO. OF ESP’ Gb] 8. ANNUAL SUBSCRIPTION | een | ANNUALLY mn PRICE $25 00 LOCATIO City, County 22 pivindte Avenue, Wore ae Massachusetts 02138 5. LOCATION OF 22 Divinity Avenue, Cambridge, Massachusett MES AND COMPLETE ADDRESSES OF PUBLISHER, EDITOR PUBLISHER ee re “Addresa) Arnold Arboretum of Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts Stephen A. Spongberg, 22 Divinity Avenue, Cambridge, Massachusetts 02138 id Elizabeth B. Schmidt, 22 Divinity Avenue, Cambridge, Massachusetts 02138 > > Be > Led, ; h en 7: , , L- ri Th if } } } } } wrers must be given f. ih I i '! i , 1 fii its sai Pan o idress ; ll } f hy Hl lual h eisinie is published ists ; A : ADORESS 2 Divinity Avenue NAME | Arnold Arboretum of Harvard University | 22 | Cambridge, Massachusetts 02138 | 4 1 PERCENT OR MORE OF 8. MORTGAGEES, TOTAL AMOUNT OF BONDS so state) NAME ADDRESS Cambridge, Massachusetts 02138 President and Fellows of Harvard University | | | TO MAIL AT SPECIAL RATES (Section 132.122, PSM) NONPROFIT The purpose, function HAVE NOT CHANGED DURING (Uf changed, PRECEDING I2 MONTHS PRECEDING 12 MONTHS with this statement.) AVERAGE NO, COPIES EACH ACTUAL NO. COPIES OF SINGLE 10. EXTENT AND NATURE OF IR LATION ISSUE OURING PRECEDING ISSUE PUBLISHED NEAREST TO 12 MONTHS FILING DATE A. TOTAL NO Ane Qn, 800 500 [B. PAIDCIRCULATION ——————— 1, SALES THROUGH STREET ~ are VENDORS AND COUNTER SALES none none . MAIL SUBSCRIPTIONS od m 2 2 L suBSC 706 708 TOTAL PAID CIRCUL 7 f d 10B2) ane at = {06 (08 D. FREE DISTRIBUTION BY MAIL SAMPLES, COMPLIMENTARY 3. 3 E, TOTAL San = [09 fli F. COPIES NOT DISTRIBUTED 1. een USE, LEFT OVER, UNACCOUNTED, SPOILED ont 89 TER PRINTING 2 ENTS F , none none G, TOTAL (Sum of E, Fi and 2 h - Qo in A) 00 ATURE AND TITLE OF EDITOR, PUBLISHER, BUSINESS I certify that the statements made by me ove are correct and complete MANAGER, Lede 6 dhomidt . ) z a T THE REGULAR RATES (Section 132.121, Postal Service Manual) 39.U.S. Cc ; this statute, y licatl In | 1 at the phased e S.C. 3626, SIGNATURE AND TITLE OF ee ISHER OR OWNER Pp: . . shorn 3526 (Page 1) * (See instructions on reverse) Aug. 1978 NOTICES Volume 61, number 2, of the Journal of the Arnold Arboretum will be published simultaneously with number 3 during the summer of 1980. Both issues will be exclusively devoted to P. F. Stevens’s ‘“A Revision of the Old World Species of Calophyllum L. (Gutti- ferae).”’ . Because of continually increasing production costs, the Editorial Committee of the Journal of the Arnold Arboretum is requesting authors to help defray publication expenses. Beginning with manu- scripts received and accepted for publication as of March, 1980, a page charge will be levied. Acceptance of manuscripts for the Journal will, however, continue to be based solely on appropriate- ness and scientific merit, not on an author’s ability to meet page costs. Information on page charges and general editorial policies can be obtained from the Managing Editor. Journal of the Arnold Arboretum a January, 1980 CONTENTS OF VOLUME 61, NUMBER | | Fruits and Seeds of the Brandon Lignite, V. Rutaceae. Bruce H. TIFFNEY Ne eee Pe I The Metasequoia Flora and Its Phytogeographic Sigsilieance. SuHiu Y1 ING PRU cat oe oa eee, hc ee eee Notes on Airosperma (Rubiaceae), with a New Species from Ee a STEVEN P. DARWIN ee issues Spa pe ee 9D. Eleutherococcus vs. ae Suu Yinc Hu Se gt ene Sena oe es 107 A New Species of Cestrum olen) from Southern Colombia. = ARMANDOGEHUNZIKER «0.0, 28 oe 13 Statement of Ownership; 3 ees gee AiG Volume 60, Number 4, including pages 403-542, was issued December V7 1979: : ae JOURNAL oF te ~ ARNOLD ARBORETUM HARVARD UNIVERSITY VOLUME 61 NUMBER 2 _K. S. Bawa - Printed at Edwards Brothers, — Ann Arbor, aoe mem : < of this BENUs. 8: — Ss. US ISSN 0004-2625 Soiienal of ee Se Arhoreiuia - Published quarterly in January, ape ys and October by the Arnold Arboretum, Harvard Univer rsity. - Subscription price $25. 00 per year. Subscriptions and remittances should be sent to Ms. E. B. ‘Schmidt, ‘Arnold _ Arboretum, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A _ Claims will not be accepted after six months from the date of i issue. POSTMASTER: send address changes to Ms. E. _ Schmidt, Arnold Arboretum, 22 Divinity Avenue, Cambridge, Macacieere 02138, U.S.A. Volumes esi ‘reprinted, and some back numbers of ne 52-56 are _ available from the Kraus Reprint SOEORS CR: Route 100, Millwood, New Yo tk 10546, USA EDITORIAL COMMITTEE S. A. ‘Spongberg, Editor CE Bs Schmidt, Managing Editor | P. S. Ashton : P. F. ‘Stevens = C-E. Wood, Jr. pte Ly eet 4 I ified = COVER: chome nearly 160 times) of a Fij ijian eraeate Calophyllum eucocarpu ACES form the basis of ue vas s stylized cover design. A row of trichomes is — in the device on the cover, while a cluster of eave appears on the offprin As in recent years, she designs, based on holot e material i in | the herbarium OE ~ _ the Arnold Arboretum, were ‘dra awn by Karen Stoutsenberger. E “Calophyllum: 5 oes (Guutiferae) isa f widesangig genus of etal forest: trees in bo th the New and Old Wo tlds, and some species are of considerable economic importance as sources of. valua e imber. Its use on the cover of this volume ae of the Journal of the Arnold Arboretu is appropriate since the ‘April a nd July numbers are devoted to P. F. sievenited Ss revision of the Old World ceeepresentalives _ Second-class postage paid at Boston, Massachusetts, and additional offices. JOURNAL OF THE ARNOLD ARBORETUM VoL. 61 ApRIL 1980 NuMBER 2 A REVISION OF THE OLD WORLD SPECIES OF CALOPHYLLUM (GUTTIFERAE) P. F. STEVENS WHEN I arriveD in Lae, Papua New Guinea, in May, 1970, as a junior and rather raw Forest Botanist, the question arose as to which economically mportant group of plants I should revise. M. J. E. Coode suggested the Guttiferae, although he later admitted that it was hardly the group to give to somebody with little taxonomic experience, or even to anyone with whom one may later wish to communicate in a civilized fashion. However, a revision of the Papuasian species of Calophyllum L., the genus of the Guttiferae with the most species attaining loggable size, duly appeared (Stevens, 1974a), and my interest in the genus as a whole was stimulated. When I moved to the Arnold Arboretum of Harvard University, I was able to extend my studies on the genus; the resulting revision of the Old World taxa is presented here. The relatively few New World species will shortly be studie Calophyllum, the largest genus in the Calophylloideae, can be readily recognized by its opposite leaves with closely and regularly parallel veins that alternate with latex canals; flowers that usually appear to be hermaphro- ditic; ovary with a single, basal, anatropous ovule; and fruit that is drupelike. These characters, apart from the second, are derived ones (although it should be noted that fruit structure in other Calophylloideae is poorly known), and Calophyllum is probably strictly monophyletic—i.e., it includes only (and all) the extant derivatives of a single lineage. Superficially, the species are all rather similar; once seen, the genus should not be subsequently misidentified. Nevertheless, there are over 200 names available for species in the Old World alone. Although the natural tendency is to work on other, more obviously tractable genera, careful study of taxa like Calophyllum 1 is needed to lay the foundations for future, more specifically evolution-orien ted studies that are, in turn, essential before evolution in tropical rain forests can be appreciated. The questions that I asked myself as I extended my studies were three. Is it or is it not possible to recognize taxa throughout the range of the genus in the Old World? If it is possible, how did the taxa that I had previously recognized in Papuasia relate to these? Are there relationships between the © President and Fellows of Harvard College, 1980. Journal of the Arnold Arboretum, 61: 117-424. April, 1980. 118 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 patterns of morphological variation in Calophyllum and ecological or other variables that suggest ideas relating to the evolutionary history of some of the species? In the course of the examination of about 7000 numbers of the genus from the Old World, as well as field work in Malesia in 1975/76, the questions were resolved in somewhat unexpected ways. Species, based on discontinuities Madagascar and Vietnam, in particular, is still poorly understood. No super- ined specific taxa, even informal ones, can be maintained. One hundred and seventy-nine species have been described. altho ough some are poorly known and are merely given a number; varieties are recognized in eight species. Although in a few cases eS aeibe dca narrow or conservative species limits have been adopted, field experience, as well as the distinctness of most taxa, suggest that the number of recognized taxa is not too high. There will be reductions, but more taxa remain to be described. Characters from all organ systems have been used in the Ses i a and preliminary observations made on lamina anatomy are mentioned in a number of cases. A few of the characters studied are of particular eh It was, perhaps, to be expected that a survey of the fruits of the genus would be of considerable systematic importance (see Smith & Darwin, 1974; Stevens, 1974a), but the variation in seedlings and young plants surpassed all expectations and provided m information of systematic significance. This variation, which is greater in the young plant than in the seedling, and greater in the seedling than in the germinating plant, is as extensive n occurs in the adult plant as well; this, too, is of considerable stematic importance and has rarely been reported before. An additional suite of highly important characters, totally ignored for the better part of a century, has been provided by a study of the often rather bizarre hair types. arr polyembryony has been discovered in a few taxa. Most of the taxa that I had recognized in Papuasia stood up to the closer and ie examination possible in this study; no Papuasian taxon has an unexpected range in West Malesia. Indeed, insofar as there is a major dividing line within Calophyllum in the Indo-Malesian region, it occurs between New Guinea and Borneo. The area from Sulawesi to Samoa forms one region while that from Borneo and the Philippines to India forms another Little that is definite can be said with regard either to the selbotive value of the characters observed or to the evolutionary history of the genus or any of its species. However, I have attempted to put our knowledge of the genus into biogeographic, ecological, and evolutionary frameworks, more in the hope of stimulating further work than in an attempt to answer specific questions. It is clear that many of the ane recognized do have particular ecological preferences and discrete and the kind and amount of variation in seedling and growth characters, in particula ar, may relate to both the current ecology of individual species and the process of diversification within the genus. The suspicion that apomixis occurs in some taxa opens o 1980] STEVENS, CALOPHYLLUM 119 another, possibly very important, line of inquiry. The importance of the question that is so apparent in the field—how several apparently similar species can grow in one place—is manifest to those interested in tropical ecology. The question remains unanswered, but it now can be phrased much more exactly. The present treatment can, however, be considered only provisional, even in the context of the recognition of morphologically definable species. The genus has been woefully undercollected in the past throughout much of its range, and we still do not know the basic morphology of many taxa, much less understand their ae variation | or details of their SF abation patterns. Study is still needed, of it in the field, and extensions of our knowledge of several suites ‘él aces especially those concerning growth and anatomy, are immediate desiderata; we must find out much more about the ecology and breeding systems of the species in the genus. Study of the other genera of the Calophylloideae is planned. With this, as well as with studies by S. W. Jones, J.-F. LeRoy, and N. K. B. Robson of other genera in the Guttiferae, variation in Calophyllum will be placed in a more general context, and phylogenetic analysis of this variation ma become possible. In the absence of this wider context, however, phylogenetic analysis is premature TAXONOMIC HISTORY Linnaeus included two species of Calophyllum, C. inophyllum' and C. began to be disentangled. Confusion over the correct use of the name still persists; C. calaba should be typified on the element from Sm Lanka in the protolog (Stevens, 1980b). Subsequent to those of Linnaeus, the first taxa still referable to Calophyllum were described by Willdenow (1811). The majority of taxa in India and Ceylon were recognized early by Wight (1839, 1840) and Thwaites (1858), while Choisy (1849, in a preprint of an article eventually appearing in 1851) described a number of taxa based on specimens from Wallich’s herbarium. Choisy’s work was not very satisfactory, and although Planchon and Triana (1862a) managed to disentangle the relationships between the names, specimens cited, and descriptions in Choisy’s treatment, the nomenclatural consequences of Choisy’s work still haunt workers on the genus. The other major early worker on Calophyllum was Miquel (1854, 1859, 1861), who described a number of taxa from western Malesia. Planchon and Triana (1862a; see discussion after Calophyllum teysmannii for publication date of their work) first revised the genus, and despite their evident trepidation in disentangling Choisy’s work and the problems caused by the poor material available to them, they more than succeeded in their ‘Authorities for Indo-Malesian taxa are given in the species accounts. 120 JOURNAL OF THE ARNOLD ARBORETUM [voL. 6] modest aim of facilitating the task of subsequent monographers. They described nine species from the Old World as new and recognized a total of 29 species from this region, not including doubtful or incompletely known taxa. Only two of the species they recognized are not maintained as formal taxa in this revision, but almost a third of the names that they used are nomenclaturally incorrect. Unfortunately, they could not incorporate some of the taxa described by Miquel in their revision, and subsequent workers have not examined the material on which Miquel based his names; not surprisingly, Miquel’s names are the earliest for three common Malesian species. After Planchon and Triana’s publication, there was little work of any note on the genus for almost thirty years. Anderson and Hooker described a number of Indian taxa (Anderson, 1874), while Pierre (1885) named and illustrated a number of taxa from mainland Southeast Asia. Vesque (1889, 1893) was the second and last person to revise the genus; he examined the anatomy and indumentum of many species (see Vesque, 1889, for illustrations), and in his contribution to Monographiae Phaneroga- marum (1893), he incorporated detailed descriptions of the hairs and lamina anatomy in the species descriptions. He recognized four sections—IJnophyl- lum Vesque, Microphyllum Vesque, Apetalum Vesque, and Hypodermata Vesque—in the Old World, in part delimited on tepal (‘‘sepal’’ plus ‘‘petal’’) number. The sections were subdivided informally on anatomical and/or morphological characters. Species from the New World were not assigned to sections. In the Old World Vesque recognized 35 species, with 17 additional species being less well known and not assigned to sections; his species limits basically followed those of Planchon and Triana. Like Clarke (1894), I do not know on what specimens Vesque (1893) based his anatomical work; it was Clearly not always or not only on those specimens designated by an exclamation mark in his publication. Subsequent to Vesque’s publications, work on Calophyllum has been local in scope. Henderson and Wyatt-Smith’s excellent study (1956) of the Malayan species is the most important in this time, although—despite the authors’ unrivaled field knowledge of the genus—they were careful to point out the provisional nature of their account. They found that bark and slash characters were often helpful in recognizing a species, that the leaves, although superfi- cially so similar, were often diagnostic of a species, and that a supraspecific classification based on tepal number was impossible to employ. They recog- nized 46 taxa, of which they described 20; 43 of these taxa are maintained here at some level, although often with different names. Whitmore (1973) basically followed Henderson and Wyatt-Smith’s work. There is little other recent work of note. Perrier de la Bathie (1951) wrote an account of the Madagascan species, while Maheshwari (1960) provided a somewhat cursory treatment of those from India. Pitard (1910) and Gagnepain (1943) dealt with the genus in the Vietnam-Cambodia area; unfortunately, the new names proposed by the latter are invalid since the descriptions are in French. Recent accounts of the members of the genus in Papuasia (Stevens, 1974a) and the South Pacific region (Smith & Darwin, 1974) paid considerable attention to the structure of the fruit for the first time; in neither were 1980] STEVENS, CALOPHYLLUM 121 sections recognized. Stevens (op. cit.) also gave brief details of the lamina anatomy of the Papuasian species and described the germination and young plant of some species. VARIATION IN THE GENUS AND ITS USE IN CLASSIFICATION GERMINATION AND YOUNG PLANT In the main part of this paper, accounts of germination, the seedling,’ and the growth of the young plant are given for 36 species (33 primarily Malesian species, two from Sri Lanka, and one from the Mascarenes), and details of some of these stages are recorded for several others. The initial indications of the taxonomic importance of these early stages of growth, initial stages (see below) can be distinguished by the use of such additional characters as leaf shape and size and the color of the young leaves—characters that are not discussed in detail here; as a result, most species are recognizable from a very early age. The variation found has helped to clarify some taxonomic problems, for example, the limits of Calophyllum rigidum and C. venulosum, the relationships between C. venulosum, C. grandiflorum, and C. waliense, botanic garden and in the wild. Some of .he characters apparent in these ow It will be realized, from reading the accounts, that for a number of species progeny of only a single tree has been observed. A further complicating factor is that in most cases the observations were made in the wild, with germination and the young plants occurring under a variety of conditions; these factors are discussed below. Despite these caveats, there seems to be little infraspecific variation in many characters of these early stages. The main exceptions are variation in the number of pairs of leaves in the seedling and in their disposition (rather less variation), as well as in the presence of scale leaves in both axillary and terminal innovations’ in the young plant. However, in Calophyllum soulattri, and perhaps also in C. tetrapterum, there is infraspecific variation both in germination and in the growth of the young plant; this situation requires more study. Interspecific variation in germination, the seedling, and the growth of the young plant can be accommodated in fifteen classes (Ficure 1, a—o; TABLE 1). The differences between some of the classes are clearly slight and may prove to be unimportant when more data accumulate. The seedlings all conform to the Heliciopsis type and subtype (de Vogel, 1980). >The seedling is defined here as the plantlet after germination when growth in length ceases for the first time. °An innovation is the extension growth produced between two successive periods of rest 122 JOURNAL OF THE ARNOLD ARBORETUM [voc. 61 Ficure 1. Germination, seedlings, and young plants. Capital letters denote germination types, lower case letters show young plant types (double letters denote older stages), the seedling types are unmarked. For further explanation, see text. 1980] STEVENS, CALOPHYLLUM 123 TaBLe 1. Species of Calophyllum known to have early growth conforming to the types shown in Figure Calophyllum sclerophyllum. Calophyilum calaba vat. bracteatum, C. grandiflorum, C. moonii. Calophyllum scriblitifolium. Calophyllum biflorum, C. euryphyllum, C. ferrugineum var. ee and var. oblongifolium, C. sundaicum, C. teysmannii var. teysman and var. ease C. tomentosum, C. wallichianum var wallichianum and var. i ra ie brassii, c oe (part), C. venulosum (only one seedling know Gato aie tiie robustum, C. rupicola ee Calophyllum gracilipes, C. recurvatu Calophyllum blancoi (2), C. obiguinervium (growth slow, but habitat dry), C. tetrapterum var. tetrapter Calophyllum rigidum (seedling ide 0.5-2.7 cm. long). Calophyllum alboramulum (seedling internodes ca. 1 cm. ), G; papuanum, C. pauciflorum, C. pisiferum (two or three pairs of seedling leaves), C. suberosum, ie oe ans oO in ee et et k. Calophyllum nodosum. 1. Calophyllum pulcherrimum. m. Calophyllum inophyllum, C. tacamahac n. Calophyllum soulattri (part), C. na C. waliense (seedling internode ca. 1.5 cm. lon 0. Calophyllum neo- -ebudicum, C. peekelii. GERMINATION. Germination in Calophyllum is cryptocotylar: the cotyledons remain more or less enclosed in the stone, and the hypocotyl is short. This germination type is likely to be constant throughout the genus. The radicle may emerge in as few as ten days after the fruit falls to the ground in species with thin stone walls, or in as much as three months in species with thicker walls. The radicle is at, or near, the base of the fruit and may emerge during germination in one of three ways. n the majority of species for which germination is known, the radicle breaks through the stone just to one side of the base (Ficure 1, A). In species with thick-walled stones, such as Calophyllum euryphyllum, the wall adjacent to the radicle is thinner than that elsewhere (Ficure 6, b), thus facilitating emergence of the radicle. Similar areas of thinner wall can be seen in other species with thick-walled stones (e.g., C. goniocarpum, C. trachycaule, and C. vexans), which presumably germinate in the same way. A few species with moderately thick to thick stone walls have a basal plug that is pushed out by the radicle during germination (Ficure 1, C). This plug can be seen both in a longitudinal section of the stone as oblique lines (Ficure 6, i) and in the surface view of the cleaned stone, usually as a circular area that is lighter in color than the rest of the stone and is sharply demarcated from it. Species with basal plugs do not necessarily have thicker stone walls than those without them; thus Calophyllum euryphyl- 124 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 lum, which lacks a plug, has much thicker walls than the variant of C. soulattri, which has one. In Calophyllum tacamahaca the basal plug is pushed out by the anaes but the adjacent wall of the stone is also broken (Ficure 1, B). Thi possibly due to the fact that the radicle faces further away from the ee of the fruit than is usual in the genus. saa Mia mupaner and aueanemeut ie the seedling’s leaves are of g stages are unlettered in Ficure 1). However, ‘there is some intraspecific variation in the number of pairs of seedling leaves: Calophyllum pisiferum has two or three pairs; C. papuanum has two, three, or four. In some species that have two pairs, the internode between these leaves is clearly shorter than subsequently developed internodes, and the four seedling leaves may appear to be verticillate. In some other species the two pairs of leaves are separated by well-developed internodes, while yet other species may be intermediate in this character, as is possibly the case in C. rigidum. Calophyllum teysmannii nearly always has two pairs of leaves pseudoverticillately arranged, but occasional seedlings have three or more pairs separated by well-developed internodes. The lowest pair(s) of seedling leaves may be small and soon fall off; this situation is especially common in taxa with three or more pairs. However, such small leaves cannot be confused with the scale leaves of species like Calophyllum pulcherrimum. Younc pLant. After a period when there is no growth, a bud on the seedling grows out and produces one or more pairs of leaves before growth stops again. There is considerable taxonomically useful interspecific variation at this stage also. The terminal bud is usually functional. The plant is erect, and the leaves are opposite; all leaves are fully expanded, each pair being separated by well-developed internodes. The single example (Calophyllum sclerophyllum) in which the young plant has alternate leaves needs confirmation. In several species the internodes of the young plants are short, often less than 1.5 cm. long; only after some time are internodes of greater length produced. The development of such short internodes is genotypically controlled in Calophyllum soulattri (Stevens, 1974) and probably also in C. brassii, C. gracilipes, C. grandiflorum, C. recurvatum, and C. venulosum, in all of these species each flush may consist of only a single pair of leaves, and the increase in height of the plant is slow. However, young plants of many species may appear to grow like this when occurring in unfavorable habitats, and the young plants of species with small fruits may initially produce rather short internodes. As a result, observation of this character is difficult. Species with either long or short internodes may have an initially more or less plagiotropic stem. The leaves tend to be arranged in two ranks along the stem because of twisting both in the internodal regions and of the petioles. Subsequent straightening of the axis generally occurs within 15 cm. of the stem apex 1980] STEVENS, CALOPHYLLUM 125 In those species in which the terminal bud does not function, growth is by the development of an axillary bud that first produces one or two pairs of scale leaves separated by short internodes, and then pairs of expanded leaves separated by longer internodes; this growth pattern is then repeated and the whole axillary shoot system is more or less plagiotropic. Although it is not clear that the axillary buds in young plants of Calophyllum pulcherrt- mum (in which the adult plant characteristically has such a growth pattern) always produce the first flush, functional terminal buds are absent in plants of slightly greater size on up to the adult. In Mesua nagassarium (Burman f.) Kostermans, also of the Guttiferae, the mature plant of which has a similar growth pattern to that of C. pulcherrimum, the terminal bud of the seedling is not functional, and growth is by development of axillary buds (pers. obs.). ARCHITECTURE Little is known about the developmental stages between the young plant and the flowering and fruiting adult plant, so the diversity of the genus in terms of architectural models (Hallé, Oldeman, & Tomlinson, 1978) is unclear. However, growth of all species appears to be discontinuous, and branching is rhythmic. Most species probably conform to Rauh’s model, in which the trunk and branches are orthotropic and the inflorescences are axillary. Species such as Calophyllum alboramulum differ only in having terminal inflorescences; they conform to Scarrone’s model. Those species in which the stem is initially plagiotropic appear to conform to two subtypes of Troll’s model, those that have a functional terminal bud (e.g., Calophyllum gracilipes) and those that do not (e.g., C. pulcherrimum). Hasit, TRUNK, AND BARK There are numerous characteristics of the trunk and bark that help one to distinguish between and recognize species in the field. Henderson and Wyatt-Smith (1956) have compiled a useful table of miscellaneous field data, including these characters, for the species from Malaya and Singapore. Species of Calophyllum are usually trees, often with a clean, straight bole (Ficure 2, a, e). Species such as C. nodosum (FIGURE 2, c) may form shrubs, but there is considerable infraspecific variation in habit. Some species, such as C. rufigemmatum, habitually reach a larger size than others. Observations on the way the branches are held are few, but there is clearly interesting variation. Species such as C. peekelii have erect branches, twigs, and leaves, while in species such as C. pulcherrimum and C. rigidum at least the smaller branches are pendulous. Throughout the genus, presence or absence of stilt roots, flying buttresses, or plank buttresses tends to be rather constant within a species. Knee roots, loop roots, or pneumatophores (Ficure 2, b) are uncommon, occurring only in swamp-dwelling taxa. Although the presence of such characters is often constant in a species, pneumatophores may be found in plants of C. soulattri growing in swamps, but do not occur in plants growing elsewhere—a not uncommon phenomenon. 126 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Ficure 2. Habit, trunk, and bark. a, Calophyllum teysmannii var. inophyl- loide; b, C. ardens (note pneumatophores); c, C. nodosum; d, e, C. aff. castaneum (d, note boat-shaped fissures). (Photographs, P. S. Ashton.) 1980} STEVENS, CALOPHYLLUM 127 The most useful bark characters include color and texture of the outer bark, color of the inner surface of the outer and of the under bark, and color, amount, opacity, and stickiness of the latex. Plants of many species of Calophyllum have distinctive, diamond- to boat-shaped fissures (FicureE 2, d), especially when young; the fissures may become deeper and confluent with age. The bark of other taxa may be smooth, smooth except for vertical lines of lenticels, or exfoliating in flakes or scales (Ficure 2, a). Unfortunately, many of these characters are qualitative. Trunks of Calo- phyllum roseocostatum and C. tetrapterum of abou t the same size have yellowish bark with discrete, diamond- to bon ee fissures, yet they are not identical; latex color confirms identification based on the subtle differences between the two. Furthermore, bark color and texture vary considerably with age in some taxa; thus the bark of C. teysmannii, although initially smooth, later becomes deeply furrowed, while the bark of such taxa as C. wallichianum var. incrassatum and especially C. collinum remains fairly smooth even in moderate-sized trees. The color of the latex in twigs and fruits often differs from that of the trunk, and this difference may be responsible for some of the infraspecific variation recorded for latex color. It is difficult to interpret other people’s field notes; hence, the description of the bark and trunk characters prefacing each species should be treated with caution. LEAF ARRANGEMENT AND SHOOT GROWTH In the arrangement of leaves and the growth of the shoots, there are numerous taxonomically important characters, practically all of which have been over- looked in earlier studies. The leaves are nearly always opposite and decussate, as is characteristic of most of the Guttiferae, but leaves of Calophyllum caledonicum may opposite, truly whorled in threes (cf. Planchon & Triana, 1862b), alternate, or adjacent (in pairs at a node, but not opposite). All these arrangements may occur on a single shoot, although the leaves are usually wholly decussate, or rarely wholly whorled. Sporadic variants with the leaves in whorls of three also occur elsewhere: I have seen seedlings of C. pulcherrimum with such whorled leaves. More important variation occurs in young plants of C. sclerophyllum, in which the leaves are borne singly and the stem is zigzag; the adult plant has decussate leaves (see also above). This observation needs confirmation since it is uncommon for any Guttiferae outside the Kiel- meyeroideae to have alternate leaves at any stage of their life history (see Engler, 1925; Willis, 1974). Exceptions occur in some West African species of Psorospermum Spach (Hypericoideae) such as P. alternifolium Hochr. N. K. B. Robson, pers. comm.) and perhaps in seedlings of Paramammea Leroy (Calophylloideae; Leroy, 1977). he internodes on each innovation are usually of similar length, and leaves at each node are fully expanded. However, there is consistent variation in internode length along each innovation and/or in the development of the leaves in some species. In a number of species the first pair(s) of leaves of the axillary innovations 128 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 are reduced to scales and are separated from the main stem and from each other by very short internodes that are usually less than 5 mm. long; the other pairs of leaves of the innovation are fully expanded and are separated Ith occur in a number of western Malesian species of Calophyllum, primarily in the C. garcinioides, C. pulcherrimum, and C. pyriforme groups. (The first and last of these groups have functional terminal buds; in the C. pulcherrimum group the terminal bud rarely functions, and growth is primarily by the development of axillary buds.) These scale leaves are caducous and are best recognized on herbarium specimens by the scars that they leave (Ficure 7, b); in species with stout stems and infrequent branching, they are difficult to observe. Infraspecific variation in the presence of scale leaves may occur (e.g., in Calophyllum macrocarpum, C. pisiferum, and C. tetrapterum). Scale leaves occur at least sometimes on axillary innovations of very young plants in several species (e.g., C. biflorum, C. rigidum, C. novoguineense, and C. tomentosum), but are lacking on adult plants of these species. They are found on terminal innovations of young plants of C. scriblitifolium, but whether or not they occur in mature plants is unknown; they are occasionally found on young plants of C. recurvatum but not in the adult plant. There are also the fairly general tendencies for the first internode of the innovation are needed, the presence of scale leaves and associated short internodes is an important taxonomic character and is very useful in identifying sterile material. The first-developed internode of axillary innovations in Calophyllum pervillei and C. milvum and its relatives (all from Madagascar), and in Calophyllum praetermissum (Borneo), is notably longer than the others; subsequent inter- nodes are approximately equal in length (Ficure 3, c). A similarly long ‘“‘basal’’ internode occurs in Calophyllum sp. 61* (Ficure 3, e), but this internode is not the first produced in the innovation. In other species the last internode of an innovation may be so reduced as to be almost absent (e.g., only 1.5 mm. long); the four leaves of the terminal group then appear to be pseudoverticil- late (C. verticillatum and C. chapelieri, both from Madagascar). Less extreme reduction occurs in such species as C. cuneifolium (Sri Lanka), C. obliquiner- vium (West Malesia), and C. ceriferum (Vietnam). The seedling of C. obliqui- nervium does not have pseudoverticillate leaves (see above), but details are unknown for the other species Some of the variation described above correlates with certain more dynamic aspects of shoot development that, unfortunately, are very poorly known. Although in many species of Calophyllum each innovation consists of several pairs of leaves (Ficure 3, a), some species, probably including all those with “Species with sea eat but no names, are not described formally; the number refers to the Species Account 1980] STEVENS, CALOPHYLLUM 129 Ficure 3. Shoot growth. Arrows denote start of most recent innovation. For explanation, see text well-marked horizontal lines at the nodes (Ficure 7, a; see also below), usually produce only one or two (rarely more) pairs of leaves per innovation (FIGURE 3, c). Such species include C. aureobrunnescens, C. calaba, C. euryphyllum, C. leleanii, C. moonii (young plant also), C. papuanum,° C. pauciflorum, pobeeancns on these species kindly confirmed by K. Damas and E. E. Henty (pers. c 130 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 C. persimile (young plant also), C. rufigemmatum, C. soulattri (young plant also), C. suberosum, C. teysmannii var. teysmannii and var. bursiculum, and C. vexans. It is interesting to note that although adult plants of C. teysmannii var. inophylloide produce many leaves each flush, this is not so in the young plant; plants of C. wallichianum var. incrassatum apparently show similar variation. The taxa that produce few leaves each flush seem to do so largely independent of the vigor of the plant. Hallé et al. (1978) distinguish two types of branch development, prolepsis and syllepsis, that are often correlated with the morphology of the shoot. Both types occur in Calophyllum, although there is no easily observable morphological correlation. At least sometimes in C. inophyllum and C. bra- siliense Camb., axillary shoots are sylleptic, developing without an obvious pause and also contemporaneously with the axis from which they arise; however, the first internode of such shoots is not notably elongated as seems to be common in sylleptic shoots. Calophyllum depressinervosum, C. nodosum (Ficure 3, b), and C. pulcherrimum all have proleptic shoots; the shoots develop from buds that have undergone a period of rest, and there are basal scales. All taxa that are recorded as having such basal scales probably show this type of growth pattern (see also Ficure 3, e). In all taxa that produce a single pair of leaves per innovation (and in many that produce several pairs), the buds are proleptic in the timing of their development, but there are no basal scales; the first internode of the axillary innovation may be slightly shorter than the first internode produced by the terminal bud (Ficure 3, a, c). Finally, in C. praetermissum and C. milvum (in the latter only probably), the first internode produced by the proleptic axillary bud is notably longer than those subsequently produced (Ficure 3, d). INDUMENTUM AND Hair Tyre With the possible exception of the poorly known Calophyllum sp. 73, the buds of which consist of imbricate scales (see below), all species have some indumentum, at least on their buds. In some species that have scale leaves at the bases of the innovations, the terminal buds tend to be glabrous, or to have hairs only at the edges of the bud scales. In these species, there can be great variation in the development of the indumentum on the terminal bud either on different specimens (C. garcinioides) or on a single specimen (C. nodosum). In specimens of C. depressinervosum from Borneo, the lateral buds also tend to be glabrous. Generally speaking, the more coriaceous and less leaflike outer pair of bud scales have obvious indumentum. A number of species characteristically have indumentum that is either very well (e.g., Calophyllum molle) or very poorly developed (e.g., C. inophyllum), or distributed distinctively on the plant (e.g., C. insularum). However, many species (e.g., C. collinum, C. rubiginosum, C. soulattri, and C. tey smannii) display considerable infraspecific variation in the development and distribution of indumentum. In C. euryphyllum and C. robustum the indumentum of young plants observed is less well developed than that of adults, although a single individual has not been followed throughout its life Of much greater taxonomic importance is hair structure. Vesque (1893) 1980] STEVENS, CALOPHYLLUM 131 described in detail the hairs of 31 of the 35 species that he treated; he had illustrated hairs of 27 species in his earlier (1889) work. Hair structure has otherwise been almost completely ignored, although, as will become apparent below, it shows a great deal of taxonomically important variation. A microscope is needed to observe hair structure, since in many species the hairs are only about 40 um. long. Hairs can be scraped from the terminal bud and mounted in water. Since they are brittle, they often break; thus Vesque did not describe the interesting basal portion of the hairs of Calophyllum soulattri (his C. spectabile) or C. pentapetalum (his C. amplexicaule, C. buxifolium, and C. pseudotacamahaca). In the present study hairs were examined by means of both ordinary and polarized light; the specimens examined covered both the morphological variation and the geographic distribution of the species. Describing the hairs is difficult because their shape is basically irregular. Terms currently used for different hair types (Payne, 1978) offer little help, yet there is no point in coining new terms. In all species the hairs are basically uniseriate, and in many they are more or less adaxially curved. Even in species with long, apparently straight hairs, the cell arrangement at the very base may give an indication of this adaxial curvature (e.g., Calophyllum cucullatum, Ficure 22, a, b). Some cells near the base are usually papillate; the papillae of individual cells project and are often restricted to, or better developed on, the abaxial surface. Rather short (less than 300 wm.), adaxially curved hairs with basal papillae (Ficure 9) are the most common type in the genus. In a few species with small hairs, the papillae cover the hair and make the cell arrangement beneath invisible (C. subsessile, Figure 29, n). Even in some species with long hairs, all the cells may be papillate, although there is often infraspecific variation in the development of these papillae (C. venulosum, Ficure 29, o-q, s—u Some species (e.g., Calophyllum bracteatum, Ficure 13, e-g; C. teysmannii, Ficure 25, 1, m) have hairs with one, or sometimes more, basal branch(es); these branches are formed from at least one cell. The apex of the hair sometimes appears to be forked because of a large apical papilla or branch. In the group of taxa related to C. undulatum, the branch at the base is often unusually prominent, and the hair may be almost mesifixed (Ficure 33, q). Some species (e.g., C. alboramulum, Ficure 9, a-d) have several branches or large papillae. In others (e.g., C. hosei, Ficure 26, p, q) the hairs may be almost stellate. It should be emphasized that the distinction between papillae and branches is not clear cut, and species that habitually have strongly papillate hairs will often have a few branched ones as well. The basal cells of the hairs tend to be narrow, while the distal cells are a occasionally, as in Calophyllum pyriforme (Ficure 27, b), all the cells have unthickened walls. In some species, especially those with longer hairs, the occurrence of thin-walled apical cells seems to be an inconstant character, but in others, such as C. coriaceum (Ficure 19, a—c) and C. tomentosum (FicureE 11, 1), the apical cells are habitually little thickened. The unthickened, 132 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 61 air-filled cells at the tips of the hairs of C. costulatum give the terminal bud a pale color, but in most cases the brown cell contents, probably tannins, make the bud dar The transition eet the thicker-walled basal cells and the thinner-walled, longer, apical cells is usually gradual. However, in Calophyllum as ps the apical cells are much broader, and in shorter hairs “‘stalk’’ and ‘‘head’’ are clearly distinguishable. Some hairs of C. insularum ae an abrupt transition between swollen, rather thin-walled, apical cells and smaller cells with thicker walls. In C. rufigemmatum, as well as in C. ferrugineum and some of its immediate relatives, the cells toward the base are short, thick walled, and as is usual, slightly birefringent under polarized light; the two apical cells are much elongated, broader, very thick walled, and strongl birefringent (Ficure 18, k, n). Calophyllum castaneum and some of its relatives are similar, although the thickening of the apical cells is less evident. walls are usually oriented ras to the long axis of the hair. However, in Calophyllum cucullatum, C. rufigemmatum, and a few other species, a single cell wall is rather sonia oblique. Apart from the papillae, the surface of the hair varies from smooth to rugulose. Surface ornamentation is especially prominent in Calophyllum undulatum and its relatives, C. thorelii, C. robustum, and some other taxa. In species such as C. robustum, the cell surface may be substriate. There may be considerable variation in the hairs from a single terminal bud, since they are often densely packed and show variable development. This factor, coupled with the general irregularity of hair structure, can cause and C. venulosum). On the other hand, C. soulattri and C. leleanii, which have been frequently confused, have basically different hair types, despite occasional reduced hairs found in the former that suggest the hairs of the atter. I have emphasized the structure of the hairs on the terminal bud, since in many species hairs are not commonly found elsewhere. However, because most species have naked buds (i.e., there are no bud scales), it is perhaps not surprising that relatively little variation is found when different parts of the plant are examined. Although the characteristically bifurcate hairs of Calophyllum collinum are more common on the terminal bud than elsewhere, this is a difference of degree only. Some specimens of C. rubiginosum have dimorphic hairs—albeit of the same basic type—on the stem; that this dimorphism is not apparent on the terminal bud is probably due to the spatial constraints there. The hairs on the inflorescence are of the same basic type as those on the terminal bud, although they tend to be smaller and less papillate or branched and to have thinner walls (e.g., Ficure 27, cf. 0, p). Hairs on the ovary and stamens are still more simple in structure. Infraspecific variation in hair type is generally variation on a theme. In some taxa, such as Calophyllum biflorum and C. teysmannii, the hairs range from a structure similar to a morulus (moruloid) ca. 30 wm. across to distinctive 1980] STEVENS, CALOPHYLLUM 133 hair types over 1 mm. long; similar reduced types occur on young plants of C. euryphyllum and C. robustum. All such moruloid hairs are superficially similar, although there may be differences in the ornamentation of the cell wall Despite the problems dealt with above, hair structure basically correlates with taxonomic groupings. In some cases where there is considerable variation in hair type (e.g., Calophyllum canum, C. scriblitifolium, and perhaps also C. polyanthum), the limits of the species are unclear. On the other hand, there is relatively little variation within C. soulattri, despite the broad limits adopted here for that species. Species believed to be related on other morphological grounds usually have similar hair structure, and infraspecific variation is not discontinuous. Hair type is thus a valuable aid in the circumscription of taxa, in identification (especially of sterile material), and in suggesting and confirming relationships. Twic AND Bup Characters of the twig, such as thickness, angling, presence of horizontal lines at the nodes (see above; Ficure 7, a), color on drying, development and persistence of the indumentum, and hair type, are all useful in characteriz- ing a species, although there is considerable variation in many of these characters. The axillary buds are usually small and inconspicuous, although in species such as Calophyllum rigidum those that give rise to inflorescences are notably plump. In a few species (e.g., C. rupicola), the axillary buds are supra-axillary in position. There are sometimes two buds in the uppermost leaf axils (e.g., C. euryphyllum, C. bifurcatum); the upper bud of the pair is the larger The shape and size of the terminal bud is fairly constant within a species and is of great help in identifying sterile material. The terminal bud is naked in most species and is nearly always densely covered with crustaceous to tomentose indumentum. In Calophyllum papuanum and its relatives the bud is strongly flattened since the blades of the leaves forming it are somewhat expanded (Ficure 4, a). In several species the bud is enclosed by scales that do not expand, and in the imperfectly known Calophyllum sp. 73, the scales are clearly decussate and glabrous. The bud may be borne a little way above the uppermost pair of expanded leaves on a short length of stem that develops before growth ceases. This underdeveloped internode is con- spicuous in some species, and it becomes of normal length when growth resumes In a few species the terminal bud dies. Axillary buds then take over, and branching tends to be profuse. Such nonfunctional terminal buds occur in Calophyllum pulcherrimum, in which the axillary innovations have basal scales, and in the very different C. savannarum, in which the axillary innovations lack basal scales (Ficure 4, d); they probably occur in additional species as noted in the descriptions, but field observations are lacking. The terminal bud and the associated pair of uppermost axillary buds are usually exposed, but they are enclosed and protected by the bases of the 134 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 £4. Bud and twigs, x 3. a, We a Ppapuanum (NGF 7277), ea bud. b-d, C. savannarum (bb 30477): b, c, node from side and front; d, apex of stem. e, C. teysmannii var. bursiculum (S 17230), node from side surrounding leaves in species with pseudoverticillate leaves. In a few taxa (e.g., Calophyllum teysmannii vars. teysmannii and bursiculum, and C. pani- culatum variants), the uppermost pair of leaves tends to remain erect, and the terminal bud is enclosed by the space formed by the opposing concave petioles. In C. savannarum the terminal buds presumably fall off very early (I have seen no specimens with buds), and the axillary buds are enclosed in a pocket formed by a concavity of the stem, which is swollen at e odes, and an opposing concavity in the petiole base (Ficure 4, b, c). This condition also occurs in the related C. articulatum, which, however, has functional terminal buds. LEAF The leaves are always petiolate, albeit sometimes very shortly so. Many species can be recognized by differences in the dried lamina, but the characters are difficult to describe accurately, and some experience is needed to appreciate amina shape is generally fairly useful; many species have basically ovate, obovate, or elliptic leaf blades, with only a moderate amount of variation in the shape of the apex. The base of the lamina may vary considerab y; for example, from cordate to acute or cuneate in Calophyllum soulattri, C. pentapetalum (Ficure 30, h-o), and C. venulosum (Ficure 30, a-g). Especially in C. moonii, C. venulosum var. tenuivenium, C. vergens, C. calaba var. 1980] STEVENS, CALOPHYLLUM 135 bracteatum, C. macrocarpum, C. rigidum, and C. lanigerum var. austro- coriaceum, leaf shape and size may differ considerably between young and adult plants. The general tendency for the leaf blades of the young plant to be longer and narrower than those of the adult is carried to an extreme in the last four taxa mentioned above (to 25 by 2 cm. in C. calaba var. bracteatum, and 50 by 5.4 cm. in C. lanigerum var. austrocoriaceum). mportant, but more or less qualitative, characters include the texture of the dried lamina, whether or not the whole blade has dried flat, the way the margins dry, the color of the upper and lower surfaces, and the prominence of the midrib, venation, and sometimes also the latex canals, which are usually hardly visible. These characters often reflect anatomical differences (Stevens, 1974a). Reference is made to the color groups in Dade’s chart (as given in Stearn, 1973, pp. 240, 241) matched by the dried blade; Ridgway’s (1912) charts were used in the matching process, but the detailed data that were collected for all species are not included here. When measuring venation density, one must take care not to confuse latex canals with veins® (Stevens, 1974a) and not to take measurements on poorly developed or damaged leaves. The course of the marginal vein(s) is usually obscured by thickening, although in some taxa (and to a varying degree) a submarginal vein may be distinguish- able, as in C. tetrapterum var. obovale. In this and other taxa the marginal thickening is often well developed and may be up to | mm. wide. In so species, notably C. garcinioides, there are obliquely and irregularly acer latex canals on the lower surface of the blade that are superimposed o the normal series of latex canals that alternate with the veins. Characters similar to those described above are useful in the identification of living material, although they may then appear different. The midrib in a dried leaf may be angled, although it is rounded in a living leaf; raised venation may be invisible. The young leaves may have a distinctive color; thus those of Calophyllum enervosum have a broad, red stripe down the middle, and those of C. rigidum are white. Variation in this character occurs within C. teysmannii var. inophylloide. Such characters are usually not mentioned in the accounts or the discussion, but attention should be paid to them by collectors in order to build up a body of reliable observations. GALLS Galls, usually on the leaf but sometimes also affecting the stem, have been described from several species in the genus, but no correlation of gall type with taxonomy has been noticed before. However, different types of gall do not occur randomly. Large, pustular galls (Ficure 5, g) occur on both Calophyllum chapelieri and the related C. verticillatum, while disc-shaped ones are found on C. vernicosum (all three species from Madagascar). Calophyllum trapezifolium, from Sri Lanka, has a spherical gall that is bilabiate *Rameji (1967) studied venation development in leaves of Calophyllum ia aula He found that the marginal veins formed first; then the secondaries, alternate ones of which differentiated to form the latex canals; and finally the tertiaries, andi linked the secondary veins and were usually invisible in the adult leaf. 136 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Ficure 5. Galls. a, b, Calophyllum ee ela (Kostermans 23599), apex of twig, xX 2: a, from outside; b, transverse section. c, d, C. teysmannii S var. inophylloide (Stevens et al. 27 5), underside of leaf: c, x 0.5; d, e, f, C. whitfordii (Williams 639), underside of leaf: e, x 0.5; f, x C. chapelieri (herb. d’Aleizette s.n.), underside - leaf, * 2h, oC. teysmannii var. teysmannii (Meijer 7057), apex of twig, at the apex (Ficure 5, a, b); this gall is also found on not-quite-typical specimens of the related C. walkeri. Both varieties of C. teysmannii have a distinctive bilabiate gall (Ficure 5, h) apparently caused by coccids; similar galls occur on C. aureum and specimens tentatively, but perhaps incorrectly, assigned to C. fraseri. Slitlike galls (Ficure 5, d) also frequently appear on C. teysmannii. 1980] STEVENS, CALOPHYLLUM 137 Small, conoid galls probably produced by gall midges occur mainly on several species from Papuasia, the Pacific, and the Philippines (Ficure 5, f). The a ; mon only on C. leptocladum, a species superficially similar to C. whitfordii, but are not known from C. leucocarpum, the other small-leaved species that occurs there. Little is known about the organisms causing these galls. Anthony (1976) described the galls and their causative agents on species of Calophyllum from Singapore, and there is some additional information in Docters van Leeuwen-Reijnvaan and Docters van Leeuwen (1926, 1941). This subject would undoubtedly repay further study. INFLORESCENCE The inflorescence is determinate (cymose) in all species, with both the main axis and any lateral axes being terminated by flowers. Most species that have terminal inflorescences often have additional inflorescences from i b flo may u Terminal inflorescences tend to have many flowers, but those of Galophyltum rotundifolium have only one or two flowers and lack an axis. Axillary inflorescences show interspecific differences in size, branching, and disposition of flowers along the axis. There may be from one (Calophyllum insularum) to numerous (C. paniculatum) flowers per inflorescence; an axis is nearly always present, but it may be absent in the poorly known C. rufinerve and some specimens of C. soulattri from Papuasia. Although the branching of the inflorescence tends to be a variable character, most species have either branched inflorescences, with or without an additional pair of flowers (developed from supernumerary buds) causing the node to have a flabellate appearance, or unbranched inflorescences, usually without additional flowers. In many species with axillary inflorescences, individual specimens have terminal ones. Solitary, axillary flowers occur only exceptionally, as in C. calaba var. calaba and C. rubiginosum; such flowers are larger than those in normal inflorescences. The variation in the arrangement of flowers along the axis is similar to developing. However, a number of Malesian species tend either to have 138 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 flowers in fours along the axis, or to have terminal groups of five flowers (e.g., C. costulatum, C. ai le and C. rigidum), but these species lack Gyhorled? > leaves on the tw Bracts are usually small and a ios but in a few species they are notably foliaceous and persistent. The length of the pedicel and whether or not it is incrassate in fruit both tend to show considerable infraspecific variation, although some species (e.g., Calophyllum castaneum and C. poilanei) have notably short pedicels, and others (for example, C. recurvatum and C. mukunense) have pedicels that always become incrassate in fruit. FLOWER The flowers of most species of Calophyllum from the Old World appear to be hermaphroditic, the exception being those of some species from Papuasia, which are dioecious and/or gynodioecious (Stevens, 1974a). In staminate or pistillate flowers, pistils or stamens, respectively, are usually present, albeit smaller than usual. However, more observations on the distribution of staminate and isan flowers—and on all other aspects of reproductive biology—are badly nee There has been ae accession in the past as to whether or not one can recognize sepals and petals in Calophyllum, or even if there are any sepals at all in some species (see, for example, Choisy, 1823; Wight, 1840; Vesque, 1893). The morphological distinction between the various members of the perianth has generally been admitted to be slight, as Vesque (1893), Engler (1923), Henderson and Wyatt-Smith (1956), and others have noted. Vesque (1893) used the term ‘‘sepals’’ for the parts of the perianth that were in opposite pairs (he thought that there were only two pairs), and ‘‘petals’’ for the parts that were whorled. Usually only the outer pair of perianth ale is notably thicker than the others, and they completely enclose the ; however, in a few species (such as Calophyllum polyanthum and some 6s its relatives) the outer pair is much smaller than the others and does not enclose the bud. In all species the outermost pair is the same color as the inner pair(s), although the color may be modified by indumentum. In size, shape, and sometimes even texture, this outer pair is no more different from the next pair than that pair is from the others. Thus, for descriptive purposes, all are called tepals. Although there are usually only four outer tepals arranged decussately in pairs, with any additional inner tepals being in more or less imbricate whorls of four, a few species (e.g., C. walkeri and occasionally C. blancoi) have three pairs. Such species would have six sepals in Vesque’s (1893) terminology. Tepal number, shape, size, and indumentum are all of some use in characterizing species; all, however, are variable in some species, and their use as diagnostic or differential features is limited by the paucity of material at anthesis and the often considerable distortion that occurs on drying. The number of stamens varies considerably in some species, and as more flowering material is collected this variability will doubtless be shown to be greater. It is a tedious (but necessary—see Davis & Heywood, 1963) job counting the up to 600 or so stamens that occur in species such as 1980] STEVENS, CALOPHYLLUM 139 Calophyllum blancoi and C. molle, and stamen number is not much use as an aid in identification, but within rather broad limits many species have characteristic stamen numbers (see the species pairs C. molle/C. rubiginosum, C. canum/C. lowii, C. recedens/C. vernicosum). The filaments are ofte somewhat connate at the base—notably so in species such as C. recurvatum. The phalangiate arrangement of the stamens, common in many Guttiferae, is not marked in Calophyllum, although phalangiate stamens have been illustrated or reported (see, for example, J. J. Smith, 1920; Robson, 1976). The tendency of the stamens to be in four groups may be due not so much to differential connation of the filaments as to the removal of the four innermost tepals during dissection of the flower, this causing four areas of weakness in the tissue on which the androecium is borne; however, studies on the floral anatomy of the genus are needed. Anther size and shape is, again within rather broad limits, characteristic for each species. The presence of indumentum on the anthers of some species is an interesting and hitherto unreported character, but this varies infraspecifically in C. canum. A single collection from Malaya has hairs on the filaments (see under C. rufigemmatum). Although the ovary is sometimes hairy, it is generally glabrous; the ovule is basal, anatropous, and almost without exception solitary. The style is nearly always glabrous, but in staminate flowers of Calophyllum vexans, al- though the ovary itself is glabrous, there are a few hairs on the style. The stigma is usually more or less peltate and radiate or irregularly lobed, al- though in a few species it is slightly expanded or more or less infundibular. FRuIT AND SEED The fruits of nearly all species of Calophyllum are drupaceous when mature, having a skin, a more or less succulent layer, and a stone; inside the stone is a single seed. In those species where there is no stone in the mature fruit (e.g., C. eputamen (Ficure 6, f), C. parviflorum), a thin stone is visible in the young fruit. Corner (1976) described the structure of the fruit of C. inophyllum; the fruits of all species appear to be similar in structure. The skin and the fleshy layer are both derived from the carpel wall, but the stone (the endocarp of most workers—see, for example, Cambesseédes, 1828; Smith & Darwin, 1974) develops from the outer part of the seed. Inside the stone is aerenchymatous tissue that is strongly developed in the young fruits of apparently all species but is generally not obvious in ripe fruits. However, in C. inophyllum and C. tacamahaca in particular, the aerenchyma- tous tissue (the spongy layer of the descriptions below) persists in the ripe fruit. The embryo fills the cavity of the stone and consists mainly of two large cotyledons that, although closely adpressed, remain readily distinguishable. The minute radicle points to one side of the base of the fruit, as woul be expected of a seed developing from an anatropous ovule. eae of the radicle facing the top of the seed, away from the ‘‘micropyle Bentham, 1863; Cambessedes, 1828, p/. 17, c), are incorrect; this mistake is due to confusion of the pointed cotyledonary end of the seed of Calophyllum inophyllum with the radicle (see, for example, Miers, 1855). In those few 140 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 species known to have tiara ae seeds, the pte are of various sizes and are much intertwined. mbryo is usually whitish, but in C. suberosum it is a rather frightful Sri Generally, however, collectors have not mentioned embryo color. Most earlier workers paid little attention to the structure of the fruit, probably because of the paucity of mature fruits. However, there is much useful 2 variation in size, shape, and surface of the dried fruit (Ficure , h), as well as in thickness of the various layers and in texture of ge layer when dry. Fruit characters are usually constant within a species; however, there is considerable intraspecific variation in Calophyllum rocarpum (FicurE 28, g, h) and C. soulattri, at least as presently circumscribed. There are large air spaces in the fleshy layer (mesocarp—the ‘‘outer layer’’ of the descriptions) of some species when dry. These spaces may be under Ficure 6. Fruits. a, b, Calophyllum papuanum (NGF 10331), stone, X 1: a, from outside; b, longitudinal section. c-e, C. goniocarpum (bb 33914), : ; : ee C. peekelii (NGF 18330), x 0.5: h, from outside; i, longitudinal section. j, C. macrocarpum (FRI 10926), transverse section, < 0.5: 1980] STEVENS, CALOPHYLLUM 141 the skin, adjacent to the stone, or throughout the outer layer and completely disorganizing it (Ficure 6, cf. d, i, j). The development of these spaces in older fruits may result in strikingly different, strongly wrinkled submature and smooth mature fruits, as in Calophyllum dioscurii and C. incumbens;, the skin of fruits with large air spaces under it is, not surprisingly, brittle. However, how the air spaces form is not known, and their taxonomic significance is not always clear, owing to the paucity of material with fully ripe fruits (see especially C. neo-ebudicum and C. vitiense). The stone furnishes numerous characters of systematic use. It is angled in Calophyllum trachycaule and a few closely related species (FicureE 6, Cc); and either rounded or angled (the latter only at higher altitudes) in C. papuanum. A number of species have two to four radiate markings coming from the apex of the stone (Ficure 17, h); these are especially noticeable in the C. venulosum complex. However, this character is not always easy to observe, especially in those taxa in which the outer layer does not detach cleanly from the stone. Other useful characters of the stone include apex shape (rounded, pointed, or retuse), surface texture (smooth or rough—Ficure 6, a), and presence or absence of a basal plug. Compared to the rest of the stone, the basal part is often paler in color and somewhat different in texture in several West Malesian species (e.g., C. rufigemmatum and C. dasypodum), but a true plug (i.e., a circular area that is demarcated by a line of weakness in the stone and that is pushed out by the radicle during germination) is, except in C. inophyllum and C. tacamahaca, so far known only from Papuasian and Pacific species. There is infraspecific variation in the presence of a plug in C. soulattri as delimited here. ANATOMY Woop. The literature on the wood anatomy of Calophyllum is rather scattered, but it appears that interspecific differences are minimal, even when very different species from widely different areas are compared. Authors studying the anatomy of species from the same area have found practically no differences between them (Moll & Janssonius, 1906; Pearson & Brown, 1932). The following description of the wood anatomy has been drawn up mainly from the references cited below (see also Vestal, 1937; Lakhanpal & Awasthi, 1965; Lakhanpal, 1970). Vessels (lacking to) 2 to 12/sq. mm., 300-1250 by (30-)150-350 jm. across, usually single, scattered, sometimes in oblique lines, perforation plates simple, pits bordered where adjacent to tracheids and simple or sometimes slightly bordered where adjacent to rays and wood parenchyma, walls ca. 4 p thick; tracheids vasicentric; fibers 450-1680 by 6-20 wm. across, with walls 2-5 wm. thick; rays 8 to 15/mm., heterogeneous, with erect central cells and procumbent marginal cells, uniseriate, sometimes partly biseriate or exceptionally triseriate, 140-500 ym. tall; parenchyma in apotracheal bands, interrupted or not, (1 or) 2 to 9 cells wide, sometimes with crystals, bands varying considerably in density (see Prakash, 1966, for interspecific variation in C. inophyllum), paratracheal to almost aliform parenchyma also reported (Fundter & Wisse, 1977). =) 142 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 (Details taken from the following species: Calophyllum milvum (Madagascar; Lecomte, 1922); C. inophyllum, C. polyanthum, C. soulattri, C. calaba var. bracteatum, and C. apetalum (India to Burma; Gamble, 1881: Pearson & Brown, 1932; Chowdhury & Ghosh, 1958): C. calaba var. bracteatum and C. inophyllum (Vietnam; Lecomte, 1926); several species of uncertain identity (Malaya; Desch, 1941); C. inophyllum, C. calaba var. bracteatum, C. dasypo- dum, and C. venulosum var. venulosum (Java; Moll & Janssonius, 1906); C. canum and C. obliquinervium (Sabah; Burgess, 1966); C. ?blancoi and C. inophyllum (Philippine Islands; Kanehira, 1924); C. papuanum and C. pauciflorum (Papuasia; van der Graaff & Baas, 1974); and C. inophyllum, C. insularum, and C. ?sil (Papuasia; Fundter & Wisse, 1977)). Lear. Vesque (1889, 1893) illustrated and described the lamina anatomy of the species that he recognized, and Stevens (1974a) gave brief descriptions for the Papuasian species; Schofield (1968) described the petiolar and nodal anatomy of two species; recently D’Arcy and Keating (1979) described the lamina anatomy of species of Calophyllum from Panama, including species introduced from the Old World. A number of characters are helpful in delimiting species, including height of the epidermal cells; presence, height, and lignifica- tion of the hypodermis; characters associated with the vascular bundles, latex canals, and midrib; width of the marginal thickening, and number of vascular bundles and latex canals that it encloses; and thickening and lignification of the walls of the spongy mesophyll. When most of the spongy mesophyll has lignified walls, the layer of cells adjacent to the lower epidermis often remains unlignified, as in Calophyllum teysmannii var. bursiculum, C. garcinioides, C. rugosum, and C. sakarium. Additional latex canals may occur: in C. pauciflorum (Stevens, 1974a) and in C. trapezifolium and related species (Vesque, 1893), they are found in the lignified tissue below the vascular bundles; in C. garcinioides they are found adjacent to the lower epidermis; and in C. sakarium the vascular tissue in alternate vascular bundles is largely replaced by a large latex canal. Details of the lamina anatomy are given for a few species below: however, although a general anatomical survey of the genus would be of interest, it is outside the scope of the present study. Anatomical characters often help in delimiting some species (Vesque, 1893; D’Arcy & Keating, 1979), but problems may result from infraspecific variation and the unwitting study of juvenile foliage (Stevens, 1974a). POLLEN Erdtman (1971) described the pollen of Calophyllum inophyllum as being (2- or) 3-colporate, suboblate to prolate-spheroidal, and with a diameter of about 38 wm. Yi-Zhen (1979) gave palynological details of C. inophyllum, C. membranaceum, and C. thorelii (probably C. polyanthum) that agree with Erdtman’s account; he described the pollen walls as being finely reticulate and at least weakly 2-layered. Preliminary observations on pollen from species from throughout the range of the genus suggest that there is little variation in basic pollen type, although there appears to be some in details of surface 1980] STEVENS, CALOPHYLLUM 143 ornamentation; Anderson and Muller (1975) found the pollen of the different peat swamp-dwelling species of northwestern Borneo to be similar. A more comprehensive survey of pollen is underway (Banerjee & Stevens, in prep.). CHEMISTRY Hegnauer (1966) has summarized the early chemical work on the genus. There has also been a considerable amount of more recent work, and species from throughout the range of the genus (America, Madagascar, India and Sri Lanka, Southeast Asia, Malesia, Australia) have been surveyed for many types of compounds. Although the broad picture of the variation of some classes of compounds is becoming apparent, the taxonomic implications of this variation is unclear. Interspecific chemical variation in the species from any one area is unknown, and the amount of infraspecific variation is also practically unknown (see also below) except for that occurring in taxa from Sri Lanka (see Sultanbawa, 1973; Gunesekara ef al., 1977 The esters of fatty acids, and also the free acids themselves, are found in large amounts in the seeds. Oleic, linoleic, stearic, and palmitic acids are the main compounds involved, and they are probably found throughout the genus. Saponins occur in the leaves of Calophyllum inophyllum and C. brasiliense, and cyanogenetic glucosides have also been reported. Tannins are fairly general in the genus, especially in the bark, and Ashton (1964) noted that the proteins occurring in gelatin solution were readily precipitated by ground-up leaves of several species of the genus that grow in Brunei. Ethereal oils also occur in the genus. A considerable variety of xanthones is found in the wood and also in the bark. (Xanthones are found in some diversity only in the Gutuferae and Gentianaceae.) One of these, jacareubin, occurs in all but one’ of the species of the genus so far analyzed, but is very uncommon elsewhere, being known only from a single species of Mesua L. (Carpenter, Locksley, & Scheinmann, 1969). Jacareubin is of interest in being a morellin analogue; morellin is an active principle in gamboge, which comes from Garcinia morella Desr. (a member of the Clusioideae - Garcinieae) and other species of that genus. There has been discussion as to whether or not jacareubin always occurs in Calophyllum inophyllum and other species, or if it appears only with age (Al-Jeboury & Locksley, 1971; Kumar, Ramachandran, & Sultanbawa, 1976); there may also be infraspecific variation in other xanthones in C. inophyllum. References to other works in which xanthones have been isolated and characterized may be found in Gunesekara et al. (1977) and Bhanu and Scheinmann (1975). The latex of Calophyllum is extremely rich in complex coumarin derivatives: 4-pheny]l or 4-alkyl coumarins, hydroxy acids, or cyclohexadienone derivatives with phenyl analogues (Gautier, Kunesch, & Polonsky, 1972). The structures of many substances of this type have been proposed (see Gunesekara et ’The exception in Calophyllum is C. moonii (Gunesekara et al., 1977, as C. soulattri). 144 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 61 al., 1977, for references). Kawazu, Ohigashi, and Mitsui (1968) found three complex coumarin derivatives in the leaves of Calophyllum inophyllum to have piscidial activity, and several 4-n-propylcoumarins found in Mammea americana L. (Calophylloideae) have considerable insecticidal activity (Crom- bie et al., 2) CYTOLOGY AND BREEDING SYSTEMS Only a single chromosome number is known from the genus, 2n = 32 in Calophyllum inophyllum (Darlington & Wylie, 1955). In the majority of species, the flowers appear to be hermaphroditic. However, dioecy or some other type of breeding system occurs in the Papuasian species Calophyllum papuanum and its relatives and in C. leleanii (Stevens, 1974a, the latter as C. solomonense). Flowers in the genus are generally reported to be sweetly scented, and to judge by seed set of an isolated tree of C. papuanum, the pollen vector may travel 50 meters or so (cf. Stevens, 1974a). The flowers are not obviously adapted to an oligolectic pollinator. Hybridization may occur; however, close analysis of the situations in which it is suspected is clearly needed. Apparent hybrids between Calophyllum inophyllum and C. paniculatum, C. inophyllum and C. chapelieri (both Madagascar), C. ferrugineum and C. teysmannii (Malaya), C. teysmannii and C. biflorum (Sarawak), and C. inophyllum and C. caledonicum (New Caledonia) are discussed under the second-named member of each species pair; see also C. verticillatum and C. parviflorum. Polyembryony occurs in three small-seeded species from West Malesia—this is the first report of its occurrence in the genus. It can be recognized by the occurrence of up to six naked embryos forming a close-knit mass in place of the normal single embryo. Polyembryony has been found in Calophyl- lum rupicola (FRI 20873, Malaya), C. depressinervosum (SAN 17093, Brunei), and C. nodosum (S 25720, Sarawak; Stevens et al. 881A, Sabah). In the case of C. nodosum, fruits from three adjacent trees were polyembryonic, and up to three embryos were germinating at one time, although one was much larger than the others. The causes of this polyembryony are unknown, but in view of the reports of apomixis elsewhere in the family (see references in Maguire, 1976; also Planchon & Triana, 1862b), apomictic polyembryony is suspected. (It may be noted that polyembryony also occurs in Mesua; e.g., in M. nuda Kostermans ex T. C. Whitmore—pers. obs.) From this summary, it is clear that observations on cytology, the breeding system, and hybridization will greatly increase our understanding of the taxonomy and evolution of the genus. CLASSIFICATION OF THE GENUS The recognition of taxa in any group in which the members have many characters in common is something of a problem, and the suspicion that some species of Calophyllum may be apomictic has not eased the situation. The heightened similarities caused by ecological convergence, particularly 1980] STEVENS, CALOPHYLLUM 145 notable in the small-leaved montane species (see C. clemensorum and C. vergens), make it very easy for one to be misled by superficial resemblances, and misidentifications are numerous in the literature. Specimens cited under a taxon described from Malesia should have been referred to four other taxa, while specimens of a fifth taxon were annotated with the same name but not cited; the five taxa are not closely related. A particularly good example of confusion caused by close superficial similarity is the recognition that two species were included in my original concept of C. suberosum (Stevens, 1974a; see C. persimile below). DELIMITATION OF SUPRASPECIFIC TAXA Vesque (1893), in part following such earlier workers as Choisy (1823, 1824) and Planchon and Triana (1862a), recognized sections and smaller, but informal, groupings in his classification of the genus; these were based mainly on tepal (sepal and petal) number, the presence or absence of a ypodermis, and other details of anatomy, hair structure, and inflorescence type. An semen characters, of course, cannot be used by themselves to delimit taxa (but see Vesque’s section Hypodermata), and none of these characters pitas particularly highly with other ones; nor, indeed, is the distinction between the various states of each character very sharp (see the preceding discussion). Hence supraspecific taxa are not recognized here. One of the more distinctive groups of taxa is that centered around Calophyllum pyriforme. All taxa in this group have similar, stipitate fruits with a fibrous outer layer and a very thin-walled stone, as well as axillary innovations and inflorescences with basal scars. However, these characters are found individually in other species or species groups. Thus, there are two other main groups of species with axillary innovations and inflorescences like those of the C. pyriforme group; one is centered on C. nodosum an the other on C. garcinioides, but neither is otherwise obviously related to the C. pyriforme group (for instance, they have very different inflorescences and fruits). Unfortunately, germination, seedlings, and young plants are known in only one of the groups, and here only in two species (C. nodosum, C. pulcherrimum); when they are more widely known, they may help to delimit these and other groups of related species. All that can be recognized above the species level are groups of taxa, the members of which are clearly more related to one another than to any other taxa or groups of taxa. DELIMITATION OF TAXA AT AND BELOW THE RANK OF SPECIES, AND Its IMPLICATIONS Studies of any widespread lowland Malesian group that are finished in under a lifetime of field work must necessarily use morphological characters as the main criteria for the recognition of species. Population size and structure is unknown in Calophyllum, and it is unlikely that there will be direct evidence bearing on the size of interbreeding populations or the location of intrinsic sterility barriers throughout the genus. It is premature to attempt to circum- scribe taxa on the basis of possession of unique, derived characters; not only is the relationship of Calophyllum to other genera of the Guttiferae 146 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 unclear (see below), but knowledge of the basic morphology of many of the taxa included in the genus is still poor. A complaint about the morphological species concept is its element of arbitrariness and inconsistency (although similar complaints can be leveled against other species concepts—see, for example, Sokal & Crovello, 1970), and this is perhaps particularly true when one is dealing with taxa that inhabit different areas. At the present level of knowledge of the genus in most of Malesia, it is easier to recognize slight differences that are correlated with geography than comparable differences between plants in a single locality. Hence, the occurrence of not very different, but non-intergrading, taxa at one locality may be overlooked. The ranking of variation that is correlated with geography has been ably discussed by van Steenis (1957), who listed four criteria to help in this endeavor: comparison of the differentiating characters of the taxa under discussion with those of well-established taxa in the same genus; careful observation; comparison of the variation between the allopatric taxa with that in allied species that have a wide, continuous range (‘‘deductive analogy’’); and experimental taxonomy. Only the first criterion helps directly in establishing the rank of such taxa. ““Deductive analogy’’ is difficult in practice, even if one has correctly identified closely related taxa; a wide-ranging species may show variation in characters the extreme states of which, if not connected by intermediates, would define perfectly ‘‘good’’ species. One cannot reliably extrapolate from one case r event of a type for which it is difficult to predict an outcome—hybridization if the two taxa were to become sympatric. If two geographically separated taxa that are recognized as subspecies (or varieties, if subspecies are not being recognized) are later found to grow together and yet to remain distinct, the tendency will be for the same two taxa to be called species. A test for a biological species is being applied in an otherwise morphological context. Darwin’s (1859) ideas on evolution did not suggest that the species was more ‘“‘real’’ than other taxonomic ranks, nor were his ideas so se by many leading botanical systematists of the day (see Bentham, 1874). Nevertheless, the temptation to treat the rank of species as ey different from other ranks, or to admit a type of evidence—that is not allowed elsewhere—as deciding specific or infra- specific rank (e.g., van Steenis, 1957; Shelter er al., 1973) has become deeply ingrained in systematics, largely because of the findings of genetics. Ss recent work increasingly suggests (see Raven, 1976; Cronquist, 1978; and Levin, 1979, for references), it may be hazardous to assume that the limits of a species that was defined on morphological criteria will correlate with data from ecology, breeding systems, evolution, or geography. However, this does not mean to say that morphology is a biologically inert and meaningless property of an organism; other species concepts face the same problem of non-correlation. A species defined as a group of individuals exchanging genes only within the group has no further necessary properties. Suggestions 1980] STEVENS, CALOPHYLLUM 147 that a species so defined has properties more like those of an individual than those of a class (philosophically defined—see, for example, Hull, 1978) do not alter this fact. The biological (Mayr, 1969) and evolutionary (Simpson, 1961) species concepts suffer from the same problems, and they, too, are impossible to apply directly to any medium- or large-sized taxonomic group of Malesian plants. The morphological species concept has the virtue of being eminently operational in a situation where the level of knowledge is not far advanced, and of course later on as well. The species recognized here are delimited by discontinuities in the variation of several of the morphological characters discussed above; varieties show comparable discontinuous variation in only one or two such characters. The extent of the discontinuities and the characters involved affect the rank at which taxa are recognized. Species are often constant over large areas, or if variable, they do not intergrade with others. The limited field observations made suggest that species have relatively invariable and often distinctive bark characters (see also Henderson & Wyatt-Smith, 1956), fruit, germination, and seedling types. Most species have a discrete geographic range (as do varieties), and many have clear ecological preferences. Species that on morphological grounds are thought to be related often differ in both geographic ange and ecological preference. Although many species grow together, hybridization is not notably common. Thus, the morphological species in many cases seems to correlate with other types of information, but it must be remembered that morphological evidence has been paramount in species delimitation. As was suggested above, correlations with these other types of information are in a way fortuitous, albeit highly satisfying, but conflicts between the taxon limits that they suggest and those implied by morphological characters must be expected as more becomes known. The variation in characters of anatomy, fruit, germination and seedling, and bark sometimes may seem not to have been treated satisfactorily. This is not willful inconsistency: in the present state of knowledge of the genus, these characters may be of limited use when unsupported by characteristics that are observable on all, or many, specimens. As will be clear from the extensive discussion after many species, the rank and even the limits of some taxa remain problematic. Although I have seen over 70 taxa in the field, and the other taxa are delimited by gaps in the variation similar, as far as is possible, to those that separate the taxa that I have seen in the field, such extrapolation is clearly hazardous (cf. Lewis, 1955, quoted favorably by van Steenis, 1959, who suggested that one should somehow allow for “‘the variation possible’’ in a species). I have kept doubtfully distinct taxa separate and have fully, but not formally, described taxa that may eventually be worth recognizing but that are poorly known and have no names at present. In all cases, problems are fully discussed, which should make any future adjustments in rank easier (see also Smith, 1979). Most characters useful in delimiting species nevertheless show considerable infraspecific variation in some taxa; these taxa can still be separated by use of other characters, which themselves break down elsewhere. However, 148 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 one cannot argue that, if a given character does not separate a given pair of taxa, or varies at a level below the rank of the taxon under consideration, it therefore cannot be used to separate any taxa at that rank. If such variation is not too frequent, the character may be treated more carefully elsewhere, although not discarded; it is the pattern of variation in the problem under immediate consideration that matters. Many of the vegetative characters (including color of the dry lamina, prominence of venation, and density of indumentum) that make even sterile material fairly easily assignable to a species, eae if the provenance of the specimen is known to within two t kilometers or so, are essentially qualitative. Other characters, such as ee thickness and venation density, are quantifiable, but they tend specimens from juvenile plants. This may make species boundaries appear to be more vague than they are. Confirmation that apomixis occurs in Calophyllum will be of little help in deciding at which rank to recognize taxa. The types of variation patterns in apomictic groups are as varied as the groups themselves, and they often cannot be incorporated satisfactorily in the normal taxonomic framework. However, it is perhaps unexpected that two of the taxa of Calophyllum in which possibly apomictic polyembryony has been detected are closely related, wide ranging, and relatively invariable; they may even grow together on occasion. Systematists working on tropical groups should of course be on he lookout for apomixis, since it may occur quite widely (see Kaur et al., 1978). ai several caveats should be borne in mind when reading the species accounts. It is clear that the inference of evolutionary processes from taxo- nomic patterns is fraught with problems and with numerous possibilities of circular argument. EVOLUTION, DISTRIBUTION, AND ECOLOGY FossiL RECORD OF THE GENUS Prakash (1975) has summarized the literature dealing with the fossil record of the Guttiferae, and that relevant to Calophyllum is evaluated here. The record of fruits of Calophyllum from the middle Eocene in Rajasthan, India (Lakhanpal & Bose, 1951), must be treated with caution, since although they were compared to those of C. trapezifolium (properly C. austroindicum), they have no characters that unmistakably mark them as belonging to Calophyllum. Leaves of Calophyllum, possibly from the lower to middle Eocene, have been reported from Assam (Bhattacharyya, 1967). Foliar remains of Calophyl- lum have been found in several places in Malesia. Calophyllum nathorstii Geyler was found near Palembang, Sumatra, in rocks of the younger Neogene (possibly Pliocene; Krausel, 1929), as well as in Miocene coal of Labuan 1980] STEVENS, CALOPHYLLUM 149 (Geyler, 1887; a second, unnamed species that was compared with C. inophyllum was also found there). Remains of what is quite possibly a species of Calophyllum occur in aaa sediments of Java (Goppert, 1854, as Musophyllum truncatum Gopper The genus Calophyllodendron Sante & Awasthi, based on fossil wood, seems similar to Calophyllum. Three species, Calophyllodendron cuddalorense Lakhanpal & Awasthi, C. eoinophyllum Prakash, and C. indicum Lakhanpal & Awasthi, have been described. Remains of the genus have been found in sediments of Upper Miocene age in Assam (Prakash, 1971), the Mio-Pliocene boundary near Pondicherry, India (Lakhanpal & Awasthi, 1965; the comparison of Calophyllodendron indicum with the extant Calophyllum apetalum is not justified), and the Mio-Pliocene boundary in Assam (Prakash, 1966, the best record). Calophyllum pollen of late Miocene age is known from Brunei (Muller, 1972; Anderson & Muller, 1975); the genus was ett an important component to that of swamps found in the same area fodny. Calophyllum pollen may occur in the Eocene in India (as Calophyllumpollenites rotundus Sah & Kar; Sah & Kar, 1974), but the record must be confirmed. There are few early records of the genus elsewhere. Calophyllum pliocenicum Krasser was reported from the Pliocene of Brazil (Krasser, 1903; Krasser thought that it was very close to C. calaba (= C. brasiliense). Calophyllum calabiformis Berry was described from the Miocene of Trinidad (Berry, 1925). Wood possibly comparable with that of Calophyllum or Mesua has been described from the lower Miocene of Egypt (as Guttiferoxylon fareghense Krausel; see Prakash, 1975) RELATIONSHIPS OF CALOPHYLLUM AND MAJor PaTTERNS OF EVOLUTION IN THE GENUS The immediate relatives of Calophyllum in the Calophylloideae - Calophylleae are Mammea L. (including Ochrocarpus Thouars), Paramammea Leroy, Mesua (including Kayea Wall.), and Poeciloneuron Bedd.; this last genus is rather distinct in details of wood anatomy (Baretta-Kuipers, 1976). Although Mammea has been placed in the Clusioideae by Robson (in Heywood, 1978) in the belief that the united cotyledons were a swollen hypocotyl,* it is correctly placed in the Calophylloideae (e.g., Planchon & Triana, 1862b; Brandza, 1909; pers. obs.; unpublished studies by S. W. Jones). All these genera have basal ovules. Mesua is an Indo-Malesian genus; Mammea extends from Africa and Madagascar (de Wilde, 1956; cf. Kostermans, 1961) to Indo-Malesia, with a single species (Mammea americana) in tropical America; Paramammea occurs on Madagascar; Poeciloneuron includes two species, both from western India. Mesua has a mid-Tertiary fossil record (Prakash, * Leroy (1975, 1977) distinguishes Ochrocarpus from Mammea by its embryo, which lacks any trace of gag ae and its latex canals; in both characters Ochrocarpus is similar to Garcinia. The circumscription of Mammea adopted here is for convenience only; as Leroy (197) notes, representatives of this group on Madagascar are still poorly understood. 150 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 1975) in India; earlier records of both Mammea (Late Cretaceous!) and Mesua need to be confirmed. The other tribe in the Calophylloideae is the Endodes- mieae (see Engler, 1925), characterized by a single apical ovule. It includes two monotypic, West African genera, Endodesmia Bentham and Lebrunia Staner (Staner, 1934). Some of the characters separating a genera are listed in TABLE 2 Although it is clear from Tae 2 that Calophyllum is easily distinguishable from the other Calophylloideae by its single basal ovule and its nearly always strictly parallel venation, its exact relationship to any of these genera, including members of the Endodesmieae, is unclear. Most chemical and palynological data are presently restricted to the three larger genera of the Calophylleae, Calophyllum, Mammea, and Mesua, but it is hoped that further studies on the subfamily will permit a resolution of the relationships of all the genera. Vesque (1889, 1893) placed the species of Calophyllum in a more or less evolutionary arrangement. He considered that C. inophyllum was ‘‘le groupe presence of a basal plug in the stone are derived characters, C. inophyllum may be a member of a not notably primitive group of taxa that diversified most in eastern Malesia (see also below). Moreover, the members of the various groups of species that Vesque recognized in the Old World often do not seem immediately related. Thus, the members of one of his groups (C. soulattri (his C. spectabile), C. walkeri, C. cerasiferum, and C. sclerophyl- lum) have a hypodermis, but little else, in common. Several methods are currently used to assign evolutionary polarity to characters; that is, to determine which character state is advanced and which is primitive. This is not the place to review the value of such methods, except to note that the one that appears to be by far the most satisfactory, out-group comparison, cannot be used here because it is not yet clear what the sister group of Calophyllum—and the sister group of this larger group—is, an all-too-common complaint in botanical systematics (Stevens, unpubl. ms.). To compare variation within the Calophylloideae or Guttiferae as a whole is a less rigorous form of out-group comparison that can be used here faute de mieux. The suggestion that the state of a character can be considered primitive within a group in the absence of other evidence (e.g., Estabrook, 1977) is unacceptable since the commonness of a derived character state in a group depends on the subsequent diversification of the monophyletic line that it characterizes (e.g., Stebbins, 1974; Judd, 1979; Stevens, unpubl. ms.). Functional considerations have been much used, as by Vesque (1893), but there is no reason why a character state that is believed to have some adaptive value should be considered derived when compared with the state that has no apparent adaptive value (e.g., Davis & Heywood, 1963). Neverthe- less, derived and primitive states are suggested for some characters in TABLE 3 by using these and other suspect criteria. This table is perhaps most useful in stimulating discussion, especially since no polarity has been suggested for the majority of characters. It would be premature to suggest detailed relationships of the taxa here. Tas_e 2. Some characters distinguishing genera in the Calophylloideae. * Calophyllum Mesua Mammea Paramammea_ Poeciloneuront Endodesmiat Lebrunia HAIRS UNISERIATE + _ - ? = = se BuD SCALES PRESENT —(+) + + + + + + (always) VENATION OBVIOUSLY RETICULATE = + ts aE + —(+) + (+, very rare) INFLORESCENCE TERMINAL —(+) are — — +,- 4+ = (+, very rare) FLOWERS PERFECT +(-) + —(+) — + + + ‘‘SEPAL’” NUMBER 4(2) 4 2 2 2,5 5 4 STAMENS CONSPICUOUSLY CONNATE —(+) - —,+ + = + = ANTHERS WITH TERMINAL INTRORSE PORE - _ _- + = = STYLES DEEPLY DIVIDED = = +(-) +4 = _ STYLES LONG +(-) + _ + + £ as OVULE, LOCULUS, AND SEED leeds ls 4-8; 1,2; 4; 1,2,4; 6-10; 6-10; 4; 2:51 I; 1; 1 Let NUMBER 1 or more | or more up to 8 OVULE POSITION Basal Basal Basal Basal Basal Apical Apical PEDICEL FLESHY IN FRUIT _ = = a = FRUIT DEHISCENT 7 —(+) —(+) _ + = = rom Staner, 1934; Perrier de la Bathie, 1948; de Wilde, 1956; Spirlet, 1966; Leroy, 1977, and pers. comm.; Stevens, Data fr genera gens Paramammea. o species of Poeciloneuron, P. indic O)h e n ao ) has two small, tEndodesmia calophylloides has um Bedd. and he pauci ciflorum B edd., may outer, persistent tepa nts wo large subpersistent tepals, and six large 1978. stipulelike structures. 1974b, and pers. obs. of all not be very closely pines nee ane eee of P. pauciflorum see I, , decid tepals; the anthers are not locellate (in both nL oe the anthers dehisce, at least instially, by introse, apical pore slits); and rns are colle ietike per in ie leaf axil. For palynological differences, see Seetharam & Pocock, WOTTAHdOTVD ‘SNAAALS [0861 IST JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 TABLE 3. Possible primitive and derived states of some morphological characters in Calophyllum 1) hwWN— oe) On —_ © oO — — SEEDLING AND YOUNG PLANT Seedling leaves separated oe well-developed vs. shortened internodes. Leaves opposite vs. alternat Growth orthotropic vs. ee Young plant initially with well- eee vs. shortened internodes. TRUNK AND BARK Trunk lacking buttresses, knee roots, or pneumatophores vs. with these structures presen LEAF ARRANGEMENT AND SHOOT GROWTH Internodes separating expanded leaves all of similar length vs. variation within an innovation. . Scale leaves present vs. absent. HAIR TYPE Hairs adaxially curved, rather short, and papillate at base vs. other types. TWIG AND BUD . Terminal bud functional vs. abor . Terminal bud strongly flattened (with partly expanded blade) vs. more or less rounded. . Terminal bud not enclosed in petiole bases of last pair of leaves vs. enclose Axillary buds not enclosed in pockets formed by stem and petiole base vs. enclosed. LEAF . Acute-cuneate-rounded at base vs. cordate or narrowly attenuate. . Midrib raised vs. broadly depressed. INFLORESCENCE . Terminal vs. axillary. . Branched and/or flabellate vs. unbranched and with flowers in pairs. . Flowers many vs. few or one . Axis present vs. absent. . Internodes of approximately equal length vs. some much shorter than others. FLOWER . Perfect vs. staminate or pistillate. . Tepals 8 vs. consistently fewer or more than 8. Hairs on inside of outer tepals absent vs. present. nt. . Ovary or anthers glabrous vs. pubesce 1980] STEVENS, CALOPHYLLUM [53 TABLE 3 (continued). FRuIT 24. Stone with basal plug lacking vs. present. 25. Embryo whitish vs. purple. Groups of taxa that can be recognized are discussed below and in the species accounts. Since there is no obviously ‘‘primitive’’ species of Calophyllum, the species accounts start with C. alboramulum, a taxon with terminal branched inflorescences and thin-walled stones, neither an obviously specialized charac- ter. If the presence of bud scales is considered unspecialized (an interesting suggestion; they are common in the Calophylloideae as a whole), C. garcinioides or Calophyllum sp. 73 might be also considered taxa with several apparently unspecialized characters. All these putatively primitive taxa are Indo—West Malesian, but the distribu- tion of ‘‘primitive’’ taxa is not necessarily the same as the place of origin of the genus. The same can be said for the equation of the present area of greatest diversity of the genus (and its immediate relatives), also Indo—-West Malesia, with place of origin. Although Calophyllum may have originated in the Indo-Malesian tropics—possibly by the late Eocene—and spread from there, this is only a tentative hypothesis and lacks critical support. GENERAL DISTRIBUTIONAL PATTERNS OF TAXA AND CHARACTERS The New World species must be mentioned briefly to place the discussion in context. Vesque (1893) recognized only four species from the New World, and there are clearly not many species there. The variation shown is correspond- ingly less than in the Old World: there is no variation in the distribution of leaves along the stem; there are no bud scales; the inflorescences are habitually axillary and often branched; dioecy probably occurs in some species—e.g., Calophyllum brasiliense (Pennington & Sarukhan, 1968); hair structure is basically similar (Vesque, 1893; pers. obs.); fruit variation is slight, although basal plugs (Fanshaw 2439—C. brasiliense) and thin areas near the base (Krukoff 7268—C. ellipticum Rusby) occur. The wood anatomy of New World species (Greene, 1932) is similar to that of Old World species. D’Arcy and Keating (1979) noted that C. longifolium Willd., from Panama, was similar to C. soulattri in details of lamina anatomy; from the data that they and Vesque (1890, 1893) give, species from the two hemispheres are similar in lamina anatomy. The New World species may be derived from a single ancestor originally from the Old hoes? Calophyllum has been in America at least since the Miocene (see above). In the Old World, Calophyllum has a ee icy Indo-Malesian distribu- tion pattern, with extensions to Madagascar and Fiji. The genus is basically restricted to humid, closed, lowland to colline or sometimes montane rainforest, although a few species grow in drier or more open habitats. Van Steenis 154 JOURNAL OF THE ARNOLD ARBORETUM [voc. 61 (1949) noted that in Malesia rather young individuals may flower and fruit when in the open, but there are no early successional species in the genus. Mar | shows the regional distribution of Calophyllum species in the Southeast Asian/ Australian region; distribution maps of individual species are included in species accounts. Calophyllum inophyllum is clearly a highly vagile strand species (which sometimes also grows inland), and it occurs throughout the range of the genus in the Old World. It shows a certain amount of geographically correlated variation. The Mascarene endemic C. tacamahaca seems to be closely related to it, and is perhaps derived from it (see also below). The origin and relationships of the other endemic species in Madagascar and the Mascarenes are problematic. Perrier de la Bathie (1948) thought that the species in Madagascar were derived from Calophyllum inophyllum; the adaptive radiation that had occurred suggested to him that Calophyllum arrived in the oldest of the seaborne invasions of the island. Raven and Axelrod (1974) consider that the genus arrived there by long-distance dispersal. The species growing in Madagascar and the Mascarenes show a moderate amount of diversity, but I know of no unique character that distinguishes them as a group from the rest of the genus. Although several other taxa have distribution sia like that of Calophyllum (see Perrier de la Bathie, 1948; Dejardin et al., 1973, for examples), little is known about the causes. Calophyllum en eas from the Mascarenes, is probably a local derivative of C. inophyllum, while the affinities of C. eputamen and the related C. parviflorum are unclear. India and Sri Lanka have no species in common apart from Calophyllum inophyllum, although C. austroindicum, from the Western Ghats, may be related to the complex of five species in Sri Lanka centered on C. trapezifolium. from Sri Lanka, is part of a complex also occurring from Vietnam to Australia; there are numerous similar examples, as a glance at Flora Malesiana will show. Many of the species from mainland Southeast Asia are poorly known. Calophyllum poilanei and C. membranaceum are distinctive species without obvious relationships; C. rugosum is perhaps related to the Malayan C. rotundifolium, and C. parkeri to C. fraseri and C. garcinioides, both Malesian. There are a few widespread species, and the local C. lineare is very closely related to one of them (C. pisiferum). The other species are poorly understood or are members of taxonomically difficult complexes that extend into India (C. polyanthum) and, perhaps, Malesia (as C. symingtonianum). he species of Calophyllum growing from the Isthmus of Kra in southern Thailand to the Pacific and northern Australia can be considered together. Map |. Distribution of Calophyllum in Southeast Asia, Malesia, and Austra- lia. Figures within areas: above line, endemic species; below line, total number of species. Figures on lines separating areas: number of species in common between two areas. 0 200400 800s wD ILOME TERS SINUSOIDAL EQUAL-AREA PROJECTION WNTIAHdOTVO ‘SNAAALS [0861 ssl 156 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Although Malesia as a phytogeographic unit may not include the area from the Solomon Islands eastward (e.g., van Steenis, 1950), the tropical Pacific flora is basically an attenuated Malesian one (van Balgooy, 1960), and the few Australian species of Calophyllum can also be included here; they, too, are tropical taxa. The central part of this region has usually been divided into three provinces (van Steenis, 1949, 1950, 1979; van Balgooy, 1960; van Steenis & Schippers-Lamertse, 1965): South Malesia includes the islands from the Tanimber Islands to Java; East Malesia is bounded on the west by Zol- linger’s line (i.e., it includes the oo and the Kei Islands); and West Malesia includes the rest of Males owever, as is clear from ee van Steenis (1950) and van Steenis and Schippers-Lamertse (1965) in particular, the South Malesian Province is separated because its flora is depauperate, although basically West Malesian, and has an admixture of species that do not grow elsewhere in Malesia but occur further to the north. The East and West Malesian provinces, on the other hand, are defined by the large number of genera that occur in only one province, and by the high species-level endemicity in both. Recently, van Balgooy (1976) has proposed combining the West and South Malesian provinces into an extended West Malesian Province (subsequent reference to West Malesia will be in this sense), returning more to the ideas of Merrill (1923) and other earlier workers. The distribution both of species and of distinctive characters within Calophyllum follows this east-west division of Malesia. Only six species (of which Calophyllum inophyllum is one) are found in both areas (Map 1). The variable C. calaba var. bracteatum grows from Timor westward, and var. australianum occurs in Australia; it is clearly a species with wide ecological amplitude that also can grow in drier areas. Calophyllum soulattri occurs throughout Malesia; it has small, dark blue to black fruits and is probably dispersed by birds. Calophyllum laticostatum, which occurs in New Guinea and possibly also in the Philippines, and Calophyllum sp. 129, found on the Moluccas and an island off southwestern Borneo, are related to one another and may have eastern affinities. Calophyllum wallichianum var. incrassatum is basically West Malesian but may occur on Sulawesi. Only nine species of Calophyllum are recorded from the erstwhile South Malesian Province. Calophyllum dasypodum, C. dioscurii (known from only a single, sterile specimen in Java), and C. venulosum are all restricted to West Malesia. Two species are endemic. Calophyllum grandiflorum is related to C. venulosum. Calophyllum archipelagi is probably related to the predomi- nantly East Malesian C. /aticostatum and joins the already fairly substantial list of Java-Lesser Sunda Islands endemics (van Steenis, 1979). The wide- spread, predominantly West Malesian C. tetrapterum is known from Bawean and Karimundjawa islands only; the plants of the latter island often have Bornean affinities (van Steenis & Schippers-Lamertse, 1965). Calophyllum widespread. Hence, the affinities of Calophyllum in South Malesia are predominantly to the north and west. The climate of the Lesser Sunda Islands is rather dry, and Kalkman (1955) showed that the West Malesian floristic 1980] STEVENS, CALOPHYLLUM ey element decreased in importance from west to east in these islands (see also van Steenis, 1979). The Sulawesi-Moluccas region has more species with affinities to the east than to the west. Calophyllum celebicum may be most similar to the West Malesian C. teysmannii var. inophylloide, but C. undulatum and C. articulatum are related to eastern species. In addition, six species reach their westerly limit here. However, many species from this crucial area are poorly understood, C. soulattri is very variable, and there may be additional undescribed taxa; recent collections from Sulawesi kindly sent to me by G. Musser proved embarassingly difficult to identify. Taxa from the Philippines show relationships with those of both West and East Malesia. The relationship with East Malesian taxa, which caused the Dickerson-Merrill modification of Wallace’s line so that it passed to the west of the Philippines (e.g., Merrill, 1926), has been emphasized recently by B. L. Burtt (e.g., in Ashton & Ashton, 1972, and in R. M. Smith, 1977). Calophyllum sp. 121 is perhaps related to the East Malesian C. neo-ebudicum. The conoid galls found on the leaves of C. whitfordii, in particular, are most common in East Malesian and Pacific species. However, the link with West Malesia is numerically stronger, since five species occur on Borneo and/or Palawan, as well as on that part of the Philippines that has always been insular. The distribution of five distinctive character states (Map 2) confirms this basic east-west division and reflects the occurrence of different groups of species in the two areas (cf. Stevens, 1974a). Species that have stones with basal plugs are probably under-recorded. Species with blue to blackish fruits are predominantly East Malesian; about 14 such species occur in New Guinea, but there are only four in West Malesia (C. polyanthum and C. austroindicum, from southwestern India, may also have blue fruits, but the observations must be reconfirmed). Although it is not known with certainty which character states are derived, none of the ‘‘East Malesian’’ character states is necessarily primitive (see above). Despite the lack of clear evidence, Calophyllum probably moved from West to East Malesia as continental drift brought the Australian plate near Southeast Asia 15 to 20 million years ago (Raven & Axelrod, 1974; van Balgooy (1976) and van Steenis (1979) suggest that there may have been earlier floristic contact). Diversification in East Malesia has been subsequent to this movement. The difference between the species of Calophyllum in the two halves of Malesia may be a partial reflection of this circumstance, East Malesia being colonized by only a few taxa. As has long been known (e.g., Merrill, 1926), many of the islands in West Malesia were interconnected at various times during the Pleistocene, and others were much less isolated then; most of the Philippine Islands remained separate, although Palawan was connected to Borneo. There was a similar extension of land in East Malesia, although islands such as New Britain, Sulawesi, and the Moluccas have always been isolated. Hence, the two halves of Malesia have remained separate, but the greater affinities of the species from Sulawesi and the Moluccas with East, 158 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61] rather than West, Malesian species and the relationships of those from the oe ag still pose problems. est Malesian area, 28 species are centered on the Sunda Shelf, Se on mainland Sumatra and/or Malaya as well as on Borneo, and several others are clearly related to this element. The relationships of the Malay Calophyllum-flora are at least as much with that of the relatively distant Borneo as with the closer Sumatra, a not uncommon pattern (Keng, 1970; see also the accounts of C. alboramulum and C. fraseri). Endemicity is high in Borneo, less high—and mainly in montane species—in Malaya, and low in Sumatra. Within the East Malesia—Pacific region, endemicity is highest in New Guinea, but local endemics occur throughout the area. Guppy (1906) suggested that Calophyllum burmannii (properly C. cerasi- ferum) and C. spectabile (properly C. vitiense), both from Fiji, were dispersed there by fruit pigeons, rather than being derived from the local C. inophyllum, also from Fiji; Smith and Darwin (1974) thought that at least some species in the Fiji-Samoa area might have arisen from C. inophyllum. Although C. vitiense and C. cerasiferum may be derived from a taxon other than C. inophyllum, all three taxa have stones with basal plugs; such taxa preponderate in East Malesia and the Pacific, so the origin of C. inophyllum itself may have been in this general area. If the suggested relationship of C. leucocarpum to Papuasian species is correct, there may be three independent lines of the genus on Fiji, probably reaching the archipelago by island-hopping in the general path Solomon Islands-New Hebrides—Fiji (Smith, 1979). EcoLocy Several taxa of Calophyllum in Indo-Malesia have wide ecological ranges. These species often, but not always, occupy a wide geographic range, but the amount of morphological variation within the taxa differs. The i asin predominantly West Malesian C. pisiferum, C. rupicola, C. biflorum, C. ca C. tetrapterum var. seuap icra, C, Soules o fe neon vars. tsa and inophylloide, and C. lation, which at least in C. canum, C. pisiferum, C. rupicola, C. soulattri, C. nee: var. teysmannii, and C. venulosum is partly correlated with ecology. Unfortu- nately, details of the ecology of the highly variable C. blancoi in the Philippines are practically unknown. The Papuasian C. sil and C. vexans also show a considerable amount of infraspecific and ecological variation and likewise inhabit a considerable area; morphological variation is correlated with geog- raphy in both species. Calophyllum sp. 151 (a local species), C. depressinervo- sum, C. pulcherrimum (all basically West Malesian), C. papuanum, C. peekelii, and C. neo-ebudicum (all basically East Malesian) grow in a variety of habitats but show less infraspecific variation Map 2. Distribution of five charac ter states in Malesian species of Calophyi- lum: ——, scales at bases of pee innovations and inflorescences; ——— X ——.,, terminal inflorescences (to India and possibly the seu —— xX , angled stones and/or flattened terminal buds; gs at bases of stones (to Fiji and Samoa, excluding widespread G eR and its derivative, C. tacamahaca; also in South America). SCALE | ° 700 400 #oo, 800.1000 ROME TE SINUSOIDAL EQUAL-AREA PROJECTION WOTIAHdOTVO ‘SNAADLS [0861 6S1 160 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 A few Bornean species (e.g., Calophyllum hosei, C. scriblitifolium, and perhaps C. sclerophyllum) grow both in peat swamps and in the apparently very different kerangas forests on leached, arid sands. Such distributions are rather common (Brunig, 1974) and reflect certain essential ecological similarities, such as extreme oligotrophy, in the at first sight rather disparate environments. our out of the eight species (Calophyllum neo-ebudicum, C. peekelii, C. soulattri, and C. vexans) from the Solomon Islands have wide ecological tolerances, perhaps a reflection of the depauperate flora there (Whitmore, 1969). Most Malesian taxa have more restricted ecological ranges. Many grow in more or less well-drained, mixed dipterocarp forest and in its East Malesian equivalent. Within this broadly defined habitat, a few species such as Calophyllum macrocarpum seem to prefer stream sides (without being rheo- phytes); others, like the four species from the Solomon Islands studied by Greig-Smith, Austin, and Whitmore (1967), may show more subtle preferences (but see Stevens, 1974a; also below). Calophyllum nodosum occurs most frequently in forests growing on rather acid soils and is locally abundant in extreme kerangas vegetation; it also grows in submontane vegetation. It shows considerable variation in foliar characters but little in those of flower or fruit. Calophyllum obliquinervium also grows most commonly in forests on rather acid soils; it grows in a number of other habitats but shows little variation. Several species (e.g., Calophyllum collinum, C. costatum, C. exiticostatum, C. incumbens, C. milvum, C. papuanum, C. polyanthum, C. symingtonianum, C. trapezifolium, C. thorelii, C. undulatum, and C. vitiense) grow in colline forest up to about 1300 meters in altitude. Another group of species, including C. cerasiferum, C. garcinioides, C. hirasimum, C. rigidulum, C. rotundifolium, C. pauciflorum, and C. walkeri, occurs predominantly above 1300 meters. As might be expected, the species that grow at high altitudes tend to have rather short, broad, thick leaves that are not much pointed at the apex. Since most of these taxa in Malesia seem to be independently derived from lowland taxa (cf. Whitmore, 1973), they have little else in common; the species growing at high altitudes in Sri Lanka are closely related to each other. There is no noticeable increase in indumentum with altitude. Some species grow in other, rather specific habitats. Several West Malesian species (e.g., Calophyllum ardens, C. sclerophyllum, C. sundaicum, C. scriblitifolium, and probably also C. praetermissum) grow in peat swamps, and others like C. lowii and C. mukunense grow in ‘“‘periodically inundated”’ forest. The only fairly well-collected East Malesian species that habitually grows in swamps or periodically inundated ground is C. suberosum. A few species that have restricted distributions grow in forest over limestone or ultramafic rock. Calophyllum aurantiacum and C. calcicola have been recorded only from the former, while C. complanatum, C. sakarium, and Calophyllum spp. 98 and 143 have been recorded only from the latter. A number of additional species sometimes grow over ultramafic rock. A few species grow in drier forests: C. calaba var. calaba from Sri Lanka is the best example, 1980] STEVENS, CALOPHYLLUM 161 and C. sil (from Australasia) is usually found in gallery forest. Three taxa are rheophytes: C. pisiferum (not always, and only in mainland Southeast Asia), the closely related C. lineare, and the typical form of C. rupicola. Calophyllum inophyllum is characteristically a strand plant; it shows a certain amount of ecogeographic variation. In many places, especially swamp forest, well-drained lowland forest, and ridges in lowland to montane forest, several species of Calophyllum may of species have been observed in the swamp forests of southern Johore (Malaya) and near Kuching (Sarawak); in ridge forests throughout Malesia— notably Malaya, Sarawak, and Sabah (even on ultramafic rock); and in mixed forests in Papua New Guinea (there is a particularly interesting area in lowland forest around Kiunga, where at least 12 species of Calophyllum occur) and New Britain. EvoLuTION The discussion in the preceding section in a way epitomizes the problems that arise in discussing the evolution of the genus. Although species may have very distinct ecological preferences, this is not always so, and obvious “‘adaptations,’’ even to very different habitats, are few. Possible exceptions are the small, broad, thick leaves of the montane species, the narrow leaves of the rheophytic species, and the stilt roots of many of the swamp-dwelling species. There are no major discontinuities in the pattern of morphological variation in the genus (see also van Steenis, 1968), and even if all the species in the genus except for two were unknown, these two would probably still be placed in the same genus. The variation in the genus is almost kaleidoscopic (Cullen, 1968) and on a rather restricted theme, yet numerous taxa can be recognized. These taxa frequently grow together (the “‘sympatry’’ of the following discussion) although they do not often hybridize. It is very difficult to understand both the evolution and the maintenance of this species diversity. Much of the variation appears to be without adaptive significance and so might seem to confirm some of van Steenis’s ideas on evolution in the tropics (see van Steenis 1977, 1978, and references therein; Fedorov, 1977, also leans toward such a view). However, this position can be supported only by a much more detailed knowledge of plant groups than is currently available, and the only reasonable position in the absence of such knowledge is that of an agnostic. One cannot expect a revision like the present one to resolve this important issue, but the suggestions made below may bear closer examination The morphological variation of the Sioa and young plant in Calophyllum is considerable; the pigmentation of the young leaves also varies at this Oo with germination, there is little variation in embryo morphology. The stem anatomy of the seedling has not been examined, but there is certainly less variation in lamina anatomy at this stage than occurs in the adult (see also 162 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Stevens, 1974a). There is no obvious correlation of the variation of these young stages with ecological parameters. Unfortunately, few genera have been studied in comparable detail. Diospyros shows much variation in germination and seedling, and Wright (1904) keyed out the species from Sri Lanka using characters of fruit and seedling. Maury (1978) studied many aspects of the fruit, germination, and young plants of even when still young plants. De Vogel (1980) does not mention any comparable examples. There is considerable selection pressure on a plant in the early stages, and the environmental requirements for germination and establishment are often very precise. Grubb (1977) has particularly emphasized the importance of the “‘regeneration niche’ (the environment of the very young plant) in the maintenance Rae species diversity, even suggesting that species in such genera as Diospyros, Rinorea Aublet, Shorea Roxb. ex its tner, and Eugenia L. eal eee may differ by only a single character in the regeneration ae it might also be argued that relatively little variation is likely at this stage, with physical factors such as temperature and humidity showing relatively less variation during times when germination is possible compared to later stages. Character complexes adapted to the conditions occurring then will not change readily; at least, there will not be a complete reorganization, since a major change would destroy the developmental pattern at a vulnerable stage. Similarly, genes acting late in development will often not show their effects at these earlier stages (Stebbins, 1950, 1974 Study of Calophyllum may produce evidence relevant to this problem. It is possible that some of the characters apparent throughout the life of the plant are important only at the younger stages, being selectively neutral later on. Young plants of C. pulcherrimum, bustum, and C. rigidum all have plagiotropic leading axes, and the adults have more or less pendulous branches. One might ask at what stage, if any, this is of functional significance (see Givnish, 1978, for discussion on ene possible Selective advantage o compound leaves; insome respects ulv to a compound leaf). Other characters, like leaf number snd disposition in the seedling, and the initial period of slow growth as well as the distinctive leaf shape in the young plant can be of significance only at early stages. In this context, the populations of C. soulattri from Manus to the Solomon 1 the only major differences between the two appear to be in fruit, seedling, and young plant. Another interesting case is C. sclerophyllum, which is sometimes superficially similar to C. teysmannii var. inophylloide, although ” White (1962, 1978) noted that species of Diospyros in Africa belong to many sections, and sympatric species are not usually closely related; species in genera like Shorea are delimited on characters other than those of the regeneration niche. 1980] STEVENS, CALOPHYLLUM 163 the seedlings, young plants, and anatomy of the two are aes different. Finally, one of the more important distinctions between C. pers mile and in the “‘regeneration niche’’ may be of considerable importance in both t development and maintenance of diversity in Calophyllum. In the adult plant, the considerable variation in the disposition of the leaves along each innovation, the branching pattern (monopodial vs. sympodial), and the way the leaves (notably erect in some taxa, such as Calophyllum peekelii, but usually spreading to pendent) and branches are held must all affect how leaves intercept light. Again, comparison with other tropical genera is difficult because such characters are not often observed, but similar variation seems to occur in tropical montane Ericaceae (Stevens, 1980a). In two cases the growth pattern of the plant may be connected with its preferred habitat. The trunk is less strongly orthotropic in Calophyllum inophyllum than in other species, and individuals may start to lean when only a few meters tall. This relatively weak orthotropy, doubtless aided by wave action may cause the trunk to become Proc cumbent on the beach, and branching is profuse. However, like other species with the same branching pattern, C. nodosum may also be a sizable tree; species with functional terminal buds may also be shrubs. The variation in hair type is considerable, although it is variation on a rather limited theme. Hairs seem to protect the resting bud, since in those species in which the buds are covered by thick perulae, or protected in pockets formed from the stem and petiole (e.g., Calophyllum savannarum), the buds tend to be glabrous. The brown contents of the hairs, more obvious in some species than in others, are probably tannins—protectants against e empty. thick walls of the apical cells of C. rufigemmatum clearly increase the physical protection of the bud, as may the small, but very papillate and thick-walled, hairs of species such as C. subsessile and C. persimile. Although the fruit is always drupaceous, variation is considerable. Possible dispersal agents are summarized here; however, careful observations are still sadly lacking. A few species are habitually or occasionally dispersed by water, ay of the seeds. Unfortunately, I do not know of other groups that have more blue-fruited taxa in East Malesia than in West Malesia. Closely related species usually differ only slightly in hair type, fruit structure, germination, and young plant. Such species are usually allopatric, and they 164 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 may also be ecologically differentiated. Examples are listed in TasLe 4, although this table should really be treated as a list of cases worthy of closer investigation. As noted above, variation within many species shows a pronounced correlation with geography, and the varieties of most species that are so subdivided usually grow in different areas (infraspecific variation in both C. teysmannii and C. tetrapterum is not quite so clear-cut). Many of the observations and suggestions made above are compatible with the idea that speciation in Calophyllum is basically allopatric; sympatry occurs only after subsequent divergence. Morphological confusion nearly always occurs between allopatric, rather than sympatric, taxa; hybridization between sympatric taxa is not common. Closely related taxa are separated by eco- geographic barriers, whether or not there are also reproductive barriers. This series of generalizations agrees with observations on West Malesian Dip- terocarpaceae (Ashton, 1977, 1978), Diospyrosin humid tropical Africa (White, 1962, 1978; the sympatric taxa often belong to different supraspecific taxa), and Drimys J. R. & G. Forster in Papuasia (Vink, 1970); it disagrees with Fedorov’s ideas on speciation (Fedorov, 1966). The four species of Calophyl- Tas_e 4. Geographic and ecological preferences of closely related taxa of MADAGASCAR C. verticillatum/C. chapelieri. Usually grow in different places, but some overlap; intermediates? Sri LANKA C. walkeri/C. trapezifolium/C. cuneifolium/C. th ee vergens. First three have largely different altitudinal roe C. ple Cc trapezifolium at least sometimes grow e general area. calaba var. calaba/var. pening Ecological ie aod but oe rare overlap. InpIA—CHINA C. polyanthum/C. touranense/?C. balansae. Largely grow in different areas; tara within C. polyanthum has pronounced geographic component. VIETNAM—MALESIA C. calaba var. bracteatum/var. cuneatum/var. E./var. australianum. Var. cuneatum ecologically separated; vars. E and australianum geographically separated. C. pisiferum/C. dispar/C. lineare. Never grow together, but no obvious ecological preferences; taxonomy of group not satisfactory. West MALESIA C. ferrugineum/C. biflorum/ C. sundaicum /C. costulatum. The three varieties of C. ferrugineum grow in different areas, var. orientale 1980] STEVENS, CALOPHYLLUM 165 TABLE 4 (continued). ecologically differentiated as well; C. sundaicum ecologically differentiated; C. biflorum largely geographically separated; C. costulatum poorly known; C. ferrugineum var. ferrugineum and C. biflorum grow together in hill forest, S. Johore; C. ferrugineum var. ferrugineum, C. sundaicum, and C. costulatum (variant) grow together in swamp forest, S. Johore. C. wallichianum var. wallichianum /var. incrassatum/ var. tahanense. Vars. wallichianum and incrassatum basically geographically separated, var. tahanense in part ecologically C. gracilipes/C. recurvatum/C. aurantiacum. C. gracilipes and C. recurvatum Saeed separated, C. aurantiacum ecologically. C. roseocostatum/C. stipi tatum/C. elegans / Calophyllum sp. 65/?C. pyriforme. No two taxa found in one place, but both C. roseocostatum and C. elegans occur near Kuching. C. venulosum var. venulosum / vat. tenuivenium /C. subsessile/C. grandiflorum. No taxa in this complex grow together, yet no major geographic isolatio C. nodosum/ C. fee re /C. pulcherrimum. Usually grow apart, although sometimes not far distant; all grow together at Sabal Tapang, Sarawak; C. ae often on more acid soil than other two, which differ notably in C. teysmannii £G: sterphy lum. mare ecologically isolated; when growing together (swamps, Johore), form of C. teysmannii involved is that least similar to C. sclerophyllum; Pee of C. teysmannii not sympatric East MALESIA C. pean parvifolium/C. caudatum/C. novoguineense/C. olor/C. confusum/C. bifurcatum/C. leucocarpum. Geographically Benen although C. novoguineense and C. bicolor grow together near Kiunga. C. trachycaule/C. goniocarpum/C. piluliferum/C. heterophyllum/ Calophyllum sp. 143/C. rufinerve. ee geographically separated, but C. trachycaule and C. goniocarpum grow together near Kiunga, Calophyllum sp. 143 and C. goniocarpum in same area near Buso; taxonomy of group poorly understood. C. laticostatum/C. insularum/Calophyllum sp. 129/C. sil/C. collinum. First three geographically separated and closely related; second two less closely related; C. sil sensu stricto ana differentiated: C. collinum and C. laticostatum sympatric at Kiun C. papuanum/C. pauciflorum/C. vexans. Vaigely geographically ipa also ecologically; C. ale and poorly understood form of C. vex grow together near Kiun Cc. _suberosum /C. Se nie. ‘Ecologically separated; considerable differences Ss. C. neo-ebudicum/C. cerasiferum/C. vitiense. C. cerasiferum usually at highest Sidiades. otherwise no major geographic separation. 166 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 lum that Grieg-Smith et al. (1967) thought were examples of closely related species having different niches are perhaps better considered as examples of not immediately related species that frequently can grow together. These species—C. soulattri, C. vexans (their C. cerasiferum), C. neo-ebudicum (their C. vitiense), and C. peekelii (their C. kajewskii)—quite frequently grow together, as can be seen from the data given by Grieg-Smith ef al., and as I have observed in New Britain. It is difficult to compare the amount of differentiation between allopatric and sympatric taxa within a genus, let alone between different genera (see also below), but in both Calophyllum and Drimys even sympatric taxa may be rather similar (variation in seedlings and young plants of Drimys is not great (pers. obs.)). However, current knowledge of Calophyllum, and of most tropical groups, rather vitiates this type of analysis. As mentioned above, subtle, but discontin- ce sympatric variation may escape recognition because of the small sample ize from most localities and the practical limits on field work in the area; Secs correlated variation may be overemphasized because of patchy collecting. The variation of C. biflorum and C. teysmannii var. inophylloide on Gunong Matang, Sarawak, is pertinent here, since the differences between the two ‘‘forms’’ of each of these taxa on this hill are similar to those that characterize allopatric varieties in other species. Although this variation seems to intergrade with that in the rest of the taxa, field experience throughout Borneo might show that all specimens could be referred to these two ‘‘forms’’ the separation of C. persimile from C. suberosum was suggested by a field observations. Hence any explanation of the patterns observed in terms of processes can only be speculative, but one can at least identify areas for future work. ACKNOWLEDGMENTS I have become deeply indebted to numerous people during the course of this study; to all those who pega queries, provided information, and elped in other ways, but who are not thanked individually, my sincere thanks. I am very grateful to the nee of the many herbaria that I have visited, or which have sent specimens on loan, for permission to examine material held by their institutions,"° and I am particularly grateful to the directors of the herbaria at Bogor, Kepong, Kuching, Lae, Sandakan, and arawak for their hospitality and for enabling me to see so many taxa in the field. Special thanks are due to Dr. L. M. Perry and Dr. S. A. Spongberg, who have consistently encouraged me in this work; the staff and students of the Harvard University Herbaria as a whole, for helpful discussion and for putting up with my periods of preoccupation; and two reviewers of the manuscript, for their very useful comments. Mrs. L. Dickinson and the staff 10 AAU, B, BISH, BKF, BM, BO, BRI, C, CAL, CANB, CHR, DNA, E, F, GH, K, KEP, KLU, LY, M, MASS, MAU P, PERTH, PNH, SAN, SAR, "SING, SYD, TAI, U, UC, US, W, WRC Gervais follow ee & Keuken, 1974). 1980] STEVENS, CALOPHYLLUM 167 of the libraries of the Arnold Arboretum and Gray Herbarium, Mr. M. Canoso, Mr. W. Kittredge, and the herbaria staff have been unfailingly helpful. To Dr. R. A. Howard and Dr. P. S. Ashton, past and present directors of the Arnold Arboretum, are due my thanks for their support, and to the latter my perhaps somewhat qualified thanks for putting at my disposal his collection of almost 300 specimens of Calophyllum collected during the course of his ecological work in Sarawak. Most of the maps are Goode’s series of base maps, © University of Chicago. A welcome grant from the Atkins Fund defrayed some of the expenses of the field work. Robin Lefberg and Karen Stoutsenberger made the illustrations, for which my thanks. Lisa Frost, Kathy Holland, Laura Sahagian, Jennifer Snyder, and Beverly Vincent have typed drafts of the manuscript and have dealt successfully with rather cryptic handwriting. Last, but by no means least, I am very grateful to Elizabeth B. Schmidt for her careful editorial work. SYSTEMATIC TREATMENT Calophyllum L. Sp. Pl. 1: 513. 1753, Gen. Pl. ed. 5. 229. 1754; Lam. Encycl. Méth. Bot. 1: 552. 1785: Juss. Gen. Pl. 286. 1791; Willd. Sitz-ber. Ges. Naturf. Fr. Berlin Mag. 5: 78. 1811; Choisy, Mem. Soc. Hist. Nat. Paris 1: 228. 1823, in DC. Prodr. 1: 562. 1824; Endl. Gen. PI. 1028. 1840; Choisy, Descr. Guttif. Inde, 41. 1849; Migq. FI. ie Indié 1(2): 509. 1859; Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 241. 1862; Bentham in Bentham & Hooker f. Gen. ae 1; 175. 1862; T. Anderson in Hooker f. FI. Brit. India 1: 271. 1874; Vesque, Epharmosis 2: 6. 1889; King, Jour. Asiatic Soc. Bengal, II. 59: 172. 1890; Vesque in C. DC. Monogr. Phanerog. 8: 529. 1893; Trimen, Handb. Fl. Ceylon 1: 98. 1893; Engler in Engler & Prantl, Nat. Pflanzenfam. 3(6): 220. 1895; Pitard in Lecomte, Fl. Gén. Indo-Chine 1(4): 316. 1910; Ridley, Fl. Malay Penin. 1: 181. 1922; Engler in Engler & Prantl, Nat. Pflanzenfam. ed. 2. 21: 192. 1925; Gagnep. Fl. Gén. Indo-Chine Suppl. 1(3): 268. 1943; Perrier de la Bathie, Fl. Madagascar Comores, Fam. 136: 3. 1951; Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 285. 1956; Maheshwari, Bull. Bot. Survey India 2: 139. 1960; Backer & Bakh. f. Fl. Java 1: 384. 1963; A. C. Sm. & Darwin, Jour. Arnold Arb. 55: 216. 1974; P. F. Stevens, Austral. Jour. Bot. 22: 349. 1974. Type: Calophyllum inophyllum L."' Ponna Rheede ex Ludwig, Defin. Gen. ed. 3. 239. 1760. Nomen superfluum. Calaba Plum. ex Adanson, Familles 2: 446. 1763. Nomen superfluum. Augia Lour. Fl. Cochinch. 337. 1790, pro minore parte. Balsamaria Lout. Ibid. 467. Tyre: Balsamaria inophyllum Lour. Apoterium ae Bijd. Nederl. Indie 1(5): 218. 1825. Type: Apoterium sulatri Blum "' Calophyllum is usually typified by C. calaba, but that species does not agree with the generic description in Linnaeus, Gen. Pl. ed. 5. 229. 1754. Fuller details will be given in a later article. 168 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Evergreen trees or shrubs, sometimes buttressed or with stilt or loop roots; bark often with characteristic diamond-shaped fissures becoming confluent with age; latex present, nearly always colored. Twigs + flattened and angled; terminal buds usually naked, sometimes with scales, rarely aborting and not functional; indumentum of uniseriate hairs usually at least on buds. Leaves decussate in adult plants, very rarely alternate in young plants, petiolate; lamina entire, + coriaceous, with close parallel venation alternating with and usually more prominent than latex canals. Inflorescences terminal and / or axillary, with (1 to) 3 to numerous flowers, branched or not, axes terminated by flowers; bracts usually deciduous. Flowers pedicellate, usually her- maphroditic; tepals 4 to 16, outer one or two pairs rarely much different from the rest; stamens numerous, at most obscurely fasciated, the filaments usually only slightly connate at base, the anthers small, basifixed, dehiscing by long, lateral slits; ovary unilocular, with single basal, anatropous ovule, the style present, the stigma + expanded, often peltate. Fruit drupelike, the pericarp consisting of exocarp and well-developed, fleshy to fibrous mesocarp, the testa consisting of stony layer and usually transient spongy layer; seed single (except in polyembryonic taxa), radicle just to one side of base of fruit, cotyledons large. Germination cryptocotylar, epicotyl short. 2n = 26 (one species). About 187 species, 179 in the Old World, predominantly Indo-Malesian, ca. 8 species in the New World, from Mexico and the Caribbean to Argentina. Most species are small to medium sized or sometimes large trees of humid, tropical, predominantly lowland rain forest, but some species grow in mid- montane or more seasonal forest The wood of the larger species is used quite often in construction; where Calophyllum is abundant (e.g., Malaya, the Solomon Islands), it is an important component of the timber trade. The fruits of a number of species yield an oil that is of some economic use, either as a medicine or in lamps (see especially C. inophyllum). Calophyllum has been used in canoe manufacture throughout its range, the seaside C. inophyllum being particularly eta in this respect, and in the era of sail several species furnished masts and s Local names piven to the species usually seem to have little speci the generic name, ‘‘binta(n)gor,’’ is much used in West Malesia, often qualified (e.g., ‘“‘bintangor batu’’ is the local species with hard wood) GuipE To Use oF Keys AND DEscrIPTIONS The characters used and their terminology have been extensively discussed in the preceding pages. However, certain aspects bear repeating and expanding here. Venation density should be measured in the middle part of the leaf away from any area of damage. Venation prominence is a qualitative character; however, Ficure 7, a and c, gives an approximate indication of the extent of the variation. Deciding whether or not the lamina immediately adjacent to the midrib is raised and is thus apparently continuous with the midrib is sometimes not easy; Ficure 7, c should help. It is easy to overlook the 1980] STEVENS, CALOPHYLLUM 169 a 2 TTT Be ey Hid 3 2 a bis es — - mEeTRic | 1 is GURE 7. Some Sr ae as shown on dried specimens. a, Calophyllum Rig nana (FRI 13796), node with transverse line, lamina at lower left, scure venation. b, Calophyllum sp. 65 (S 23854), scars left by scales at hah & k u na C. tetrapterum var. pee € (S 27894, leaf on right), midrib sharply demarcated from surrounding lamina, thickened lamina margin (double arrow). d, C. tetrapterum var. pre e (S 23286), nodes lacking transverse lines or markings (arrows). 170 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 scales that are sometimes present at the bases of the axillafy innovations (Ficure 7 The species descriptions are generally neha: Parentheses enclose less commonly encountered variation. The descriptions of the bark are uneven, with much information having been ne from field labels and should be read with caution; descriptions for the species that I have seen are somewhat more comparable. Measurements were taken from dried specimens, unless otherwise mentioned, although flowers were boiled in water before measure- ments were taken. It is not easy to see the obscure horizontal lines that occur at most nodes in some species (FicureE 7, a, qd), sas they are noted only when they have been seen; since I became aware of them only at a rather late stage in the revision, they are probably hadenee sed (as perhaps in the Madagascan taxa). The color of the dried petiole is mentioned only when it contrasts very strongly with that of the stem, and the whitish, probably waxy, covering of the dried leaves of a few species is mentioned only when it occurs. A few species have leaves that dry longitudinally concave, or sometimes boat shaped; this is mentioned only when it is prominent. Unusually well-developed marginal thickening (Ficure 7, c), prominence of the latex canals, and notably thin petioles or pedicels are mentioned only when they occur. The number of flowers in an inflorescence is usually an uneven one, and the number of tepals in a flower is usually even, but this fact is not mentioned in the descriptions for the sake of brevity. The pedicel generally thickens somewhat in fruit, but in a few species this is exceptionally notable e can be assumed that all tepals are fringed with hairs but are otherwise eee that all stamens are glabrous and have filaments that are slightly one at the base, and that all ovaries are glabrous. Plugs at the base of the stone are recorded only when they occur; these, too, are probably underrecorded. Full literature citations have been given for each name (except for Calophyl- lum inophyllum) so that the reader can find his way through the often confusing earlier literature. All specimens cited have been seen, unless otherwise nee 1 . is added only in the citation of type collections. The abbreviations of these institutional series basically follow those suggested by Jacobs (1968).'* Local '? However, I have cited CF numbers as KEP (Malaya; they are part of the same SAN). Other abbreviations used here are FHI (Forest Herbarium, Ibadan, Nigeria), SF (Service Forestiére) and RN (Resources Naturelles) (both Madagascar), MAU (Mauritius), CP (Ceylon Plants), FHP (Flora of Hassan Project, India), BSI (Botanical 1980] STEVENS, CALOPHYLLUM 171 names are usually not given because of the problems mentioned earlier; people interested in such names should write to the author for a list. Herbarium abbreviations follow those in Holmgren and Keuken (1974). Keys The keys presented below have been made as practicable as possible, hence the emphasis on vegetative characters and also the rather frequent keying out of a number of taxa more than once. A single key to all the species recognized has not been presented, since if the characters used were those on the ordinary specimen the key would be so long due to multiple entries that it would be very unpleasant to use. Characters of inflorescence, flower, and fruit, which often most readily characterize a species, are not observable on most specimens, and a key using such characters would allow identification of only a minority of the specimens. A multiple-entry key could be produced, although many characters do not lend themselves readily to the either / or scoring necessary for such a key and many taxa are incompletely known; in addition, its use would be difficult if the user treated it as a key with single character dichotomies, as I suspect would be the case. Therefore, seven keys are presented. The areas they cover are: 1) Africa, Madagascar, and the Mascarenes; 2) Sri Lanka and peninsular India; 3) mainland Southeast Asia (Burma, Thailand, and the Nicobar Islands, to China); 4) Malaya and Sumatra to the Lesser Sunda Islands; 5) Borneo; 6) the Philippine Islands to the northern Pacific; and 7) Sulawesi to Australia and the southern Pacific. Survey of India), CCC (Canton Christian College, China), KLU (University of Malaya, Kuala Lumpur), NGBF (New Guinea Biological Foundation), and FDA and FDF (Fiji Departments of Agriculture and Forestry, respectively—see A. C. Smith, 1979). A complete identification list can be obtained from the author. —_" 1. Key to Species of Calophyllum in Africa, Madagascar, and the Mascarenes . Innovation with terminal 2 (or 3) pairs of leaves almost pseudoverticillate. 2. Lamina (2-)3-5.3(-8) cm. long; inflorescence usually 5-flowered See eae eae ee eee 34. Ae verticillatum. Lamina at least 8.5 cm. long; inflorescence with 7 to 11 flowers. .............. 0.00200. . C. chapelieri. — flower ik 4 tepals [introduc : Innovation with each pair of leaves clearly separated by inte rode. a rather thinly coriaceous, drying dull greenish San. closely undulate at margin, with at least 13 veins/5 mm.; Ala. ei Ue dels ge Ge ae et a we a ee Be 21. soulattri. 3. Lamina rarely thinly coriaceous, if so then venation density less, otherwise not as above; flower (where known) with (7 or) 8 or more tepals. 4. Lamina shallowly cordate at base; petiole almost absent... ..................200.4 27. C. lingulatum. 4. Lamina at most rounded at base; distinct eda ale presen 5. Axillary innovation with leaves + restricted to top half, basal internode much longer than others [lamina less Nn than 7 cm. long, venation usually + ies 6. Terminal bud 5-7 mm. lon ng oe ey ee ee Bo es a ee 26. C. milvum. mm. lon 7. Lamina broadly Paced. at base [stone wall less than 0.3 mm. across] 7. Lamina cuneate to acute a 8. ae + spherical, stone wall ¢ ca. 2 mm. thick; terminal bud 1-2 mm. long. ......... 30. C. pervillei. ovoid, or stone wall less than 0.5 mm. thick, or terminal bud 3-4 mm. lon . 30. C. aff. pervillei. ; Axillary Gestion with leaves + regularly scattered along its length, basal internode not notably longer than other o; Terminal bud 3.5-4 mm. long; lamina 4.3-7.8 cm. long; inflorescence apparently terminal [imperfectly known; Comore islands)|;.2-2:3425 4.6 Be ee sch bet eb ede Se bee ee See ee tees ws 40. C. comorense. 9. Usually not this combination of vegetative characters; inflorescence rarely terminal. 10. Terminal bud less than 4 mm. long; lamina usually less than 6 cm. lon . Wenation on lower surface of lamina + flat to een edges of midrib subimpressed; fruit with strongly fibrous outer layer, fibers persisting around stone. ................ 35. C. fibrosum. . Venation on lower surface of lamina usually + es ae of midrib raised; fruit lacking persistent ibers — —" — — CLI WOLAYOdUV ATONYV AHL JO TVNANOL 19 “Toa] 12. Lamina 0.9-3.7 cm. long, with 13 to 19 veins/5 mm. ................ 31. C. drouhardii. 12. Lamina usually more than 2.5 cm. long, with up to 12 (to 14) veins /5 mm 13. Lamina 5-11 cm. long, acuminate at apex, angle of divergence of venation G0 210] walle er en 3 g..% Rene mgr at ephe Te ee Sieh Fe cA wei Eade, Eimear a eee TA ote Ee te ae Ao ah dee Jeena 42. C. vernicosum. 13. Lamina to 6 cm. long, acute to rounded at apex, angle of divergence of venation 30—60(—65 14. Angle of divergence of venation 20-40°; lowest internode of inflorescence ca. 2.5 cm. long Piaget Win ka wih |i ae oe tht pecs oa ony on aa ay te ras eet cts eats ee 29. C. laxiflorum. 14. Angle of divergence of venation at least 50°; lowest internode of inflorescence at most 1(-1.5 MOTs die ing wes Oy ke eae Se ee Sa eke Gal tin Sct One cat C. aff. pervillei. 10. Terminal bud at least 4 mm. long; lamina often more than 6 cm. long. 15. Lamina obovate to elliptic, with 4 to 8 (to 10) veins /5 mm., venation clearly raised; aa (1.5-)2.5-4.5 em: Jone: fuistally seaside plant)... «452 4 eh 6 SG-53446S AS he EEE E SG 8 . C. inophyllum. 15 . Lamina usually ovate to elliptic or aes: ta : to) 7 or more veins/5 mm., ieee often not m. lon very clearly raised; pedicel up to 2.5(-3.5 16. Lamina acute to acuminate at apex 17. Terminal bud 1.1-2.4 cm. long: lamina somewhat more than 3 times longer than broad....... Gh St ae ah A a Ge eS ee ee Coe oe es ee ee Cc a al Ly, Terminal bud less than 8 mm. long; lamina somewhat less than 3 times longer than broa thick, the stone wall 0.3-0.6 mm. thick, lacking basal plug. ......... C. recedens 18. Lamina acute at apex, with (6 to) 8 to 11 veins/5 mm.; fruit with outer layer 2.5-4 m thick, the stone wall 0.9-1.5 mm. ia with basal plug. ......... 39. C. tacamahaca. — On Lamina obtuse to rounded or retuse at a 19. Lamina drying + flat, midrib depressed fruit with stone wall at most 0.2 mm. thick. rae 5.7-17.5 cm. long; twigs 2.5-5 mm. across; outer layer of fruit at least (?2-)3 mm. TONS cline seen Rae Ban sce Bete nA an ct Gatien teed enemas tc is Wana Uae da Mes 36. C. eputamen. 20. Tata 2-4.8(—7.5) cm. long; twigs 1.3-3 mm. across; outer layer of fruit less than | mm. PIG ee ig 2 oh as a, Se hae pt oss he ew a Ye OS DO ee 37. C. parviflorum. 19. Lamina usually drying with recurved margin, often longitudinally concave toward center, midrib above usually with at least margins raised; fruit with stone wall at least | mm. thick........ . Sse See ee ee See Se ee eee ee ee ae 32. C. aff. paniculatum. [0861 WOTIAHdOTVO ‘SNAAXRLS 2. Key to Species of Calophyllum in Sri Lanka and Peninsular India 1. Inflorescences terminal [lamina at least 6 cm. lon 2. Lamina rounded to cordate at base; indumentum tomentose. bo: Get ee ode Ge ee ee a 89. C. cordato-oblongum. mina acute at base; indumentum PUBCIUIEN, goo 43a e oe SE oR BE we 4. C. polyanthum. — , Inflorescences axillary. 3. Terminal bu d 1.2-2 cm. long; fruit with stone retuse at apex. .. 2... .0000000.0.... 04 bey 25. C. moonii. 3. Terminal bud less than | cm . long; fruit with stone rounded to acute at apex. . Leaves varying greatly 1 in size along each twig [internodes less than 2 cm. long]. a ee ae ee 20. C. bracteatum. 4. Leaves not varying greatly i in size along each twig. 5. + Persistent, conspicuous, tomentose indumentum on terminal bud, stem, and inflorescence axis Fp eares ovate to elliptic, with 5 to 11 (to 13) veins/5 mm., angle of divergence 45-65°]. ............ 11. omentosum 5. Transient to + persistent, puberulent to subtomentose (very rarely tomentose) indumentum, teers on stem and usually restricted to base of inflorescence axis. 6. Lamina at least 7 cm. long, with 5 to 8 (to 10) veins/5 mm., veins prominent on both surfaces; pedicels 1.5—5 n cm . long; stone with basal plug... 2.2... ee C. inophyllum. : Lamina less than 7(-10) cm. long, venation density and prominence variable; pedicels 0.4-2. 6(- 3.5) cm. long; stone lacking basal plug. De ~ Twigs strongly flattened; internode often more than 2 cm. long; uppermost pair of axillary buds often erect and conspicuous, ca. 1/4-1/2 length of terminal bud; internodes often more than 2 cm. long. 8. Lamina rounded (slightly retuse) at apex, shallowly cordate at base. ............. 17. C. vergens. 8. Lamina acute to retuse at apex, acute to cuneate (rarely rounded) at base. 9. Pedicel at anthesis less than 0.5 mm. across, in fruit less than 1 mm. across; flower with 4 tepals; fruit with thin outer layer becoming + disorganized Dy air SPACES. be et oe we We ew 19. C. calaba. 9. Pedicel at anthesis more than 0.7 mm. across, in fruit ca. 1.5 mm. across; flower with ca. 8 tepals; fruit with well-developed, compact outer layer... ........2...2.2.2.2.. 14. C. trapezifolium. . Twigs usually at most slightly flattened; uppermost pair of axillary buds rarely erect or conspicuous; internode usually less than 2 cm. long. 10. Lamina with 3 to 7 (to 9) veins/5 mm., venation prominent, angle of divergence (25- Fa a [terminal internode of innovation about !/2 length of DUNEIS | 4 os ow & He eo a ee es Sw C. cuneifolium. 10. Lamina often with more dense venation, venation prominent or not, angle of ee aay more than (40-)5S0°. vLl WOLAYOPUV GIONUV AHL JO TYNYNOL 19 “10A] 11. Lamina with up to 6 (rarely 8) veins/5 mm., venation subapparent above, subobscure below, latex canals often impressed; inflorescence with 5 to EL OWELS™ hore et ae oy esa 13. C. thwaitesii. 11. Lamina with 5 to 14 (rarely to 17) veins/5 mm., venation not more soe on upper surface than on lower, latex canals rarely impressed; inflorescence with 7 to 19 flower 12. Twig + 4-angled; flower with 10 to 15 tepals; pedicel in fruit (2—-)2.5- 3 mm. across. ........ 12. Twig strongly 4-angled to subalate; flower with 4 to 10 tepals; pedicel in fruit ca. 2 mm 3. Lamina usually rounded at base, midrib on upper surface abruptly narrowed from base; all internodes of innovation approximately same length; fruit less than 1.5 cm. long. ...... . poshcein a et des Ea te ee a ee ee sds eS i Be ee | 18. C. apetalum. . Lamina usually narrowly cuneate to acute at base, midrib on upper surface narrowing gradually from base; terminal internode of innovation sometimes about 1/2 length of others; fruit ca. 3 Cit ION. 2 ao os DER EE Ee ee CSR SOE ESS Se Ca SS x 16. C. austroindicum. — Ww 3. Key to Species of Calophyllum in Mainland Southeast Asia (Burma, Thailand, and the Nicobar Islands, to China) 1. Twigs (1.5-)2.5-5 mm. across, very strongly vaca to subalate, + nitid [indumentum often tomentose]. . Twigs with prominent leaf scars; petiole 2-5 mm. long. .. ©. 6 1 ee ee 174. C. poilanei. 2. Twigs lacking prominent leaf scars; petiole at me 101 mm. long. 3. Twigs drying blackish; internode 0.5-2.5(-5) cm. long. . 2... ee es 83. C. macrocarpum. 3. Twigs drying brownish; internode 2-7 cm . Lamina with 6 to 11 veins/5 mm.; fruit spherical, ca. 1.5 cm. long [inflorescences terminal and axillary]. ..... Pele aon MR Secor ibe. in Sk GO hs es GR Gesccleae wae Reena Get EN de. Lhe VE Ne Oe pies Sea a . C. dongnaiense. 4, Lamina with 10 to 16 veins/5 mm.; fruit ovoid to ellipsoid, 2.5-3 cm. long, sharply pointed at apex [inflorescence Osition UNKNOWN) ¢ 64 4s wb ee Ee ee eh ee we A A ee ee eee ee aoe eS 170. Calophyllum sp. 1. Twigs often less than 2.5 mm. across, if more then not strongly 4-angled or subalate, rarely nitid. 5. Inflorescences terminal (and sometimes axillary). 6. Lamina retuse to subacute at apex. WOTIAHdOTVO ‘SNAAALS L086! SLI an et te ees ae es Ge em Ae Be Se Mee, a ee ge eh oe SO A ek ee ee 71. arkeri. 7. Twigs 1.2- - mm. across, usually drying brownish or grayish; basal internode of axillary inflorescence a 5- Ml cm. or more lon 8. Terminal bud 2.5-5 mm. long; lamina with + obscure venation usually more obvious on upper surface. .... . Psa te SN Ae Gong te be oe Saye alee Mees Cee ee A GEG 4k gs eG. Be la ee a ee 154. C. rugosum. 8. Terminal bud 1.3-2 mm. long (unknown in C. balansae); lamina with apparent venation not notably more obvious on upper surface. 9. Lamina flat at margin, angle of divergence of venation 30-55S°................. 97. C. ceriferum. 9. Lamina + undulate and = at margin, angle of divergence of venation 60-70%. ..... 6. C. balansae. . Lamina acute to acuminate at a 10. Lamina often drying bic ae [gray-olivaceous above, sabelline below], midrib on upper surface surrounded by raised blade, indistinct; fruit ellipsoid, usually deeply wrinkled. .........2......... 3. C. thorelii. 10. Lamina usually not drying notably bicolored, midrib on upper surface surrounded by raised blade or not; if fruit deeply wrinkled then not ellipsoi 11. Venation density on upper surface of lamina apparently twice that on lower. .... 3. C. thorelii group ‘‘s’’ 11. Venation density same on both surfaces of lam 12. Midrib on upper surface of lamina sinounet by raised aoe indistinct. 13. Terminal bud 3-7 mm. long; lamina with (6 to) 8 to 11 v ns TS os ue es oe 8 7. C. symingtonianum. 13. Terminal bud 2.3-4 mm. long; lamina with 4 to 8 vena bs Ae ie Eo eke Me, Sa a 8. C. dryobalanoides. 12. Midrib on upper surface of lamina not surrounded by raised blade. distinct. 14. Lamina with 9 to 16 (to 21) veins/5 mm., attenuate at base [fruit finely wrinkled, outer layer 1.5-3 Hac CHNGK de eek a ee ea te ee eh ee he ee Be 4. C. polyanthum. 14. Lamina with 6 to 10 _ 14) veins /5 mm., acute to cuneate at bas 15. Petiole 0.7-2.5 cm. long; lamina coriaceous, margin slightly ndulsie but not recurved or slightly so; fruit finely wrinkled, outer layer 3-5.5 mm. thick. .............. 5. C. touranense. 15. Petiole 0.3-0.9(-1.3) cm. long; lamina thinly coriaceous, margin undulate and a fruit coarsely wrinkled, outer layer ca. 1 mm.thick...........0.0.0..20....004. . C. balansae. 5. Inflorescences axillary. 16. Axillary innovation with basal scars; lamina with indistinct venation and + impressed latex canals. ee ee ee a | bs ee oe ae BA ee ee ee ee ee ee a ee Oe Oe ee es a ee . C. depressinervosum. WOLAYOdANV GCTIONAV AHL AO TIVNYUNOL "10A] 19 16. Axillary innovation nearly always lacking basal scars; lamina with + distinct venation, latex canals impressed or not. 17. Lamina obovate to elliptic, at least 7 cm. long; pedicel (1.5-)2.5-4.5 cm. long [lamina with 4 to 8 . ne veins /5 mm. stone with basal plug: seaside tree] 2 fa ke ks Be A ee ee eS eee 8. nophyllum. ee Lamina ovate to elliptic (rarely obovate), often less than 7 cm. long; pedicel less than 1.5(-2.5) cm. oo 18. Lamina narrowly elliptic, ca. 10 times longer than broad. ..................4. 103. C. lineare. 18. Lamina broader, less than 5 times ae than broad. 19. Lamina with 12 to 25 veins/5 20. Lamina often more than 7 cm. long, rather closely undulate at margin; inflorescence eee: outer k. layer of Truit at least. 1mm. thick. 6 4 ah Bok he ee GE ee ew we SOR oulattri. 20. Lamina often less than 7 cm. long, rather distantly undulate at margin; oe anbenehed: outer layer of fruit less than a TM COCK. ee ail ee. ahs ce ee 19. C. calaba. 19. Lamina with up to 12 veins/5 m 21. Lamina drying notably esinicdl coriaceous, latex canals impressed or not; bud often supra-axillary. Sgt hh ed, A a es Sk ae a eo ee OY Be ee oe . rupicola. 21. Lamina not drying notably bicolored, thinly coriaceous, latex canals rarely impressed; bud not supra-axillary. Lamina rounded to acute at apex; stem often short-tomentose. ......... 102. C. pisiferum. 22. Lamina acute to aaa at apex; stem at most puberulent. 23. Terminal bud 2-4 mm. long; lamina ca. 2 times longer than broad; bracts inconspicuous, deciduous; ses with 4 (to 9) tepals. ALG: Grae Bees is a we ee Re ee 100. C. tetrapterum. 23. Terminal bud 3.5-7 mm. long; lamina ca. 3 times longer than broad; bracts often foliaceous and persistent; flower with 8 (to 12) tepals. ..........202.. 175. C. membranaceum. 4. Key to Species of Calophyllum in Malaya, Sumatra, and the Sunda Islands 1. Axillary innovation with basal scars; twigs less than 3 mm. across [species sometimes with basal scars, but with thicker twigs, key out in second half of couple 2. Lamina with indistinct venation, drying almost completely flat, attenuate at base... .......... 74. C. enervosum. 2. Lamina with + distinct venation, often drying definitely undulate and/or recurved at margin, usually other than attenuate at base. [0861 WOTTAHdOTVD ‘SNAAALS 3. Terminal bud glabrous, clearly composed of several pairs of decussate scales. ........... 73. Calophyllum sp. 3. Terminal bud with indumentum, only one pair of reduced leaves or scales visible. 4. Lamina (5-)8-15 cm. long, latex canals not impressed; terminal bud 2.5-7 mm. long. 5. Lamina often drying + broadly concave, with 10 to 19 veins/5 mm.,; inflorescences terminal (and also axillary). age. este tecTe AE a ae eB cee ee a a a ge ee ee Ge ee ee he ec 72. C. flavoramulum. Se ae Ne Sie ae ee SE Oe ee eee we Gee eR Ge ae ws el ss a ee 70. C. fraseri. . Lamina 3.2-8 cm. long (rarely to 13 cm., then latex canals on lower surface impressed); terminal bud less than 4 mm. long. 6. Midrib on upper surface of lamina usually surrounded by raised blade, spe aaa aaa on lower surface inconspicuous. 7. Lamina drying cinnamon-sabelline on lower surface, latex canals often impress BSS 7. Lamina drying umber to sabelline on lower surface, latex canals not impressed. 8. Lamina usually bluntly acuminate at apex; fruit at least 1.4 cm. long, + ovoid. ¢ 4D; . es eel 8. Lamina more or less rounded at apex; fruit less than 1 cm. long, s fe C. nodos 6. Midrib on upper surface of lamina not surrounded by raised blade, distinct, on lower surface ae 9. Twigs less than 2 mm. across; lamina drying bicolored [gray-brown above, yellow-brown below]; ‘fal less PATE PaO CU ONES 9). tcc ca, eck, ee atte yl. Soe Be Tie cas ae hie 9 nee @ i Se ok 101. C. rupicola. . Twigs (1.5-)2 mm. or more across; lamina drying + same color on both surfaces; fruit at least 1.5 cm. lon \o &. 10. Lamina retuse to subacuminate at apex, with (9 to) 11 to 17 (to 22) veins/5 mm.; fruit ellipsoid. oi ek a Me es ee eG ce a a GS le ewe es a 88. C. hosei. i 1. Axillary innovation lacking basal scars, or twigs more than 3 mm. across. 11. Lamina cordate to auriculate at base 12. Lamina suborbicular, venation more prominent on upper than lower surface; inflorescence reduced to 1 or 2 flowers. Pe ea ee ee ee ee ee ee ene re eee ae ee eee ee ee 155. C. rotundifolium. 2. Lamina nearly always at least twice as long as broad, venation not more prominent on upper than lower surface; inflorescence with at least (1 to) 3 flower 13. Lamina 15-36 cm. long; pedicel at anthesis 3-7.5 cm. long. — eee Te Se ee eee ee ee 91. C. grandiflorum. 8LI WOLAYOPUV ATONYV AHL dO TVNYNOL 19 ‘10A] 13. Lamina to 15(—23.5) cm. long; pedicel at anthesis less than 3 cm. long. Twigs up to 1.5 mm. across, drying brown; fruit less than] cm. long. ............. 19. C. calaba. 14. Twigs at least 2 mm. across, drying yellowish to grayish; fruit at least 1.5 cm. lon 15. Midrib strongly depressed only at base of lamina; inflorescence with (3 to) 5 (to 11) flowers. ...... we eh sale ae Bis Si sd vas a aS ase! aa Bg id Sein a mo 0. C. venulosum. . Midrib strongly depressed for at least the lower 1/4 of lamina; inflorescence with 5 to 17 flowers 16. Twigs rounded to obscurely 4-angled; basal internode of inflorescence to2 mm. long. ........ bh eB & & SS OEE ee Se EE So. Be Se 93. C. mukunense. 16. Twigs oe 6-angled; basal internode of inflorescence 2-4 cm. long [cultivated specimens of m from. Bogor key Out here). 244. tne Sa Sawa e ee 6S eS 92. C. subsessile. 11. are — rounded to oe at base. a at maturity tomentose or sparsely tomentose on entire lower surface 18. eee terminal (sometimes also from uppermost leaf axils); fruit furfuraceous. _ wa 19. Terminal bud 4-8 mm. long; twigs drying yellowish... ...............2.. 158. C. rubiginosum. 19. Terminal bud ca. 1.4 cm. long; twigs es dark Drown: 2-25.45 % 2 4.444 4824 54. Calophyllum sp. 18. Inflorescences axillary; fruit not furfuraceous 20. Lamina with (8 to) 11 to 17 veins/5 mm.; ovary tomentose. ............. 2000. 159. C. molle. Lamina with (3 to) 5 to 8 (to 11) veins /5. mm.; ovary glabrous.................. 104. C. dispar. 17. Lamina at maturity at most puberulent on lower surface. Zi; ig rtcenee terminal (sometimes also arising from uppermost leaf axils). na retuse at apex, 2-5 cm. long; uppermost pair of axillary buds erect, ca. 1/2 length of terminal bud; Gracie PollaceGus, PCISISIENt. so ga te ee Be ee he eS Ee ee ee ee 157. C. aureum. 22. Lamina usually acute to acuminate at apex, at least 3.5 cm. long; uppermost pair of axillary buds relatively short; bracts not foliaceous and persistent. 23. Lamina 3.5-6.7 cm. long, with (9 to) 11 to 16 pe Mee ie ee 156. C. aureobrunnescens. 23. Lamina 5-19.5 cm. long, with 5 to 12 veins/5 m 24. Petiole ca. 1 mm. across; lamina ovate to subslliptic, 10°9 Cm. Jone... 4 3 C. symingtonianum. 24. Petiole at least 2 mm. across; lamina elliptic to oblong or subobovate, at least a )8.5 cm. . Lamina very coriaceous, broadly rounded toward apex, finally shortly acute; terminal bud Leiehe2. 7) Cm) ION Ga ‘6: ane e oa HB wea oe Oe ee ek |, 47. C. coriaceum. 25. Lamina coriaceous, acuminate at apex; terminal bud 6.5-10 mm. long. .............. [0861 WOTTIAHdOTVD ‘SNAAALS Pa ap eli axillary (rarely terminal). 26; na very strongly revolute at margin when dry, thickly coriaceous. ........ 49. C. scriblitifolium. 26. ere not very strongly revolute at margin, thickly to thinly coriaceous. 27. Terminal bud (0.8-)1-3 cm. lon 28. Lamina drying closely undulate at margin; uppermost pair of axillary buds erect; ar ee ese with Strone lateral branches. «sw. 46.4% & HSK WR oe ES SS Sw ewe De OES 1. oulattri 28. Lamina not drying closely undulate at margin; uppermost pair of axillary buds rarely erect; felon scone usually without strong lateral branches. 29. Terminal bud (3-)4.5-10(-11) mm. long; lamina with veins diverging at 40—70°; axillary bud often notably plump, sometimes supra-axillary................0.0. 0004 . rigidum. . Terminal bud (0.8-)1-3 cm. long; lamina with veins diverging at (60-)70-85°; axillary bud rarely plump and/or supra-axillary. 30. Midrib on upper surface of lamina strongly sulcate, margins raised, narrowing gradually NO \o from b oie Lamina often drying nitid, with (10 to) 12 to 17 (to 21) veins/5 mm.; inflorescence rarely branched or flabellate; fruit pointed BUAPCRe 2 a BAR He Gas ‘5 . canum. 31. Lamina drying dull, with 10 to 15 veins/5 mm. ; inflorescence usually branched and/or flabellate; fruit rounded at apex. ................ 2.000004 30. Midrib on upper surface of lamina raised (not notably sulcate) and/or abruptly narrowing near base. 32. Lamina with 5 to 11 veins/5 mm., venation very distinct. ...............0. ee Ae ie, ee ee ee 45b. C. lanigerum var. austrocoriaceum. 32. Lamina with 11 to 18 veins/5 mm., venation usually rather indistinct. 33. Inflorescence axis 0.3-1.3 cm. ‘long; hairs with large, thick-walled apical cells; fruit with stone wall (0. 8—)1.3- - MMC UNCK: 66k 4 Bk de eR OG 44. C.r ufigemmatum. 33. Inflorescence axis 1.7-7 cm. long; hairs lacking large, thick-walled apical cells; fruit with stone wall 0.3-0.8(-1) mm. thick. .........0.02., 43. C. wallichianum. 27. Terminal ae less than 0.9 cm. long 34. Lamina retuse to obtuse at a 35. Twigs drying shiny; ain “will 3 to 6 veins /5 mm.; flower with (8 to) 12 tepals; fruit + beaked at apex. ........ CAGES Gig eet Gee Baek ee ee ee lee ee eS ee eee 169. C. subhorizontale. WOLAYOIAV GIONUV AHL AO TVNYNOL 9 ‘10A] I 35. Twigs not drying shiny; lamina usually with more than 6 veins/5 mm.; flower with 4 to 8 (to 12) tepals; fruit + rounded at apex. 36. Venation on upper surface of lamina clearly denser than on lower [indumentum brown, usually tomentose; basal internode of inflorescence (0.2-)1-5 cm. long; inner surface of outer layer of fruit shiny, striate]. .......... 36. Venation same density on both surfaces of lamina. 37. Midrib on upper surface of lamina surrounded by raised blade, indistinct. . Lamina to 6(-8.3) cm. long, coriaceous; tree without stilt a Se et le OAS 151. Calophyllum sp. 38, Lamina 5.7-21 cm. long, very coriaceous; tree with stilt roots. ............ 1. C. sclerophyllum. 37. Midrib on upper surface of lamina not ane eae by raised blade, distinct. 39. Lamina with (3 to) 5 to 11 veins/5 mm., usually + thinly coriaceous. 40. Indumentum subtomentose, ep a on young stem and inflorescence axis; lamina less than 6 ONG: «cy dnb es OE Se A ee EER ae eee ee eee eee ee OS 102. C. pisiferum. Indumentum puberulent, inconspicuous; lamina at least 7 cm. long. ......... 38. C. inophyllum. 39. ae with (8 to) 10 to 18 veins/5 mm., (thinly) coriaceous . Twigs less than 1.5 mm. across; lamina with midrib on ‘upper surface often disappearing below apex, 7 + flat: fruit ca. 7 mm. long, smooth. . 2... we ee es . C. exiticostatum. 41. Twigs usually more than 1.5 mm. across; lamina pales midrib on upper surface continuing to apex, usually raised; fruit more than | cm. long, wrinkled or 42. Lamina often with either clearly thickened aca ee least 0.3 mm. wide or submarginal vein; fruit smooth or faintly striate. 2. 6 6. ee es 78. C. teysmannii. 42. Lamina not as above; fae aadied. 43. Lamina with 8 to 12 veins / 5 mm., drying grayish sepia, the margin discolored, brownish, inflorescence with flowers in pairs. ©... 2. 2 ee ee 87. C. biflorum. 43. Lamina with 10to 18 veins /5 mm., not ae grayish sepia or with discolored margin; inflorescence with flowers tending to be in groups o 44. Plant with pale brown, tomentose aanatsarane basal internode of inflorescence 1.7-5.2 86. cm. long; fruit spherical, drying closely wrinkled. ............ . costulatum. 44. rer with brown, subtomentose to puberulent indumentum; basal internode of inflorescence 15-)0.4-1.8 cm. long; fruit ellipsoid, drying distantly wrinkled. .85. C. sundaicum. 34. Lamina acute to acuminate ee cuneate) at apex. WNTIAHdOTVO ‘SNAAALS [0861 I 45. ee on upper surface of lamina depressed, narrowing gradually from base [lamina elliptic, with (4 or) 5 to 9 veins/5 ee Bue ee Gee ip eae Res we ce ee es ee ce, a ee eg ee ek ee 130. C. archipelagi. 45. Midvib on upper surface of lamina with at least margins raised, sometimes narrowing gradually from ce 46. Twigs 2.5-7 mm. across, drying blackish or shiny. 47. Twigs 4-alate, drying blackish (rarely brown); petiole (1.2-)3.5-7 cm. long. ........ 83. C. macrocarpum. 47. Twigs with raised lines, but not alate, drying brown to yellowish; petiole 0.6-1.5 cm. long. ........... Pe ae ae ee ee ae ee ee ee ee a ee er ee a re ee 169. C. subhorizontale. . Twigs less than 3.5 mm. across, if more then drying neither blackish nor shin 48. Lamina thinly coriaceous, often drying greenish brown, closely undulate at margin; uppermost pair of axillary buds erect, conspicuous; inflorescence branched...................0202840. 21. C. soulattri. 48. Lamina not as above; uppermost pair of axillary buds rarely erect and conspicuous; inflorescence rarely branched. 4 oN ON 9, Lamina elliptic to oblong, venation on upper surface apparently twice as dense as on lower surface; basal internode of inflorescence (0.2—)I-5 cm. long; inner surface of outer layer of fruit striate. .......... Ste. eae Gree eareeoa sae eee @ oe Hoe ae A we eae ee ee ee ewe ee 84. C. ferrugineum. 49. Lamina variously shaped, the venation on both surfaces usually of same density, if not then basal internode of inflorescence less than | cm. long; inner surface of outer layer of fruit not striate 50. aie drying yellowish, ee or grayish. . Petiole (0.8-)1-2.5 cm. long; lamina drying + concave, with very distinct venation. ......... Ue ee ah ae ch a OE ee, Woe ee aw, ete de ee 90b. en Mrs var. tenuivenium. . Petiole to 1.5 cm. long; lamina not drying concave, venation distinct or not. 52. Lamina abruptly acuminate at apex; blackish-drying petiole conta sharply with pale twig; outer layer of fruit 1-1.8 mm. thick. ...................00, 106. C. dioscurii. . Lamina acute ie apex; petiole and stem not strongly contrasting in color; outer layer of fruit less than 1 mm. thick. 53. Lamina as 12 to 20 veins/5 mm.; inflorescence with (5 to) 7 or more flowers. ...... Keane Ace oe ee Ee SO ee ee ee ee ee 19. C. calaba. 53. Lamina with 6 to 13 veins /5 mm.; inflorescence with up to 5 (to 7) flowers. = wa —_ a) N 54. Lamina with 6 to 9 veins/5 mm.., midrib on upper surface not surrounded by raised blade; anthers with hairs. ..........-.......2.200. 150. C. gracillimum. 54. Lamina with 8 to 13 nae 5 mm., midrib on upper surface surrounded by raised blade; Alters PIADLOUSe. x 2:2: x.e eu the 6 ae ee Se Se a ee ae 151. Calophyllum sp. WOLAYOPdUVY ATONYV AHL JO TVNUNOL 781 19 “T0A] 50. Twigs drying brownish. 55. Lamina usually drying bicolored [olivaceous- ies aa and cinnamon-honey below]; inflorescence axis and pedicel puberulent, the latter incrassate in fruit. ............ 105. C. incumbens. . Lamina rarely drying pee eet inflorescence ae glabrescent toward apex, pedicel usually glabrous, rarely incrassate in fru 56. Midrib on upper surface of an narrowing gradually from base, strongly sulcate at first; inflorescence axis 3-5.5 cm. long. 57. Hairs curved, papillate on one side. .............2.2.... 101. C. aff. rupicela. 1» ddamnS S: ereet; Mranched:. s 2. & oe & fw 4 oo atte Ne ble Gis Oo we 48. C. dasypodum. . Midrib not as above; inflorescence axis usually less than 2 cm. lon 8. Lamina with (3 to) 5 to 9 (to 11) veins/5 mm., rather thinly coriaceous 59. Sere bud (3.5-)4.5-7 mm. long; lamina 6-15 cm. long; outer layer of fruit (0.5-)1-1.8 HOM, “he ete a Be a Ae ae he ee 104. C. dispar. 2. cena bud 2-4.5 mm. long; lamina 3-7.5 cm. long; outer layer of fruit 0.2-0.5 mm WICK gs hich sk oe ae Get ee ee ee ee Ee ee, & Hee we 102. C. pisiferum. Loa) LF) 1S a) nN 58. Lamina with at least 7 veins/5 mm., thinly coriaceous to coriaceous 60. Lamina with 10 to 17 veins/5 mm., veins sometimes apparently twice as dense on upper surface as on lower, angle of divergence 40-70°; terminal bud with dark brown, tomentose indumentum. .....................004. 23. C. rigidum. 60. Lamina not as above; terminal bud with grayish to brown, crustaceous to short-tomentose indumentum. 61. Lamina with 12 to 20 veins/5 mm., drying + sepia, dull, eee te BAe oe ade > ss fs as ek ee Mae oth, a i Se ee no eae le a ae De C. calaba. 61. Lamina with up to 14 veins/5 mm., drying other than dull sepia, aoe bicolored or with pale margin. . Lamina drying bicolored [gray-brown above, yellow-brown below], lower surface often puberulent; bud often clearly supra-axillary; fruit often ovoid. ...... Cree oe ae oe Gg a a Bp at eee EE ee ee 101. C. rupicola. . Lamina drying bicolored or not, lower surface glabrous; bud rarely clearly supra-axillary; fruit usually spherical. 63. Terminal bud 4.5-7 mm. long; lamina rather thickly coriaceous. ..... . 70 nN i) lon) NO [0861 WNTIAHdOTV) ‘SNAHAALS Li ie 63. Terminal bud 2-4 mm. long; lamina coriaceous or thinly coriaceous. 100. usually pale; fruit less than 1.2 cm. long. ..... C. tetrapterum. 64. Twigs 1.2-1.5 mm. across, not an 4-angled; margin of lamina not notably pale; fruitca. 1.7 cm. long. ........ 9. Calophyllum sp. 5. Key to Species of Calophyllum in Borneo Lamina at least sometimes cordate or auriculate at base. 2. Midrib on upper surface of lamina depressed; inflorescences terminal and often axillary, axillary inflorescence with basal intermode less than 2mm, long, .<.2is eves cawedbiegegene bed e eh RS GRE LEEK EHS 93. C. mukunense. 2. Midrib on upper surface of lamina not igs ia inflorescences occasionally terminal, usually axillary, axillary inflorescence with basal internode more than 3 m 3. Axillary shoot lacking basal scars; ae usually with 4 tepals; outer layer of fruit more than 1.5 mm. thick. ..... LeeLee es ee ea oe ee ee eae eee a ee eS eae ee 90a. C. venulosum var. venulosum. 3. Axillary shoot nearly always with basal scars; flower with 4 or 7 or more tepals; outer layer of fruit ca. 1 mm or less k. 4. Lamina 9.5-14.8 cm. long, midrib on upper surface surrounded by raised blade; flower with 4 tepals. ........ Pe a ae eS ee ee he a ee ee ee ee . C. complanatum. 4. Lamina 1.2-8.5 cm. long, midrib on upper surface not surrounded by raised blade; flower with at least 7 tepals. 5. Lamina very coriaceous, venation on upper surface rather obscure; inflorescence axillary; flower with 8 tepals. gn! Ig Ae devas aa se oe ee ee ee ae ee ee ee 94. C. calcicola. a cnet coriaceous, venation on upper surface prominent; inflorescence terminal and axillary; flower with (8 (Oy AEG 1S Oss, 4 ke dee oe eee ee ee ee eee ee oe eee 8 EAA 99. C. clemensorum. Lamina broadly rounded to acute or decurrent at bas 6. Terminal bud at least 1 cm. long (if no bud ae inflorescence terminal, see couplets 17(2), 23(1)-25(1), 36(1), 42(1), 47(1), 48(2), 61), and 67(1)). Lamina broadly recurved toward margin [lamina very coriaceous; plant of peat swamps].... . 49. C. scriblitifolium. 7. Lamina not broadly recurved toward margin . Lamina usually thinly coriaceous and closely undulate at margin; inflorescence branched; pedicel relatively long and slender; fruit less them 5 Cm, One. « eae eee ee ae OR ee ee Se ee BS 21. C. soulattri. 8. Lamina usually coriaceous, not closely undulate at margin; inflorescence rarely branched; pedicel not Aas long and slender; fruit (where known) usually more than 1.5 cm. long. WOLAYOIUV GTONYUV AHL JO TVNUNOL 19 “T0A] POE Ses ae Me Tae SOTO a See rac ciiS AubOS oaaey fe oe ReD eM nth, (mar ey Dee Ueth pa EC SL See cae 129. Calophyllum sp. 9. Midrib on upper surface of lamina not broadly depressed, narrowing gradually from base or not. 10. Hairs with 2 to 4 much-expanded apical cells, the walls birefringent under pepuae light. 11. Lamina not cucullate at base, with 7 to 15 veins/5 mm.; pedicels 0.42.5 c Bee as ae it nah GA AA Ane ek al ea rh ate wy ete en cen ae eae Po 45b. C lanigerum var. austrocoriaceum. 11. Lamina slightly cucullate at base, with 11 to 22 veins /5 mm.; pedicel 0.2-0.5 m n 12. Twigs 1. 5-3 mm. across; terminal bud to 1.5 cm.long. .......... 32. Cc castaneum (variant). 12. Twigs 3.5-5.5 mm. across; terminal bud at least 2 cm. long. ............ 52. C. castaneum. 10. Hairs lacking much-expanded apical cells, the walls at most slightly birefringent under polarized iene 13. Anthers and ovary with hairs; fruit (not yet known in Borneo) with walls at least 1 mm. thick [lamina Slightly cucullate Al DaSe) a: we ao oe oy Gee eS BE Se ew 159. C. molle (variant). 13. Anthers nearly always and ovary always glabrous; fruit with stone walls less than 1 mm. thick. 14. Lamina often slightly cucullate at base, venation notably fine and clear; hairs not branched, all Celis t+) papillate. 62-46% k's eee ee ESS EO ee a ee ae OS 52. C. woodii. 14. Lamina not slightly cucullate at base, venation not notably fine and clear; hairs various, but not as above. 15. Twigs 1.8-3 mm. across; margin of lamina (+) closely undulat — 16. Hairs papillate at base; lamina not drying notably led. usually shiny on upper oe — On flowers with 4 (to 8) tepals. 2... 50. C. ca . Hairs strongly branched; lamina drying notably bicolored, not shiny on upper surface; aigwers with 8 to 12 tepals. 17. Twigs drying brown, puberulent when young; inflorescence axillary. ........... eee eae ae ae ae oe ee eee nee ee ee ee ae eee 2. C. blancoi ae ia 17. Twigs drying blackish, rather persistently subtomentose; inflorescence terminal... .. . hte se em dete gk eaten GREP era Bare remain as ae eae pHa Ess ery group ‘‘f’’. . Twigs 3-5 mm. across; margin of lamina distantly undulate. 18. Inflorescence branched; fruit smooth, stone wall less than 0.2 mm. thick. .......... Sho dyed. ae es ade ee Ges eee Gs eee we ee es SE ee OS ae eto en ewe eee 51. C. lowii. 18. Inflorescence nearly always unbranched; fruit wrinkled, stone wall more than 0.2 mm. thick. 43. peey ee a ee cg TEs eens ee eee en ees ke ep eee es ee C. wallichianum. [0861 WOTIAHdOTV) ‘SNHAALS 6. Terminal bud less than 10 mm. long. 19. Twigs with two prominent V-shaped transverse lines at nodes, otherwise subterete; lamina broadly rounded at base. Ed ee Ge Raa ive Oe we oo oe Ae BE ee ee 78b. C. teysmannii var. bursiculum. 19. Twigs with at most obscure transverse lines at nodes, flattened and/or angled; lamina rarely broadly rounded at ase. 20. Uppermost internode of innovation at most = as long as others. 21. Axillary innovation with (sub)basal sca Lamina subacuminate at apex, midrib on upper surface sharply raised; terminal bud sharply pointed; 67.-G fruit (2 poimted at apex. 4.2024 nb ee dk wt eee SEES ERAN SER ES Oe . C. elegans. 2. Lamina acuminate at apex, midrib on upper surface not sharply raised; terminal bud not sharply pointed; frei Tounded Al apex. «iene bee Se REPRE EEE ASE CHES ERE EO . C. stipitatum. 21. Axillary innovation lacking basal scars 23. Lamina with pale marginal band at least 0.4mm. wide. ........ 100b. C. tetrapterum var. obovale. 23. Lamina lacking marginal ban 24. Lamina acute to acuminate ee CORUNA ys % 4-4: 4 ee 4 Ga 160. C. obliquinervium. 24. Lamina rounded to retuse at a poy oe terminal; on on upper surface of lamina apparently twice as dense as on bn fe ae ee ee ee oe ee ee ee EE oe 161. C. sakarium. 25; oe axillary; venation same density on both sides of lamina. 26. Twigs drying finely reticulate; lamina with 3 to5 veins/5mm........ 75. C. confertum. 26. Twigs not drying finely reticulate; lamina - 6 to 9 veins/S mm... .76. C. praetermissum. 20. Uppermost internode of innovation about same length as other 27. Midrib on upper surface of lamina surrounded by aaa blade, midrib itself rarely much further raised, midrib on lower surface depressed or not. 28. Lamina very coriaceous, often spotted on lower surface; fruit more than 2.4 cm. long [trunk with stilt TOUCSES 46. pe OS ed et ee Se ee ES eee ae ee ES 1. C. sclerophyllum. Lamina coriaceous, ia not spotted on lower surface; fruit (where known) less than 2 cm. long [stilt roots rare at most 29. Lamina 7-15 cm. long, midrib on lower surface raised, 25 veins/5 mm. [fruit not known]. ..... .- re ee ee ee ee ee ee ee ee eee ee ee 56. C. glaucescens. 29. Lamina up to 8(-10.5) cm. long, midrib on lower surface nearly always + depressed, fewer than 13 (to 17) veins/5 mm. 28. oo WOLAYOPUV GIONYV AHL AO TVNYNOL 19 104] 30. Lamina drying orange-brown on lower surface, veins invisible below in both fresh and dried leaf, latex canals often + impressed above. ............... 8. C. depressinervosum. 30. Lamina usually drying grayish brown (rarely orange-brown) on lower surface, veins visible below in both fresh and dried leaf, latex canals not impressed above. 31. Fruit spherical, less than 1 cm. long, stone wall ca. 0.2 mm. thick [throughout Borneo]. de cB cg? ie ge icy eae: Gs uh series a ae tay Serctuts eu toe GR Gy ek Ge es Gs ee ce nee ; nodosum. 31. Fruit ovoid, more than 1.2 cm. long, stone wall 0.3-0.5 mm. thick [mostly western Borneo]. 0. Coos ee ee a ee eee Ee ge te at ete oe Be Ee, Kas ee . pulcherrimum. 27, Midrib on upper surface _ lamina not surrounded by raised blade, midrib itself raised or not, midrib on lower surface usually rais 32. Lamina usually ovate or oblong-elliptic, rounded to retuse (rarely bluntly pointed) at apex. 33. Leaf inserted on stem at + right angle; lamina very coriaceous, drying flat, venation obscure. ... . se ss Ge es BL oe He EG, Bd es, he HB ee Se le ee es ee Ee 162. C. ardens. 33. Leaf usually ascending; lamina variable in texture, usually not drying flat, venation usually apparent 34. Lamina with obvious (0.3-1 mm. wide) band of marginal thickening, discoloration, or distinct submarginal vein; trunk often with spurs or stilt roots 35. Lamina with discolored margin, veins visibly continuing to margin; fruit wrinkled, inner surface 87. C wo sn of outer layer striate and usually SINy. “6 aed oe oe we eee we we . biflorum. Lamina with thickened margin, or with submarginal vein, veins not visibly yo oana to margin; fruit smooth or not, the inner surface of outer layer rarely striate, never s iny. 36. Twigs less than 2(-2.5) mm. across; terminal bud 1.5-4 mm. long; fruit less than 1.5 cm. long, outer layer less than 0.5 mm. thick... .. . 100b. C tetrapterum var. obovale. . Twigs (1.3-)2.5 mm. or more across; terminal bud (2-)4 mm. or more long; fruit more than 1 cm. long, outer layer more than thick. 37. Terminal bud enclosed by petiole bases; ‘lamina with 4 or 5 (to 7) veins/5 mm.; lowest internode of inflorescence 3-7.5 cm. long [spurs and stilt roots apparently ADSCRUl Sk a. -k as @ Sos FG we aed dO ad eee ee es Be 82. C. havilandii. : Terminal bud enclosed by petiole bases or not; lamina with (4 to) 7 or more veins/5 mm.; lowest internode of innovation less than 3 cm. long [spurs and/or stilt roots often present]. Ww nN we ~ [0861 SNHAHLS WO TIAHdOTVO L81 38. Older twigs drying rather pale yellowish brown and contrasting with darker petioles; ovoid, less than 1.7 cm. long, sharply pointed at apex, sharply wrinkled WHER OEY: 5 ike GG eet ee eee eee ee hae eS 77. C. andersonii. . Older twigs not drying notably paler than petioles; fruit usually subspherical to ellipsoid, at least 1.7 cm. long, rounded at apex, + smooth when mina often punctate below, drying olivaceous; trunk with stilt roots up to 3 m. or so [seedling with one pair of leaves; young plant with alternate mete en eo eee ee ee ee ee 81. C. sclerophyllum. . Lamina not punctate below, drying brown; trunk with stilt roots up to 1 m or aa roots lacking [seedling with two pairs of leaves; young plant with Cpposile (eaves): oi 44446 eta e He eee eee eS 78. C. teysmannii. 34. Lamina lacking hes aes or discolored margin or distinct submarginal vein; trunk rarely with spurs or stilt r 40. Axillary shoot Sith (sub)basal scars; lowest internode of axillary inflorescence usually less than 5 mm. 41. Lamina 1-4 cm. long; fruit unknown, probably less than 1.5 cm. long. .......... 2s EE ee ee ee eee eee 61. Calophylium sp. 41. Lamina more than (2.5-)4 cm. long; fruit more than 1.5 cm. long. 42. Inflorescence terminal and axillary; flower with 12 or more tepals; fruit subspherical, bs ie aoa ee es ee ee ee ee a ee . garcinioides. 42. Inflorescence axillary; flower with 4 or 8 tepals; fruit ellipsoid or pyriform, smooth or shallowly wrinkled. 43. rae with (9 to) 11 to 17 (to 22) veins/5 mm.; flower with 4 tepals; fruit 88. C. hosei. Ww oo eS) \o ee Ma ee al 0 i ee ee ae ee ee ee ee ipsoid. 43. ine with 6 to 11 veins/5 mm.; flower with 8 tepals; fruit pyriform. ..... 7 oR eee ke ae ee a ee ees ee eee ee ee . C. elegans. 40. Axillary shoot lacking (sub)basal scars; lowest internode of axillary inflorescence usually more than 5 mm. long. 44. ae (2.5-)3.5-6.5(-9) cm. ee terminal bud to 3 (very rarely -7) mm. long; fruit either furfuraceous or | cm. or less lon 45. Twigs and midrib short- eens lamina with at least 14 veins/5 mm., midrib often disappearing ca. 3 mm. below apex 881 WNLAYOIUV ATONYUV AHL JO TVNYNOL 19 ‘104] 46. Twigs slightly flattened; terminal bud 2-3 mm. long; Cane on lower surface lamina only slightly raised. .. 2... ....000042., C. exiticostatum 46. pes strongly flattened; terminal bud 3-6.5 mm. long; ae on lower surface amina strongly raised. .................0004 19c. C. calaba. 45. ies and midrib furfuraceous to puberulent; lamina with 3 to 9 veins/5 mm., midrib not disappearing below apex. 47. Venation on upper surface of lamina apparently twice as dense as on lower liniflorescence terminal |: 2-2 cone eee eee es ee a 161. C. sakarium. 47. Venation same density 0 on both sides of lamina. 48. Lamina + cuneate oe base, with rather obscure venation; inflorescence axillary, 167 axis less than 1.7 cm. long. ............... . C. aurantiacum. 48. Lamina broadly Pouniled at sn with distinct venation; inflorescence terminal and axillary, axis at least 2 cm. long. ........... 98. Calophyllum sp. 44. Lamina ei pal over 5 cm. long; terminal bud more than 3 mm. long; fruit not furfuraceous, over | cm 49. Lamina ae thinly coriaceous, with 5 to 10 veins/5 mm.; fruit at least 2.5 cm. long [usually seaside tree; trunk often not vertical]. ........ 8. C. inophyllum. 49. aries usually coriaceous, with (7 to) 9 or more veins /5 mm.; fruit less than 2.5(-3) long [not seaside tree; trunk usually vertical] . 50. "Venation on upper surface of lamina apparently twice as dense as on lower 51. Lamina sharply recurved at margin; inflorescence axis at least 2 cm. "Jong. Bde ae ee Seer aes tee eee Os eRe ae ae en ci 84c. C. fer rrugineum var. orientale. 51. Lamina not sharply recurved at margin; inflorescence axis less than 2 cm 0) da aa ee eae eS er EY On 23. C. rigidum (variant). 50. Venation same density on both sides of lamina. 52. Lamina rather closely undulate at margin, usually drying yellowish brown; inflorescence with relatively short basal internode and relatively long lateral pn [pedicel notably long (up to 3 cm.) and slender; fruit spherical, OU ace -& Ae tats i os Oe ee ee. SE oe ee 21. C. soulattri. : Lamina not closely undulate at margin, usually drying other than yellowish brown; inflorescence lacking relatively short basal internode, rarely branched. Nn N WNTIAHdOTVO ‘SNAAALS [0861 681 53. Indumentum tomentose, apne (on stem, terminal bud, and often Ns uw midrib of lower surface of lam 54. Lamina 3.8-12.5(-15) cm. cn drying chestnut brown; apical cells of hairs notably expanded. .. . . 45a. C. lanigerum var. lanigerum. 54. Lamina 12.5-18 cm. long, drying olivaceous-brown; apical cells of airs not notably CADANCEG: 5 aw ae % 34 GS 53. Calophyllum sp. . Indumentum i inconspicuous, crustaceous to subtomentose. 55. Lamina often hs fruit spherical, smooth, outer layer not detaching cleanly TIOM StONC) w..6..6:<. oce-9 tow Heo ee . teysmannil. Lamina oblon eelliptic: fruit + ellipsoid, wrinkled, outer layer detaching cleanly from stone 56. Lamina drying + flat; lowest internode of inflorescence 0.4-1.8 cm. long [usually in peat swamps, rarely in kerangas; latex yellow]. ee ee ee ee ee ae ee ee 85. C. sundaicum. 56. Lamina usually undulate at margin; lowest internode of inflores- cence (0.35-)0.7-3.2 cm. long [rarely in peat swamps, usually in kerangas or mixed dipterocarp forest; latex usually white eee et a Ge eo oR ge ee i eae On biflorum. a; oi) 32. Lamina ovate to oblong-elliptic (very rarely subovate), + acuminate at apex. 57. Twigs at least 3 mm. across, 4-subalate to alate, drying black, rarely brown; petiole (1.5-)3 cm. 83 OIMOre 1OGE. 61 Mt oe a ee oe a ee, a ke Se . C. macrocarpum. 57. Twigs often less than 3 mm. across, rarely 4-alate or la ae petiole shorter Lamina margin with clear thickening at least 0.3 mm. wide or with distinct submarginal vein. 59. Lamina thinly coriaceous; fruit less than 1.5 cm. long, sai: layer with air spaces developing. 100b. a ae ee a eS eS Se Co oC a C. tetrapterum var. obovale. 59. Lamina coriaceous; fruit over 1.5 cm. long, outer layer compact. .............02. df Shee eh is eden SR IPR ee Ee Ge at ee ee ee 8c. C. teysmannii var. inophylloide. 58. age margin sepa a Paaeoeny nor with distinct submarginal vein. 60. Axillar seat h (sub)ba ne ay Se across veins on lower surface of lamina; inflorescence aoe pr? Sak ee oe Ge, eh dO . C. garcinioides. WOLAYOPdUV ATONYV AHL JO TVNYNOL 19 “10A] 61. Lamina lacking irregularly ascending latex canals on lower surface; eee amma 62. Lamina 11.5-23 cm. long, lower surface subpersistently and shortly tomento bo AP tas Ashe Sh oe, ©. Ue ede a ae GE, ee ge Sted, GR ate, 63. C. ee 62. Lamina less than 11.5(-13) cm. long, lower surface glabrous. 63. Terminal bud and ae tomentose. 64. Terminal bud (3.5-)4.5-7 mm. long; lamina 6-13 cm. long; outer layer of Pruits(0.3=) 15 mm. hk. ys a ie he we ws oO 104. C. dispar. 64. Terminal bud 2-4.5 mm. long; lamina 3-7.8 cm. long; outer layer of fruit O,2-0,5 Mm, WICK. .g. owe eee ES De MR ee Ee eS a C. pisiferum. 63. Terminal bud and stem with puberulent to crustaceous indumentu . Venation on both surfaces of lamina inconspicuous (also in living leaf), latex canals sometimes impressed above; fruit spherical. .............. B GetS-oh.G-e Sip Bee Ge Se gear ee ae ae ce ao. iG. depressinervosum. 65. Venation on at least one surface of lamina prominent (also (?always) in living leaf), latex canals not impressed above; fruit stipitate 66. Midrib on upper surface of lamina not sharply raised, often drying reddish 64. C brown; internode 1-5 cm. long... ......... . roseocostatum 66. Midrib on upper surface of lamina sharply raised and/or not drying reddish brown; internode 1-3.5cm.long............ 65. Calophyllum sp. 60. Axillary shoot lacking (sub)basal scars. Inflorescence terminal, sometimes also from uppermost leaf axils. 68. Twigs drying yellowish; anthers with hairs; fruit ellipsoid. . 158. C. rubiginosum. 68. Twigs drying blackish, brown, or gray; anthers ce fruit subspherical. 69. Lamina with 9 to 19 veins/5 mm.; fruit less than 1.5 cm. long. .......... bh eo US es a GO ee eee ee 160. C. obliquinervium. 69. Lamina with 4 to 10 veins /5 mm.; fruit more than 1.5 cm. lon 70. Lamina drying grayish brown above; inflorescence axis and pedicel puberulent. l Peo ae be a ee eee ee eS oe a ee Gee og ke . C. griseum. 70. Lamina not drying grayish brown above; inflorescence axis and pedicel glabrous. Sd ee ee a oe a a ee ee 1. C. alboramulum. 67. Inflorescence axillary, if exceptionally terminal then also from axils along twig. WONTIAHdOTVO ‘SNSAALS [0861 161 ~ _ . Lamina eh drying yellowish brown, thinly coriaceous, margin closely undulate; inflorescence branched and ‘ or flabellate; POC SMOG S. G ee. ce ee oe eG ne HG as pe oh ee Oe A es ee DO ee ee . soulattri. . Lamina See drying other than yellowish brown, often coriaceous, margin variably undulate; inflorescence ne branched nor flabellate; fruit nearly always wrinkled or furfuraceous. 72. Terminal bud, and sometimes also twigs, with (dark) brown, tomentose indumentum. 73. Lamina coriaceous, margin not undulate, venation on upper surface apparently twice as dense as on lower [fruit 1.2-1.6 cm. long, the outer layer 1-2.5 mm. thick, compact]. ................000. 23. C. rigidum. 73. Lamina (thinly) coriaceous, margin + undulate, venation same density on both surfaces. 74. Twigs little flattened, strongly 4-angled to subalate; fruit furfuraceous 75. Lamina strongly recurved at margin; twigs 2.5-3.7 mm. across; filaments + connate. . .166. C. recurvatum. 75. Lamina not strongly recurved at margin; twigs 0.6-2.5 mm. across; filaments connate for up to 0.8 mm OUYe. tee eee hee ee OA Che Oke Sheba wee ee hoe ee ee oe ee 65. C. gracilipes. a | — 74. Twigs not as above; fruit not furfuraceous. 76. Lamina 9-22 cm. long; fruit at least 1.3 cm. long. 77, Lamina thinly coriaceous, closely undulate at margin; fruit pyriform.......... 63. a sae adaes 77, Lamina coriaceous, not closely undulate at margin; fruit + spherical .......... oodii. 76. Lamina less than 13 cm. long; fruit less than 1 cm. long. 78. Terminal bud (3.5-)4.5-7 mm. long; lamina 6-13 cm. long; outer layer of fruit (0.5-)1-1.5 mm. thick. Pl ce ge tee Ge SAD ee er Sy eee ae oe Gee ee a ee ee ee ee 104. C. dispar. 78. Terminal bud 2-4.5 mm. long; lamina 3-7.8 cm. long; outer layer of fruit 0.2-0.5 mm. thick. ...... Ges gists Be He eh ee Bae RR A ee Ee ee ee ee ee 102. C. pisiferum. 72. Terminal bud and twigs lacking tomentose indumentum 9. Lamina coriaceous, often shiny, venation often subobscure below; pedicel puberulent, usually incrassate and to mm. across in fruit; outer layer of fruit 1-1.8 mm. thick, with air spaces developing under skin........... Gna a Le Be ee. See eae ee a, a gee eg eR EN ee wn He ee 105. C. incumbens. 79. Lamina coriaceous or not, rarely shiny, ies + apparent below; pedicel glabrous to subtomentose, incrassate in fruit or not; outer layer of fruit not asa 80. Twigs drying ae to yellowish; te . -9(-10.5) cm. long, strongly acuminate at apex 81. Petiole 3.5-6 mm. long; lamina with 12 to - (to 28) veins/5 mm.; fruit strongly wrinkled, at least when young, outer ae developing large air spaces. ..................0000, 106. C. dioscurii. 81. Petiole 7-18 mm. long; lamina with 8 to 14 ae mm.; fruit almost smooth, outer layer compact. He oe St eee ae ae ae gy te ee eS ee Ee we a ee 107. C. banyengii. QOLAYOIUV ATIONYUV AHL AO TVNUNOL c61 19 “10A] 80. Twigs drying brownish to blackish; lamina usually not as a 82. Lamina (3.3-)9-22 cm. long, ‘with 10 to 18 (to a veins ny 5 mm.; inflorescence axis, pedicel, and back of at least outer pair of tepals puberulent; fruit ovoid. ............0......008.4 50. C. canum. Lamina usually less than 12.5 cm. long, with up to 12 (to 18) veins/5 mm.; upper part of inflorescence axis, pedicel, and back of outer pair of tepals usually glabrous; fruit spherical to ellipsoid (not known in Calophyllum sp. 164). 83. Lamina drying bicolored, tawny brown below; fruit spherical, ca. 1.2 cm. long, the outer layer detaching l oo Ss cleanly from stone, 1.3-2 mm. thick, compact.................... 63. Calophyllum sp. 83. Lamina not drying bicolored, other than tawny brown below; fruit not as above. 84. Terminal bud 1.3-2 mm. long, laser eeen uate internode beneath bud 1-6 a ee lamina with 13 to 18 veins /5 mm.; anthers less than0.4mm.long. ............ 4. Calophyllum sp. 84. Terminal bud usually longer, underdeveloped internode usually shorter a ae lamina usually with fewer than 12 veins /5 mm.; anthers at least 0.5 mm. long 85. Venation often rather obsee fruit furfuraceous. .............. 165. C. gracilipes. 85. Venation rather clear; fruit not furfuraceous. 86. Lamina ca. 3 times longer than broad, the midrib narrowing gradually from wide base, depressed; pedicel and back of outer pair of tepals puberulent. . 48. C. dasypodum. . Lamina usually less than 2!/2 times longer than broad, the midrib narrowing quickly near base and/or not very wide at base, not depressed; pedicel and back of outer pair of tepals glabrous. 87. Lamina thinly coriaceous, clearly acuminate at apex; fruit spherical .........., Sf te 70. eas Se Se Sy ea eee we 100a. C. tetrapterum var. tetrapterum. 87. Lamina coriaceous, + obscurely acuminate at apex; fruit ellipsoid. . . 87. C. biflorum oo ON 6. Key to Species of Calophyllum from the Philippine Islands to Botel Tobago and the Northern Pacific 1. Axillary innovation with basal scars; axillary inflorescence with basal internode less than 2 mm. long [terminal bud 1-2.5 ee ae ee ee ee ee ee ee ee ee ee ee a ee ee ee 62. C. pelewense. WOTIAHdOTVD ‘SNAAALS [0861 £6 1. Axillary innovation lacking basal scars; inflorescence with basal internode usually more than 2 mm. lo 2. 2. long. Terminal internode of innovation nearly always much shorter than others; lamina obovate to elliptic, acuminate at apex, 3 to) angle of divergence of venation 35-50"... 2... 160. C. obliquinervium. Terminal internode of innovation about same length as others; lamina not as above, angle of divergence of venation usually more than 50°. ee Soe ee Be Aik me et eo ee 171. C. brachyphyllum. sorganized by air spaces [lamina eG ae Abr ae Cakbae es ue. des eae SL, Ge cede a . pentapetalum var. pentapetalum. 6. Lamina with (3 or) 4 to 8 veins/5 mm., venation clear............ 90a. C. venulosum var. venulosum. 6. Lamina with at least 8 veins/5 mm., venation rather obscure. 7. Uppermost pair of axillary buds less than 1/s length of terminal bud, inconspicuous; lamina sparsely tomentose over entire lower surface... 2.2... 2... 95. Calophyllum sp. 7. Uppermost pair of axillary buds ca. Y/3 length of terminal bud, conspicuous; lamina tomentose on (also Ge Se hc ee A ag, a Se ee oe oe hg a ee We Ses ue he Sa 21. C. soulattri. 3. Lamina rounded to attenuate at base; petiole often more than 6 mm. long. b at first broad but abruptly narrowed at base: twigs drying A ta aw ee ee aah te oe, Os a ee Wa, Oe. S es a a. ae 55. C. cucullatum. 8. Lamina rarely broadly rounded and not cucullate at base, midrib at most subabruptly narrowed at base; twigs drying blackish to yellowish; indumentum tomentose or various. 9. Lamina 1.4~-3 cm. long, very coriaceous, with 13 to 16 veins /5 mm., margin plane, midrib on lower surface a OEDICSSCD oa he 6 Bb Od eGo Ge ho oe wo S ES oe OAR deh ee 172. C. rigidulum. sually not \O tt fet] 3. i=] jet) c ie) oa be) = a p = ge oO ot ee =) ~” ~” B lh > @) 5 oF o o> oh < oO Lan] oe i=" p 5 we) < Se i=] ~”n Baa (A 3 8 8 = ian ot. ion 2) 5 oO = oO =} wn c <5 oo e) oO (= depressed. 10. Lamina thinly coriaceous, closely undulate at margin, with 5 to 8 veins/5 mm., the venation prominulous above, obscure below, the latex canals impressed above; inflorescence with 3 to 5 flowers; fruit furfuraceous. Oe ET ee Miva AS GB a ag ee, Seas Bian sg. A St A Ses Bred eG de te eR ss en te . C. gracilipes. 10. Lamina + coriaceous, otherwise not as above; inflorescence usually with more than 5 flowers; fruit not S WOLAYOUUV ATONYV AHL AO TVNUNOL 19 Toa] 11. Inflorescences terminal (also axillary). 12 _ No Termi nal bud 1-4 mm. long; lamina acute to rounded at apex, drying flat; twigs not drying notably VClOW ISO ttt he yaar oy Se. a oh a we ee A eee Oe RK SRS . pentapetalum. : Terminal bud 2.5-20 mm. long; lamina + acuminate (rarely rounded) at apex, not drying flat; twigs sometimes drying notably yellowish. 13. Terminal bud 2.5—4.5(-6) mm. long; lamina drying concolorous, often nitid; hairs oie {lamina 153 often with conoid galls]. ©... 0. ee ee . C. whitfordii. 13. Terminal bud 6-20 mm. long; lamina usually drying bicolored, rarely nitid; hairs usually branched [lamina very rarely Wilh CONOIG-CaNS)s- invade un owe hewa dae be eS ae S . C. blancoi. 11. Inflorescences axillary. 14. Midrib on upper surface of lamina surrounded by raised blade, rather indistinct; twigs drying blackish; — aN terminal bud (1.5-)2-3 mm. long, perhaps not functional... .........0.., 68. C. oliganthum . Midrib on upper surface of lamina not surrounded by raised blade; twigs rarely drying blackish: terminal bud more than 3 mm. long, probably functional. 15. Midrib on upper surface of lamina narrowing gradually from base, + strongly sulcate or depressed. 16. Twigs strongly angled, short-tomentose. ...............0.0. 48. C. dasypodum. 16. Twigs rarely strongly angled, inconspicuously puberulent 17. Lamina elliptic, midrib on upper surface depressed; fruit Ca. 1.9 Ci, NOOR: x. ve 6 oa dee eo ace See I Boel ie a ae Oe Gs Se ee ee ee 128. 4G; oe 17. Lamina ovate to elliptic, midrib on upper surface sulcate to almost flat; fruit 1.9-2. Ci ONE. « gre & od. o...6 bee bee ee Ee eo oe es 121. Calophyllum Sp. 15. Midrib on upper surface of lamina not narrowing gradually from base, + strongly raised. . Terminal bud ca. 4 mm. long; lamina with 5 or 6 veins /5 mm., the \ venation apparent above, obscure below, the latex canals + impressed on lower surface; inflorescence with ca. 5 DIO WiCT Sy) io :xitm: ce Bah ee oe ee a as le a Oe RG ae ke ae 178. Calophyllum sp. 18. Terminal bud usually longer than 4 mm.; lamina usually with denser venation, the venation + apparent on lower surface, whether or not latex canals impressed; inflorescence usually with more than 5 flowers. 19. Uppermost pair of axillary buds erect, 1/3-1/2 length of terminal bud; lamina often closely undulate at margin; inflorescence usually branched and/or with flowers arranged flabel- lately; flower with 4 tepals; fruitsmooth.................. 21. C. soulattri. [0861 SNHAALS v WO TIAHdOTVO —_— — 19. Uppermost pair of axillary buds less than 1/3 length of terminal bud, usually + spreading; amina not usually closely undulate at margin; inflorescence rarely branched or with flowers arranged flabellately; flower with (4 to) 8 or more tepals; fruit only rarely smooth when ripe. 20. Lamina rounded to retuse or subacute at apex, drying + concolorous, with 4 to IO°VEINS/ SINR 64.8 aS ER EE EE eee ea Bs. ey inophyllum. 20. Lamina acuminate at apex, often drying bicolored, with 5 to 18 a veins /5 fee Woe! as ee Seek ees Seve eon ue ce ee ak de ho eae Gee a C. blancoai. 7. Key to Species of Calophyllum from Sulawesi to Australia and the Southern Pacific . Lamina 32-45 by 9-11 cm., with 3 to 5 veins/5 mm.; twigs strongly angled [poorly known species]. 179. C. macrophyllum. . Lamina smaller, usually with less dense venation; twigs usually not strongly angled. 2. Terminal bud strongly flattened transversely, leaf blades clearly visible [hairs much branched to stellate] . 3. Lamina (2.4-)6.5-17(-22) cm. long; ovary tomentose; outer pair of tepals at least 5 mm. long; anthers in staminate DOWwer 5455 fim, lone. 26-25 24 Bh EEE PAA EEE ERE GE ODER OHS , papuanum. 3. Lamina to 12.5 cm. long; ovary glabrous; outer pair of tepals less than 5 mm. long; anthers in staminate flower .5-2.7 mm. lon 4. Lamina subacute to rounded at apex, 2.5-5.5(-9.5) cm. long; tepals (rarely 7 or) 8; anthers in staminate flower O;teier Mate IOWES a.c koe Be tee ye ee Ge el a a eS eC oe Se 136. C. pauciflorum. 4. al apie to acuminate at apex, 3.4-12.5 cm. long; tepals 4 to 8; anthers in staminate flower 1.8-2.6 mm. 1GGR, Deke eee BEER ER eee eee eh eee Be tee EE ER eee 137. C. vexans. 2. Terminal “bud not strongly flattened transversely, leaf eae not aay visible. 5. Terminal bud less than 4 mm. long, or sometimes appa 6. Terminal bud apparently absent, underdeveloped aeaiede 3 "it 13) mm. long; lamina with 5 to 7 veins/5 mm., ancle of divergence of venation 45-60". 2:4 nk eh eR RR EROS Ee ES 117. C. savannarum. . Terminal bud present, underdeveloped internode inconspicuous; venation not as above. 7. Lamina Saari long-acuminate—caudate at apex . Lamina with 6 to 8 veins/5 mm.; axillary innovation with basal Scars. i.6 «4 dee 4 95 112. C. bifurcatum. 8. ae with (7 to) 9 to 16 veins / 5 mm.; axillary innovation lacking basal scars. . Lamina + nitid, margin closely undulate, midrib on upper surface raised, surrounding lamina also raised, nN 961 WOLAYOdUV ATONYV AHL JO TVNUNOL 19 “T0A] midrib on lower surface with + depressed edges. .................-.4.4 111. C. undulatum. 9. Lamina dull, margin not closely undulate, midrib on upper surface raised, but surrounding lamina not raised, midrib on lower surface not with eens CORES..6, 5:-3 eeeeeene SS: 4 ae ae eS 110. C. caudatum. 7. Lamina retuse to subabruptly acuminate at a 10. Lamina broadly rounded to cordate at Bee Lamina a very coriaceous; axillary bud recessed into stem and petiole base....... 116. C. articulatum. 11. Lamina coriaceous; axillary bud exposed. 12. Lamina with 4 to 7 veins/5 mm.; pedicel 2.2-3.5 cm. long............. 113. C. parvifolium. 12. Lamina with 12 to 16 veins/5 mm.: pedicel 0.3-1 cm. long. ......... 19e. C. calaba var. E. 10. Lamina other than broadly rounded to cordate (an occasional leaf broadly rounded) at base. 13. Lamina with 5 to 9 (to 11) veins/5 mm.; fruit 5-7 mm. long, the stone wall ca. 0.1 mm. thick, GAPlea! Boe SRS ee ee eee hee OE ee oe ee ee ee ee Seo 108. C.n as ee 13. Lamina with (8 to) 10 or more veins/5 mm.; if fruit less than 1 cm. long then the stone wall ca 0. thick, angled. 14. Lamina 2.2-5.5 by 0.5-1.5 cm.; fruit with angled stone. ......6..4654% 143. Calophyllum sp. 14. Lamina 2.7-9 by 1.2-4.5 cm.; fruit with rounded stone 15. Twigs drying yellowish brown: fruit ca. 1.6 cm. long, outer layer less than 1 mm. thick. ..... Siew: Se doe BG Ga RE Bh ee Ba Oe eS 152. C. leptocladum. 15. Twigs drying dark brown to blackish; fruit 2.3-2.8 cm. long, outer layer 2-3.5 mm. thick. . Lamina ca. 2 times longer than broad; fruit drying wrinkled, stone wall 1.3-1.5 mm. thick. eG ee ae eas Je Ee ee ee ee a Se Saree Gee eee Be ee Be eee 146. C. morobense. 16. Lamina ca. 3 times longer than broad; fruit drying smooth, stone wall i a Me 3 mm. thick. Be ee Me hee es aU OT oe Ge Se eet de es ee A ee a es ee Se ae Oe . C. confusum. 5. Terminal bud more than 4 mm. long, always present. 17. Midrib on upper surface of lamina raised, often not clearly distinct from lamina, surrounding lamina also raised. 18. Lamina (thinly) coriaceous; twigs up to 3.5 mm. across. i Uppermost pair of axillary buds erect; fruit less than 2 cm. long... ............ 21. C. soulattri. ppermost pair of axillary buds spreading: fruit ca. 5 cm. long. 18. oe thickly coriaceous; twigs (1.5—)3-7 mm. across 20. Twigs 5-7 mm. across; lamina ovate, 2—2!/2 times longer than broad; trunk with stilt roots, the sap clear yellow, sticky. bees Se He Ge, exe Gos Ge Ge AE te Ge wet oe, ee Se Ss he ee ee es a 148. C. suberosum. [0861 WOTIAHdOTVD) ‘SNHAALS 20. Twigs (1.5—)3—5 mm. across; lamina elliptic to oblong (rarely subovate), (2—)2!/2-3 times longer than broad; trunk without stilt roots, the Sap Opaque yellow, Tluid: 64444448) 8s bods ee od 149. C. igi 17. Midrib on upper surface of lamina raised or not, clearly distinct from lamina, surrounding lamina not rais oN, nN pa Lamina usually more than 9 cm. long, with (3 or) 4 to 9 (to 11) veins/5 mm., venation apparent; fier edeace axis (1.3-)3 cm. or more long; pedicel (0.4-)1-4.5(-6.3) cm. long. 22. Lamina less than 2(-2!/2) times longer than broad, drying other than sepia on upper surface and smooth even under strong magnification, midrib on upper surface often rather quickly narrowed at base. ..... be eb ee bk OR ee ooo, ee ee ee ee ee BOR we ae ee Bye ree OF inophyllu m. 22. Lamina usually ca. 3 times longer than broad, drying sepia on upper surface and minutely bullate under strong magnification, midrib on upper surface gradually narrowing from base........ 118. C. vitiense. . Not as above 23. Midrib on upper aes of lamina broadly depressed, at least in bottom !/s, margins not raised; terminal bud plump, conspicu 24. Midrib on upper ne of lamina usually narrowing in lower ca. 1/3; flower with (9 to) 12 (to 16) Ve oe ee oe oe ee ee ee eee eae ee ee 122. C. caledonicum. 24. Midrib on upper surface of lamina narrowing gradually from base; flower with 4 to 8 (to 12) tepals. 25. Lamina narrowly ovate to elliptic, usually drying nitid, midrib and venation notably richer brown than rest; stone with basal plug. ...........0.02..020....2000.2. 12. C. neo-ebudicum. Lamina narrowly elliptic to obovate, midrib venation not notably different color from rest; stone without basal plug [fruit not known in C. c 26. Lamina obovate, + retuse at apex, ered: see 0.3-1 mm. wide; tepals (6 to) 8. ..... pe Meee gece A tee ae ees et @ ce ee ee | Go ect ee ay ee ee ee 123. C. carrii. 26. Lamina usually + elliptic, + rounded to acute at apex, thickened margin variable; tepals (where known) 4 to 6. 27 oma often less than 21/2 times longer than broad, midrib on upper surface 0.25-0.6 m. wide at midpoint; terminal bud 0.5-1(-1.3) cm. long. ..........2.. Wt Fae Gras 15 ; ae at least 21/2 times longer than broad, midrib on upper surface 0.2-1.6 mm. wide at midpoint; terminal bud (0.8-)1-2.7 cm. long 20; wn No oa 61 WOLAYOPdUV GATIONYV AHL AO TYNUNOL 19 ‘10A] 28. Lamina with 6 to 11 veins / 5 mm.; pedicels 0.7-1.3 cm. long; outer layer of fruit 1 thin, disorganized by air spaces [flower unknown]. ....... 29. Calophyllum sp. 28. Lamina with (9 to) 11 to 20 veins/5 mm.; pedicel 0.2-0.8(-1.1) cm. long; outer layer of fruit (?0.5-)1-3 mm. thick, not disorganized by air spaces. . . 128. C. laticostatum. 23. Midrib on upper surface of lamina with raised margins even near base; terminal bud plump or not. . Inflorescence reduced to solitary flower; flower bud obpyriform; outer pair of tepals with indumentum on both surfaces [lamina ovate, long-acuminate, with (13 to) 16 to 19 (to 22) veins/5 mm.]....... C2 Gene Bee ee eae ee ee ee ee Pee bee eee es hee 3. C. insularum. 29. Inflorescence with more than one flower; flower bud (where known) spherical to ellipsoid; outer pair of tepals nearly always glabrous inside. 30. Lamina usually broadly rounded to cordate at base; petiole less than 8 mm 31. wa — Terminal bud to 5.5 mm. long, probably not functional: axillary innovation ae basal scars; lamina ste: caudate at apex [flower not known]. .......... 112. C. bifurcatum. . Terminal bud at least 5 mm. long, functional; axillary innovation lacking basal scars; lamina acute at apex a2. Uppermost pair of axillary buds 1/s-1/2 length of terminal bud, erect and conspicuous; inflorescence branches conspicuous, ca. 1/3 or more length of main axis; fruit less than i 22 1.5 cm. long, stone wall less than 0.3 mm. thick... .........002.. . C. brassii. 32. Uppermost pair of axillary buds shorter, + oe ore eae inflorescence branches, if any, relatively shorter; fruit more than 1.5 c ong, stone wall at least 1 mm. thick A aed panera: sterile material from Tae Jaya and westward keys out here with 4 to 7 veins/5 mm., venation apparent; stone not angled, with basal eke ee Ge ?P ae ee ey aie ae ee eee ce ois ee Sena ee eee On enna ee waliense. 33. Lamina with 9 to 17 veins/5 mm., venation subobscure; stone angled, Moline basal PUR. 6.46 f.2 kk oP OS hee Se oe de hee we eG aS 142. C. goniocarpum. WOATIAHdOTVO ‘SNAAALS [0861 661 30. Lamina rarely broadly rounded at base; petiole often more than 8 mm. long. 34. Twigs 3.5-6.5 mm. across, almost square or + 4-alate, often drying blackish; lamina thickly coriaceous 35. Terminal bud 0.7-0.9 cm. long; fruit less than 2.5 cm. long, stone wall less than 0.1 mm. thick, basal plug lacking. Leah eee hee eee Ge hee hee te Pee ee ee ea ee ee eee eee 0. C. celebicum. oh bee ek ee ee ee oe be ee re ee ee a ee ee ee ee ee 131. C. peekelii. . Twigs usually thinner, if as thick then not almost square or alate, usually drying brownish; lamina thickly coriaceous Ww BSN 36. Lamina retuse to acute at apex 37. Lamina usually less than 8 cm. lon 38. Tepals 8, outer pair puberulent on back; fruit 2.8-3.5 cm. long [Australia]... ........ 125. C. costatum. 38. Tepals 4 to 6, outer pair glabrous on back [flowers of C. collinum not known] and/or fruit less than 2.2 cm. long [Papuasia]. 39. Lamina with 6 to 11 (to 14) veins/5 mm.; inflorescence with terminal flowers + in groups of five; ripe fruit not known, but probably with angled stone... ..........2....... 140. C. heterophyllum. 39. rag with 9 to 20 veins/5 mm.; inflorescence with terminal flowers in groups of three; fruit with rounded 40. east 0.4-2 cm. long; hairs on terminal bud with several branches, often bifurcated at apex; inflorescence lacking persistent foliaceous bracts; fruit with outer layer 1.5-3 mm. thick, + compact. 24 40. Internode 1-6 cm. long; hairs on terminal bud with at most one or two branches, not bifurcated at apex; inflorescence usually with persistent foliaceous bracts; fruit with outer layer thin and disorganized MALE ACG. 4 ee th, ee he Gs ws “Ds ic So Se OS 138. C. hirasimum. 37. Lamina usually more than 8 cm. lon 41. aa much branched, often + persistent over entire lower surface of lamina; uppermost pair of axillary buds 3-10(-14) mm. long, spreading, conspicuous; fruit 2.8-6 cm. long. ............. 134. C. euryphyllum. WOLAYOUUV AGTONYUV AHL JO TVNYNOL 19 “104 ] 41. Hairs rarely much branched, very rarely persistent on lower surface of lamina; uppermost pair of axillary buds erect to spreading, rarely so conspicuous; fruit less than 3.2 cm. 42. Lamina with 4 to 10 (to 12) veins/5 mm., venation usually apparent on at least one surfac 43. Lamina broadly rounded (also apiculate) at apex, margin sharply and rather Scene recurved; fruit ca. 2 cm. long, stone wall less than 0.05 mm. thick. ..............2.2022.. 79. C. aerarium. 43. ia not broadly rounded at apex, margin slightly recurved; fruit variable in size, stone wall more than 0.2 mm. thick [not known in C. heterophyllum 44. pupae bud 0.6-1(-1.2) cm. long; lamina less than 10 cm. long; inflorescence with terminal flowers ETA PROMS Ol TtVGs Gp. ae co asd- we eud e. & Ge esgs Mead, Seah AT wg Mn 140. C. heterophyllum. 44. Terminal bud 0.8- 2 cm. long; lamina usually more than 10 cm. long; inflorescence with terminal WEIS I -PrONpS Ol UNECE: foe ok Ge GS oe ee ad wr eS 32. C. leleanii. 42. Lamina with (6 to) 10 or more veins/5 mm., venation often + obscure on one or both surfaces. . Lamina usually rather thinly coriaceous, margin closely undulate; uppermost pair of axillary buds + adpressed to terminal bud, conspicuous; inflorescence branched and/or flabellate; fruit with thick, COMpPACt-OULEL 1AVEr.. g ¢ ou ue wig ea we dae Hee RL ee R&A ee ee 1. C. soulattri. 45. Lamina not as above; uppermost pair of axillary buds variable; inflorescence unbranched or with inconspicuous lateral branches: outer layer of fruit often not compact. . Inflorescence lacking axis; fruit ca. 5 mm. long [poorly known species]. .... 141. C. rufinerve. 46. Inflorescence with axis; fruit at least 1.5 cm. lon 47. Inflorescence axis, pedicel, and back of outer tepals farinose-puberulent; flower with 8 tepals; 2 fruit with stone neither angled nor with basal plug. ..........2022. 126. C. obscurum. 47. par eee glabrous or pubescent, back of outer tepals glabrous; flower with 4 to 12 tepals; fruit with stone either angled or with basal plug. 48. ee with 8 to 12 tepals; stone rounded, basal plug present. .... . 120. C. cerasiferum. 48. Flower with 4 (rarely 5) tepals; stone angled, basal plug absent 49. Lower surface of lamina subpersistently tomentose; pedicel tomentose, 2.5-6 mm. long. hate hee ae Be aa ek ae oe ea ee oe ee 139. C. trachycaule. WNTIAHdOTVD) ‘SNAAALS [0861 102 49. Lower surface of lamina with inconspicuous indumentum; aay Se 5-9 mm. long (0-18: Mim. ta frit). wk see a ee Sw ee Se ee . C. goniocarpum. 36. Lamina acute to acuminate at apex 50. Terminal bud 1.6-3.3 cm. long; midrib on lower surface of lamina strongly angled. ........ 24. C. robustum. 50. Terminal bud to 1.5 cm. long; midrib on lower surface of lamina rounded to subangled. 51. Lamina usually thinly coriaceous, margin closely undulate; inflorescence branched and /or flabellate; fruit spherical, the outer layer thick, compact, the stone thin, unmarked [uppermost pair of axillary buds usually erect and COOSPICUOUSI« « = 3-544 fon he See ee eee Shee Ee EEE PA EAAS CDRS 21. C. soulattri. . Lamina not as above; inflorescence usually unbranched; fruit not as abov 52. Inflorescence without axis; pedicel pilose; fruit ca. 0.5 cm. long ese known species]. .141. C. rufinerve. 52. Inflorescence with axis; pedicel usually glabrous, fruit (0.7-)1 cm. or more long. 53. Indumentum dense, tomentose, persisting on terminal bud, stem, lower surface of lamina (frequently), inflorescence axis, and pedicel. 54. Lamina with 4 to 9 veins/5 mm., usually drying bicolored [brown to vinaceous-olivaceous above honey to sabelline below]; fruit less than 1 cm. Jone, <2 49 4 24 440. 404/44 109. C. bicolor: 54. Lamina with 12 to 20 veins/5 mm., not drying notably bicolored; fruit more than 2 cm. long. oe be eae Ae ae oe ee ae el See oe Pe oe eee ete ee ee 139. C. trachycaule. . Indumentum usually other than tomentose, pedicel and lower surface of lamina glabrous or almost 17a) — Ns WwW O. 55. Lamina very coriaceous, margin almost plane, midrib narrowed gradually from base, sharply raised, venation obscure on both surfaces; fruit ellipsoid, stone sharply pointed at ae ee ee ee Vee eee eee ee eee we eee oe ae ee ee ee 147. C. acutiputamen. 55. Lamina not as ere piraner Neyo stone at most obtusely pointed at apex. 56. mane bud 4 ong; a with 7 to 9 veins/5 mm., venation — latex canals mpressed aa ee at ie ca S tiny LOMB vec ee ow ee Se oe C. le arpu 56. Terminal bud usually longer than 5 mm.; lamina with (6 to) 8 to 22 veins 7 5 mm., venation apparent, latex canals not impressed below; pedicel at anthesis (where known) more than : mm. long. COT WOLAYOUUV GIONUV AHL AO TVNUNOL 19 “T0A] 57. Flower (where known) with 8 or more tepals and/or fruit with basal plug. 58. Terminal bud 2-5 mm. long; lamina with midrib 0.1-0.2(-0.3) mm. a at midpoint. Mee see eS tas Gee nny cf ese etd Cane Near n eaar R A Te e eeNe Gh ae morobense. 58. Terminal bud (0.4-)0.6-1.6 cm. long; lamina with midrib (0.1-)0.2-0. ecu 8) mm. wide at midpoint. 59. Midrib on upper surface of lamina rather narrow and + raised even at base... . . eg ee eh ar ne ee Oe a ea ae oe ee ae aera 145. C. streimannii. 59. Midrib on upper surface of lamina broad and + depressed at base . Twigs not often drying coarsely Striate; lamina usually drying olivaceous- umber and nitid, midrib and veins notably browner than rest of lamina. ........ ran ie Gade Ga eee ee oe ae ee we Qe se Oe EY, C. neo-ebudicum. 60. Twigs soon becoming coarsely striate; lamina drying dull sepia, midrib darker, veins not discolored. 2.4 4444 48 eevee ee Pee es 120. C. cerasiferum. 57. Flower (where known) with ca. 4 tepals and/or fruit lacking basal plug. 61. Lamina with 12 to 22 veins/5 mm.; outer layer of fruit developing air spaces, stone Clearly ANPleG. , collections with stones that have basal plugs; Map 10. Distribution of Calophyllum soulattri (stars in Southeast Asia—Australia. 283 284 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 5394 (L, sING), Tjiakar, 500 m., Meijer 4765 (cANB, K, L, SING), base of Mt. Sago, Kampong ge Panjang, 600 m., Meijer 5648 (L); Priaman, aa aie HB 2359 (Bo, L, u). Utara: Padang Si Dimpoean, Padang Lawas, Sosopan, Aek Si Olip, Toroes 5447 (a, G, K, ec Is. Morsala, Silabua, 5 m. ee collector 16, anno 1951 (a, Bo, K, L, SING); Sibolga, Baroes, Koboen (M. Tapoes), O m., bb 31028 (a, Bo, L, Ny, SING); Zuid Nias, Hili Batoe, 10 m., bb 5754 (Bo). ATJEH: Karolanden, Lao Pengoeloe, bb 12514 (Bo), Lao Soeloe, 200-300 m., bb 9287 (Bo); Labeohan Batoe, Bila, Goenoeng Panjaboengan, Toroes 4357 (a, NY, US), Pasar Baroe, bb 8890 (Bo), Goenting Sago, bb 7725 (Bo); Simaloer Is., Tapah (Defajan), Achmad 1691 (Bo, L). BANGka: Lobok Besar, Perlang near Kabu, 50 m., bb 34191] (a, k, sinc); Rias, 10 m., bb 15396 (Bo, Sarawak. Ist Division: Kuching, S 21597 (kK, L, SAR, SING); Lundu, S 9558 (K, L, SAR, SING); Semengoh F.R., 30 m., Stevens et al. 285 (a); Mattang, Beccari P. B. 1299 (F1); Telok Paku, Bako N [atl.] P[ark], S 1475] (a, K, SAN, SAR sING). 3rd Division: Batang Lassa, S 33634 (f) (KEP, SAN, SAR). 4th Division: Batang Bintulu, 0 m., S 12098 (k, L, SAN, SAR, SING). BRowet, Andalau F.R., 60 m., Ashton s.n., July 1959 (sar); Sungei Samit (Belait), KEP 30429 (x, KEP). SABAH. Ranau: Mile 9, Telupid—Ranau road, SAN 73828 (k, L, SAN). Sandakan: Ulu Dusun F.R., 5 m., Stevens et al. 350 (f) (A). Kinabatangan (Lamag): Bintang Mas Karamuak, 21 m., SAN 81641 (a). Lahad Datu: Matrid, A 2493 (kK, KEP, L, SING); Silam, 90 m., SAN 47680 (k, L, SAN, SAR, SING); Pulau Sakar, 12 m., SAN 25363 (kK, KEP, L, SAN, SAR, SING); Kelumpang F.R., Mostyn Lor Malang, 30 m., SAN 40883 (k, L, SAN, SING); Tembaran Is., 30 m., KEP 80511 (KEP); Kunak, 15 m., SAN 68098 (k, L, SAN); Madai F.R., 90 m., SAN 46117 (kK, L, SAN). Semporna: Semporna, near Agr. Station, SAN 26321 (?f) (BO, K, KEP, L, SAN, SAR, SING). KALIMANTAN. Timur: Salimbatoe, Sungei Roemah, 25 m., bb 11237 (f) (Bo); Mara, 150 m., bb 10833 (f) (Bo); Mt. Ilas Bungaan, m., Kostermans 13937 (f) (Bo, K, KEP, L, LAE, NY, SING); Tandjong Redeb, Kelai R. near Long Lanuk, 10 m., Kostermans 21129 (f) (Bo, CANB, G, K, L); Tundjung Plateau, near Djohan Asa, 100 m., Kostermans 12623A (so, CANB, K, KEP, L, SING); Tidoengsche Landen, Malinay (Kabiran), 10 m., 17855 (a, Bo); Bloe-oe, Jaheri 1489 (Bo); W. Koetei, Mendom (Sei Atan, Klindjan), 100 m., bb 29268 (f) (Bo, L, Mo); Boven Makaham, Laham, 40 m., bb 20607 (a, Bo); E. Koetei, G. Tepian Lobang, on Menubar, NE. of Sangkulirang, 200m., Kostermans 6034 (f) (A, BO, K, SING). Selatan: G. Pamatton, Korthals s.n. (L); Doesson, Korthals s.n. (Lt); Martapoera, Manoekapan, Labohm 207]! (Bo), Djoengoer, 350 m., bb 10413 (Bo); Afd. Pleihari, Kampong Kintap, bb 2154 (Bo, L), Bentok Darat, 300 m., bb 10723 (Bo); a Alang, 200 m., bb 14199 (Bo). Tengah: Sampit, Samra ang, 30 m., bb 10540 (Bo), Koeling, 10 m., bb 9858 (f) (Bo); Afd. Beneden Dyak., ye Kladan eneN bb 2097 (Bo, t). Barat: Moera Kojong, 10 m., bb 7479 (Bo); Palo, 5 m Becking 35 (Bo); Mempewa, Toho, 25 m., bb 6326 (+f) (Bo). Philippine Islands. PaLawan: Binohan Mt., near Puerto Princesa, Ebalo 395 (a, NY, UC). CALAMIAN: Busuango Is., Malbato, Marche B305 (rp). Minporo: Mt. Yagaw, 500 m., PNH 19091 (1, pnu). Luzon. Mountain: Bontoc subprov., Vanoverbergh 1888 (7h) (P), 3280 (7h) (F). Cagayan: sine loco, FB 22061 (us). Nueva Vizcaya: Caraballo Mt., Loher 13636 (mM, us). Pangasinan: sine loco, FB 8298 (ny). Nueva Ecija: sine loco, FB 22182 (BM, Bo, Pp, us). Aurora: Baler, Principe, Merrill 1049 (k, Ny, us). Bulacan: Angat, Species Blancoanae 336 (A, BM, BO, 1980] STEVENS, CALOPHYLLUM 285 GH, s, w). Zambales: sine loco, FB 5841 (?c) (Ly). Rizal: Montini, one 12299 (h) (a, M, uc); Manila, San Mateo, idan 976 (FI, k). Bataan: Lamao R., Mt. Mariveles, FB 1812 (k, ny, us). Laguna: Famy, aca eae 500 m., PNH 39376 (= Lagrimas Softy 23 sean 959) Se “ ioe NG); San Antonio, FB 13197 (Bm, Bo, K, Ly, P Bice BS 28632 (h) (A, BO, K, P, us); Lucban, 1000 m., O iiner ee (s) (A, BO, E, F, FI, G, K, L, LY). Camarines Norte: sine loco, FB 30616 (ny). Camarines Sur: Sipaco, FB 30331 Cae Albay: Monito, FB 27383 (c) (a); Bataan Is., BS 80689 (kK, MICH, NY). Sorsogon: Cumadcad R., 150 m., PNH 9776 (c) (a, PNH); Irosin (Mt. Bulusan), Elmer 17385 (s) (A, BISH, BM, BO, C, F, FI, G, GH, K, MO, NY, P, U, UC, US). CATANDUANES: Sine loco, BS 30293 (A, US). SIBUYAN IsLaNnpD: Capiz, Macellanes (Mt. Be: Giting), 310 m., Elmer 12293 (nisu, BM, BO, E, F, FI, G, K, MO, NY, W). Ticao IsLanp: sine loco, FB 1085 (K, Ny, Us). Samar: sine loco, BS 17493 (Bm, kK, us). Leyte: sine loco, Wenzel 1613 (A, BM, F, G, MO, NY). NeGros: Negros ne Dumaguete, Cuernos Mts., 1067 m., Elmer 9837 (h) (A, BM, E, FI, G, K, LY, MO, US, W). PANay: Guimaras Is., FB 317 (Ny, us). MINDANAO. Surigao: sine ec FB 25074 (a, kK, Us). Agusan: oe (Mt. ace 610 m., Elmer 13989 (a, BISH, BM, C, E, F, FI, G, NY, P, U, W). Lanao: Lake Lanao, Camp Keithley, Clemens s.n., Feb. "1907 a0 June, 1907 (xk). Bukidnon: Mt. Candoon, BS 38937 (h) (a, K, P, US); Mt. Dumalucpihan, BS 38967 (a, p, us); Kimantri Plain, Malaybalay, FB 30279 (B, BO, P, US). Davao del Sur: Sete Cruz, Williams 2770 (a, ny, us). Java and adjacent islands. Barat: Buitenzorg, Tjikeumeuh, 250 m Bakhuisen f. 3563 (Bo, L, uv); Pakantjilan, Bakhuisen f 276 (vu); SE. van Depok., Tjilong, Backer 31212 (Bo); Bodjong Eyot., 200 m., Bakhuisen f. 6351 (so, L); Tjitjadas, SE. of Batavia, 15 m., Lanjouw 8 (A, BO, K); Kampong Baru, 200 m., Schiffner 2245 (a, Bo, K); Kampong Nangrang, 250 m., Schiffner 2244 (Bo, L); Tjamplong, Kuhl & van Hasselt s.n. (L); ee Kulon Nature Reserve, Mt. Pajung, 300 m., Kostermans s.n., 17 Dec. 1960 (a, Bo, L, sING), Tjiuluran- Sangiang Sirah, 100-300 m., Wirawan 344 (f) ns K, L, SING). TENGAH: Djokja- karta, Sleman, Kalioerang, 1000 m., Ja 2569 (Bo, L); Karel bij Karangmodjo, Boerrigter 85S (Bo); Semarang, Koorders 2873 (Bo, G); Kedoengdjati (Djam- bosschen), Koorders 25403 (po); Djapara, Soemanding, Ja 1837 (Bo); Kedoe, Weetoetoe, W. van Kedjadjar, 500 m., Plokheim s.n., 9 Nov. 1921 (so). Timur: montis Ngebell, prope Pondrongo, Waitz s.n. (1): Kediri, G. Pandan, Thorenaar 204 (Bo). Kartmoenpsawa: Tandjong Gelam, Karta 305 (Bo). BAWEAN IsLanp: G. Besar, 400 m., Buwalda 3192 (= Ja 4234) (f) (4, Bo, L). Lesser e : Batulanteh, 500-600 m., Kostermans 18611 (f) (Bo, G, L, P), 19076 (A, BO, C, CANB, G, LAE, NY, P, SING); Matemaga, 650 rm., bb 14050 (po); Kawowo, 700 Mt. Ndebi, 400 m., Kostermans s.n., April 1965 (f) (L); Werang (Kempo), 350 m., bis Berg Beliling, 1600 m., ‘Schmutz 2018 (?f) (L); Denge to Wai Rebo, 900 m., Schmutz 3537 (f) (i): Nunang, 900 m., Schmutz 663 (?f) (1); Ngada, Nderoe, 800 m., bb 11399 (po); Maoemere, G. Egon, 1703 m., bb 6897 (Bo). Sotor: Belodoea, 750 m., bb 15975 (Bo). Wetar: Kali, N. v. Ilwaki, 700 m., bb 27200 (?f) (Bo, K, L, sinc). Celebes. TaLaup: Karakelang, G. Duata, 200 m., Lam 2925 (a, Bo, x), G. Piapi, 400 m., Lam 3263 (Bo, K). SULAWESI: Manado, Amoerang, Loban Kolai, 250 m., bb 17133 (A, BO); Paloe, Negilalaki, 286 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 1250 m., bb 28259 (Bo), Banggai, Pongian, 100 m., bb 31876 (r) (Bo); Minahassa, Kajoewatoe, 200 m., Koorders 17292 (so, P), bij Tondano, 900 m., Koorders 17302 (Bo, P), Pinamorongangebergte, 500 m., Koorders 17301 (Bo, P): Boleang Mongondow, Rangagon, 250 m., bb 32482 (Bo); B. Boalemo, Popaja, 250 m., bb 15698 (Bo); Mamasa, Pena, 1500 m., bb 33066 (?) (Bo, L); Enrekeng, 2033 (r) (Bo), nabij La Rona, bb 2370 (30), Boschafd. Bataemati, 600 m., bb 23570 (po), B. Takale Kadjoe, 1500 m., bb 24074 (?) (Bo, L), Oesoe, Cel. IIT /63 (r) (Bo), Kawata, 200 m., Cel. V/318 (r) (aA, Bo); Sungei Tolewonu, Musser et al. 528 (a); Sungei Sadaunta, Musser et al. S-19a (a); oe 100 m., Kjellberg 864 (Bo); Lepo-Lepo, presso Kandari, Beccari s.n. (FI, herb. Becc. 1138); Mengkoka, Kolaka, 0-100 m., Elbert 3229 (r) (A, BISH, BO, CANB, FI, G, SING, US). KABAENA: Balo, Eempuhu, Elbert 3305 (r) (Bo). Muna: Onsoeme, 100 m., bb 21773 (r) (A, Bo). Butunc: Bone, Bonteriloe, 1100 m., bb 29018 (Bo). Moluceas. SANANA: Molboefa, 200 m., bb 29819 (Bo, L, sING),; Soelabesi, Hulstijn & Atje 410 (Bo, L). HALMAHERA: Tasoa, G. Sembilan, 300 m., Pleyte 266 (A, BO, K, SING). Morotai: G. Sabatas, 180 m., Lam 3567 (A, K). BATIJAN: sine loco, de Vries (795) (L). Buru: Kak Toea, 800 m., bb 22831 (L, Mo); Wai Geren Olon, 800 m., bb 21506 (Bo, L). AMBon: Waai, Mt. Salahatu, 200 m., Kuswata & Soepadmo 255 (a, BO, K, LAE, P, SING); Benteng, Rant 533 (so); Amboina, Pl. Rumph. Amboin. 482 (Bo, F, K, L, MO, P, SING, US), Hitoe Messen, 150 m., Pl. Rumph. Amboin. 481 (BM, BO, GH, K, NY, P, US); Latoera, Boerlage 415 (po). SERAM: Kairatu, Gemba, 0-5 m., Kuswata & Soepadmo 110 (A, BO, L, NY, P, SING), Waiselang, 150 m., Kuswata & Soepadmo 232 (a, BO, CANB, K, P, SING); Roembatoe, 800 m., bb 23035 (Bo); W. of Piroe, 1-100 m., Rutten 1645 (po, kK, L); Porong Mountains, Makina, 1000-1200 m., Rutten 2246 (a, Bo); Wae Kali, 100-200 m., Kornassi 1017 (Bo, kK, L); Kiandarat, 60 m., bb 25839 (Bo). Papuasia—see Stevens (1974). Also, Kar Is-anps: Keteil a Tual, Beccaris.n. (Fi, herb. Becc. 1139-1141). Tanimpar IsLanps: Makatian, 30 m., bb 24403 (a, Bo); P. Jamdena, Weri Ranarmoje, Buwalda 4676 (a, BO, K, SING); near resthouse between Kp. Ingei and Otember, Buwalda 4216 (A, BO, K). Aru Istanps: P. Kobroor, Dosinamalaoe, Buwalda 5093 (a, Bo, kK). IRIAN Barat. Geelvink Bay: Japen, Seroei, bb 30629 (a, sinc). Papua New Guinea. Manus: Derimbat, 50 m., LAE 52459 (a, Laz); Kaguli Ridge, 30 m., LAE 52940 (Lae); Mt. Dremsel, 600 m., LAE 59153 (Lak). Australia. NoRTHERN Territory: Melville Island, Dunlop 3435 (pNa, cans); Port Darwin, anon., anno 1890 (sri). Caroline Islands. Patau: Urukthapel, Dutton 80 (us). Ecotocy. Small tree of lowland or colline rainforest, rarely dominant in canopy (Great Nicobar Island); to 1700 m. alt. Prefers well-drained habitats but sometimes in swamp forest, where it develops small knee roots (Malaya— Corner, Joc. cit.; Sabah—pers. obs.). In Sabah, Papua New Guinea, and the Sol Islands (Santa Isabel), also over ultramafic rock. Also in secondary forest. Flowering throughout year, but in Java mostly September to December. (In Cambodia flowering in October, fruiting December to March—Dy Phon 2012). Entire inflorescence white (except yellow anthers and reddish stigma); flower scented. Ripe fruit purplish black. Galls—conical protuberances ca. | mm. tall—occur just in from the edge 1980] STEVENS, CALOPHYLLUM 287 of the lamina on Pleyte 266 (from the Moluccas). Similar galls on a specimen from Sumba Island were caused by gall midges (Docters van Leeuwen- -Reijn- vaan & Docters van Leeuwen, 1941). GERMINATION. The radicle breaks the stone to one side of the base, and the seedling has two pairs of leaves usually separated by a very short internode, although in Stevens et al. 285A the internode is up to 1 cm. long. Subsequent internodes are ca. | cm. long; their length is not affected by the light or nutrient status of the plant. The terminal bud is functional, and growth is erect. Eventually internodes more than 5 cm. long are produced (Stevens, 1974, loc. cit.; Stevens et al. 285A, 358, 916). Burger’s account (1970) of the germination does not agree with this. On Manus Island, and probably also on Bougainville and the Solomon Islands (Papuasia), the radicle pushes out a well-developed basal plug 5-7 mm. across during germination. The seedling has two pairs of leaves separated by a y short internode; the first internode is up to 5 cm. long (LAE 52458, "$2459: NGBF 1219). Locat uses. Although the wood of Calophylium soulattri is not very durable, it is used for masts and spars and in house construction throughout its range. In Bangka “‘getah malang-malang”’ is used to poison dogs. The bast, ‘‘babakan slatri,’’ is given to horses in Djakarta once a month to keep them in good condition. An infusion of the root is rubbed on to alleviate rheumatic pain. Oil from the seeds is used like that of C. inophyllum (q. v.); the sourish fruits can be eaten, but if eaten in excess they cause severe stomachache and diarrhea (Heyne, Joc. cit.). In the Caroline Islands fresh bark from the shoot is used as medicine for women who have just given birth (Lauterbach, as C. cholobtaches). On the Huon Peninsula (Papua New Guinea) the leaves are reported to make a durable thatch (Clemens 184). Although a variable species, Calophyllum soulattri is usually readily recog- nizable by the well-developed uppermost pair of axillary buds that are nearly always tightly adpressed to the terminal bud, and by the somewhat thinly coriaceous lamina that is rather closely undulate at the margin and that often dries a characteristic olivaceous-buff on the upper surface, the venation being fine. The inflorescence, which nearly always has lateral branches half the length of the main axis or longer, is also distinctive. The fruits are usuall subspherical and dry blackish and smooth; the outer layer is well developed and compact, and the stone wall is thin. The hairs show little Variation; they typically have a single basal branch. The epithet soulattri is derived from a Javanese local name for the plant. Calophyllum soulattri has been confused with C. kunstleri (= C. rigidum). However, the venation on the dry lamina of C. kunstleri is apparently twice as dense on the upper surface as on the lower, the inflorescence is unbranched (the flowers tend to be arranged in pseudowhorls), and the fruit usually dries brown and wrinkled. In Malaya, sterile material of C. soulattri and Gc il has been confused; for differences between these species, see TABLE 6. 288 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Variation within Calophyllum soulattri The greatest variation within Calophyllum soulatiri is in the Philippines- Celebes-Moluccas area. Although some of this variation is rather trivial (in such characters as indumentum, leaf texture, and midrib type—this perhaps rather surprisingly) and is neither correlated with other characters nor always constant within a collection, field work in this area may lead to some modification of the limits of C. soulattri. More important variation occurs in the fruit (e.g., the larger-fruited specimens from the Tanimbar Islands). he rather remarkable variation in germination, seedlings, and fruits in C. soulattri from the easternmost part of Papuasia (see below) may eventually lead to the plants there being given formal recognition. In the area from Java and eastern Borneo to the Moluccas and the Aru Islands, there are a number of specimens with strongly flattened twigs that are rounded at the edges (specimens ‘‘f’’ in the list above). Such specimens also tend to have a thicker lamina that dries a browner color than is usual; the indumentum is often, but not always, poorly developed; and the pedicels are strongly incrassate in fruit. In specimens of this type from the Lesser Sunda Islands, the submature fruits are about 1.5 cm. long and dry brown and wrinkled: in Borneo (e.g., Kostermans 6034, Kalimantan) the fruits dry brown and smooth, with the stone walls about 0.7 mm. thick—thicker than is usual in C. soulattri. Kuswata & Soepadmo 253 (Ambon), which also has flattened twigs, has a chartaceous lamina and stones with walls only about 0.4 mm. thick. In a number of specimens from Sulawesi (‘‘r’’ in the list), the midrib on the upper surface of the lamina is surrounded by raised blade. However, such specimens are not otherwise unusua Fruiting specimens from the Tanimbar islands (e.g., Kuswata & Soepadmo 110) have fruits up to 2.3 by 1.8 cm. that dry blackish and wrinkled; the stone walls are about | mm. thick. Such specimens have not been incorporated in the description and are included here faute de mieux; they possibly approach Calophyllum leleanii. Two specimens from Sulawesi (bb 24074, 33066) have large, ellipsoid fruits 2-3.3 cm. long with ellipsoid stones that are pointed at the apex and have walls ca. 1.5 mm. thick. Specimens collected by Kurz from the Nicobar Islands also have rather large fruits; in this case the wall of the stone is ca. 0.8 mm. thick, and the specimens are more typical. Specimens similar to those from Papuasia that have well-developed tomen- tose indumentum on the vegetative parts and on at least the lower part of the inflorescence axis (see Stevens, 1974a, loc. cit.) occur elsewhere in the range of the species. However, there are intermediates between such specimens and those with inconspicuous indumentum, and it is unwise to recognize taxa based on indumentum characteristics. Calophyllum spectabile var. ceramicum was described from a robust specimen with leaf blades up to 27 by 10.2 cm.; such specimens (€.g., Pleyte 266) are quite common in the Moluccas. Similar specimens have also been collected in central Sulawesi, where the local people consider that the plants belong to a species distinct from the small-leaved form (compare Musser 1980] STEVENS, CALOPHYLLUM 289 et al. 528, the large-leaved form, with Musser et al. 507, the smaller-leaved form). However, it seems premature to recognize a taxon based on these large-leaved specimens. Elmer based his invalidly published name, Calophyllum sorsogonense, on a plant that has the inflorescence, flower, and fruit characters of C. soulattri, but has a lamina that is shallowly cordate at the base and dries a rather dark, dull brownish gray, and a petiole that is short (specimens “‘s’’ in the list, from the Philippine Islands); at first sight this is a very distinctive form. However, the leaves of some specimens from Sulawesi have well-developed petioles and blades that are more or less acute at the base and more or less truncate at the apex, being finally mucronate; in characters like indumen- tum and color of the dried leaf, they are similar to C. sorsogonense. Such specimens (e.g., Koorders 17300, to which the name C. oblongum Koorders has been given) also approach some from Java (e.g., Koorders 28105). Sterile specimens recently collected in Sulawesi (e.g., Musser et al. 1044) also have cordate leaf bases, but are otherwise quite different. The type specimen of Calophyllum hibbardii (Elmer 9837) has leaf blades that dry a rather dark brown color; although the lateral branches of the inflorescence are long, the inflorescence is not flabellate, and its lowest internode is up to 1.7 cm. long. (The isotype of C. hibbardii at Lyons has a terminal inflorescence.) The pedicels in young fruit are up to 3.2 cm. long, and the fruits are immature—when mature, they might be rather large for C. soulattri. Calophyllum hibbardiiis a fairly distinct variant of C. soulattri, but it cannot be recognized formally (specimens matching the type of C. hibbardii are denoted by ‘‘h’”’ in the above list; they are known only from the Philippine Islands). Other specimens from the Philippine Islands (‘‘c’’ in the list) have coriaceous leaf blades with relatively distant venation, but they are otherwise like Calophyllum soulattri. Stevens (1974a, loc. cit.) cited NGBF 1089A (from Manus Island, Papua New Guinea; consisting only of fruits) as Calophyllum pseudovitiense (= C. neo-ebudicum), albeit with some hesitation: the stones have basal plugs (as in C. pseudovitiense) but were picked up underneath a tree of C. soulattri. Further collections from Manus Island by D. B. Foreman clearly show that such fruits come from a tree that in vegetative and floral characters agrees with C. soulattri. However, the relatively thick-walled stones have basal plugs that are pushed out during germination, and the young plant has long internodes from the very beginning; in all young plants seen of C. soulattri sensu stricto (in Sarawak, Sabah, various places on the Papua New Guinea mainland, and New Britain), the internodes are initially short. Similar fruits occur probably on Bougainville (NGBF 1219) and certainly on the Solomon Islands (New Georgia group: BSIP 192, 1249, 2812, 5953, 6144, 11597, 15808); they may be up to 2.5 cm. long. The only other specimen of C. soulattri sensu lato with mature fruit seen from the Solomon Islands (Choiseul: BSJP 17507) had stones with a discolored area ca. 2.5 mm. across at the base, as in ‘‘normal’’ C. soulattri, but no plug. Latex of specimens from the Solomon Islands is frequently reported as being yellow(ish) in color, sometimes clear 290 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 (e.g., BSIP 192), but not always (BSIP 3833 and 2388 apparently had white latex). NGBF 1219, taken from the tree under which the stones with basal plugs were collected on Manus, had white latex. Calophyllum soulattri on the mainland of Papua New Guinea sometimes has yellow latex (e.g., Hartley 11906), although it is usually white. Fruits of both types turn blackish purple when ripe. Clearly much careful field work is needed to understand this variation; it should be remembered that C. paludosum C. T. White was described from a specimen (BSIP 192) that has stones with a basal plug. Synonymy and Nomenclature In the following discussion, reasons are given for the reduction of several species to synonymy under Calophyllum soulattri, and a number of names are lectotypified. Further details of the synonymy of C. lanceolatum and names based on specimens collected in Papuasia are given in Stevens (1974a). Calophyllum soulattri was described from material originating from Java. At Geneva there are three specimens of this species originally in Burman’s herbarium and subsequently in that of Delessert; two are annotated ‘“‘soulattri”’ (by different hands, one apparently by Burman). Mermill (1917, loc. cit.) was correct in his equation of Bintangor montana Rumphius (Herb. Amboin. 2: 216. ¢. 72. 1741) and B. montana secunda Rumphius (ibid., p. 217) with Calophyllum soulattri: Rumphius mentioned the characteristically drying leaves. Bintangor montana secunda was described as having leaf blades ‘‘bifid’’ at the apex; specimens of C. soulattri with retuse apices are uncommon, even in Sulawesi. Apoterium sulatri is lectotypified on a specimen apparently annotated by Blume held at Leiden (herb. Lugd. Bat. 903, 343-183). The original publication of Calophyllum tetrapetalum must be ascribed to G. Don, not Roxburgh. Don described C. tetrapetalum from a stove plant, and the description fits C. soulattri: “leaves ovate or oblong lanceolate, obtusely acuminated. Racemes axillary, short, corymbose, 2-3 together... . Native of the East Indies.’’ I have not seen authentic material of C. tetrapetalum Roxb. ex G. Don. Roxburgh’s description (1832, loc. cit.) of a plant with very finely serrulate leaves is at first glance that of a different species (or genus!). However, a Roxburgh specimen from the East Indies at the British Museum, although annotated C. tetrapetalum, is clearly C. soulattri: because of the way that the specimen was pressed, the rather closely undulate leaf margin appears to be serrulate. The name Calophyllum suriga is a superfluous name for C. soulattri, since the latter is cited as a synonym in the protologue. Whether or not the species described by Carey (in Roxburgh, 1832, Joc. cit.) is to be assigned to C. soulattri is another matter (see also Planchon & Triana, Joc. cit.). The description reads rather like that of Mammea longifolia (Wight) Planchon & Triana;'* for instance, the flowers are said to be borne below the leaves. '? Kostermans (1961) used the name Mammea suriga (Buch.-Ham. ex Roxburgh) Kosterm. for this species. 1980] STEVENS, CALOPHYLLUM 291 I have not seen material which could be assigned to the original C. suriga. Calophyllum hirtellum, C. cymosum, and C. diepenhorstii have all previously been reduced to synonymy under C. soulattri or C. spectabile; they are based on specimens representing a common form of the species. Calophyllum cymosum was based on two specimens, one collected by Junghuhn from northern Sumatra, the other collected by Teysmann from western Sumatra; a sheet of the latter at Utrecht is designated the lectotype. Boerlage (loc. cit.) described two varieties of Calophyllum spectabile (= C. soulattri as now interpreted) and reduced C. diepenhorstii to a variety of that species, but none of these varieties can be maintained. The type specimen of C. spectabile var. miquelii is unremarkable. A few more details of the synonymy of some Papuasian names can now be given. I have seen syntypes (or fragments of them) of Calophyllum kiong, and it is clear that Whitmore was correct in reducing this to synonymy under C. soulattri (Whitmore, 1967, loc. cit.). The name ‘‘kiong,’’ used 75 years ago in the Finschhafen area and taken by Lauterbach and Schumann for the epithet of their C. kiong, is still used there for this plant. The reduction of C. versteegii to C. soulattri is not yet definite. The inflorescences of the isotypes may have a short axis to 2 mm. long, but the inflorescence is neither flabellate nor branched. The leaf blades dry a dark, rich, shining brown above. These are rather unusual characters for C. soulattri. Details of the typification of C. solomonense are discussed under C. leleanii. Calophyllum lancifolium was described from Philippine material with very narrow leaf blades. However, there are intermediates between it and broader- leaved specimens (e.g., Williams 2770 and FB 24736). The specimen of Elmer 13266 at the Arnold Arboretum is designated the lectotype of C. lancifolium. Calophyllum zschokkei, also from the Philippines, was described from speci- mens having rather small leaves and lacking well-developed indumentum; they link the larger-leaved specimens from the Philippine Islands with two particularly small-leaved specimens, BS 38967 (Mindanao) and BS 30293 (Luzon). The sheet of Elmer 12129 at the Gray Herbarium is made the lectotype of C. zschokkei. Vanoverbergh 1888, determined by Merrill as Calophyllum vanoverberghii and collected from the type locality of that species, matches the type of C. hibbardii (see above) fairly well. Calophyllum vanoverberghii was described as having an oblong, very coriaceous leaf blade that dried the same color on both surfaces, while the duplicates seen of Vanoverbergh 1888 have an ovate, subchartaceous leaf blade that has not dried concolorous. Although the description of the inflorescence of C. vanoverberghii suggests that it is similar to that of C. soulattri, I hesitate to reduce C. vanoverberghii to synonymy under C. soulattri without seeing type material. Of the five specimens cited in the original description of Calophyllum cholobtaches, fragments of three (Ledermann 14251, Raymondus 128 and 334) have been seen; Raymondus 29 and 293A have not. Ledermann 14251 was the only fertile specimen (Lauterbach, Joc. cit.), and the fragment of this number at Wrocklaw (designated the lectotype) has the branched inflores- cence and almost chartaceous, rather greenish-drying lamina characteristic 292 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 of C. soulattri. Lauterbach compared C. cholobtaches to C. kiong (also = C. soulattri) but noted that the former had less dense and more steeply ascending venation. These differences are of no significance. The leaves seen of Raymondus 128 and 334 both have only 6 to 8 veins /5 mm.—very distant for C. soulattri. However, only the former specimen is probably to be referred to C. soulattri, while the latter is C. pelewense (q. v.). Early authors almost without exception used the name Calophyllum spec- tabile Willd. for C. soulattri. Calophyllum spectabile was described from Mascarene material and is a synonym of C. tacamahaca (for more details see Stevens, 1976, loc. cit.). 22. Calophyllum brassii A. C. Smith, Jour. Arnold Arb. 22: 352. 1941; P. F. Stevens, Austral. Jour. Bot. 22: 359. 1974. Type: Netherlands New Guinea [Irian Jaya], Idenburg River, 4 km. SW. of Bernhard Camp, 850 m., 7 March 1939, Brass & Versteegh 13122 (holotype, a; isotypes, BO, BRI, L). Tree 20-30 meters tall, d.b.h. to 68 cm.; trunk without spurs or buttresses; outer bark yellowish brown to dark brown, with few to many shallow fissures, inner surface brown-orange and orange mottled; under bark brown-red and dark red mottled; inner bark reddish; latex white, rather curdy, slightly sticky (also reported to be light yellow or brownish). Twigs (strongly) flattened, 2.5-6 mm. across, + 2-, 4-, or 6-angled, with obscure raised horizontal lines at node, drying brown (yellowish), + persis- tently subfarinose to short-tomentose; axillary innovations lacking basal scars; internodes 0.5—6(-8) cm. long; uppermost pair of axillary buds + pointed, 2.5-7 mm. long, erect; terminal bud subconical, 0.5-1.4.cm. long, with brownish gray, short-tomentose to (?)subcrustaceous indumentum, the hairs unthickened except at base, often with short, basal branches, the underdeveloped internode to 1.5 mm. long. Petiole 1-6(-8) mm. long, broadly concave above, convex to angled below, subpersistently tomentose or subglabrous; lamina elliptic to oblong, (3-)6-17 by (1.9-)4-6 cm., rounded to shallowly retuse or apiculate at apex, (narrowly to) broadly rounded or cordate at base, usually only slightly undulate and strongly recurved at margin, (thickly) coriaceous, drying brick or cinnamon to greenish olivaceous above and near sabelline below, subpersis- tently puberulent to short-tomentose on midrib below (also above), the midrib above usually narrowing fairly quickly near base, raised, 0.25-0.6 mm. wide at midpoint, below strongly raised, usually angled, the venation subobscure (rarely subapparent) above and below, depressed to raised above, raised below, 10 to 16 veins/5 mm., angle of divergence 65-80°. Inflorescences from foliate axils, with numerous flowers, flabellate, branched, branches to 1.3 cm. long and with 5 flowers, the axis 1.2—5 cm. long, + tomentose, especially at nodes and toward base, lowest internode 1.5—4.5 mm. long, upper internodes to 1.7 cm. long; bracts suboblong to semiorbicular, to 3 mm. long, deciduous; pedicels (0.5-)0.7-2 cm. long, glabrous. Flower (?)her- maphroditic; tepals 4 (rarely 5), the outer pair lingulate, 3.8-4.5 by 1.7-2 mm., the inner pair elliptic to obovate, 4-4.7 by 2.1-2.7 mm.: stamens 65 1980] STEVENS, CALOPHYLLUM 203 to 105, the filaments to 2.3 mm. long, the anthers subelliptic, 0.4-0.7 mm. long, retuse at apex; ovary 0.6-0.9 mm. long, the style 1.2-1.7 mm. long, the stigma peltate-infundibular, 0.3-0.5 mm. across, not radiate. Submature fruit broadly ovoid, ca. 1.4 by 1.3 cm., apiculate, drying brown, deeply wrinkled; outer layer not detaching cleanly from stone, 0.7-1.5 mm. thick, compact; stone ovoid, 1-1.1 by 0.8-1 cm., apiculate, the walls less than 0.2 mm. thick, smooth, unmarked; spongy layer initially thick. DistriBuTION. Western portion of the New Guinea mainland (Map 9). ADDITIONAL SPECIMENS SEEN. Papuasia. IRIAN Jaya. Vogelkop: N. slopes of upper Aifat Valley, between Senopi and Aifatfekaan, W. of Kebar Valley, 920 m., BW 12843 (L, LAE), 870 m., BW 12848 (L, Laz). Fakfak: Tavoei [Onin Peninsula], bb 22317 (probable) is, BO, = LAE, sinc). Digul: 5 km. NE. from junction Iwoer R. and E. Digul, Sub. Div. Moejoe, 400 m., BW 8530 (L). Papua New Guinea. Western: Kiunga, ie m., Stevens et al. 838 (A ). Eco.ocy. Primary colline (often with Agathis Salisb.) or sometimes lowland rainforest, (20-)400-920 m. alt. Flowering and fruiting in March; ripe fruit blacki Stevens et al. 838 has subspherical, eventually crateriform galls in rows along the midrib of the lower surface of the lamina, or sometimes just in from the margin. GERMINATION AND YOUNG PLANT. The radicle breaks the stone wall to one side of the base. The seedling has two pairs of leaves separated by an internode less than 5 mm. long. Subsequent internodes are less than 1.5 cm. long. The terminal bud is functional, and the plant is erect. Only after the plant is some 30 cm. tall are internodes of ca. 3 cm. long or more produced (pers. obs.). Calophyllum brassii can be recognized by its strongly flattened twigs and its very coriaceous leaves, which are usually broadly rounded to cordate at the base, strongly recurved at the margin, and with rather dense venation. The branched inflorescence has a short basal internode and numerous flowers with four tepals. The epithet commemorates L. J. Brass, one of the greatest botanical collectors in New Guinea. A few fruits occur on Stevens et al. 838; all lack an embryo. They are spherical, about 7.5 mm. long and across, with the outer layer about 1.8 mm. thick, and the obovoid stone about 5.5 by 3.2 mm. across and strongly apiculate (all measurements from fluid-preserved material). The wrinkled fruits of Brass & Versteegh 13122 are still immature, but it is interesting to see that there, too, the stone is apiculate; it is probable that mature fruits, when dried, will be almost smooth. Calophyllum brassii is close to C. soulattri (Smith, loc. cit.; Stevens, loc. cit.). However, the leaf blades of C. soulattri are rarely so coriaceous or cordate at the base, the inflorescences have fewer flowers, and the stone is not apiculate. BW 12843 and 12848 (from the Vogelkop) differ from the other specimens 294 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 of this species in their yellowish-drying twigs and their relatively thinner leaves. Stevens et al. 838 tends to have short internodes, and like the specimens from the Vogelkop, its indumentum is subtomentose. Although this specimen was found at much lower altitudes than the others, it should be noted that at Kiunga genera and species commonly found at higher elevations grow at less than 30 meters altitude. In addition to Calophyllum brassii, Di- morphanthera F. Mueller (two species), Vaccinium L., Lithocarpus B1., and Podocarpus L’ Her. ex Pers. all grow in the same area. BW 8530 is superficially similar to the form of Calophyllum goniocarpum that grows at Kiunga, since it has the most narrowly rounded leaf bases of the specimens included in C. brassii (cf. also bb 22317). Calophyllum brassii and C. goniocarpum are not at all closely related and differ in inflorescence (much longer axis with short branches (if any) in C. gonio- carpum) and fruit (angled, much larger, thicker-walled stone in C. gonio- carpum). The uppermost pair of axillary buds is erect and usually well developed in C. brassii, and suberect and inconspicuous in C. goniocarpum; this difference helps to distinguish sterile material of the two taxa. 23. Calophyllum rigidum Miq. Fl. Indiae Batavae, Suppl. 1(3): 497. 1861; Mueller in Walp. Ann. Syst. Bot. 7: 357. 1868; Vesque in C. DC. Monogr. Phanerog. 8: 606. 1893; H. Keng, Gard. Bull. Singapore 28: 255. 1976. Type: Sumatra, ad littoram Siboga, Teysmann, HB 642 (holotype, u; isotypes, Bo, K, L, MEL, P, W). Calophyllum kunstleri King, Jour. Asiatic Soc. Bengal, II. 59: 174. 1890; Ridley, Fl. Malay Penin. 1: 182. 1922, pro parte; I. H. Burkill & M. R. Henderson, Gard. Bull. Straits Settl. 3: 347. 1925; M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 320. 1956; Maheshwari, Bull. Bot. Survey India 2: 141. 1960, pro parte; I. H. Burkill, Dict. Econ. Prod. Malay Penin. ed. 2. 1: 416. 1966, pro parte; T. C. Whitmore, Tree Fl. Malaya 2: 187. 1973; Corner, Gard. Bull. Singapore Suppl. 1: 104. 1978. Synrypes: Malaya, Perak, Larut, 300 feet [90 m.], Dec. 1883, Sie eee peas, 5328 (A, BO, Fl, G, K, SING, US), 100 feet [30 m.], Dec. 1883, 5374 (G, k, sinc, uc, us), 100 feet [30 m.], Jan. 1884, 5459 (FI, P). Tree 6-24(-36) meters tall, d.b.h. to 45 cm.; trunk without buttresses, ‘loop roots’’ sometimes present in plants from swamps; outer bark gray-brown to dark brown, shallowly fissured or with vertical lines of lenticels, often hoopmarked, inner surface blackish; under bark dark red; inner bark red; latex clear yellow and fluid or sticky, opaque yellow and not very sticky, or yellowish brown. Branches often pendulous, at least in Malaya. Twigs slightly to strongly flattened, (0.9-)1.2-3.5 mm. across, + 4-angled, soon becoming rounded, usually drying blackish (mid-brown), subpersistently puberulous to short-tomentose; axillary innovations lacking basal scars; internodes (1.3—)2—4(-6.5) cm. long; uppermost pair of axillary buds subacute, to 4mm. long, suberect, plump and very conspicuous just before flowering; terminal bud plump to narrowly conical, (3—)4.5-10(—-13) mm. long, with usually castaneous, short, tomentose indumentum (hairs, Ficures 13, 0; 14, n-s), 1980] STEVENS, CALOPHYLLUM 295 underdeveloped internode to 2.5(-6) mm. long. Petiole 0.6-1.5(-2.3) cm. long, concave to + flat above and convex (rarely subangled) below, subpersistently puberulo-tomentose; lamina ovate (subelliptic to suboblong), (4-)6.7-17.5 by (1-)2.2-5.6 cm., gradually (short-)acuminate at apex with acumen to 1.7 cm. long, (rarely retuse (Sarawak)), cuneate or attenuate (rarely rounded) at base, not undulate or barely so and not recurved at margin, coriaceous, drying bay to umber and usually shiny above, umber and + shiny below, + persistently puberulent to subtomentose on midrib below (also above), the midrib above gradually narrowed from base, level or slightly raised, + strongly sulcate, 0.2-0.4(-0.7) mm. wide at midpoint, below usually strongly raised, rounded or slightly striate, the venation subobscure to apparent above, latex canals and veins often equally clear especially in older leaves, apparent below, latex canals less prominent than veins, raised, (7 to) 10 to 17 veins/5 mm., angle of divergence 40-70(-80)°. Inflorescences from foliate axils along twig, with 3 to 11 flowers, usually unbranched (rarely with 3-flowered branches to 3 mm. long), the axis 0.2-2.5 cm. long, short-tomentose toward base, lowest internode 1-5 mm. long, flowers often congested (with 2 pseudo- whorls of 4 and 5, or 4 and 3, flowers); bracts ovate, 2-6.5 by 1-2.5 mm., short-tomentose below, subpersistent; pedicels 0.5-2.7 cm. long, slender, glabrous, to 1.2 mm. thick in fruit. Flower (?)hermaphroditic; tepals 4, the outer two ovate, 4-5.5 by 3-4 mm., the inner two elliptic to obovate, 5-9 by 2.3-3.5 mm.; stamens ca. 75, the filaments to ca. 4.5 mm. long, the anthers elliptic-oblong, 0.5-0.8 mm. long, retuse at apex; ovary 1.5-2.5 mm. long, the style 2.8-4.3 mm. long, the stigma peltate, ca. 0.5 mm. across. Fruit spherical, 1.2-1.6 cm. long and across, minutely apiculate, when young usually ovoid and sharply pointed at apex, drying brown, broadly wrinkled; outer layer eventually detaching cleanly from stone, |-2.5 mm. thick, compact; stone ellipsoid to subspherical, 1.1-1.4 by 1-1.25 cm., rounded at apex, the walls 0.15-0.5 mm. thick, smooth, unmarked; spongy layer thin. DistripuTION. Southern Malay Peninsula to western Borneo, scattered; excluding Java (Map 11) SELECTED SPECIMENS SEEN. Malaya. Perak: Larut, 90 m., King’s collector 45 m., KEP 71466 (ker); Port Dickson, P[asir] Panjang F.R., KEP 66506 (KEP); ‘Senaling Inas F.R., SFN 1753 [= KEP 628 fide Henderson & Wyatt- Smith, loc. cit.] (sinc); Tampin F.R., S. slope of G. Tampin, 540 m., FRI 14233 (K, KEP, SAN, SING); G. Angsi, Pedas, 457 m., FRI 14586 (k, KEP, SAR, sING); Kuala Pilah, Serting, KEP 62876 (ker). PAHANG: Ulu Sungai Anak Endau, 180 m., FRI 8109 (atypical) (A, KEP, SAR, SING); Tasek Bera State Land, KEP 35850 (KEP): Rompin, KEP 6538 (sinc). JoHorE: Mawai, Sungei Tementang, SFN 34719 (a, B, K, KEP, LAE, P, SING); G. Panti E., Jumali & Kuswata 218 (Bo); Kota Tinggi, Ridley 4185 (k, sinc); Kluang F.R., 225 m., KEP 76289 (KEP); Renggam F.R., 90 m., KEP 71264 (kep); Ulu Sedili, G. Sumalayang, 610 m., FRI 14104 (atypical) (A, K, KEP, SAN, SING). Singapore: Jurong Road, 17th mile, Baker 5732 (Gc, Nsw); Bukit ee Ridley 1955 vas Sumatra. Riau: Upper Riauw, Pakanbaru, Tenajan Soepadmo 40 BO, C, E, L, SING). SELATAN: Banjoeasin en ee anne Bajoeng Lintjir, 15 m., 148 296 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 | Map I1. Distribution of Calophyllum rigidum in Malesia. TI P 15 (po, L). Utara: Siboga, Teysmann, HB 642 (Bo, K, L, MEL, P, U, W). Bancka: Djebus, Teysmann s.n. (Bo). BeLirunc: Tandjong Pandan bb 10244 (po); Mangar, Teysmann s.n. (Bo). Borneo and adjacent islands. Sarawak. Ist Division: Bako Natl. Park, 75 m., S 16203 (?) (A, K, SAN, SAR); Lundu, G. Pueh, 457 m., S$ 13704 (a, Bo, kK, ao SAR, SING); G. Matang, 305 m., Stevens et al. 250 (?) (A); Semengoh Arboretum, 30 m., Stevens et al. 139 (a); Semengoh F.R., Stevens et al. 288 (?) (A). KALIMANTAN. Tengah: Beneden Dajak, Kapoeas, Moeroei, Danau Rawah, | m., bb 2602 (L). Barat: Karimata Archipelago, Teysmann, HB 11379 (po, Fl, K, L); Soengai Kenepai, Hallier 2138 (a, Bo, K, P); Smitau, P. Madjang, 30 m., bb 7677 (Bo). EcoLtocy. Well-drained mixed dipterocarp forest (throughout its range), swamps (Malaya, Sumatra), and sub-kerangas vegetation (Sarawak); 0-610 m. alt. Flowering December and February (Malaya), June (Sumatra); fruiting March to May, August (fruit whitish (KEP 35850)). GERMINATION AND YOUNG PLANT. The radicle probably breaks through the stone to one side of the base. The seedling appears to have two pairs of leaves separated by an internode 0.5-2.7 cm. long (/48 T J P 15). Each later-produced internode is successively longer. The terminal bud is functional, but the young plant is arched, with the leaves being held in one plane; the stem only later straightens. (Young plants seen in both Malaya and Sarawak: Stevens et al. 53, 247.) Loca uses. Calophyllum rigidum is used in ship building and for general construction (Malacca). Calophyllum rigidum can be readily distinguished from other species of the genus by its rather coriaceous lamina, which is usually gradually acuminate at the apex and which often dries a dark, bay brown. The venation density on the upper surface of the lamina is often apparently twice that on the lower (due to the latex canals as well as the veins being raised on the former), 1980] STEVENS, CALOPHYLLUM 27 and the midrib on the lower surface is very prominent, yellowish, and rounded. As King noted in his description of C. kunstleri, ‘“‘the nervation is closer than in any other species that I have seen, and the surfaces of the leaves have a peculiarly lustrous sheen.’’ The terminal buds usually have a very dark brown tomentum, and the axillary buds are often plump, conspicuous, thick outer layer. The epithet rigidum refers to the leaf blades, which are usually thick and rigid when dr The inflorescence of Calophyllum rigidum is commonly described as being subumbellate, but it is unlike the subumbellate inflorescence of C. soulattri. The inflorescences of C. soulattri are usually flabellate and branched; those of C. rigidum are never flabellate and are only rarely branched. In some specimens up to seven flowers from three nodes may arise together on a condensed axis (e.g., King’s collector 5328, Malaya), while in others four flowers from two nodes may arise together and be separated from the terminal triad, or triad plus one or two pairs, by an internode ca. | cm. long (e.g., SFN 30877 and Ridley 4185, Malaya; Soepadmo 40, Sumatra). In specimens such as Teysmann, HB 642 (Sumatra) and Hallier 2138 (Kalimantan), the inflorescences commonly have a subbasal group of four flowers and a terminal group of five flowers. The variation in leaf—especially in Calophyllum rigidum—is rather great. The type specimen has leaf blades somewhat more coriaceous than those of other specimens, although with similar details of midrib and venation. Its twigs dry brown, while those of the other specimens usually dry blackish. Most of the specimens collected from Malacca and Negri Sembilan (Malaya) are at first sight very different from the others, having slender twigs (less than 1.3 mm. across), small leaves (less than 8 by 2.2 cm.), and flowers usually in pairs on the inflorescences and separated by well-developed internodes. In Derry 151 the lowest internode is almost absent. However, in other details these specimens are not unusual. FRI 8109 and 12304 (Malaya) have more or less elliptic leaf blades with rather obscure venation, but in details of infructescence and fruit they agree well with the other specimens. Some specimens from around Kuching, Sarawak and arched seedlings are also typical of the species; their flowers are unknown. Stevens et al. 288 (Sarawak, sterile) is a perplexing specimen; it is rather robust and has something of the facies of Calophyllum castaneum, but its indumentum is quite different, since the rel — not have the enlarged, birefringent apical cells characteristic of that s The hairs of Calophyllum rigidum are rather variable, Bega ck the variation is less than might be expected (Ficures 13, 0; 14, . There is frequently a basal branch, often made up of only a a. cell. The cells, at least at the base, have notably thick and rather rough walls, and the hairs are often entirely composed of such cells. If the apical cells are thin walled, they 298 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 lack the brown contents found in the thick-walled ones. The hairs are erect or curved; if curved, the curvature is either notably abrupt and at the middle or near the apex of the hair, or more gradual and from near the base. 24. Calophyllum robustum P. F. Stevens, Austral. Jour. Bot. 22: 392. 1974. Tyre: Papua, Northern District, Dobodura Plain, Samboga River area, 100 feet [30 m.], 13 March 1945, NGF 2077 coll. Cavanaugh & Fryar (holotype, LAE; isotypes, A, BRI, CANB, E, K, L). Tree 18-30 meters tall, d.b.h. to 65 cm.; truck (?)without buttresses; outer bark yellowish at first, becoming gray to brown, deeply fissured; under bark pink; inner bark brown; latex milky or clear yellow; sapwood yellowish red, extremely hard, merging to red heartwood. Crown spreading, smaller branches and leaves + pendulous. Twigs flattened, 3.5-5 mm. across, 4-angled and with a + prominent raised line decurrent from middle of petiole, sometimes with obscure horizontal raised lines at nodes, drying dark brown to blackish, subpersistently farinose- puberulent; axillary innovations lacking basal scars; internodes 1.5-6 cm long; uppermost pair of axillary buds + pointed, 1-4(-12) mm. long, spreading: terminal bud narrowly conical, 1.6-3.3 cm. long, with brown, subcrustose to puberulent indumentum (hairs, Ficure 13, n, also moruloid), underdeveloped internode 4-7 mm. long. Petiole 1.3-2.3 cm. long, deeply concave above, convex to angled below, drying blackish, glabrous to persistently puberulent; lamina oblong to elliptic, 13-31 by 5-8.5 cm., abruptly acuminate at apex, rounded to shortly acute at base, rather closely undulate and narrowly recurved rgin, coriaceous, drying gray-olivaceous above and sabelline below, subpersistently farinose-puberulent on midrib below, the midrib above nar- rowing gradually from base, raised, center sulcate, 0.4-0.6 mm. wide at midpoint, below strongly raised, angled, the venation subapparent above, apparent below, raised, 13 to 16 veins/5 mm., angle of divergence 70-80°. Infructescences from foliate axils, with scars of 5 to 7 flowers, unbranched, the axis 0.4-1.1 cm. long, with farinose to puberulent indumentum at least near base, lowest internode 4-8 mm. long; bracts unknown; pedicels 1.2-1.5 cm. by 2.5-3 mm., glabrous. Flower unknown. Fruit spherical to ovoid-ellip- soid, 2.5-3 by 1.8-2.3 cm., acute to subrounded at apex, drying brown, densely and shallowly wrinkled; outer layer not detaching cleanly from stone, 1.2-2.6 mm. across, compact, strong; stone ca. 2.5 by 1.7 cm., obtusel pointed at apex, the walls less than 0.1 mm. thick, smooth, (?)unmarked; spongy layer thin. DistTripuTION. Eastern New Guinea (Map 9). SPECIMENS SEEN. Papuasia: see Stevens, Joc. cit. Eco.tocy. Well or poorly drained lowland rainforest, 30-180 m. alt. Fruiting March and November. GERMINATION. The radicle probably breaks through the stone to one side of the base. The seedling has two pairs of pseudoverticillate leaves. Subse- 1980] STEVENS, CALOPHYLLUM 299 quently produced internodes are long, but the stem is arched at first, becoming erect only some 30 cm. or more behind the apex. The terminal bud is functional. (Based on two seedlings only; see also Stevens, Joc. cit.) Calophyllum robustum can be recognized by its long, narrowly conical terminal bud, and by its densely veined, more or less oblong leaf blades 13-31 cm. long that have a strongly angled midrib on the lower surface and that dry an olivaceous color on the upper surface. The wrinkled fruits have a tough outer layer and appear to have a very thin-walled stone. (The 1.5 mm. thick stone mentioned earlier (Stevens, op. cit., p. 394) may refer to this tough outer layer, but more collections are needed to confirm fruit structure.) The epithet robustum refers to the large leaves and terminal buds and the thick stems of this species. 25. Calophyllum moonii Wight, Ic. Pl. Indiae Orient. 1: p/. 1/1. 1839, Illus. Indian Bot. 1: 129. 1840; Walp. Rep. Bot. Syst. 1: 397. 1842; Thwaites, Enum. Pl. Zeyl. 52. 1858; Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 269. 1862; Bedd. Fl. Sylv. 3: xxii. 1871. Type: Ceylon [Sri Lanka], eastern Korle, Moon s.n. (n.v.). IGURE 15, i. C. spectabile auct., non Willd.; T. Anderson in Hooker f. Fl. Brit. India 1: 271. 1874, pro parte; Trimen, Handb. Fl. Ceylon 1: 99. 1893; Vesque in DC. Monogr. Phanerog. 8: 583. 1893, pro parte; Lewis, Descr. Catal. Ceylon, 20. 1902. C. soulattri auct., non Burman f.; Alston in Trimen, Handb. FI. Ceylon Suppl. 6: 22. 1931. Sastri et al Wealth India 2: 20. 1950, pro parte; Worthington, Ceylon Trees, 32. 1959. Tree 20-30 meters tall, d.b.h. to 35 cm.; no buttresses; outer bark yellowish, thin and smooth, or very dark brown and smooth, or dark yellow-gray, thick, and with wide panels (Worthington, /oc. cit.), sometimes deeply fissured; inner bark dark red; latex clear (Kostermans 24 Twigs flattened, 3-5 mm. across, + 4-angled, eer blackish, with brown, puberulent to tomentose indumentum, soon glabrescent or not; axillary innovations lacking basal scars; internodes (1-)3-12 cm. long; uppermost pair of axillary buds + pointed, 2-12 mm. long, spreading to suberect, very obvious; terminal bud plump, 1.2-2 cm. long, with brown, tomentose indumentum (hairs, Ficure 14, t), underdeveloped internode to 2 mm. long. Petiole 1-1.8 cm. long, concave above, convex below, sometimes persistently puberulent below; lamina ovate to suboblong or elliptic, (7-)9-16(-20.5) by (3.7-)5-6.5 cm., rounded to retuse (acute) at apex, broadly rounded to acute at base, distantly undulate and slightly or not recurved at margin, coriaceous, drying hazel above and sabelline below, soon glabrescent or persistently puberulo- tomentose on midrib below, the midrib above narrowing quickly near base, raised, 0.3-0.5 mm. wide at midpoint, below strongly raised, angled toward apex and striate toward base, the venation subapparent above and apparent below, raised, 8 to 13 (to 16) veins/5 mm., angle of divergence 70-85°. Inflorescences from foliate axils toward ends of twigs, with 5 to 11 flowers, flabellate, unbranched, the axis 1-3.5 cm. long, + tomentose at base, lowest 300 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 f, g, fruit, x : , from outside; g, longitudinal section. h, terminal bud, x i moonii (Kostermans 246 ), X 1, arrow at indented stone apex. j-l, C. ieee ee (7929 SF). j, terminal bud, x 3. k, 1, fruit, x 1: k, from outside; |, longitudinal section. m-o, C. paniculatum. m, terminal bud (1297 RN), X 1. n, 0, fruit (11078 SF), 1: n, from outside; o, longitudinal section. p, q, C. aff. paniculatum | (23120 SF), fruit, x 1: p, longitudinal ; 55 : T aps Sea section; s, from outside. t, C. vernicosum (de Cary 14522), terminal bud, 1980] STEVENS, CALOPHYLLUM 301 internode 0).3-2 cm. long; bracts ovate, ca. 3 mm. long, deciduous; pedicels (0.3-)0.6-1.8 cm. long, glabrous, in fruit to 3 mm. thick. Flower known only in late mae (?)hermaphroditic; tepals (?)4 or 6 to 8, the outer pair ca. 6 by 5.5 mm., the inner ones 6-7.5 by 2.5-4.5 mm.; stamens 75 to 110, the filaments to ae mm. long, the anthers oblong, 1-1.5 mm. long, retuse at apex; ovary ca. 2 mm. long, the style ca. 2.5 mm. long, the stigma peltate, ca. 1.2 mm. across, 2-radiate. Fruit ovoid to ellipsoid, 1.8-2.5 by 1.6-1.9 cm., apiculate or not, sometimes with fibrous stipe to 3.5 mm. long, drying cinnamon, smooth or shallowly and broadly wrinkled; outer layer not detaching cleanly from stone, ca. 1 mm. thick, notably fibrous, air spaces developing; stone subellipsoid, 1.1-1.4 by 0.9-1.2 cm., shallowly retuse at apex, the walls ca. 0.2 mm. across, smooth, unmarked; spongy layer thin. DistriBuTIon. Southwestern Sri Lanka (Map 7). SELECTED SPECIMENS SEEN. Sri Lanka: Kalutura Distr., Hallawakellae forest, Welipenna, Balakrishnan 1178 (x, us); Kuttapitiya Jungle, Pelmadulle, 4 m., Worthington 6437 (?) (L); Ratnapura Distr., Moropitiya logging area, Koster- mans 24670 (a, E, K, L, US); Athweltota near Moropitiya, Jayasuriya 904 (aau K); Mahane-Makalane hill forest, near Moropitiya, 400 m., Kostermans 24701 (L); Kalawana—Moropitiya road, close to culvert 29/3, 100 m., Cramer 4159 (us); Hiniduma F.R., Kostermans 24704 (?) (L, us), Hinidumkande (Heycock), near Hiniduma, 500 m., Kostermans 25502 (k, L); Heycock Forest, sea level, Cramer 3084 (?) (us); Kunneliya Forest S. P., 60 m., Worthington 5262 (bm, F); Naunkilla ela, Kaneliya F.R., 30 m., Worthington 3678 (BM); sine loco, CP 3402, pro parte (kK, Us). Eco.ocy. Lowland rainforest, to 500 m. alt. Fruiting April to June; fruit green, acid. GERMINATION AND YOUNG PLANT. The radicle probably breaks the stone near the base. The seedling has a single pair of leaves. Subsequent pairs of leaves (which are produced singly) are separated by internodes of 0.7-1.3 cm. The terminal bud is functional, and the plant is probably erect. (Kostermans 25560, four plants.) Calophyllum moonii is a distinctive species recognizable by its long terminal ud and conspicuous, often spreading, uppermost pair of axillary buds; its large, coriaceous leaf blades that are often rounded to retuse (at most acute) at the apex; and its few-flowered, flabellate but unbranched inflorescences. The fruits have a fibrous outer layer and a thin-walled stone that is shallowly retuse at the apex. The epithet commemorates F. Moon. There is some variation in the apex of the lamina in Calophyllum moonii. Wight (1839, Joc. cit.) illustrated a specimen with rather narrowly elliptic leaf blades that are acute at the apex and base, but many specimens have broader, subovate leaf blades that are more or less retuse at the apex and rounded at the base. I thought at first that there were two species involved, but there is no difference in characters such as terminal bud, indumentum, and fruit in specimens differing in leaf apex. Wight’s illustration shows the flower as having four tepals; in the flowers dissected I have found six to eight. 302 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 The specimens from Hiniduma and Kuttapitiya Jungle are sterile; they are included here with some hesitation. The retuse apex of the stone of Calophyllum moonii is rather remarkable. It was noticed in all fruiting specimens examine There is little similarity between Calophyllum moonii and C. soulattri (or C. spectabile auct., non Willd.), with which it has been confused; Kostermans (1976) noted that the two were distinct. Although the inflorescences of C. moonii are flabellate, they are unbranched (vs. practically always branched in C. soulattri). In C. soulattri the uppermost pair of axillary buds is characteristically closely adpressed to the terminal bud, the lamina is thinner and dries closely undulate, the fruit dries blackish and has a compact outer layer that practically lacks fibers, and the stone is rounded at the apex. The relationships of C. moonii are obscure, but the hairs are similar to those of C. tomentosum (cf. Ficures 14, t, and 11, 1). 26. Calophyllum milyum P. F. Stevens, sp. nov. Ficure 15, a-c. C. parviflorum auct., non Baker f.; Drake in Grandid. Hist. Phys. Nat. Polit. Madagascar 35: pl. 357. 1894; Lecomte, Madagascar Bois Forét d’Analamazaotra, 102. p/. 39. 1922; H. Perr. Mém. Mus. Nat. Hist. Nat. Paris, n.s. 24: 78. 1948, in Humbert, Fl. Madagascar Comores, Fam. 136: 8. 1951. A speciebus aliis Calophylli in internodio infimo innovationis circa duplo longiore quam altis, foliis parvis coriaceis basibus cuneatis vel in petiolis decurrentibus et nervis lateralibus subobscuris, floribus cum 8 tepalis (duobus exterioribus interdum subpersistentibus), et parietibus fructus 1.2-2 mm. crassis, differt. Tree 12-20 meters tall, d.b.h. to 60 cm.; trunk without buttresses; outer bark gray, brown, or whitish, fissured; inner bark yellowish or red; latex yellow or biackich (Lecomte, loc. cit Twigs somewhat flattened, |.8-3(-3.5) mm. across, 4-angled or rounded, drying dark brown to blackish, puberulent when young; axillary innovations lacking basal scars; lowest internode of an axillary innovation (1.8-)2.8-5.5 cm. long, others 0.5-3 cm. long; uppermost pair of axillary buds rounded, to 1.2 mm. long, suberect; terminal bud plump, (3.5-)5-8.5 mm. long, with compact, grayish to brown indumentum (hairs, Ficure 13, m), underdeveloped internode to 2(-6) mm. long. Petiole 0.3-1.1 cm. long, shallowly concave above and convex below, glabrescent; lamina elliptic to obovate (rarely subcuneiform), (1.8-)2.3-7 by (1.1-)2-4.2 cm., rounded to retuse at apex, cuneate to decurrent at base, slightly undulate and narrowly recurved at margin, very coriaceous, drying umber to sabelline on both surfaces, + transiently puberulent on midrib below, the midrib above gradually narrowed from base, at least margins raised, 0.2-0.6 mm. wide at midpoint, below raised, striate, the venation obscure on both surfaces, slightly raised to flat, 9 to 15 veins/5 mm., angle of divergence 55-65°. Inflorescences from foliate axils, with 7 to 11 flowers, unbranched, the axis 3-6 cm. long, puberulent, especially toward base, lowest internode 1-4 cm. long; bracts ovate, ca. 1980] STEVENS, CALOPHYLLUM 303 3 mm. long, deciduous; pedicels 0.5-1.3 cm. long, + glabrous. Flower (?)hermaphroditic; tepals 8 (rarely 7), the outer pair broadly ovate, 6-7.5 by 6-6.5 mm., sometimes persisting in fruit, the inner ones + obovate, 7.5-12 by 3-8 mm.; stamens 115 to 180 (to 255), the filaments to 4.5 mm. long, the anthers oblong, 1-2.1 mm. long, + retuse at apex; ovary 2-2.5 mm. long, the style 3-3.5 mm. long, the stigma peltate, 1-1.4 mm. across, 3- or 4-radiate. Fruit spherical to ovoid, 1.8-2.7 by 1.5-2.1 cm., rounded to acute at apex, drying brown, broadly and shallowly wrinkled, perhaps smooth when young; outer layer probably not detaching cleanly from stone, 1.5-3.5 mm. thick, compact; stone subspherical to ovoid, 1.4-1.8 by 1.2-1.65 cm., obtuse (rounded) at apex, the walls 1.3-2 mm. thick, smooth, unmarked; spongy layer thin. Type: Madagascar [Moramanga], forét d’Analamazaotra, 800 m., Perrier de la Bathie 5328 (holotype, P; isotypes, c, K). DistripuTion. Central Madagascar (Map 12). SELECTED SPECIMENS SEEN. Madagascar: Ambatondrazaka, Manakambahiny- est, Nonokambo, 26556 SF (pr), Manaka, 6901 RN (p); Ambodipaiso, Toby-Am- bodipaiso, 900 m., 26663 SF (p); massif de |’ Andringitra, forét d’ Ambodipaiso, Manakanbahiny, 1200 m., Cours 2333 (pr); Perinet, Analamazaotra, 25673-25677 SF (all p); Bevetraka, 21266 SF (p), 21267 SF (pr); Sahahamy, 2530 SF (e), Ambotrafanga, 27-B-R-172 SF (rp); Manjakandriana, Antiahambavy, 15879 SF (Pp); Ambatondrazaka, Manaka[na] Est, 6901 RN (pr); Fianarantosoa, Fan- drandava, Sambalahy, 13531 SF (p); Ajozorobé, Tsaralahy, Fenéol 15 (pr); Anosibe, ouest du village d’Antandava (P.K. 45 de la route Moromanga—Ano- sibe), 28423 SF (p), Ankazomanitra, 26809 SF (p); sine loco [?E. Imerina], Baron 2235 (£, K pro parte, p), 2244 (E, K, P), 2662 (k), 4433 (BM, K, Mo, P), 7045 (k). Ecotocy. Locally common in colline /lower montane, mossy, and dry forest, sometimes on summits, 800-1200(-1500) m. alt. Flowering December and January. Fruiting November to January; fleshy fruit eaten by lemurs and potomacheres, the former rejecting and the latter ingesting the seeds (?stones: Perrier de la Bathie, 1948, loc. cit.). (Perrier de la Bathie (1951, Joc. cit.) recorded flowering in January and February, fruiting from February to June.) GERMINATION. The seeds apparently do not germinate unless they are surrounded by the pericarp (Perrier de la Bathie, 1948, loc. cit.). Locat uses. The wood is used in carpentry and construction. Calophyllum milvum is a typical montane species of the genus, having small, rather thickly coriaceous leaf blades with obscure venation, shoots with short internodes, and inflorescences with relatively long axes and large flowers. However, the lowest internode of the innovation, which is clearly about twice as long as the upper internodes, and the thick-walled stone immediately distinguish it from other species, including C. parviflorum, with which it has been confused. The epithet milvum alludes to the characteristic, almost kitelike lateral shoots. 304 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 The specimen 29179 SF (Massif d’ Anorimbato, a l’ouest d’Ambaravaranala leap is from a locality to the west of the others cited, and may be related to Calophyllum milvum. However, the lamina has much eae venation and is broadly revolute at the margin. The specimen was in flower when collected in May; flowering in C. milvum seems to be restricted to December and January. Other specimens with lateral shoots like those of C. milvum but with clear venation are discussed under C. pervillei. 27. Calophyllum lingulatum P. F. Stevens, sp. nov. Ficure 15, f-h. A speciebus aliis Calophylli quibus laminis cordatis habent in inflorescentiis axillaribus haud ramosis, floribus cum 8 tepalis, et fructibus apicibus acutis putaminibus parietibus crassis haud signatis vel angulatis, differt. Tree; trunk and bark not known. Twigs slightly flattened, 1.7—-2 mm. across, strongly 4-angled when young, drying dark brown, + glabrous; axillary innovations lacking basal scars; lowest internode of axillary innovation 2.2-3.5 cm. long, others 1-2 cm. long; uppermost pair of axillary buds rounded, less than | mm. long, suberect; terminal bud plump, 3-4.5 mm. long, with brown, subcrustaceous-puberulous indumentum, underdeveloped internode absent. Petiole 1-1.5 mm. long, obscure; lamina ovate or lingulate to oblong, 4.7-8.2(-12.5) by 2.2-4.5 cm., rounded to obtuse at apex, cordate at base, strongly undulate and slightly recurved at margin, coriaceous, drying shiny and umber on both surfaces, subfarinose on midrib below, the midrib above abruptly narrowed near base, soon becoming raised, 0.2-0.4 mm. wide at midpoint, below raised, angled toward apex and striate toward base, the venation subapparent on both surfaces, slightly raised, 7 to 10 veins/5 mm., angle of divergence 60-70°. Inflorescences from foliate axils on ends of twigs, with 5 to 9 flowers, unbranched, the axes 1.9-5 cm. long, farinose-puberulent toward base, lowest internode 1.3-2.5 cm. long; bracts elliptic, ca. 3 mm. long, deciduous; pedicels .35-1.4 cm. long, sparsely farinose. Flower (?)hermaphroditic; tepals 8, the outer pair ovate, 5.5-7.5 by 4.5-6 mm., the inner ones elliptic to obovate, 8.5-1l1 by 2.4—-5 mm.; stamens 85 to 100, the filaments to 4.5 mm. long, the anthers oblong, 1.2-1.6 mm. long, slightly retuse at apex; ovary 1.5-2.5 2.5-3 mm. thick, compact; stone ellipsoid, ca. 1.65 by 1.4 mm., rounded at apex, the walls |-1.7 mm. thick, smooth, unmarked; spongy layer thin. Type: Madagascar, Ambanja, Marovato, 18 Nov. 1952, 4500 RN coll. Sajy (holotype, Pp; isotype, P). DistrRIBUTION. Northern Madagascar (Map 12). ADDITIONAL SPECIMEN SEEN. Madagascar: Mandritsara, Antsirabe, Antsiatriaba, 3-R-388 SF (Pp) EcoLocy. Flowering and fruiting in November. 1980] STEVENS, CALOPHYLLUM 305 44 n Maps 12, 13. 12, distribution of Calophyllum milvum Bee circles), C milvum var. (long, erect triangle), C. lingulatum (empty stars), C. vernicosum (half-circles), C. fibrosum (erect triangles), C. humbertii Greed triangles), C. verticillatum (solid stars), and C. chapelieri (empty circles; overlapping distribution with C. verticillatum, stars in circles) in Madagascar. Inset: C. comorense (Comores Islands). 13, distribution of C. recedens (solid ee C. paniculatum (erect triangles), C. aff. paniculatum | (solid stars), C. paniculatum 2 (empty stars), C. aff. paniculatum 3 (inverted triangles), me C. drouhardii (empty circles) in Madagascar. Inset: C. tacamahaca (Masca- renes). Locat uses. The wood is used in construction and for joinery. Calophyllum lingulatum can be recognized by its cordate-based lamina, axillary inflorescences, flowers with eight tepals, and fruits with rather thick-walled stones that are neither marked nor angled. The epithet refers to the rather tonguelike shape of the leaves. Calophyllum lingulatum is perhaps related to C. milvum, which has similar inflorescences, flowers, and fruits. Both species apparently also have axillary innovations with the basal internode much longer than the others, although this must be confirmed for C. lingulatum. There are, however, numerous 306 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 vegetative differences between the two. The terminal bud of C. lingulatum is shorter, and the lamina is thinner and cordate at the base. The midrib on the upper surface of the lamina is sharply — the venation is more distant and is easily visible, rather than subobscur Although 3-R-388 SF is sterile, it is almost as Calophyllum lingulatum. The measurement of lamina length in parentheses in the description is taken from this specimen. 28. Calophyllum humbertii P. F. Stevens, sp. nov. Ficure 15, d, e. A Calophyllo milvo, quo aliter similis est, in gemma terminali 2-3 mm. longa (in C. milvo (3.5-)5-8.5 mm. longa), folia basi obtusa vel rotundata (cuneata vel decurrenti), inflorescentia bracteis foliaceis subpersistentibus praedita (parvioribus, haud persistentibus), et putamine parietibus circa 0.2 mm. crasso (1.3—2 mm. crasso), differt. Small tree; trunk and bark not known. Twigs slightly flattened, 1.7-2 mm. across, slightly 4-angled, drying blackish, glabrous; axillary innovations lacking basal scars; first internode of an axillary innovation 2-4.5 cm. long, others 1-3 cm. long; uppermost pair of axillary buds rounded, ca. 0.7 mm. long, erect; terminal bud plump, 2-3 mm. long, brown-puberulent, underdeveloped internode to 0.8 mm. long. Petiole 2-3 mm. long, broadly concave above and convex below, glabrous; lamina obovate to suborbicular, 1.9-3.9 by 1.2-2.9 cm., rounded or slightly retuse at apex, obtuse to rounded at base, not undulate but slightly recurved at margin, coriaceous, drying umber to fulvous on both surfaces, glabrous at maturity or persistently subpuberulent on midrib below, the midrib above gradually narrowed from base, slightly depressed to flat, 0.3-0.5 mm. wide at midpoint, often disappearing ca. 3 mm. below apex, below raised, + rounded, the venation obscure on both surfaces, + flat, 8 to 11 veins /5 mm., angle of divergence 45-60°. Inflorescences from foliate axils, with 7 to 9 flowers, unbranched, the axis 3.2-4.5 cm. long, glabrous or sparsely subfarinose toward base, lowest internode 1-2.5 cm. long; bracts leafy, cuneiform, to 2.7 b 2.9 cm., subpersistent; pedicels 0.6-1.1 cm. long, glabrous. Flower (?)her- maphroditic; tepals 8 (rarely 9), the outer pair ovate, 4.5-5.5 by ca. 3.5 mm., the inner ones obovate to elliptic, 7-8 by 3-6.5 mm.; stamens 100 to 115, the filaments to 3 mm. long, the anthers oblong, 1- 14 mm. long, retuse at apex; ovary ca. 1.5 mm. long, the style ca. 2 mm. long, the stigma peltate, 1.3-1.5 mm. across, + 3-radiate. Fruit broadly ovoid, ca. 2.3 b 1.8 cm., obtuse at apex, drying brown, sublongitudinally wrinkled; outer layer detaching cleanly from stone, ca. 2 mm. thick, compact; stone broadly ovoid, ca. 1.7 by 1.4 cm., rounded at apex, the walls ca. 0.2 mm. thick, smooth, unmarked; spongy layer thin. Tyre: Madagascar, sommét oriental du massif de Marojejy (nord-est), a l’ouest de la haute Manantenina, affluent de la Lokoho, 1850-2137 m., anno 1949, Humbert & Cours 23728 (holotype, P; isotypes, P). Distripution. Northern Madagascar (Map 12). 1980] STEVENS, CALOPHYLLUM 307 ADDITIONAL SPECIMENS SEEN. Madagascar: massif de Marojejy, a l’ouest de la haute Manantenina, 1500-1700 m., Humbert 22592 (p); vallee de la Lokoho (nord-est), mont Beondroka, au nord de Maroambihy, 1000-1450 m., Humbert 23516 (P Ecotocy. Small tree, montane (sometimes ericoid) forest, 1000-2137 m. alt. Flowering and fruiting in March. Calophyllum humbertii can be recognized by its short terminal bud; axillary innovations in which the first-produced internode is much longer than subse- quent ones; small, broad leaf blades; and fruits, which have a relatively thick, compact outer layer and a thin-walled stone. The epithet commemorates the noted French botanist, H. Humbert. The facies of Calophyllum humbertii is somewhat similar to that of C. milyum; in both the axillary shoots have basal internodes that are much longer than the others. However, the shorter terminal bud, broadly obovate to suborbicular lamina, subpersistent and foliaceous a aia thin-walled stone of C. humbertii immediately distinguish it from C. m while those of the latter are clearly immature; in neither is the stone wail well developed. 29. Calophyllum laxiflorum Drake, Bull. Soc. Linn. Paris 2: 1220. 1896; H. Perr. Mém. Mus. Nat. Hist. Nat. Paris, n.s. 24: 79. 1948, pro parte, in Humbert, Fl. Madazascar Comores, Fam. 136: 12. fig. 2, 6-8. 1951, pro parte. Type: Madagascar, sine loco, Chapelier s.n. (holotype, P; isotype, P). C. sorapa Drake in Grandid. Hist. Phys. Nat. Polit. Madagascar 35: pi. 358, 2. 1896. (?)Tree; trunk and bark not known. Twigs rather strongly flattened, 1.7-2.4 mm. across, strongly 4-angled, drying blackish, glabrous when mature, axillary innovations lacking basal scars; internodes (0.2-)0.5-5 cm. long; uppermost pair of axillary buds rounded, less than 0.5 mm. long, inconspicuous; terminal bud plump, 1.7-2.4 mm. long, with short, grayish indumentum. Petiole 2-3.5 mm. long, broadly and shallowly concave above, convex below, glabrous; lamina obovate, 2.7-4.8 by 1.4-2.4 cm., acute at apex, cuneate or + rounded at base, slightly recurved but barely undulate at margin, coriaceous, drying dark brown and nitid above, dark brown below, glabrous when mature, the midrib above rather abruptly narrowing at or near base, slightly raised, 0.15-0.25 mm. wide at midpoint, below raised, not very conspicuous, obscurely angled, the venation above + apparent, slightly raised, below subobscure, not raised, 4 to 7 veins/5 mm., angle of divergence 20-40°. Inflorescences from foliate axils, with 5 to 9 flowers, unbranched, the axis 4.5-6 cm. long, glabrous, lowest internode ca. 2.5 cm. long; bracts unknown; pedicels 1.1--1.9 cm. long, glabrous. Flower and fruit unknown (ovary ca. 2 mm. long). 308 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 DistripuTION. Known only from the type collection, made in (?eastern) Madagascar. Calophyllum laxiflorum is a poorly known species that can be recognized by its short terminal bud, its obovate leaf blades that dry dark, shiny brown and have steeply ascending venation, and its long, few-flowered inflorescence axis with a long basal internode. The epithet /axiflorum alludes to the inflorescence, in which the flowers are separated by relatively long internodes. Perrier de la Bathie (1951, Joc. cit.) included three species in Calophyllum laxiflorum: C. laxiflorum sensu stricto, C. pervillei, and C. verticillatum (the last described here as new). 28357 SF has leaves similar to those of the preceding species, but its fruits are quite different; there are yet other small-leaved specimens with more widely ascending venation. Since the taxonomic Situation is so unciear, the two early names have been narrowly circumscribed. Calophyllum verticillatum, described below, is very different from the other small-leaved taxa with which it has been confused. Other small-leaved specimens are discussed after the treatment of C. pervillei. Calophyllum laxiflorum is known only from the type specimen, which i just past flowering. The leaves differ from those of C. pervillei anid pee SF in their steeply ascending venation (angle of divergence 20-40°, vs. at least 50° for the other species) and in the dark, shining, brown color of the dried lamina (with the midrib noticeably paler). The lowest flowers on the inflorescence axis of C. laxiflorum are borne ca. 2.5 cm. from the base. The lowest pedicel scar in infructescences of C. pervillei is less than 1 cm. from the base of the axis; the type specimen of C. pervillei, which is in very young bud, has inflorescences less than 7 mm. long. Calophyllum laxiflorum has more oo twigs than C. pervillei—|.7-2.4 mm. across, rather than 0.8-1.5 m Although one sheet Chapelier s.n. at Paris that was labeled Calophyllum laxiflorum is a fruiting specimen of C. verticillatum, it cannot serve as type for C. laxiflorum. Fruits are not mentioned in the protolog, and the specimen does not have the inflorescences described for C. laxiflorum. Calophyllum verticillatum can be immediately distinguished from C. laxiflorum, C. pervillei, and the other small-leaved specimens by the pseudoverticillate arrangement of the leaves at the termination of each innovation The illustration of Calophyllum sorapa in Grandidier (loc. cit.) is clearly to be referred to C. laxiflorum; it is very similar to the type specimen of C. laxiflorum, except that tepals are illustrated. It is possipie that the epithet sorapa is a misreading of the local name, ‘‘fouraha.’’ Perrier de la Bathie (1948, loc. cit.) thought that C. sorapa was to be equated with C. pervillei; later (1951, op. cit.) it was placed in synonymy under C. chapelieri, with C. sorapa and C. chapelieri typified by the same specimen. However, in lea arrangement and inflorescence type the illustration of C. sorapa is very different from C. chapelieri, which is another species with pseudoverticillate leaves at the end of each innovation. Perrier de la Bathie (1948, Joc. cit.) was rather confused over the whole complex. He thought that Drake del Castillo separated C. pervillei from C. 1980] STEVENS, CALOPHYLLUM 309 laxiflorum by the umbelliform inflorescence of the former. Drake did not mention such inflorescences in his description of C. pervillei; he noted only that the inflorescences were cymose and shorter than the leaves. Perrier de la Bathie may have been confused by the inclusion of specimens of C. verticillatum in C. pervillei. 30. Calophyllum pervillei Drake, Bull. Soc. Linn. Paris 2: 1220. 1896, in Grandid. Hist. Phys. Nat. Polit. Madagascar 35: pl. 358, 3. 1896. Type: Madagascar, Ste. Marie, 23 April 1849, Pervillé 229 (holotype, P; isotypes, K, P). C. laxiflorum auct., non Drake; H. Perr. Mem. Mus. Nat. Hist. Nat. Paris, n.s. 24: 79, 1948, pro parte, in Humbert, Fl. Madagascar Comores, Fam. 136: 12. 1951, pro parte. Tree; trunk and bark not known. Twigs flattened, 0.8-1.5 mm. across, strongly 4-angled, drying dark brown, glabrous when mature; axillary innovations lacking basal scars; basal internodes of innovation 1.5-4 cm. long, others (0.3-)0.5-3 cm. long; uppermost pair of axillary buds rounded, less than 0.5 mm. long, erect, inconspicuous; terminal bud rather narrowly conical, 1-2 mm. long, with short, grayish indumentum to subrhombiform or subelliptic, 0.9-6 by 0.5-2.4 cm., bluntly pointed to rounded at apex, narrowly cuneate to acute at base, slightly undulate or recurved at margin or slightly recurved at very base, coriaceous, drying sepia and nitid above, umber below, margin often discolored at base, glabrous, the midrib above gradually narrowed from base, raised, strongly sulcate, at least in basal part, 0.15-0.25 mm. wide at midpoint, disappearing ca. 2 mm. short of apex, below slightly raised, obscurely angled, the venation obscure to subapparent above, slightly raised, below usually subobscure, 7 to 11 veins/5 mm., angle of divergence 50-60(-65)°. Infructescences from foliate axils along stem, with scars of 5 to 9 flowers, unbranched, the axis ca. (?)2 cm. long, lower part brown-puberulent when young, lowest internode at most 6 mm. long; bracts ovate, ca. 2 mm. long; pedicels 4-8 mm. long, stout, to 1.8 mm. across. Flower unknown. Fruit spherical, 1.7-2.2 cm. long and across, obscurely pointed at apex or not, drying smooth, dark brown; outer layer detaching easily from stone when ripe, 2—2.5 mm. thick, compact; stone spherical, 1.3-1.8 cm. long and across, rounded at apex, the walls ca. 2 mm. thick, less than | mm. thick to one side of base, smooth, unmarked; spongy layer probably thin. DistRIBUTION. Madagascar. ELECTED SPECIMENS SEEN. Madagascar: eae ee Efetz [Efatsy], Ana- lazaha, 15384 SF (Pp); sine loco, Boivin s.n Eco.ocy. Fruiting in August. 310 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Calophyllum pervillei is a poorly known taxon that can be characterized by its short terminal bud; strongly 4-angled twigs; small, obovate leaf blades with steeply ascending venation and a raised midrib disappearing just short of the apex; few-flowered, rather short inflorescence with a basal internode less than 6 mm. long; and thick-walled stone. Calophyllum pervillei is perhaps related to C. laxiflorum; for the differences between the two, see the latter. 8773 SF, from the Col d’Ambatondradama, between Mahalevona and Sahafihitra, 700 m., may belong to Calophyllum pervillei, but the fruit is ovoid, ca. 2.2 by 1.5 cm., and sharply pointed at the apex; pedicels of this specimen are up to 2.3 cm. long. However, the fruits are quite immature, since the stone is very weak and the embryo is undeveloped, so the shape differences of the fruit may be unimportant. Calophyllum aff. pervillei There are a number of small-leaved collections of Calophyllum from Madagascar that cannot presently be assigned to species. Some of the variation shown by these specimens is described below. 28357 SF, from the Col du Maningotry, 300-600 m., is vegetatively almost identical to the specimens described as C. pervillei, but its ovoid fruits are ca. 4 by 2.4cm., prominently beaked, and drying sharply and densely wrinkled. The outer layer of the fruit is 2-2.8 mm. thick, and the ovoid stone is ca. 2.7 by ca. 1.8 cm., with the walls 2.7 mm. thick. There are three specimens that have the same fruit type, with the outer layer moderately thick (1.2-2.5 mm.) and compact; however, the stone walls are rather thin (0.2-0.9 mm.). Although the lamina varies considerably in size (2.2-6.4 by 1.2-3.4 cm.), variation in venation is less (7 to 11 veins/5 mm., angle of divergence 55-70°), and the terminal bud is always plump, being 3-4 mm. long. The inflorescence axes and pedicels are puberulent. The three specimens differ greatly in fruit size and shape: the fruit may be spherical or ovoid, 1.6-3.3 by 1.4-2.5 cm., and either beaked or rounded at the apex. The status of this group is uncertain. (Specimens: 28905 SF, Amborompotsy (a 1’O. d’ ea is 5217 SF, Ambositra, Itremo, Lamamby; //563 SF, Mont Ambatomenaloha, al’E. d’Itremo.) The relatively small, spherical fruits of 11563 SF (above) are similar to those of 15037 SF (from Sarolamby, Marolambo), which, however, have very short internodes less than 1 cm. long, a terminal bud less than 2 mm. long, and a different facies. There is a group of flowering specimens from around Ambohimirahavavy, to the north of the localities of all the other specimens discussed above, which have the growth habit of Calophyllum milvum, the lowest internode of an innovation being much longer than the others, although they have a terminal bud 3-4 mm. long. Otherwise, in such characters as their clear venation and their prominently raised midrib, they at least superficially approach 15037 SF. (Specimens: 992 SF, Humbert 24890 and 25155.) 1980] STEVENS, CALOPHYLLUM 311 . Calophyllum drouhardii H. Perr. Mem. Mus. Nat. Hist. Nat. Paris, n.s. 24: 79 8, in Humbert, Fl. Madagascar Comores, Fam. 136: 10. fig. 2, 1, 2. 1951. Type: Madagascar, forét Orientale [Andrebo, E. Imerina], Perrier de la Bathie 14635 comm. Drouhard (lectotype, P; isolectotype, P). Tree 10-15 meters tall; trunk and bark not known. Twigs slightly flattened, 0.7-1.5 mm. across, narrowly 4-alate, drying mid to dark brown, sparsely brown-farinose; axillary innovations lacking basal scars; internodes 0.7-1.7(-2.5) cm. long; uppermost pair of axillary buds less than 1 mm. long, inconspicuous; terminal bud plump, 2.5-4 mm. long, brown-puberulent, underdeveloped internode inconspicuous. Petiole 2-5 mm. long, deeply concave above, convex below, fugaceously farinose; lamina elliptic to subrhombiform or obovate, 0.9-3.7 by 0.5-2 cm., rounded to obtuse at apex, + cuneate to acute at base, slightly recurved and not undulate at margin, coriaceous, drying near umber on both surfaces, often shiny above, sparsely and subpersistently farinose especially on midrib below, the midrib axils along stem (rarely also terminal), with (?)1 to 3 to 15 flowers, unbranched, the axis 0.7-6.5 cm. long, subglabrous or brown-farinose, lowest internode 0.7-1.5 cm. long; bracts sometimes foliaceous and persistent at anthesis, to 10 by 7 mm.; pedicels 0.35-2 cm. long, glabrous or sparsely farinose. Flower (?)hermaphroditic; tepals 8 or 9, the outer pair broadly ovate, 4.5-6 by 3.5-5 mm., the inner ones elliptic to obovate, 7.5-8 by 3-4 mm.; stamens 135 to 180, the filaments to 3.5 mm. long, connate for up to 0.5 mm., the anthers suboblong, 0.8-1.2 mm. long, retuse at apex; ovary 1.3-1.5 mm. long, the style ca. 4 mm. long, the stigma peltate, 0.7-0.8 mm. across, not lobed. Fruit ovoid, 2-2.4 by ca. 1.5 cm., acute at apex, drying brown, with shallow, + longitudinal wrinkles; outer layer not detaching cleanly from stone, ca. 0.8 mm. thick, compact; stone ellipsoid, ca. 1.5 by 1.1 cm., rounded at apex, the walls ca. 0.3 mm. thick, smooth, (?)unmarked; spongy layer (?)thin DisTRIBUTION. Central Madagascar (Map 13). SELECTED SPECIMENS SEEN. Madagascar: Manerinerina sur le Tampoketsa, entre l’Ikopa et la Betsiboka, 1600 m., Perrier de la Bathie 16740 (pr); Ambatondrazaka, E. Mambatovositra, Manaka, 10561 RN (Pp); Moramanga, Savoka, 900 m., 26776 SF (Pp); Ambatovy, 28373 SF (pr); Autanditra, Perinet, 8338 SF (p); a Vouest d’Itremo (O. Betsileo), 1500-1700 m., Humbert 28355 (p); pres d’Ankaloza, Boiteau s.n. anno 1968 (Pp); Ambohimitombo forest, 1350-1440 m., Forsyth-Major 341 (sm, xk); Fianarantsoa, Fandrandava, Ampamaherana, 13638 SF (re); Sambalahy, 13530 SF (pr); Andramabovata, 13270 SF (Pp), Tolongoina, Fort Carnot, 13538 SF (p). oi2 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 Eco.ocy. Colline or montane forest, 900-1700 m. alt. (Morat 3258 taken from tree growing in area with podzol pan; base of tree under water (‘‘sur curaisse, pied dans l’eau’’)). Flowering mostly December to February, rarely March and October; fruiting November and January. Loca. usE. The wood is used in construction. Calophyllum drouhardii can be recognized by its slender, narrowly 4-alate twigs, small leaf blades with dense venation and obtuse to retuse apices, and relatively very large, beaked fruit with a thin woody layer. The epithet commemorates E.-J. Drouhard, whose daughter married Perrier de la Bathie. A sheet of Perrier de la Bathie 14635 (collected by Drouhard) at Paris has been designated the lectotype of Calophyllum drouhardii. This specimen has some terminal inflorescences, and the venation on the underside of the lamina is subimpressed. 32. Calophyllum paniculatum P. F. Stevens, sp. nov. Ficure 15, m-o. A speciebus aliis Calophylli in lamina magna (oblongo-)ovata in siccitate concava marginibus valde recurvatis venulis lateralibus subdensis, inflores- centiis axillaribus ramosis, et fructu plerumque ellipsoideo, differt. ree ca. 20 meters tall, d.b.h. to 50 cm.; trunk and bark not known. Twigs slightly flattened, 3-4 mm. across, 4-angled, sometimes with 4 additional raised lines, drying dark brown to blackish, when young sparsely arinose; goad innovations probably lacking basal scars; internodes 1-4 cm. long; uppermost pair of axillary buds subacute, 1.5-7.5 mm. long, spreading; sere bud plump, 1.1-2.4 cm. long, with gray to brown, subcrustaceous indumentum, underdeveloped internode 3—10(-14) mm. long. Petiole 1.7-2.4 cm. long, broadly and deeply concave above and convex below, glabrescent; lamina narrowly ovate to oblong, 9.5-30 by 3.3-7 cm., acute at apex, acute to attenuate at base, distantly undulate and recurved at margin, drying broadly convex, sepia or olivaceous above and sabelline to fulvous-umber below, glabrous at maturity, the midrib above generally quickly narrowed near base, at first only margins raised, 0.2-0.3 mm. wide at midpoint, becoming raised toward apex, below raised, striate, the venation subobscure to subapparent on both surfaces, raised, sometimes impressed above or flat below, 11 to 16 veins/5 mm., angle of divergence 75-80°. Inflorescences from foliate axils, with 15 to numerous flowers, usually with branches to 3 cm. long and with 5 flowers, the axis 4.5-14 cm. long, farinose toward base, lowest internode 1.8-4 cm. long; bracts not known; pedicels 1-2.5 cm. long, glabrous or sparsely farinose. Flower (?)hermaphroditic; tepals 8 (rarely 9), the outer pair broadly elliptic to ovate, 5-6.5 by 4.5-6 mm., the inner ones elliptic to obovate, 7.5-11 by 3.5-7 mm.; stamens 380 to 400, the filaments to 5 mm. long, the anthers oblong, 0.6-1.2(-1.4) mm. long, retuse at apex; ovary ca. 2 mm. long, the style ca. 3.5 mm. long, the stigma peltate, 0.8-1 mm. across, + 3-radiate. Fruit ellipsoid (rarely ovoid), 2.4-2.6(-4.2) by 1.9-2.2(-2.6) cm.; + acute at apex, drying vinaceous-brown, 1980] STEVENS, CALOPHYLLUM 313 irregularly striate; outer layer detaching cleanly (?always) from stone, 2-2.3(-3) mm. thick, compact; stone ellipsoid to ovoid-ellipsoid, 1.9-2.4(-3.5) by 1.6-1.75(-1.9) cm., rounded to obtuse at apex, the walls 0.8-0.9(-1.2) mm. thick, smooth, unmarked; spongy layer thin. Type: Madagascar, Maromiandra, Ambanja, 6 Oct. 1954, 11078 SF (holotype, P). DistripuTion. Northwestern Madagascar (Map 13). ADDITIONAL SPECIMENS SEEN. Madagascar: Nossi-Bé, 5518 RN (P), 8278 RN (P), Boivin s.n., March 1851 (r); Ambanja, Ambodimanga (3 km. de Renavony), 1297 RN (P); Andampy, 10673 SF (pr); Sambirano, Massif du Manongarivo, 500 m., 11484 SF (p). Ecotocy. Light forests, to 500 m. alt., sometimes near sea. Flowering January, February, and June; fruiting in November. Loca use. The wood is used in carpentry. Calophyllum paniculatum can be recognized by its long terminal bud; long, narrowly ovate lamina with rather sharply recurved margins; axillary and usually branched inflorescences; and usually ellipsoid fruits. The epithet paniculatum alludes to the axillary, branched inflorescences of this species, an unusual combination in the genus. There are three specimens with fruits: /0673 SF, 11078 SF, and 11484 SF. 11484 SF is the only specimen with ripe fruits; they are very much larger than those of the other specimens (measurements in parentheses in the description above), although they are of the same type. The fruits of 10673 SF and 11078 SF have a well-developed woody layer, although the embryo is still small. The specimen cited above collected by Boivin from Nossi-Bé was included in Calophyllum spectabile by Perrier de la Bathie (1951). A sterile specimen collected by Boivin from ‘‘forét de Ravine-Tsara, Ile Ste. Marie”’ is superficially like Calophyllum paniculatum, although it has more distant venation (only 9 veins /5 mm.). There are a number of other specimens with large leaves, long terminal buds, axillary, sometimes branched inflorescences, flowers with usually more than eight tepals, and rather large fruits. The specimens may provisionally be called Calophyllum aff. paniculatum, but much careful field work is needed here, as with the other Madagascan species. They are discussed below. Calophyllum aff. paniculatum 1 Ficure 15, p, q. Tree 25-40 meters tall, d.b.h. to 100 cm.; outer bark brown, slightly fissured; latex yellow. Twigs 3.5-5 mm. across, with ca. 8 raised lines; terminal bud 1.4—-2.5 cm. long. Petiole 1-2 cm. long; lamina elliptic to suboblong, (8.5—)10-13 by (3-)4-6.5 cm., shallowly retuse to obtuse at apex, (shortly) acute at base, strongly recurved at margin, entire lamina + concave, the midrib above abruptly 314 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 narrowed at base, very narrow and obscured by lamina, the venation raised, 15 to 19 veins/5 mm., angle of divergence 70-80°. Inflorescence branched, with farinose indumentum; pedicels 0.5-1.6(-2) cm. long. Flower with (10 to) 12 to 14 tepals; stamens 185 to 325, the anthers oblong, 0.8-1.4 mm. long; stigma peltate, ca. 1.3 mm. across. Fruit spherical, ca. 3.2 cm. long and across, apiculate, drying brown, wrinkled; outer layer not detaching very cleanly from stone, 4.5-6.5 mm. thick, compact; stone spherical, 2.3-2.7 by 2.1-2.3 cm., apiculate, the walls ca. 2 mm. thick, smooth, unmarked, developed at base. DistripuTION. Northeastern Madagascar (Map 13). SPECIMENS SEEN. Madagascar: Massif de Marojejy, pres du col de Doan- yanala, 800-1200 m., Humbert 23120 (p); Massif de 1’ Anjanaharibe, haute An- dramonta, bassin de la Lokoho, 700 m., Humbert et al. 24526 (p); vallée de l Andramonta, 917 SF (p); Andapa, Ampontsilahy, 14578 SF (P). EcoLocy. Large tree, 700-1200 m. alt. Flowering and fruiting in December (flower scented). Calophyllum aff. paniculatum 2 Ficure 15, 1, s. Tree 15-20 meters tall; trunk and bark not known. Twigs 2.5-3 mm. across, strongly 4-angled or with ca. 8 raised lines; terminal bud 8-10 mm. long. Petiole 0.8-1.1 cm. long; lamina oblong to subelliptic, 5-12 by 2.2-4.1 cm., rounded to retuse (rarely mucronate) at apex, acute to + attenuate at base, broadly recurved at margin, entire lamina + concave, the midrib above abruptly narrowed at base, ca. 0.25 mm. across at midpoint, the venation raised, 7 to 12 veins/5 mm., angle of divergence 70-80°. Inflorescences unbranched [rarely flabellate toward base], subglabrous [or densely subfarinose]; pedicels [0.6-]1.5-3 cm. long. Flower with [8 to 10 or] 12 to 14 tepals; stamens [250 to 305 or] 690 to 720, the anthers oblong, 1-1.3 mm. long; stigma peltate, [1-1.5 or] 2-3 mm. across. Fruit ellipsoid, 3-3.7 cm. long, ca. 1.5 cm. across, rounded at apex, drying brown, with broad, rounded wrinkles; outer layer not detaching cleanly from stone, 2-3 mm. thick, compact; stone ellipsoid, 2.5-2.7 by 1.2-1.3 cm., obtuse at apex, the walls ca. 1.2 mm. thick, smooth, unmarked, not developed at base. DistripuTION. Northeastern Madagascar (Map 13). SPECIMENS SEEN. Madagascar: Nantoraka, au sud de Maroantsetra, 22855 SF (pe); Antalaha, Ambohitralana, 5290 RN (pe); Ampanavoana, 7254 RN (Pp); Fénérive, Analalava, circa 4 km. ouest de Foulpointe, 20-50 m., 20158 SF (p); Tampolo, 15212 SF (p). Ecoocy. Sublittoral forest on sand (22855 SF), also on ‘‘montagne’’ (5290 RN). Flowering June and July; fruiting October and November. 1980] STEVENS, CALOPHYLLUM 315 Calophyllum aff. paniculatum 3 Tree ca. 12 meters tall, d.b.h. ca. 40 cm.; bark not known. Twigs 3-3.5 mm. across, usually 4-angled to 4-subalate; terminal bud (0.7-)1.2-1.5 cm. long. Petiole 0.6-1.2 cm. long; lamina elliptic to oblong, 5.5-14 by 2.7-5.1 cm., retuse (rarely subacute) at apex, cuneate to attenuate at base, strongly to slightly recurved at margin (entire lamina + concave), the midrib above abruptly narrowed at base, 0.3-0.8 mm. wide at midpoint, the venation raised, (6 to) 12 to 15 veins/5 mm., angle of divergence 70-75°. Inflorescences branched or unbranched, puberulent; pedicels 0.7-2 cm. long. Flower with (211 or) 12 to 15 tepals; stamens ca. 390, the anthers oblong, 0.6-1.1 mm. long; stigma peltate, ca. 0.7 mm. across. Fruit ellipsoid, 2.9-3.3 by 1.9-2.1 cm., + acute at apex, drying pruinose-brown to dark brown, + longitudinally wrinkled; outer layer not detaching cleanly from stone, 2.5-3.2 mm. thick, compact; stone ellipsoid, 2.1-2.4 by ca. 1.4 cm., obtuse at apex, the walls ca. 1 mm. thick, smooth, unmarked, developed at base. DistRiBUTION. Eastern Madagascar (Map 13). SPECIMENS SEEN. Madagascar: forét d’Anche-Diego, 178-R-6 [SF] (small leaves) (Pp); Antanimenabaka, 26-R-202 [SF] (P); vallée de la Manampanihiny, environs d’Ampasimena, Humbert 20607bis (p); bassin de la Manampanihiny, mont Vohimano, au nord d’Ampasimena, 600-700 m., Humbert 20682bis (pe); Antalaha, Ampanavoana, Vinanivao, 19183 SF (P); Maroantsetra, Farofandina, 15745 SF (rp), Moramanga, forét ouest d’Ampandroantraka, 16954 SF (pr); Marolambo, Androrangavola, Sandranamby, 15036 SF (pr); forét Infanadiana, 13819 SF (?) (P), 14239 SF (?) (Pp); Fianarantsoa, Rindry, Fort Carnot, 7301 SF (P), 4804 SF (pr); Fort Dauphin, Mahatalaky, forét ee 14545 SF (Pp); sine loco, anon. 279 (P). Ecoocy. Flowering September and December; fruiting June and September. The specimens cited as Calophyllum aff. paniculatum I are the most distinct group. They are characterized by robust twigs and terminal buds, branched, puberulent inflorescences, and spherical fruits with a thick outer layer and stone. The specimens collected by Humbert cited under C. aff. paniculatum 3 are in some ways similar, but have thinner, 4-angled twigs and less dense venation (8 to 10 veins/5 mm Calophyllum aff. paniculatum 2 is less robust than C. aff. paniculatum 1, and its leaves have only seven to twelve veins/5 mm. (vs. 15 to 19 veins /5 mm.). The inflorescence is generally less robust and is unbranched, but the flowers have more numerous stamens. The fruit of C. aff. paniculatum 2 looks somewhat like a superannuated sausage and differs in numerous particulars from that of C. aff. paniculatum I (Ficure 15, p-s). The often or less enclosed in a pocket formed from the concave petioles and lower part of the midribs of the rather persistently erect terminal pair of leaves. The specimens from Feénérive cited under Calophyllum aff. paniculatum 2 may not belong there; both are in flower and differ from the other specimens 316 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 in the description above by the measurements enclosed in brackets. Calophyllum aff. paniculatum 3 is not a cohesive taxon. Four of the specimens cited are in fruit; in all of these the stone lacks the fibrous base characteristic of C. aff. paniculatum 2. The fruits of 4804 SF and 15036 SF are rather sharply wrinkled and have a 2.5-3.2 mm. thick outer layer with obvious latex canals; the stone is obtuse at the apex. In 15745 SF the outer layer is only ca. 1 mm. thick, although the fruits are younger. The outer layer of the fruit of 19/83 SF is ca. 2.5 mm. thick but lacks obvious latex canals. The last two specimens have a more or less flat-drying lamina, and the terminal bud is not enclosed by the erect terminal pair of leaves. Possible Hybridization with Calophyllum inophyllum 19184 SF (ve; from Antalaha, Ampanavoana, Maraontsoro; collected near a river on sandy, clayey soil) was possibly taken from a hybrid between a member of the Calophyllum aff. paniculatum complex and C. inophyllum. Although the specimen has the facies of C. inophyllum, the color of the dry leaf, its strongly recurved margin, more or less invisible midrib, and high venation density (10 to 14 veins/5 mm.) are atypical, as is the terminal bud, which is ca. | cm. long. The pecidels in fruit are up to 2.8 cm. long. The fruit is ovoid, ca. 2.6 by 1.9 cm., and beaked at the apex; the outer layer is 1.5 mm. thick; the stone has walls ca. 1.2 mm. thick and a basal plug. Apart from the absence of a thick, persistent, spongy layer, the fruit is very similar to that of C. inophyllum. cee eee eer iene Drake, Bull. Soc. Linn. Paris 2: 1220. 1896; Drake n Grandid. . Phys. Nat. Polit. Madagascar 35: pl. 358, I. 1896; a Perr. ee ve us. Nat. Hist: Nat. Paris, n.s. 24: 79. 1948, in Humbert, Fl. Madagascar Comores, Fam. 136: 10. fig. 2, 4, 5. 1951. Type: Madagascar [environs de Tamatave], Chapelier s.n. (holotype, P). Tree 3-15 meters tall, d.b.h. to 80 cm.; outer bark grayish. Twigs flattened, 3-4(-5) mm. across, rounded or with 4 obscure raised lines, becoming striate, drying brownish black, glabrous when mature; axillary innovations lacking basal scars; internodes in lower part of innovation (2—)3-7.5 cm. long, penultimate internode 0.5—1.5(-—2) cm. long, ultimate internode ca. 3 mm. long; uppermost pair of axillary buds very small and inconspicuous; terminal bud plump, 2-2.5 mm. long, with brown, + crustose indumentum (hairs, Ficure 13, 1), underdeveloped internode absent. Petiole 2.2-4 cm. long, deeply concave above, convex below, glabrous; lamina obovate (rarely elliptic), 8.6-12.7 by 1.7-6.5 cm., rounded or slightly retuse at apex, narrowly acute at base, not undulate but strongly recurved at margin, coriaceous, drying umber above and cinnamon-sabelline below, glabrous when mature, the midrib above gradually narrowed from base, raised, strongly sulcate at first, 0.3-0.5 mm. wide at midpoint, below raised, angled toward apex, striate toward base, the venation + apparent above, apparent below, raised, 4 to 1980] STEVENS, CALOPHYLLUM 317 7 veins/5 mm., angle of divergence 35-55°. Inflorescences from uppermost foliate axils (sometimes two together), with 7 to 11 flowers, unbranched, the axis 3-8 cm. long, glabrous, lowest internode 2-4.5 cm. long, the flowers tending to be in fours, with two pairs separated by a short internode, the groups of four separated by a longer internode; bracts ovate, to 3 mm. long, deciduous; pedicels 1-2.5 cm. long, glabrous. Flower (?)hermaphroditic; tepals 8 (rarely 9), the outer pair suborbicular, 7.5-9 by 7.5-8 mm., strongly concave, sometimes persisting in fruit, the inner ones elliptic to oblong, 9-14 mm. by 4.5-6.5 cm.; stamens 230 to 360, the filaments to 5 mm. long, the anthers oblong, (0.8-)1.3-2 mm. long, slightly retuse at apex; ovary ca. 1.5 mm long, the style 3.5-4 mm. long, the stigma peltate, 1.5-1.8 mm. across, + 4-radiate. Fruit ellipsoid, ca. 1.8 by 1.5 cm., apiculate or not, drying each brown, rather broadly wrinkled; outer een. separating + cleanly from stone when ripe, 2-3.5 mm. thick, compact except for air spaces developing under skin; stone ellipsoid, 1.2-1.4 by 1-1.2 cm., apiculate or not, the walls 0.4—0.6 mm. thick, smooth, unmarked; spongy layer thin. DistripuTion. Madagascar, the eastern coast (Map 12). SELECTED SPECIMENS SEEN. Madagascar: Mananara nord, Ant [an] aa apne SF (p); Fenérive, Amposina, Tanambao-Tampolo, Tampolo, 16/01 SF (pr); pres de Brickaville, Perrier de la Bathie 14077 (rp); Ambila-Lemaitso, ae manala, 8311 SF ha Mahanoro, Tambola, 4901 SF (pr); Ambilabe, Ambinany, 21530 SF (pe); Mananjary, F. Marohita, 19538 SF (p); Pangalona, 9511 SF (Pp); Andranomanry, 5628 SF (p); Faraony, Perrier de la Bathie 11571 (r). Ecotocy. Sandy soil near coast, low elevations. Flowering January, Sep- tember, November, and December; fruiting July, September, November, and December. Prominent, elliptic, crateriform galls (projections ca. 3.5 by 3 by 2 mm. sometimes occur on the lower side of the lamina, either irregularly distributed or in a row often near the margin (Ficure 5, g). Loca uses. The wood is used in construction, and oil is sometimes extracted from the see Calophyllum chapelieri can be recognized by its moderate-sized leaves more or less grouped together at the end of an innovation, and its flowers, which are borne in fours along the inflorescences. The epithet commemorates L. A. Chapelier, a collector who died in Madagascar at the age o Calophyllum chapelieri is closely related to C. ls the differences separating the two species are discussed under the latte There are two collections from Ambila-Lemaitso (4929 § SF (pr), Boiteau 1127 (p)) that appear to be intermediate between Calophyllum inophyllum and C. chapelieri. Their leaf blades are thinner than those of C. chapelieri and are more acute at the apex, the midrib on the upper surface of the lamina is rather abruptly narrowed at the base, and the margin of the lamina is less strongly recurved. The internodes at the end of the innovation are longer, and the fruits (Boiteau 1127) have stone walls ca. 0.8 mm. thick 318 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 and a thin, but well-marked, spongy layer. In all these characters, apart from the leaf apex, these specimens approach C. inophyllum. However, they differ from that species in having rather narrow leaves that are loosely aggregated at the ends of the innovations and narrowly recurved at the margins, and in the fruits, which have only a thin spongy layer and apparently lack a basal plug. A sterile specimen (Geay 7582 (mo (s.n.), Pp), from Mananjary), cited as Calophyllum spectabile by Perrier de la Bathie (1951, op. cit.), also has pseudoverticillate leaves. However, this specimen has a terminal bud up to 2.5 cm. long enclosed in a pocket formed by the concave petiole and the lower part of the midrib, and the lamina is ca. 25 by 5-7.5 cm. with a rounded to subcordate base. 34. Calophyllum verticillatum P. F. Stevens, sp. nov. Ficure 15, j-l. C. laxiflorum auct., non Drake; H. Perr. in Humbert, Fl. Madagascar Comores, Fam. 136: 12. fig. 2, 6-8. 1951, pro parte. A speciebus aliis Calophylli (C. chapelieri excepto) in paribus duobus vel tribus foliarum apice innovationis pseudoverticillatis dispositis et floribus inflorescentiarum necnon pseudoverticillatis dispositis differt, eta C. chapelieri in ramulis gracilioribus, foliis inflorescentiis floribusque minoribus, staminibus 120 ad 150 (non 230 ad 360), et parietibus putaminis 0.2-0.35 mm. crassis (non 0.4-0.6(-1) mm. crassis), differt. e 10-20 meters tall, d.b.h. to 40 cm.; bark not known. sae slightly flattened, 1-1.5 mm. across, 4-angled when young, soon ecoming rounded, drying brown, glabrous; axillary innovations lacking basal scars; internodes 1-4 cm. long, terminal internode of innovation ca. 2 mm. long; uppermost pair of axillary buds very small and inconspicuous, terminal bud plump, |.3-1.7 mm. long, with short, brown indumentum (hairs, Ficure 13, i), underdeveloped internode absent. Petiole 0.3-1.5 cm. long, concave above and convex below, glabrous; lamina obovate, (2—)3-5.3(-8) by (0.45—)1-2.2(-3.1) cm., rounded to slightly retuse at apex, acute to cuneate at base, not undulate and not recurved to narrowly so at margin, coriaceous, drying fulvous-umber above and sabelline below, glabrous or transiently farinose on midrib below, the midrib above gradually narrowed from base, + raised, 0.2-0.35 mm. wide at midpoint, below slightly raised (margins sometimes slightly depressed), substriate, the venation apparent on both surfaces, raised, 5 to7 veins /5mm., angle of divergence 30-50°. Inflorescences from uppermost foliate axils, with 5 to 7 flowers, unbranched, axis (0.5-)1.5-5.5(-7) cm. long, glabrous, the lowest internode (0.5-)l- cm. long, the next internode not developing, the uppermost, if ae well developed; bracts ovate, ca. 2.5 mm. long, soon deciduous; pedicels 0.7-2.7 cm. long, glabrous. Flower (?)hermaphroditic; tepals 8, the outer pair broadly ovate, 5.5-6.3 by 4-5.5 mm., strongly concave, the inner ones elliptic to suboblong, 7-10.2 by 2.6-5 mm.; stamens 120 to 150, the filaments to 4 mm. long, connate for up to 0.5 mm., the anthers oblong, 1.6-2.2 mm. long, 1980] STEVENS, CALOPHYLLUM 319 retuse at apex; ovary 1.8-2 mm. long, the style ca. 3.7 mm. long, the stigma peltate, 1.2-1.4 mm. across, + 3-lobed. Fruit ovoid to subspherical, 1.3-2 by 1.2-1.6 cm., mucronulate, drying pale brown, drying closely and sharply wrinkled when young, smooth when older; outer layer not detaching cleanly from stone, 2.5-3.3 mm. thick, compact apart from air spaces developing under skin; stone ellipsoid, 0.9-1.4 by 0.8-1.2 cm., rounded at apex, the walls 0.2-0.35 mm. thick, smooth, unmarked; spongy layer thin. Type: Madagascar, Fenérive, Ampasina, Tampolo, 8 Jan. 1957, 16487 SF (holotype, P; isotypes, P). DistripuTion. Madagascar, the eastern coast (Map 12). Pres SPECIMENS SEEN. Madagascar: Antalaha, Ambohitralanana, mai N (r), Vavasaha, 9109 RN (p), 9125 RN (p); Anfanavoana, Ajanazana, tania 7490 SF (Pp); Anandrovola, S. de Rantabe, 8933 SF (pr); Mee cee Ambodipaka, see SF (Pp); Soanierana Ivongo, 2456 SF (pr), Forét Sahavo- lawena, 7929 SF (pr); Fénérive, Amposina, Tampolo, 3 m., 17702 SF (p); Forét de Mangalimaso, a!’ ouest de Foulpointe, 22104 SF (r); Ambila- Lemaitso, Tampila, 6476 SF (p); Farafangana, Thorombe, Analazaha, 4833 SF (p); vallée de la Manampanihy, aux environs d’Ampasimena, 20-100 m., Humbert 20607 (Pp); sine loco, herb. du Petit-Thouars (p), Chapelier s.n. (P). Ecotocy. Lateritic or sandy soils, low altitudes. Flowering October, De- cember, and January; fruiting July (fruit not quite mature), September to December. Ellipsoid, crateriform galls ca. 2 by 1.5 mm. occur on the lower surface of the lamina on some specimens. Loca. use. The wood is used in construction. Calophyllum verticillatum and C. chapelieri are similar in their distinctive inflorescence type, with the flowers in pseudowhorls or at least close together, and in the arrangement of leaves at the ends of the twigs, and they have similar fruits and crateriform galls. However, C. verticillatum is consistently smaller in all its parts, its lamina has an at most slightly recurved margin, the arrangement of flowers in fours along the inflorescence axis is more regular, the thick sepals are rarely persistent around the base of the fruit, the stamens are only half as numerous, and the ovoid to subspherical fruit has a stone about half the thickness of that of C. chapelieri. (The almost whorled leaves and flowers of C. verticillatum suggested its specific epithet.) The distribution of C. verticillatum is somewhat more northerly than that of C. chapelieri, but it has also been collected to the south of the known range of C. chapelieri. 7490 SF, from Antalaha, is the only specimen more or less intermediate between the two species. It has larger leaves than is usual for Calophyllum verticillatum (the measurements in parentheses in the description above), ellipsoid fruit, and subpersistent sepals. However, the facies and infructescence type are more those of C. verticillatum; C. chapelieri is not known from Antalaha. 320 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 The relationship between Calophyllum verticillatum and C. chapelieri, although undoubtedly close, needs to be investigated in the field; specific rank for the two taxa seems most appropriate at present. 35. Calophyllum fibrosum P. F. Stevens, sp. nov. A speciebus aliis Calophylli in lamina parva nervis lateralibus infra planis vel leviter impressis marginibusque costarum depressis, et in fructu strato exteriore fibris prominentibus subpersistentibus circa putaminem, differt. Tree 6-30 meters tall, d.b.h. to 35 cm.; bark not known; latex yellow. Twigs slightly flattened, 0.7-1.7 mm. across, strongly 4-angled, drying brown, glabrous; axillary innovations lacking basal scars; internodes 0.5—5 cm. long; uppermost pair of axillary buds rounded, ca. 0.7 mm. long, + spreading; terminal bud plump, 1-1.5 mm. long, with short, brown hairs, underdeveloped internode to 1 mm. long. Petiole 0.3-1.5 cm. long, somewhat concave above and convex below, glabrous; lamina obovate to subelliptic, 2.9-5.8(-8.5) by 0.9-3.3(-4.5) cm., subacute to subretuse at apex and narrowly cuneate to acute at base, slightly undulate and strongly recurved at margin, coriaceous, drying brown to greenish brown and shiny above, paler brown and subpruinose below, glabrous at maturity, the midrib above narrowing gradually from base, raised, center + sulcate, 0.1-0.25 mm. wide at midpoint, below slightly raised, margins depressed, angled, somewhat inconspicuous, the venation apparent above, raised, subobscure below, + impressed, (4 to) 6 to 12 veins/5 mm., angle of divergence (55-)60-70°. Inflorescences from uppermost foliate axils (rarely from lower down), with 5 to 11 flowers, (flowers flabellately arranged), unbranched, the axis (0.5—)1-4.2 cm. long, glabrous or puberulent toward base, lowest internode (0.2-)0.5—-1.8 cm. long; bracts not known; pedicels 0.7-1.5 cm. long, glabrous, slender, to 2.7 cm. by 1.5 mm. in fruit. Flower (?)hermaphroditic; tepals 8 (rarely 10), glabrous, the outer pair suborbicular to broadly ovate, 4.7-5.7 mm. long and across, the inner ones oblong to elliptic, 7.5-9 by 2-4 mm.; stamens 140 to 155, the filaments to 3.5 mm. long, the anthers oblong, 0.9-1.3 mm. long, slightly retuse at apex; ovary 1-1.4 mm. long, the style 3.5-3.7 mm. long, the stigma peltate, 0.7-1.2 mm. across, + 3-radiate. Fruit ellipsoid, 1.6-2 by 1.3-1.6 cm. (rarely obovoid, ca. 2.3 by 1.25 cm.), rounded or mucronulate at apex, drying brown to pale brown, smooth; outer layer not detaching cleanly from stone, 3-4 mm. thick, subcompact, very fibrous; stone + ellipsoid, 1.3-1.6 by 0.8-0.9 cm., + obtuse at apex, the walls 0.4-0.9 mm. thick, smooth, unmarked, fibrous at base; spongy layer thin. Type: Madagascar, Antalaha, Ambohitralanana, 16 Nov. 1956, 8594 RN coll. Ranzokiny (holotype, p; isotype, P). DistripuTion. Northeastern Madagascar (Map 12). ADDITIONAL SPECIMENS SEEN. Madagascar: Vohemar, Anahomana, 4215 SF (rp); Sambava, 8076 RN (pr), Antsahovy, Antongondriha, 859 SF (pr); entre Tsaratanana et Analamanara (route de Sambava et Vohemar, entre Nosiarina 1980] STEVENS, CALOPHYLLUM BPA et Antsirabe-N.), 27644 SF (rp); S. d’Analamanara (pres de Tsaratanana), entre Sambava et Antisirabe-N., foréts entre la Bemarivo et la Mahanara, 27177 SF (pr); Antalaha, Ankorodomo, 7041 RN (Pr); Maroantsetra, Andrano- fotsy, Farankaraina, 15526 SF (pr), 15710 SF (rp), 16357 SF (v), 15 m., 17726 SF (r); Mananara, Ambatomiloma 5802 SF (Pp); cote est de Madagascar au Neme, Chapelier s.n. (P). Eco.ocy. Flowering March and December (flower scented); fruiting March, July, August, November, and December (outer layer of fruit quite pleasant to eat (859 SF)). LocaL uses. The wood is used in construction and canoe building. Calophyllum fibrosum is a distinctive species, even when sterile, because of its small, usually obovate lamina. The venation on the lower surface is subobscure and often impressed, and the midrib is rather inconspicuous since it is raised and has depressed edges. Fibers of the outer layer of the fruit persist around the stone after the epicarp has fallen away, hence the specific epithet. Branching is profuse, and the terminal bud may not be functional. 36. Calophyllum eputamen P. F. Stevens, Jour. Arnold Arb. 57: 168. 1976. Tyre: Mauritius, Pétrin, 600 m., 3 Dec. 1973, Coode 4263 with Guého & Badré (holotype, k; isotype, a). Ficure 6, f, g. Tree 5-6 meters tall, d.b.h. to 30 cm.; trunk without buttresses; for details of bark see varieties. Twigs not flattened or slightly so, 2.5-6.5 mm. across, 4-angled, drying brown to dark brown, glabrous; axillary innovations lacking basal scars, although often with pair of scars ca. 1 cm. from base; internodes 0.3-2.5 cm. long; uppermost pair of axillary buds unknown; terminal bud plump, 5-10 mm. long, with grayish, crustaceous indumentum (hairs, Ficure 16, a, b), underdeveloped internode absent. Petiole 0.5-1.5 cm. long, broadly and shallowly concave above, convex below, glabrous; lamina elliptic to suborbicular or obovate (rarely ovate), 5.7-17.4 by (3.3-)3.9-9.8 cm., obtusely pointed to retuse at apex, cuneate to acute at base, not undulate to distantly so, slightly recurved at margin, coriaceous, drying cinnamon to honey color above and below, glabrous when mature, the midrib above narrowed gradually from base, flat to slightly raised, striate, 0.9-3 mm. wide at midpoint, often disappearing ca. 5 mm. short of apex, below flat to raised, striate, the venation obscure to apparent above, apparent below, raised, 6 to 13 veins/5 mm., angle of divergence 60-75°. Inflorescences from foliate axils (very rarely terminal), with 9 to 17 flowers, unbranched, the axis 4.511 cm. long, glabrous, lowest internode 0.5-3 cm. long; bracts broadly ovate to suborbicular, 0.3-2 cm. long, subpersistent, subfurfuraceous below; pedicels 0.9-3.4 cm. long, glabrous. Flower (?)hermaphroditic; tepals 8 to 12 (or 13), the outer pair suborbicular to broadly ovate, 6.5-10 by 5-9 mm., sometimes glabrous, the inner ones elliptic to obovate, 9-13 by (1.5-)3.5-9 mm., innermost tepals much smaller than outer ones; stamens 270 to 410, the filaments to 7.5 mm. long, connate for up to 1 mm., the anthers oblong, 1.3-2 mm. long, + truncate LPy JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 at apex; ovary 2-2.7 mm. long, the style 4-5.5 mm. long, the stigma peltate, 1.2-1.7 mm. across, (2- to) 4-radiate. Fruit ellipsoid to ovoid, 2.4-4 by 2-2.5 cm., + acute at apex, drying brown, finely wrinkled; the outer layer (?2-)3-3.5 mm. thick, at apex and base to 7.5 mm. thick, compact; stone and spongy layer absent; embryo to 2.2 by 1.5 cm. Key to the Varieties of Calophyllum eputamen — . Terminal bud 4.5-7 mm. long; petiole 5-10 mm. long; lamina 5.7-10.5 3 CM). 1ODB ss go ee ee a A ew we we A a. var. eputamen. l. dre bud 8-10 mm. long; petiole 8-15 mm. long; lamina (7-)9.5-17.5 MOTI couse 5.42 ee Za “tio”? (Horne Islands); en a a Tahiti, New Caledonia); ‘“‘Alexandrian laurel’’ (formerly widely used, transferred and inappropriate name’’—Burkill, loc. cit.). Calophyllum inophyllum is a very useful plant. ‘‘To boil bark with water can be used as dye for fish net. Fruit produces oil called Bitter Oil used as oil for lighting purposes, fuel for automobile, airplane, and generating electricity. To mix with acid makes electric oil. It can also be used to make candle and medicine’ (Fung 20261, from Hainan). Although some of these uses are apocryphal, most parts of the plants are important to local peoples throughout the range of the species. The wood is moderately heavy and fairly strong and has a closely interlocked grain; itis especially durable under water. Canoes and small boats are frequently made of Calophyllum inophyllum, and it provides wood for ship building (mostly keels, knees, and pulley blocks). The wood is used in various other aspects of construction, as well as in making cart wheel hubs, food vessels, and bowls (the latter apparently mostly in the Pacific). Oil can be extracted from the seeds by heating them either cut and with water or finely crushed. The oil contains resin; the purified oil contains approximately 49.7% oleic acid, 23.8% linoleic acid, 16.8% palmitic acid, and 9.7% stearic acid (see Sastri et al., loc. cit.). It is used in soap making skin infections when applied externally. Injected into the muscles, the refined oil relieves the pain in leprosy. In parts of Oceania the oil is used to anoint the body, as either a substitute for, or a pleasant-smelling additive 1980] STEVENS, CALOPHYLLUM 333 to, coconut oil. Mixed with the resin of Vateria Roxb. (Dipterocarpaceae), the oil is used for caulking boats (India). Defatted protein from the seeds is of some nutritional value (Venkatasan & Rege, 1973). The round stones are used as marbles. The pounded bark may be used in cases of orchitis; the juice as a purgative; a decoction of the bark against indolent ulcers. The aromatic latex exuding from the bark is an emetic and a purgative, or it may be used as a scent (Tahiti); it has also been used as a dye and in the treatment of wounds. An infusion of the leaves is frequently used as an eye wash, but it has many other uses (e.g., against chicken-pox). The juice exuding from squeezed leaves has been used against hemorrhoids; the Cambodians inhale over leaves when they have headaches. Calophyllum inophyllum contains saponins, hydrocyanic acid, and also poisonous coumarin derivatives (Kawazu et al., 1968), and is sometimes used as a fish poison. There are numerous other medicinal and quasi-medicinal uses of the plant (see especially Sastri et al., loc. cit.; Heyne, loc. cit.; Burkill, loc. cit.; Perry, loc. cit.). Galophiitan aonhylbans is quite widely planted as an ornamental tree and is known as ‘‘Alexandrian laurel.’ Calophyllum inophyllum is a distinctive species readily recognizable by its medium-sized to large, oval to elliptic leaf blades that are more or less rounded at the apex and have relatively distant and distinct venation; its large, long-pediceled flowers that nearly always have eight tepals; and its fruits, which dry grayish and sharply wrinkled. The only species of Calophyllum habitually found on sandy beaches, it is a characteristic species of such habitats in the Indian and western Pacific oceans. The epithet inophyllum means “‘fiber leaf’’ and refers to the closely set veins that are characteristic of the genus as a whole. The major infraspecific variation in Calophyllum inophyllum is in fruit size. Calophyllum inophyllum var. takamaka was described from a small-fruit- ed form from Aldabra Island (western Indian Ocean) with fruits less than 25 mm. across; fruits from the Pacific were supposed to be 27-40 mm. across (Fosberg, Joc. cit.). All 19 fruiting specimens of C. inophyllum from Madagascar seen at Paris (except for a possibly incorrectly localized sheet in the Richard herbarium) have fruits less than 27 mm. across; the fruits of C. inophyllum in the Mascarenes may be similar in size (Stevens, 1976, loc. cit.), while those from the Seychelles are less than 3 cm. across. Although the fruits of specimens from Ceylon and India eastward are usually more than 3 cm. across, there are numerous exceptions with fruits that are ca. 25 mm. or less in diameter (e.g., Vietnam (Pierre 3652), Sabah (S 1644, A 726—the latter cited by Fosberg, Joc. cit.), Ryukyu Islands (Wilson 8020), Hainan (Lau 5631), Java (Koorders 28793), and Timor (Kooy 837)). Thus, although it seems best not to recognize the small-fruited variant of C. inophyllum formally, it is nevertheless of considerable interest that this small-fruited form should be predominant in the western Indian Ocean, when it is less frequent elsewhere. 334 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Throughout the range of Calophyllum inophyllum, some specimens may have more or less Strongly stipitate dried fruits with the stipe up to 12 mm. a specimens from the Moluccas (e.g., Pleyte 131) have a lamina that is suboblong and rather abruptly narrowed at the base, with the midrib on the upper surface also abruptly narrowed at the base. Although such specimens are very distinctive at first sight, all intermediates between them and more typical specimens occur. Although Calophyllum wakamatsui is clearly referable to C. inophyllum, its status is unclear. Fruits of specimens of C. wakamatsui growing inland on the Caroline Islands have a remarkably thick (to 3 mm.) outer layer; the leaves are also more coriaceous than is usual. If this taxon is recognized as a variety, the name C. inophyllum var. wakamatsui Fosberg & Sachet (Smithson. Contr. Bot. 45: 12. 1980) is available. The exact boundaries of the natural distribution of Calophyllum inophyllum are unclear. It was earlier suggested (Stevens, 1976, loc. cit.) that C. inophyllum might be introduced in the Mascarenes; both seedlings and regeneration occur there, and at least seedlings are found in places where introduction is more obvious, as at Abidjan, Ivory Coast. Calophyllum inophyllum seems to be native on the East African mainland (see Robins, 1976; Robson, Joc. cit.). Hillebrand (/oc. cit.) noted that C. inophyllum was planted in groves on Hawaii and was probably an early introduction; elsewhere in the Pacific it has been recorded as occurring in sacred groves (often cut down by the early missionaries). The wide distribution of C. inophyllum in the Pacific may in part have been caused by man—see Map 14. Nomenclature Roxburgh’s Calophyllum gel is based on the plant illustrated and described by Rumphius (Herb. Amboin. 2: 211. ¢. 7/. 1741). This plant, Bintagor maritima, is clea ae C. inophyllum, and Rumphius gave much interesting information about it Blume mentioned in his desorption of Calophyllum inophyllum that the flowers had twelve tepals (four sepals and eight tepals; Blume, loc. cit.). Blume’s description. Hasskarl (loc. cit.) reduced C. blumei to a variety of C. inophyllum (taxa that Hasskarl designated by lower-case Greek letters appear from his discussion to be varieties). Although C. inophyllum usually has eight tepals, an occasional flower has up to thirteen, and infraspecific taxa based on tepal number cannot be maintained. Miquel described two forms of Calophyllum inophyllum, formae B obovata and y oblongata. There are several sheets of C. inophyllum (obovata) at Leiden and Utrecht; sheet 903,343-104 at the former herbarium is designated the type of forma obovata. One shoot on this sheet has the leaves damaged and strongly retuse just to one side of the apex (perhaps leading to Miquel’s mention of the apices being ‘‘perspicue emarginatis’’); on a label both the 1980] STEVENS, CALOPHYLLUM 335 height of the tree (‘‘40’’’) and the type locality are written. The plant was almost certainly collected by Junghuhn, although this is not mentioned on the sheet. I did not see any sheets of C. inophyllum y (oblongata) from Utrecht; sheet no. 903,343-55 at Leiden is made the lectotype of this name. It agrees with the original description and was collected by Junghuhn. Balsamaria inophyllum was correctly considered to be the same as Calo- Phyllum inophyllum by Willdenow in his edition of Loureiro’s Flora Cochin- chinensis. There is a specimen labeled Balsamaria inophyllum in the British Museum collected by Loureiro from Cochinchina; it is C. inophyllum References to Calophyllum inophyilum in the literature are numerous; only a few are cited above. Anyone interested in a more complete summary of the literature should write to the author 39. Calophyllum tacamahaca Willd. Sitzungsber. Ges. Naturf. Fr. Berlin Mag. 5: 79. 1811; Choisy, Mém. Soc. Hist. Nat. Paris 1: 228. 1823, in DC. Prodr. 1: 562. 1824, pro parte; Sprengel, Syst. Veg. ed. 16. 2: 571. 1825; Camb. Mém. Mus. Hist. Nat. Paris 16: p/. 17C. 1828; G. Don, Gen. Syst. 1: 622. 1831, pro majore parte; Bojer, Hortus Maurit. 52. 1837; Wight, Illus. India Bot. 1: 128. 1840; Choisy, Rev. Guttif. Inde, 43. 1849, Mem. Soc. Phys. Hist. Nat. Geneve 12: 423. 1851, pro parte; Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 286. 1862; Vesque, Epharmosis 2: ¢. 2. 1889, in DC. Monogr. Phanerog. 8: 547. 1893, pro parte; Cordemoy, FI. fle Réunion, 333. 1895; P. F. Stevens, Jour. Arnold Arb. 57: 177. 1976. Tyre: Bourbon [Réunion], du Petit-Thouars s.n. (holotype, B, herb. Willd. 10115). C. spectabile Willd. Sitzungsber. Ges. Naturf. Fr. Berlin Mag. 5: 79. 1811; Gen. Syst. 1: 622. 1831, pro parte; Bojer, Hortus Maurit. 52. 1837. Type: Ile de France [Mauritius], du Petit-Thouars s.n. (holotype, B, herb. Willd. O11 C. lanceolatum Blume, Bijd. Fl. Nederl. Indié 4: 217. 1825; Miq. Fl. Indiae Batavae 1(2): 511. 1859. Type: described from cultivated material originat- ing from Mauritius. C. lanceolarium Roxb. Fl. Indica. ed. 2 (W. Carey, ed.). 2: 608. 1832; Hooker & Arnott, Bot. Beechey Voy. 173. 1833; Wight, Illus. Indian Bot. 1: 129. 1840. Type: described from cultivated material originating from Mauritius Saas C. inophyllum auct., non L.; Lam eek Meth. Bot. 1: 552. 1785, pro parte; Baker, FI. Vener 16. 1877, pro syn Tree 10-15 meters tall, d.b.h. to 25 cm.; trunk without buttresses; outer bark yellowish or gray in patches, smooth, inner surface brown; under bark brownish red; inner bark pink; latex yellow, clear, sticky (Coode 4174). Twigs slightly flattened, 1.5-3.5 mm. across, + 4-angled at first, soon striate, drying brown to dark brown when young, grayish when older, glabrous or sparsely brown-farinose when young; axillary innovations lacking basal 336 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 scars; internodes 0.5-2.7(-4.5) cm. long; uppermost pair of axillary buds rounded, to 1.5 mm. long, spreading; terminal bud plump, 4-8 mm. long, with gray to brown, furfuraceous to subcrustaceous indumentum (hairs, Ficure 16, p), underdeveloped internode absent. Petiole (0.7—)1.1-2.5 cm. long, elliptic (ovate to oblong), (6.3—-)8-18 by (2-)3.4-5.5(-6.8) cm., + acute (rarely subrounded) at apex, broadly cuneate to acute (narrowly decurrent) at base, undulate and slightly recurved at margin, coriaceous, typically drying greenish olivaceous above and olivaceous below, midrib and margins brownish, shiny (rarely near umber on both surfaces), glabrous (rarely sparsely subfarinose) on midrib below, the midrib above gradually narrowed from base, depressed, 0.3-1 mm. across at midpoint, becoming raised toward apex, below raised, + striate, the venation apparent above and below, raised, with (5 to) 7 to 12 veins /5 mm., angle of divergence 60-80°. Inflorescences from foliate axils, base, lowest internode (0.2-)0.6-2.6 cm. long; bracts unknown; pedicels (0.6-)1.3-2.1 cm. long, glabrous or almost so. Flower (snenaaphicdiic: tepals 7 to 9, the outer pair broadly ovate, 7.5-11 by 6.5-9.5 mm., the inner ones obovate to elliptic, 11-17 by 4.5-9 mm.; stamens 200 to 355, the filaments to 7 mm. long, connate for up to 1 mm., the anthers oblong, 1.5-1.9 mm. long, + retuse at apex; ovary 2-3 mm. long, the style 4.5-5 mm. long, the stigma peltate, 1-1.3 mm. across, 3- or 4-radiate. Fruit ovoid, 3.7-4.3 by 2-2.3(-3.7) cm. across, + acute at apex, drying grayish to light brown, deeply wrinkled; outer layer not detaching very cleanly from stone, 2.5-4 mm. across, compact; stone ovoid to ellipsoid, 1.8-2.6(-3.1) by 1.4-2.1(-2.5) cm., (obtusely) pointed at apex, the walls 0.9-1.5 mm. thick, smooth, unmarked, with basal plug 4(-8?) mm. across; spongy layer (?)rather well developed, to | mm. thick. DistripuTIon. Mauritius and Réunion (Map 13). SPECIMENS SEEN. See Stevens, Joc. cit. Eco.ocy. Rocks, recent lava flows, or primary or secondary forest, (0—)100- 50. 500) m. alt. Flowering February and March (reported April and May, Bojer, Joc. cit.); fruiting May, October, and November (fruit tastes like an apricot—Maillot & Jules 3). Coode 4174 has galled leaves; the galls are pustular outgrowths of the lamina and sometimes completely deform it. GERMINATION AND YOUNG PLANT. The radicle emerges just to one side of the basal plug, at a point where the stone is about half as thick as it is elsewhere. The basal plug is pushed out; the area of thin stone lost during germination—similar in size and shape to the basal plug —may also be delimited by a line of weakness. Three to five pairs of seedling leaves are produced, each pair separated by internodes (0.5-)1-4 cm. long. The lowest pair of leaves, which is probably sometimes reduced, had been lost in two of the 1980] STEVENS, CALOPHYLLUM Bar five seedlings seen. The seedling has a prominently four-angled stem. (Details from Coode 4175 and MAU 2851.) aa ee LocaL NAMES. ‘““Tatamaca, tatamaca rouge,’’ ‘“‘tatamaca des hauts”’ (Mauritius and Réunion). Calophyllum tacamahaca can be recognized by its striate, grayish-drying older twigs; its ovate-elliptic lamina that is more or less acute at the apex and that usually dries a characteristic olivaceous color and with a depressed midrib on the upper surface; and its fruit. The fruit is pointed at the apex and has a thick (2.5-4 mm. across) outer layer; the pointed stone has a plug at the base. The epithet tacamahaca is taken from the Mauritian name for the plant. The relationship between Calophyllum tacamahaca and C. inophyllum has been discussed earlier (Stevens, Joc. cit.); it should be noted that the length of the lower internode of the inflorescence is similar in the two species (cf. loc. cit., table 1). The two species are clearly related to one another. (The name C. inophyllum var. takamaka has been given to the small-fruited form of C. inophyllum that is common in the western Indian Ocean; for further details, see the discussion after C. inophyllum). Details of the synonymy and infraspecific variation of Calophyllum taca- mahaca are given in Stevens (loc. cit.). MAU 14546 has fruits similar to those characteristic of C. tacamahaca, and although they are less deeply wrinkled (due to their immaturity), they have a well-developed woody layer. However, the leaves are more coriaceous, the apex of the lamina is subrounded, and the midrib and petiole are broad; as a result, the specimen approaches C. eputamen in appearance. 40. Calophyllum comorense H. Perr. Mem. Mus. Nat. Hist. Nat. Paris, n.s. 1948, in Humbert, Fl. Madagascar Comores, Fam. 137: 8 fig. 1, 1-3. 1951. Type: Comoro Islands, Grande Comore, May 1886, Humblot 1542 (holotype, P; isotypes, BM, P, Ww). Tree ca. 20 meters tall; trunk and bark unknown. Twigs slightly flattened, 2-3.5 mm. across, strongly 4-angled when young, soon becoming striate, drying brown, farinose-puberulent at first; axillary innovations apparently lacking basal scars; internodes 0.5—2 cm. long; upper- most pair of axillary buds rounded, ca. | mm. long, + spreading, inconspicuous; terminal bud plump, 3-4 mm. long, with grayish brown, crustose indumentum, underdeveloped internode absent. Petiole 3.5-10 mm. long, broadly and shallowly concave above, convex below, glabrous; lamina elliptic to obovate, 4.3-7.8 by 2.2-3.8 cm., obtuse to rounded at apex, attenuate at base, distantly undulate and slightly recurved at margin, coriaceous, drying + fulvous-umber on both sides, glabrous when mature, the midrib above gradually narrowed from base, depressed, 0.6-0.8 mm. wide at midpoint, below raised, striate, the venation above and below apparent, raised, 8 or 9 veins/5 mm., angle of divergence 65-70°. Inflorescences terminal and (?)axillary, with ca. 15 flowers, branches 3-flowered and ca. 1 cm. long, the axis ca. 3.5 cm. long, 338 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 puberulent toward base, lowest internode ca. 8 mm. long; bracts + elliptic, o 6 mm. long, foliaceous, not persistent; pedicels 5-6 mm. long, sparsely puberulent when young. Flowers known only in bud, (?)hermaphroditic; tepals 8, the two outer broadly ovate, ca. 6 by 4.5 mm.; stamens ca. 120, the anthers oblong, ca. | mm. long, strongly retuse at apex; ovary ca. 1.5 mm. long, the stigma peltate, ca. 0.6 mm. across, 3-radiate. Immature fruit subellipsoid, ca. 1.6 by 1.3 cm., obtuse at apex, drying smooth, brown; at apex, the walls less than 0.2 mm. thick, smooth, (?)unmarked; spongy layer unknown. DistripuTion. The Comoro Islands (Map 12); known only from the type specimen EcoLocy. Type specimen has much moss on twigs, so was perhaps collected at 1000-1200 m. alt. in montane forest (see below). Calophyllum comorense is a poorly known species best characterized by its small terminal bud, elliptic-obovate leaves separated by short internodes, terminal inflorescences, and ellipsoid fruits with a thick outer layer and an apparently thin stone wall. The epithet comorense is derived from the Comoro Islands, where this plant grows. Calophyllum comorense is perhaps related to C. tacamahaca, which, however, has a longer terminal bud, axillary inflorescences, and a thick-walled stone. If C. comorense has ripe fruit with a thin stone wall, as seems probable, C. eputamen may prove to be its closest relative, although the terminal bud of that species is longer, its midrib and lamina are broader, its inflorescences are usually axillary, and its fruit is much larger. Perrier de la Bathie cited Humblot 1442 as the type specimen of Calophyllum comorense and thought that it was collected at 1000-1200 meters on Mt. Karthala, Grand Comore, since the twigs were covered with mosses. Humblot 1442 seems to be a misprint for 1542; the duplicate of this number at the British Museum has a stamped label with the locality given as ‘‘Angouan, Comoro Is.’ (probably Anjouan Island). 41. Calophyllum recedens Jumelle & H. Perr. Ann. Sci. Nat. Bot. IX. 11: 281. 1910; H. Perr. Mem. Mus. Nat. Hist. Nat. Paris, n.s. 24: 78. 1948, in Humbert, Fl. Madagascar Comores, Fam. 136: 2. fig. 1, 4-7. 1951, pro parte. Type: Madagascar, Firingalava, sur l’Ikopa, entre Maevatanana et Andriba, Sept. 1897, Perrier de la Bathie 327 (holotype, P; (?)isotype, K (s.n.)) C. tacamahaca auct., non Willd.; O. Hoffmann, Fest. 200 Jarh. Werder’s. Gymn. Berlin, 319. 1881; Vesque i in C. DC. Monogr. Phanerog. 8: 547. 1893, pro parte; Engler in Engler & Prantl, Nat. Pflanzenfam. ed. 2. 21: fig. 81, g—j. 1923. Tree 3—15(—?30) meters tall, d.b.h. to 30 cm.; outer bark blackish, becoming fissured; inner bark reddish; latex yellow, odorous (19291 SF, Perrier de la Bathie 327ter). 1980] STEVENS, CALOPHYLLUM 339 Twigs somewhat flattened, 1.5-2.8 mm. across, not or obscurely 4-angled, drying brown to dark brown (grayish and striate when old), glabrous; axillary innovations lacking basal scars; internodes 0.5-2.5 cm. long; uppermost pair of axillary buds subrounded, less than 1 mm. long, + spreading; terminal bud plump, 4-8 mm. long, with grayish (rarely subferrugineous), + crustaceous indumentum (hairs, Ficure 16, n), underdeveloped internode to 8 mm. long. Petiole 1-1.8(-2.6) cm. long, broadly and rather shallowly concave above, convex below, glabrous when mature, slender; lamina ovate to suboblong, 4-11.7 by 1.6-4.4 cm., acuminate at apex, abruptly attenuate or rounded to acute at base, undulate but slightly or not recurved at margin, coriaceous, drying + olivaceous-sepia and nitid above, sepia below, glabrous when mature, the midrib above gradually narrowed from base, + sulcate at first (edges usually + clearly demarcated from rest of blade), 0.2-0.35 mm. wide at midpoint, becoming raised toward apex, below raised, + angled, venation above and below + apparent, slightly raised, 8 to 10 veins/5 mm., angle of divergence 60-75(-85)°. Inflorescences from foliate axils along twigs, with 7 to 15 flowers, unbranched, the axis 2.2-7.5 cm. long, sometimes transiently brown-puberulent, especially in lower part, lowest internode (0.35-)0.7—2 cm. long; bracts ovate, ca. 2.5 mm. long, puberulent below, soon deciduous; pedicels 0.6-2 cm. long, glabrous or sparsely puberulent. Flower (?)her- maphroditic; tepals 8 (to 13), the outer pair suborbicular, 5-6 by 4.5-5 mm., the inner ones elliptic to obovate or oblong, 9-12 by 2.7-6 mm.; stamens 170 to 280, the filaments to 5.5 mm. long, connate for up to 2 mm., the anthers ellipsoid to oblong (0.5-)0.7-1(-1.5) cm. long, rounded to retuse at apex; ovary 1.2-2 mm. long, the style 3.5-5 mm. long, the stigma peltate, 0.5-0.9 mm. across, obscurely 3-radiate. Fruit ovoid to ellipsoid, 2.4-2.8 by 1.8-2.2 cm., + obtuse at apex, drying pruinose vinaceous-brown, + wrin- kled (smooth when young); outer layer not detaching cleanly from stone, 1.6-2.5 mm. thick, compact; stone ellipsoid, 1.4-2 by 1.4-1.7 cm., rounded at apex, the walls 0.3-0.6 mm. thick, smooth, unmarked; spongy layer thin. DistripuTion. North-central Madagascar (Map 13). SELECTED SPECIMENS SEEN. Madagascar: vallée d’Ifasy, Perrier de la Bathie 8153 (P); eee [Ambavatobe] , Hildebrandt 3324 (a, M, P, us); Andranofasi- ka, Ampasimatera, Mampikony, 19291 SF (vr); Maevatanana, bassin moyen du Sere (Boina), Perrier de la Bathie 3274 (rp); Beritsoka, E. du Maevatan- ana, Perrier dela Bathie 327bis (p); Ambalanjanakomby, Bentaly-Maevatanana, 14942 SF (pr); Besalampy, Bekodoka, Sahondra, 15235 SF (pr); Ankorika, Ankarafantsika, 150-200 m., Ursch 147 (pr); Morataitra, pres de Maevatanana, Perrier de la Bathie 5308 (rp); forét Bitomandry, 5349 SF (pr); Maintirano, forét Andranomena, 14781 SF (pr); forét Ambatomanga, N. du Anjozorobe, Boiteau s.n. (P). Ecotocy. Locally common in seasonal forest, usually along river banks on siliceous ground; low alt. Flowering February to April and August, September, and November; fruiting September and January (fruit somewhat fleshy; eaten by lemurs (Perrier de la Bathie 3274), despite a somewhat tere- binthlike taste). 340 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 TABLE 8. Differences between Calophyllum vernicosum and C. recedens. C. vernicosum C. recedens WHITISH WAXY COVERING Usually present Absent TERMINAL BUD LENGTH 2.5-4 4-8 mm.) TERMINAL BUD Reddish brown, Usually grayish, INDUMENTUM short-tomentose subcrustaceous LAMINA BASE Acute ae ieee or unded to a MIpRIB ON UPPER Indistinct Usually distinct SURFACE OF LAMINA VENATION DENSITY (6 to) 8 to 12 (to 14) 8 to 18 (veins /5 mm.) ANTHER LENGTH (mm.) 0.9-1.8 (0.5-)0.7-1(-1.5) OUTER LAYER OF FRUIT, 0.6-1 1.6-2.5 THICKNESS (mm APEX OF STONE Apiculate Rounded Loca uses. The wood is used in the manufacture of furniture and canoes, and the latex has apparently been used in the local pharmacopeia (Perrier de la Bathie, 1951, loc. cit.). Calophyllum recedens can be recognized by its medium-sized leaf blades that are acuminate at the apex, often attenuate at the base, and with moderately dense venation. The terminal bud has subcrustaceous indumentum; the twigs are slightly angled. The inflorescences are angled, and the flowers usually have eight tepals. The ovoid to ellipsoid fruits are 2.4-2.8 cm. long, subobtuse at the apex, and with a compact outer layer 1.6-2.5 mm. thick. The epithet recedens means ‘‘receding”’ or ‘‘differing,’’ possibly a simple allusion to the fact that the species was distinct. Calophyllum recedens is perhaps related to C. tacamahaca, from the Mascarenes; Vesque (Joc. cit.) cited Hildebrandt 3324 under the latter species. However, there are numerous differences between the two in twig, leaf, and fruit (Stevens, 1976, rable 1). It should also be noted that C. tacamahaca has longer anthers (1.5-1.9 mm. vs. (0.5-)0.7-I(-1.5) mm.). The stone of the fruit of C. tacamahaca has a basal plug; such a plug has not been noticed in C. recedens. Calophyllum recedens has also been confused with C. vernicosum, for the differences between the two, see the latter species (TABLE 8 42. Calophyllum vernicosum P. F. Stevens, sp. nov. Ficure 15, t. C. recedens auct., non Jumelle & H. Perr.; H. Perr. in Humbert, FI. Madagascar Comores, Fam. 136: 2. 1951, pro parte. 1980] STEVENS, CALOPHYLLUM 34] A speciebus aliis Calophylli in gemma terminalis 2.5-4.5 mm. longa indu- mento subfulva praedita, innovationes axillaribus cicatricibus basalibus haud ornatis, ramulo tegenti albescenti subpersistenti praedito, pagina superiore laminae nitida costa indistincta, et fructu ovoideo vel ellipsoideo apice plerumque acuto strato exteriore 0.6-1 mm. in transverso, differt. Tree 9-12 meters tall; outer bark smooth at first, becoming similar to that of a pine (?fissured). Twigs flattened, 1.3-2.7 mm. across, + rounded, drying blackish, color obscured by whitish (?)waxy covering, transiently rufous-puberulent; axillary innovations lacking basal scars; internodes 0.5-2.5(-4) cm. long; uppermost pair of axillary buds rounded, ca. | mm. long, spreading; terminal bud plump, mm. long, with short-tomentose, rufous indumentum (hairs, Ficure 16, 0), underdeveloped internode to 2 mm. long. Petiole 1.2-2.3 cm. long, broadly concave above and convex below, glabrous; lamina ovate (elliptic), 5-11 by 1.6-3.5(-4.7) cm., acuminate at apex, acute at base, strongly but distantly undulate and not recurved at margin, coriaceous, drying + concave, shiny, greenish brown above and brownish below, glabrous at maturity, the midrib above gradually narrowed from base, at least margins raised, surround- ing lamina also raised, sulcate, 0.25-0.7 mm. wide at midpoint, below slightly raised, + striate, not very conspicuous, the venation subobscure to apparent above and subapparent below, raised, (6 to) 8 to 12 (to 14) veins/5 mm., angle of divergence 60-75°. Inflorescences from foliate axils, with 7 to 15 flowers, unbranched, the axis 2.9-9 cm. long, puberulent, especially toward base, lowest internode (0.1-)0.4-1.5(-2.3) cm. long; bracts ovate, ca. 2. mm. long, soon deciduous; pedicels 0.5-2.4 cm. long, glabrous. Flower (?)her- maphroditic; tepals 8, the outer pair broadly ovate, 3.8-4.5 by 4.2-4.5 mm., the inner ones elliptic to lingulate, 6-9 by 2.5-6 mm.; stamens 75 to 180, the filaments to 4 mm. long, the anthers oblong, 0.9-1.8 mm. long, subretuse at apex; ovary 1.2-1.4 mm. long, the style 3.5-4 mm. long, the stigma peltate, 0.7-1 mm. across, obscurely 3- or 4-radiate. Fruit ellipsoid to ovoid, (1.9-)2.5- 3.4 by 1.6-1.9 cm., acute (rarely rounded) at apex, drying brown, wrinkled (almost smooth at maturity); outer layer detaching + cleanly from stone, 0.6-1 mm. thick, air spaces developing at maturity; stone + ellipsoid, 1.7-2.5 by 1.4-1.65 cm., apiculate, the walls 0.3-0.9 mm. thick, smooth, unmarked; spongy layer thin. Type: Madagascar, Ambilobe, forét degradée au nord d’Ambilobe, 10 Nov. 1958, 18934 SF coll. Capuron (holotype, P; isotype, P). DistRIBUTION. Northern Madagascar (Map 12). ADDITIONAL SPECIMENS SEEN. Madagascar: Ambilobe, Mt. Ambohipiraka, pres d’Ambilobe, Perrier de la Bathie 18754 (rp), 300-400 m., Humbert & Cours 32881 (pr), 32903 (Pp), village d’Ankatoto, Cours 5650 (p), 100-300 m., 5671 (p); Ambilobe, de Cary 14842 (c, p), 5408 SF (rp); plaine d’Ambilobe, 10507 SF (p); Beramanja, route Tafiana-Ambohimanjy, a) RN (pr), 1287 RN (P); Massif de l‘Ankarana, de Cary 14522 (BM, kK, P, US), 14563 (L, Mo, P); Diego Suarez, Ursch 264 (pr); Sambirano, Morat 1416 a sino loco, Baron 6258 (BM, K, P). 342 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Eco.ocy. Locally fairly common in light, sometimes rather dry colline forest, 100-400 m. alt. Flowering July, August, October, and November; fruiting January, August, October, and November (fruit yellowish green). Elliptic, disk-shaped galls to 4.5 by 3 mm. with a slightly raised margin and a raised center sometimes occur on the lower surface of the lamina in a line at, or a little in from, the margin Calophyllum vernicosum can most readily be recognized by its short terminal bud; dark-drying twigs with a more or less transient whitish covering; leaf blades that dry very shiny, especially on the upper surfaces, and with a midrib not sharply distinguished from the rest of the lamina; and ellipsoid-ovoid fruits with a rather thin (less than | mm.) outer layer in which air spaces develop. The epithet vernicosum alludes to the shiny, varnished appearance of the upper surface of the dried lamina; this may also be noticeable in the living leaf, as the field label of Cours 5650 suggests. I originally included the specimens described above as Calophyllum vernico- sum in C. recedens, but the resultant ‘‘taxon’’ was uncomfortably variable. However, when the specimens were re-sorted on the bases of bud size and color, such other characters as leaf type, anther size, and fruit type were found to show correlated variation. In addition, the uppermost internode of the innovation in C. vernicosum tends to be notably shorter than the others (although this is not nearly as marked as in C. verticillatum); there is not this size disparity in C. recedens. The differences between the two taxa are listed in TaBLe 8; although there is some overlap in a few of the characters, many characters are involved and even sterile material can be identified easily. The two species have distinct geographic ranges (cf. Maps 13) b) 43. Calophyllum wallichianum Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 277. 1862. Type: Malaya, Penang, Wallich dist. 4843, pro parte (holotype, G; isotypes, BM, FI, G, K). Tree (3-)15-36 meters tall, d.b.h. toca. 160 cm.; trunk usually unbuttressed; outer bark brown or gray to yellow-gray or mottled yellow, with vertical lines of lenticels or shallowly fissured, hoop marked or not, inner surface brown-orange to orange or yellowish and greenish mottled; under bark reddish orange to red; inner bark red; latex white, often turning yellowish (rarely clear yellow), rather sticky. Twigs slightly flattened, (1.5-)2.3-6 mm. across, obscurely to strongly 4-angled, drying blackish to yellowish brown, rather inconspicuously gray to brown farinose-puberulent (rarely subtomentose) when young; axillary innovations lacking basal scars; internodes (0.7-)1-5 cm. long; uppermost pair of axillary buds rounded to pointed, to 4.5 mm. long, + erect to spreading; terminal bud plump to conical, (1.2-)1.6-3 cm. long, with gray to brown, crustaceous to subtomentose indumentum (hairs Ficures 16, q-cc; 18, a-j), underdeveloped internode 1-6 mm. long. Petiole 1.2—2.5(-3.5) cm. hanes broadly concave above, convex below, often drying blackish, glabrous or with indumentum obscure; lamina oblong to ovate, (5.3-)8.5—25(-32.5) by (2.1-)3- 1980] STEVENS, CALOPHYLLUM 343 6.8(-8) cm., acute to subacuminate at apex, acute or cuneate to rounded at base, distantly undulate and not recurved to moderately so at margin, coriaceous to thickly coriaceous, drying bay to sabelline above and fulvous to olivaceous below, subpersistently farinose-puberulent on midrib below and often above, the midrib above narrowing gradually or fairly quickly from base, often + depressed at first, becoming raised (surrounding lamina some- times raised), 0.2-0.8 mm. wide at midpoint, below strongly raised, striate to angled, the venation subobscure to apparent above and below, + raise (8 to) 10 to 17 veins/5 mm., angle of divergence 65-85°. ilorescences from foliate axils, with 7 to 15 (to 19) flowers, unbranched (rarely with 5-flowered branches to 1.5 cm. long), the axis 1.7-7 cm. long, usually puberulo-tomentose toward base, puberulent toward top, lowest internode 4-11 mm. long; bracts ovate to elliptic, 4-10 mm. long, sometimes subpersistent, with dense, fawn-colored indumentum on both surfaces; pedicels 0.5-2(-2.5) cm. long, subglabrous to puberulent, slender in flower, to 4 mm. thick in fruit. Flower Sopigt tepals 4 (rarely 6), the outer pair ovate to elliptic, 5-7 by 3-3.5 mm., puberulent on back, the inner pair elliptic to obovate, 5.5-6 ey; a 5-4 mm., sparsely puberulent in band down back, or glabrous, additional ones, if any, much smaller; stamens 90 to 185, the filaments to 3.5 mm. long, the anthers oblong, 0.5-1 mm. long, + retuse at apex; ovary 1.3-1.7 mm. long, the style ca. 1.7 mm. long, the stigma peltate, 0.6-0.8 mm. across, 2- or 3-lobed. Fruit ellipsoid, + ovoid or spherical, or obovoid, 2-3(-3.5) by 1.5-2.5(-3) cm., strongly apiculate or not, drying brown to blackish, sharply and shallowly wrinkled; outer layer detaching cleanly from stone, 1-3(-4) mm. thick, compact; stone broadly ovoid to subspherical or ellipsoid, 1.3-2.3 by 1.2-1.9 cm., rounded to apiculate at apex, the walls 0.3-1.1 mm. thick, smooth, usually unmarked (with discoloration at base to 4 mm. across); spongy layer thin. Key to the Varieties of Calophyllum wallichianum 1. Lamina usually very coriaceous, the midrib on upper cane surrounded by raised blade, indistinct. ...........0.0.0. 43c. var. tahanense. 1. Lamina usually coriaceous, the came on upper surface ie never surrounded by raised blade, dis 2. Petiole (1-)1.2—2.2 mm. across; rae (1.5-)2- oF a - mm. across. ees gol she So et Se Se ME IN ees var. wallichianum. 2. Petiole (2—-)2.5—5 mm. across; twigs 2.8-6 mm. across.......... ee er a rg 43b. var. incrassatum. 43a. Calophyllum wallichianum Planchon & Triana var. wallichianum C. wallichianum Planchon & Triana; T. Anderson in Hooker f. Fl. Brit. India 1(2): 273. 1874, pro parte; Vesque in C. DC. Monogr. Phanerog. 8: 599. 1893, pro parte; M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore Oo 303. 1956, pro parte; Kochummen, Malayan Forest Rec. ed. 2. 17: 216. 1965, pro parte; T. C. Whitmore, Tree Fl. Malaya 2: 193. 1973, pro parte. 344 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 C. spectabile auct., non Willd.; Curtis, Jour. Straits Branch Roy. Asiatic . - 1894, pro majore parte; Ridley, Fl. Malay Penin. 1: 185. 1922, p oO parte C, etrapetalu Wall. Catal. 4843. 1831, non Roxb. ex G. Don (1831). Tree 12-30 meters tall; latex milky white, turning yellow on exposure, or opaque lemon-yellow. Twigs (1.5—)2-3(-3.5) mm. across. Petiole (1-)1.2-2.2 mm. across; lamina ovate, coriaceous, the margins undulate, the midrib on upper surface usually raised, even at base, distinct, 0.15-0.3(-0.35) mm. wide at midpoint. Bracts not persistent. DistripuTIon. Northwestern Malay Peninsula (Map 15). SELECTED SPECIMENS SEEN (‘‘f?’? denotes specimen with fruits). Malaya. KEDAH: Kuala Muda, Puah Reserve, KEP 59630 (ker); Kuala Nerang, Kampong Kuala, 120 m., KEP 73532 (?) (KEP). Pinanc: Government Hill, 366 m., Curtis 1153 (f) (BM, K, SING); Pantai Acheh F.R., 150 m., KEP 72564 (KEP); Pulau Pinang, 610 m., FRI 20534 (KEP); Telok Bahang F.R., KEP 66368 (Kep); Pulau Boetong Reserve, Curtis = 1153 (SING). PERAK: Panekor F.R., 120 m., FRI 3021 (a, K, KEP, SAN, SING); Pangkor Is., S. Pinang F.R., 75 m., FRI 3057 (a, k, KEP, SAN, SING); Kelidang Saiong F.R., KEP 33634 (kep, sinc); Kinta, ete F.R., KEP 63253 (ker); Telok Kopia F.R., Dindings, 150 m., FRI 3105 (a, K, “ SAN, SING); Dungun, Sagari- Melintang F.R., KEP 69418 (f) ts SING); Sembilan Is., 30 m., KEP 76530 (f) (a, kK, KEP, SING); Telok Muroh F. KEP 76733 (KEP); Batu Undan F.R., KEP 54225 (KEP); Gunong Sin F.R., 914 m., FRI 17585 (L, sar); G. Bub 396 m., FRI 11748 (k, KEP, L, SAR, SING); Sumpitan, KEP 10407 (k). Necri SemsBiLan: Kuala Pilah, KEP 93800 (KEP). KELANTAN: Kuala Mersing, Sungei Brok, 210 m., FRI 5450 (f) (A, K, KEP, SING). EcoLocy. Colline forest, 30-610(-914) m. alt. Flowering and fruiting in January. p 15. Distribution of Calophyllum wallichianum var. ee (cir- Jean and C. wallichianum var. incrassatum (squares) in Malesi 1980] STEVENS, CALOPHYLLUM 345 GERMINATION AND YOUNG PLANT. The radicle breaks the stone to one side of the base. The seedling has two pairs of leaves separated by an internode less than 1 cm. long. (KEP 10407, from Perak; FRI 6166, from Kelantan.) 43b. Calophyllum wallichianum Planchon & Triana var. incrassatum (Hender- son & Wyatt-Smith) P. F. Stevens, comb. nov C. incrassatum M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 302. pl. 2. 1956; Wyatt-Smith, Malayan Forest Rec. 17: 113. 1952, nomen; Kochummen, Malayan Forest Rec. ed. 2. 17: 216. 1965: T. C. Whitmore, Tree Fl. Malaya 2: 183. 1973; H. Keng, Gard. Bull. Singapore 28: 244. 1976; Corner, Gard. Bull. Singapore Suppl. 1: 104. 1978. Type: Singapore, Bukit Timah, ‘“‘low’’ alt. a Dec. 1937, SFN 34647 coll. Ngadiman (holotype, sinc; isotypes, a, BO, K, KEP, LAE, P, SING). C. griffithii auct., non King: Ridley, ie Straits Branch Roy. Asiatic Soc. 33: 47. 1900. Tree (3-)15-36 meters tall; latex white, often turning yellowish (perhaps rarely initially yellow). Twigs 2.8-6 mm. across. Petiole (2-)2.5-5 mm. across; lamina oblong to subovate (rarely ovate), coriaceous (rarely thickly coriaceous), the margins usually notably undulate, the midrib on the upper surface often more or less depressed at first, becoming raised, usually distinct, 0.3-0.8 mm. wide at midpoint. Bracts often subpersistent. DistriBuTION. Eastern Malaya to Borneo and perhaps Sulawesi (Map 15). SELECTED SPECIMENS SEEN. Malaya. SeLancor: Bukit Lagong F.R., 455 m., KEP 83729 (kep); Semangkok, 915 m., KEP 115248 (ker). Matacca: Pulau Besar, 13 km. from Malacca, Stone 9033 (ku); Bukit Sadanan, Derry 515 (sinc); S. side of Selandar Forest, Burkill 2010 (Bo, kK, SING). KELANTAN: 79859 (keP); along Sungei Pelong, foothill, FRI 14818 (k, KEP, siNG); 59th Mile K. Trengganu-Khota Bharu road, FRI 2524 (?) (k, KEP, SAR, SING); Ulu Dungun, 270 m., FRI 9652 (kep); Bukit Kajang, Kemaman, 150 m., Corner s.n., 2 Nov. 1935 (sinc); G. Padang Exped., Ulu Brang, 1219 m., FRI 12646 (?) Ge K, KEP, SING). PAHANG: Bukit Santong, Rompin iron mine, 365 m., FRI 3605 (A, K, KEP, SAR, SING); Lesong F.R., 180 m., FRI 20008 (xep, SAR); Kuantan, Bukit Goh Reserve, KEP 3628 (x, sinc); Kallam Ayer Reserve, KEP 17250 (kep, sinc); G. Tapis, 575 m., FRI 17692 (ker); Tasek Bera F.R., 75 .m., FRI 16960 (ker); Baloh F.R., KEP 66645 (kep); Temerloh, KEP 5473 (sING). JoHoRE: Mawai-Jemaluang road, SEN 34911 (a, BO, K, KEP, LAE, P, sinc); Kluang F.R., 90 m., FRI 7536 (a, KEP, SAR, SING); Ulu Endau, summit of Bukit Kendok, 396 m., KEP 105015 (a, : KEP, SING); Banang F. R., KEP 79420 (KEP); G. Ledang, 396 m., FRI 19202 (a, KEP, SAN); ae ‘Arong F.R., KEP 84582 (xep); Bukit Jelaboi, ay 9831 (ker); G. Pulai, 610 m., FRI 9919 (KEP); Jemaluang F.R., KEP 69971 (xep); Labis, 60 m., KEP 104953 (kep); Ma’Okil F.R., 6 m., KEP 71279 (KEP); Renggam F.R., 75 m., KEP 71269 (KEP); G. Panti F. R., 305 m. , KEP 93775 (kep). Singapore: ‘Sungei Morai, Ridley 5071 (BM, sING); Bukit Timah, SFN 38846 (LAE, SING); Changi, Ridley S.n., anno 1893 (p); Botanic Gardens, SGN 1692 (sinc). Sumatra and adjacent islands. Riau: Koeantan, Poelau Kedondong, 100 m., bb 24801 (Bo, L, sING); 346 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Upper Riauw, Pakanbaru, Tenajan R., Soepadmo 71 (Bo, c, L, SING). UTara: Hoogl. van Toba, Pandoemaan, 900 m., bb 5695 (Bo); Silindoeng, 1040 m., bb 6620 (Bo). Borneo. Sarawak. Ist Division: Semengoh F.R., 30 m., Stevens et al, 282 (a); Serian, G. Penrissen, S 16375 (a, L, SAN, SAR, SING). 3rd Division: Hose Mts., Bt. Lumut, Ulu Amau, 950 m., S 21255 (a, BO, K, KEP, L, SAN, SAR, SING); Ulu Kapit, Pelagus Protected Forest, S 33191 (KEP, KLU, SAN, SAR) SABAH. seer ineraes Karamuak, Bukit Meliau, 150 m., SAN 39310 (san, SAR, SING); Lamag, E. of Sungei Meliau, 30 m., SAN ‘53.252 (sAN), Bukit Tavai, 240 m., SAN 53316 (SAN). KALIMANTAN. Timur: Tidoengsche Landen, 7m., bb 18354 (a, Bo); Tandjong Redeb, Labanan, bb 11511 (Bo). Celebes. SULAWESI: Manado, Bol. Mangadan, Bonko, 50 m., bb 13751 (Bo). EcoLocy. Usually in well-drained, mixed dipterocarp lowland rainforest, sometimes in swamp forest or in mossy heath forest (S 2/255); 6-400(-1220) m. alt. Flowering November to February (flower scented); fruiting March to May, sometimes September and November (fruit orange (S 2/255)). GERMINATION AND YOUNG PLANT. The seedling has two pairs of leaves separated by an internode less than 1 cm. long. Subsequently produced internodes are much longer, the terminal bud is functional, and the plant is erect. (Stevens et al. 103, 106 (Johore, Malaya), 282 (Sarawak).) The epithet incrassatum (‘‘thickened’’) was coined because the pedicel becomes thickened in fruit. 43c. Calophyllum wallichianum Planchon & Triana var. tahanense (Henderson & Wyatt-Smith) P. F. Stevens, comb. nov C. tahanense M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 305. pl. 4. 1956; T. C. Whitmore, Tree Fl. Malaya 2: 168. 1973 Type: Malay Peninsula, Pahang, Gunong Tahan, Wray’s Camp, + 3500 feet [1067 m.], 27 Aug. 1928, SFN 20601 coll. Holttum (holotype, sinc; isotypes, A, BO, UC). C. spectabile auct., non Willd.; Ridley, Jour. Fed. Malay States Mus. 2: 110. 1909. Tree 6-20 meters tall; latex yellow or pale yellow, clear. Twigs 2-5 mm. across. Petiole 1.5-3 mm. across; lamina ovate, slightly undulate at margin, thickly coriaceous, the midrib on upper surface slightly sunken at base, surrounded by raised lamina, indistinct, 0.2-0.4 mm. wide at midpoint. Bracts not persistent. DistripuTIoN. Malay Peninsula (Map 16). SELECTED SPECIMENS SEEN. Malaya. Necri SEMBILAN: G. Angsi, 455 m., F. 17345 (K, KEP, SING). KELANTAN: Ulu Kelantan, Reali [Relai] F.R., FRI 7256 (A, K, KEP, SAR, SING); path to G. Rabong, 762-975 m., Shah 2510 (a, BRI, KEP, SING); Sungei Kerbut, Jeram Keteh, ca. 2 km. below K. Trenggan, 335 m., FRI 20281 (k, KEP, L, SAN). PAHANG: G. Tahan, 1067 m., SFN 8020 (Bo, K, SING). Eco.ocy. Colline or montane forest, 335-1067 m. alt. Flowering August 1980] STEVENS, CALOPHYLLUM 347 and September; fruiting February, March, June, and October (fruit dull yel- low-green). The type specimen was collected on Gunong Tahan, and this suggested the epithet Calophyllum wallichianum can be recognized by its plump terminal buds over 1 cm. long; its rather large, elliptic to ovate to oblong lamina with moderately dense venation; its unbranched inflorescence with farinose- puberulent to subtomentose indumentum; its flowers that usually have only four tepals, with the backs of the outer pair puberulent; and its medium-sized fruits that dry wrinkled and have a well-developed, compact outer layer 1-3(-4) mm. thick and stone walls 0.3-1.1 mm. thick. The hairs usually have a single, basal branch. The specific epithet commemorates the indefatigable botanist, N. Wallich. Circumscription of Calophyllum wallichianum The taxa and their limits in the Calophyllum wallichianum /incrassatum / ta- hanense complex have been quite troublesome. My original intention was to follow Henderson and Wyatt-Smith’s treatment, although it was clear that, as they realized, C. wallichianum could be separated into northern and western (Kedah, Pinang, Perak) and southern and eastern (Trengganu, Johore, Singapore) parts, the former having basically opaque, white to yellow exudate and sparsely tomentose terminal buds, and the latter with clear, yellow exudate and more tomentose terminal buds. There were also slight differences in Map 16. Distribution of Calophyllum wallichianum var. tahanense — C. scriblitifolium (circles), and C. lowii (half-circles) in Malesia. Inset: C. lowii. 348 JOURNAL OF THE ARNOLD ARBORETUM [vo. 61 leaf shape and terminal bud. Calophyllum incrassatum was separated from C. wallichianum by its more persistent bracts, usually white latex, smooth bark, and more prominent midrib. The northern populations of C. wallichianum are clearly intermediate between the southern populations and C. incras- satum in these characters, except those of the leaf, in which they agree with the southern populations. However, the northern populations of Calophyllum wallichianum have basically spherical, moderate-sized fruits with the walls of the stones usually less than 0.8 mm. thick, like those of C. incrassatum, while the southern populations (with the partial exception of those in Trengganu) have large, often ellipsoid fruits with stone walls at least 1.3 mm. thick. Indumentum studies confirmed this relationship: the northern populations and C. incrassa- tum have hairs that are slightly birefringent, with apical cells little longer than the others; the southern populations have highly birefringent hairs, with the apical two cells much enlarged and very thick walled (Ficure 18, a, b). Specimens referable to Calophyllum tahanense have been variously identi- fied as C. incrassatum, C. wallichianum, C. flavo-ramulum, or C. austrocoria- ceum. Calophyllum tahanense is separated from the first two species by the key in Henderson and Wyatt-Smith (op. cit.) only because it grows at generally higher elevations than do these species. In both indumentum and fruit type it is similar to the northern populations of C. wallichianum and to C. incrassatum. Hence, the complex must be reorganized taxonomically. The small-fruited form must bear the name tance wallichianum, because the type of that name came from such a specimen (see below). The erstwhile C. incrassatum can be aahacucied varietally from C. wallichianum sensu stricto mainly by its broader twigs and petioles; it also has a more southerly distribution in Malaya. The erstwhile C. tahanense is also reduced to varietal rank. It can be distinguished by its more coriaceous leaves, its midrib type, and possibly also by bark differences; it is basically a montane form (but see below). For the plants that formed the southern group of C. wallichianum, I have taken up Henderson and Wyatt-Smith’s manuscript name, C. rufi- gemmatum, C. wallichianum and C. rufigemmatum are readily distinguished by morphology, indumentum, and bark characters (TaBLeE 9). In the southern part of the Malay Peninsula (e.g., Renggam Forest Reserve, Johore) and on Singapore (e.g., Bukit Timah), the two species grow together. It is clear that much work still has to be done on this complex, both to ascertain the variation within Calophyllum wallichianum as here defined, especially in Sumatra and Borneo, and also to establish more clearly the relationships between C. rufigemmatum and its probable relatives, and C. lanigerum and C. ferrugineum and their relatives, since all taxa have similar, very distinctive hairs. Seedling type and germination, practically unknown in C. rufigemmatum and in C. lanigerum and its relatives, is potentially significant. Growth patterns may also be important (does C. rufigemmatum produce one or two pairs of leaves per flush, and C. wallichianum more?), as is, of course, a better knowledge of the basic morphology of many of the taxa involved. 1980] STEVENS, CALOPHYLLUM 349 TaBLe 9. Differences between Calophyllum rufigemmatum and C. wallichianum. C. rufigemmatum C. wallichianum LaTEXx Clear yellow, rarely Opaque white, often brown turning yellowish, rarely clear yellow INDUMENTUM Short-tomentose to Crustaceous to tomentose short-tomentose, rarely tomentose HAIR TYPE Two apical cells much Apical cells practically elongated, c thick-walled, elongated, but if so, birefringent not thick-walled Twics Often with obscure Obscure transverse transverse line at lines not seen nodes INFLORESCENCE AXIS LENGTH (cm.) 0.3-1.3 1.7-7 NUMBER OF FLOWERS 5 to7 7 to 15 (to 19) PEDICEL LENGTH (CM.) AT 0.35-0.7 0.5-2 ANTHESIS FRUIT LENGTH (cm.) (2.6-)3-4.4 2-3(-3.5) STONE WALL THICKNESS (1-)1.3-2 0.3-1.1 (mm.) Variation within Calophyllum wallichianum var. incrassatum The majority of specimens of Calophyllum wallichianum vat. incrassatum from Sumatra and Borneo have subconical, rather than plump, terminal buds (they are plump in Soepadmo 71); better-developed indumentum (it is even tomentose in bb 2971); a midrib with at least the edges raised, even at the very base; and large leaf blades (9.5-)15-32.5 by (2.8-)3.7-8 cm. In seedling type and general bark characters Stevens et al. 282, from Sarawak, was similar to plants seen in Malaya and Singapore, although the latex was pale yellow and opaque, rather than white (as might be expected, the latex of the twigs was white). Other specimens from the Sumatra-Borneo area are reported to have yellow (Soepadmo 71; S 16375), whitish (SAN 53316), or white (SAN 39310) latex. The fruits of this form are similar to those of Malayan specimens, although those of S 16375 are reported to be yellow. An interesting specimen from the western coast of Sumatra, bb 2971, has young, branched infructescences with tomentose indumentum; the branches have up to seven flowers and are up to 3.5 cm. long (see also below). Specimens of C. wallichianum vat. incrassatum from Sumatra and Borneo tend to have 350 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 the cells at the apex of the hairs notably larger than those toward the base (Ficure 18, g—k). The specimens of Calophyllum wallichianum var. incrassatum from Malac- ca, particularly Stone 9033, are superficially similar to Calophyllum lanigerum C. lanigerum. However, C. lanigerum has better-developed indumentum, the hairs having strongly birefringent, expanded apical cells; a plumper terminal bud; usually less dense and clearer venation; an inflorescence that is glabrous toward the apex; and bark with yellow latex. Two specimens of Calophyllum wallichianum var. incrassatum from G. Padang, Trengganu, which were collected at rather high altitudes for the species, are very robust and have a midrib approaching that of C. wallichianum var. tahanense. The fruits are spherical and sharply wrinkled and have an outer layer ca. 3 mm. thick and stone walls ca. 1 mm. thick. Henderson and Wyatt-Smith (op. cit.) cited SFN 31076 as C. tahanense, albeit with some hesitation; Whitmore determined FRI 12646 as C. coriaceum, which has similarly coriaceous leaves but is not otherwise comparable. The specimens from Selangor are included with some hesitation. One specimen, anon., 19 June 1951 (KEP), has slender petioles up to 3.5 cm. by up to 2.5 mm. FR/ 2524 (Trengganu, Malaya) has ovate leaf blades that are rather narrowly acute at the apex; the inflorescences are more robust than those of other Malayan specimens and are tomentose; the pedicels are only 7 mm. long. The record of Calophyllum wallichianum var. incrassatum from the Celebes also needs to be confirmed; bb 13751 is a sterile specimen. Typification of Calophyllum wallichianum Wallich dist. 4843, the type of Calophyllum wallichianum, was collected at Pinang and Singapore; thus, as seen from the discussion above, it might have been taken from the small- or the large-fruited element in C. wallichianum as previously delimited. However, as Henderson and Wyatt-Smith (Joc. cit.) noted, the type came from Pinang; the many-flowered inflorescences, as well as the hair type (Ficure 16, x, y), confirm this. When Planchon and Triana (loc. cit.) described Calophyllum wallichianum they noted that Wallich dist. 4843 was a mixed collection, including also C. spectabile (properly C. soulattri) and a small piece of C. polyanthum (which I have not seen under this number). There is yet another species under this number, represented by leaves and fruits of C. lanigerum var. austrocoriaceum; this element, at least, was probably collected in Singapore. Planchon noted on a sheet at Geneva that the fruits did not belong to ‘‘C. insigne,’’ and Planchon and Triana (loc. cit.) suggested that Choisy had confused ‘‘C. insigne’’ with C. wallichianum. I have not found the name “‘C. insigne’’ elsewhere, and it appears to be an error for C. spectabile (= C. soulattri). 44. aon ta rufigemmatum Henderson & Wyatt-Smith ex P. F. Stevens, sp. Ficure 17, b-h. 1980] STEVENS, CALOPHYLLUM 351 Ficure 17. a, Calophyllum sp. 54 (bb eee hee bud, x 3. b-h, ane tite b-e, . b-d, fruit, x : b, from outside; c, ‘fans: verse section; d, longitudinal section, e, eae bud, Fe ee nae RI 3921, —h, fruit, x 1: f, from outside; g, longitudinal section; , from apex. i, terminal seat x 3. j, Calophyllum sp. 53 (S 2 4675), gee bud, x . um ey et al. 133), terminal bud, x 0.75. I-n, C. wo , C. castan z ‘terminal bud (SAN 42949), x 3. m,n, fruit (SAN 16693), < 2: m, iy ee ection; n, from outside. 352 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 C. wallichianum auct., non Planchon & Triana; M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 303. pls. 1C, 3. 1956, pro parte; Maheshwari, Bull. Bot. Survey India 2: 146. 1960, pro parte; Kochummen, Malayan Forest Rec. ed. 2. 17: 216. 1965, pro parte; T. C. Whitmore, Tree Fl. Malaya 2: 193. 1973, pro parte; H. Keng, Gard. ae eee 28: 245. 1976; Corner, Gard. Bull. Singapore Suppl. 1: 105. A speciebus aliis Calophylli in gemma terminalis (0.8—)1.2-1.8 cm. longa indumento tomentoso praedita, lamina mediocra venatione densa subobscura, inflorescentia brevi, et fructu plerumque 3-4.4 cm. longo putamine parietibus 1.3-2 mm. in transversis, differt Tree 20-48 meters tall, d.b.h. to 240 cm.; trunk usually without buttresses or spurs; outer bark brown to gray, becoming deeply fissured, the inner surface blackish, brown and orange mottled, or dirty orange-brown; under bark red; inner bark red to pink or dull brown; latex clear, yellow, sticky, or light brown (Sumatra). Twigs flattened, 1.5-2.5(-3.5) mm. across, obscurely 4-angled, often with inconspicuous transverse raised line at nodes, drying blackish, subpersistently rufous-puberulent to -short-tomentose; axillary innovations lacking basal scars; internodes (0.5—)1-5(-6.5) cm. long; uppermost pair of axillary buds + pointed, to 3 mm. long, erect to + spreading; terminal bud plump to narrowly conical, (0.8-)1.2-1.8 cm. long, with rufous-puberulent to -tomentose indumen- tum (hairs, Ficure 18, k, 1), underdeveloped internode to 6 mm. long. Petiole (1-)1.2-2.7 cm. long, slender, concave above, convex below, subpersistently tomentose, drying black; lamina ovate to subelliptic, 7.3-14 by 2-5 gradually (sub)acuminate to retuse at apex, acute or cuneate to broadly rounded and often slightly asymmetrical at base, rather distantly undulate and not recurved to slightly so at margin, coriaceous, drying umber to sabelline above and umber to khaki below, subpersistently puberulent to short-tomentose on midrib on both surfaces, the midrib above narrow at base or narrowing fairly quickly near base, + raised (surrounding lamina obscurely raised), 0.25—0.35 mm. wide at midpoint, below raised, striate, the venation on both surfaces usually subobscure (rarely subapparent), raised, (11 to) 13 to 18 veins/5 mm., angle of divergence 70-80°. Inflorescences from foliate axils, with 5 (to 7) flowers, unbranched, the axis 0.3-1.3 cm. long, puberulent, ad bracts ovate, ca. 4 mm. long, often persistent at anthesis; pedicels 3.5-7 mm. long, to 12 mm. in fruit, short-tomentose. Flower (?)hermaphroditic; tepals 4 (to 8), the outer pair elliptic to subobovate, 5-7 by 2.2-4 mm., subtomentose on back, the next pair obovate, 8-8.5 by 3.5-4.5 mm., usually with strip of indumentum down back, additional ones, if any, to 9 by 3.1 mm.; stamens 140 to 250, the filaments to 4.5 mm. long, connate for up to 0.5 mm., the anthers oblong, 0.5-1.2 mm. long, + retuse at apex; ei, 1.2-2 mm. long, the style 2-2.2 mm. long, the stigma peltate, 0.5-0.7 m across, 3- or 4-radiate. Fruit ellipsoid (spherical or subobovoid), (2.6—)3 me 4 by 2.4-3.2 cm., + roun at apex, drying brown or grayish brown, with broad, longitudinal ridges and /or fine wrinkles; outer layer detaching cleanly 1980] STEVENS, CALOPHYLLUM 353 =! Ficure 18. Hairs (from terminal bud). a-j, Calophyllum wallichianum. a b, var. wallichianum (FRI 3105). c-h, var. incrassatum: c, d, bb 18354; e, f, FRI 3605; g, h, S 16375. i, j, cf. var. incrassatum (bb 24801). k l C. rufigemmatum: k, SFN 34642; 1, KEP 74159. m, C. aff. castaneum (S 16559). n-q, C. lanigerum. n, 0, var. austrocoriaceum: n, FRI 7846; 0, aba eed 5560. p, q, var. lanigerum: p, Kostermans 9591, hair to 600 pm. lo q, Kostermans & Anta 490, hair ca. 550 um. long. 1, C. castaneum (HLB 909, 68-72), hair ca. 540 pm. long. seen aie cells in k-q all strongly birefringent in polarized Hight. Scale = 120 wp 354 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 from stone, 1.3-3.5 mm. thick, compact, + striate on inner surface; stone ovoid to ellipsoid, 2.2-3 by 2-2.4 cm., apiculate or not, the walls (1-)1.3-2 mm. thick, smooth, often with triradiate marking at apex; spongy layer thin. Type: Singapore, Bukit Timah, ‘‘low’’ alt., 24 Dec. 1937, SFN 34632 coll. Ngadiman (holotype, a; isotypes, K, LAE, SING). DistriBuTION. Southeastern Malaya, Singapore, Sumatra (Map 17). SELECTED SPECIMENS SEEN (‘‘f’’ denotes specimens with fruits). Malaya. TRENGGANU: Ulu Dungun, FRI 9616 (KEP); Jerengau State Land, 285 m., KEP 76677 (£) (kK, KEP, L, sinc); Bauk F.R., 210 m., FRI 392] (f) (A, K, KEP, SAR, SING). PAHANG: G. Lesong F.R., Rompin, 15 m., KEP 80955 (f) (a, k, KEP, L, SING); Aur F.R., 15 m., KEP 75881 (f) (KEP); Mile 17, Jalan Pekan-Batu Balek, KEP 82702 (KEP); Baloh F.R., KEP 77974 (ker). Jonore: G. Panti, Panti F.R., 305 m., FRI 13796 (f) (a, K, KEP, SAN, SING); Renggam F.R.., KEP 71265 (kep); Jalan Endau, Mersing, 21 m., KEP 74159 (f) (a, BO, K, KEP, SING); Jemaluang, KEP 73457 (f) (kK, KEP); G. Arong F.R., KEP 77889 (f) (KEP); 15 m., Stevens et al. 80 (a); Kluang F.R., KEP 69919 (KEP); G, Pulai, SFN 7803 (f) (KEP); 6th Mile, Kota Tinggi- Mawai road, SFN 30987 (sinc); Mersing F.R., KEP 77814 (f) (KEP, sinc); Ulu Sedili F.R., KEP 63058 Timah, SFN 34546 (Bo, sar, sinc), Ridley 6333 (f) (sinc, w); Gardens Jungle, SEN 39451 (f) (A, K, KEP, SING); Mandai Forest, SFN 39605 (k, sING). Sumatra and adjacent islands. Riau: Afd. Lingga, P. Lingkep, Oetam Njakur (Bakong), 100 m., bb 2040 (xo, 1); bij Oetan djalan Setawar, bb 2705 (Bo); Sei Tokak, bb 3861 (L), Djago, 10 m., bb 3939 (f) (Bo); Tanjong Batang, 8 m., bb 4015 Map 17. Distribution of Calophyllum ee (solid circles), C. cas- taneum (stars), C. aff. castaneum (open circles), C. woodii (squares), and Calophyllum sp. 54 (triangles) in Malesia. Tasch Calophyllum sp. 53. 1980] STEVENS, CALOPHYLLUM 355 Bo); B. Poempoeng, bb 4047 (Bo); upper Riauw, Pakanbaru, Tenajan R., Soepadmo 222 (f) (A, BO, C, E, LAE, NY, SING). Ecotocy. An often large tree of ridges, slopes, or other well-drained habitats in mixed dipterocarp forest, 5-305 m. alt. Flowering February, September, November, and December (flower scented); fruiting January, March to August, and October (fruit pale or brownish green). SFN 33561, bb 3939, and Soepadmo 222 have neat holes bored in the stone wall and the seed eaten; these appear to be the exit holes made by some insect. GERMINATION AND YOUNG PLANT. The radicle breaks the stone to one side of the base (KEP 77989). The seedling has two pairs of leaves separated by an internode ca. 1.4 cm. long (one seedling, from Singapore, SFN 39451). Subsequent internodes are somewhat longer, the terminal bud is functional, and the plant is erect. LocaL usE. The wood is an important timber since the trees attain large dimensions. Calophyllum rufigemmatum can be recognized by its relatively slender twigs and petioles, and its ovate to subelliptic lamina that is obscurely and rather distantly undulate at the margin and has a more or less raised midrib, even at the base, and close venation. The inflorescences are short and congested, and the bracts are subpersistent; the short pedicels elongate markedly after anthesis. The fruits are distinctive; they are typically large, longitudinally corrugated as well as closely wrinkled, and with a stone wall usually at least 1.3 mm. thick (but see below). The epithet rufigemmatum (‘‘reddish brown buds’’) is quite appropriate. The circumscription of Calophyllum rufigemmatum is discussed under C. wallichianum, in which it had previously been included. The rather appropriate manuscript name that Henderson and Wyatt-Smith coined for this taxon when they thought it was specifically distinct is retained. Although Calophyllum rufigemmatum is generally distinguishable from C. wallichianum by numerous, albeit sometimes rather inconspicuous, characters (TaBLE 9), specimens from Trengganu blur this distinction somewhat in Malaya. Fruiting specimens from the Bukit Bauk Reserve have spherical, reticulate (but not longitudinally corrugated) fruits, with the outer layer and stone walls 1-1.5 mm. thick (Ficure 17, f-h). The single fruiting collection from Jerengau is intermediate between the specimens from Bukit Bauk and the typical form (see above). In other characters, these specimens agree well with C. rufi- gemmatum (see also the discussion after C. wallichianum and C. castaneum). I collected an interesting specimen (Stevens et al. 48) from the Renggam Forest Reserve, Johore, thinking it to be Calophyllum rufigemmatum although the latex was slightly cloudy. However, the dried specimen has a facies similar to that of C. lanigerum var. austrocoriaceum and also, but less mark- edly, that of C. molle. The inflorescences are those of C. rufigemmatum, but the stamens have hairy filaments and glabrous anthers. The hairs on the terminal bud are like those of C. rufigemmatum At first sight saplings of Calophyllum rufigemmatum from Johore look 356 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 very different from the adult trees. There is dense, tomentose indumentum on the terminal bud and the stem, as well as on the midrib on the lower surface of lamina; the lamina itself is suboblong and measures up to 30 by 9 cm. 45. Calophyllum lanigerum Miq. Fl. Ind. Bat. Suppl. 1(3): 498. 1961. Type: angka, prope Djebus, Teysmann, HB 3466 (holotype, u; isotypes, BO, K (?, 8.n.), L) Tree (or shrub) 2-21 meters tall, d.b.h. to 48 cm.; trunk usually without buttresses (present in montane forest in Johore); outer bark ochre to yellowish (grayish or light brown), with vertical rows of lenticels or with boat-shaped fissures, inner surface orange to dark brown; under bark deep reddish to greenish; inner bark deep red to pale pinkish; latex usually clear yellow, sticky (+ opaque, sulfur yellow, or milky). Twigs moderately to strongly flattened, 1.5-5 mm. across, + rounded to strongly 4-angled, drying dark brown to black (rarely yellowish), + persistently tomentose; axillary innovations lacking basal scars; internodes |—5(—7.5) cm long; uppermost pair of axillary buds rounded, to 3 mm. long, + spreading; terminal bud plump, 0.5-3 cm. long, with brown tomentose or grayish brown subtomentose indumentum (hairs, Figure 18, n-q), underdeveloped inter- node to 5 mm. long. Petiole 0.4-3 cm. long, broadly and shallowly (rarely narrowly) concave above, convex and persistently tomentose below; lamina subovate to oblong or oval, 3.8-20 by 1-6(—7.3) cm., rounded to retuse or subacute at apex, rounded to cuneate or acute at base, not undulate to distantly so and flat to slightly recurved at margin, coriaceous, drying chestnut to olivaceous (greenish) above, umber to olivaceous (greenish) below, subpersis- tently tomentose (soon glabrescent) on midrib on both surfaces, the midrib above gradually to fairly quickly narrowed at base, raised, (0.15—)0.2-0.35 mm. wide at midpoint, below raised, rounded to striate, the venation subob- scure to clear above and below (latex canals above as prominent as veins), 6 to 13 (to 15) veins/5 mm., angle of divergence (60-)70-85°. Inflorescences from foliate axils, with 3 to 21 flowers, unbranched (rarely with branches up to 2 cm. long and with 5 flowers), the axis 0.3-7 cm. long, tomentose at base, rest of axis subglabrous (rarely with indumentum its entire length), lowest internode (0.2-)0.4-2 cm. long; bracts to 10 by 2 mm., tomentose below, subpersistent or not; pedicels 0.4-2.3 cm. long, glabrous. Flower (?)hermaphroditic; tepals 8 (rarely 6), the outer two ovate, 3.5-7 by 2.5-5 mm., sometimes with dorsal tuft of hairs at apex, the next pair elliptic to obovate, 7.5-12 by 3.5-6.5 mm., the inner ones + lingulate to obovate, 7-12 by 2.5-5 mm.; stamens (65 to) 105 to 165, the filaments to 6 mm. long, connate for up to | mm., the anthers suboblong, 0.5-1 mm. long, rounded to slightly retuse at apex; ovary 1.4-2 mm. long, the style 3-3.5 mm. long, the stigma peltate, 0.6—-0.8 mm. across, 2- or 3-radiate. Fruit usually + spherical (ellipsoid to obovoid), 1.2-2.9 by 1.1-2.4 cm., usually apiculate, drying brown, sharply, closely, and shallowly wrinkled; outer layer detaching cleanly from stone, |-3.5 mm. thick, compact; stone + spherical, 1.1-2.1 cm. long and 1980] STEVENS, CALOPHYLLUM a5) across, rounded at apex, the walls 0.7-1.3 mm. thick, smooth (with obscure triradiate marking at apex), discolored area at base 3.5-8 mm. across; spongy layer thin. Key to the Varieties of Calophyllum lanigerum 1. Terminal bud 5-12(-16) mm. long; twigs 1.5—2.5(-3.5) mm. across; lamina 3.8-12;9@15)cmy long: «6c 2 a ke ee he me eS 45a. var. lanigerum. 1. Terminal bud 0.9-3 cm. long; twigs 3-5 mm. across; lamina 8.5-20 cm OID, Sarees tee 8.78 x we 5 eee Go OR oe os 45b. var. austrocoriaceum. 45a. Calophyllum lanigerum Miq. var. lanigerum C. lanigerum Miq.; Kurz, Natuurk. Tijdschr. Nederl.-Indie 27: 192. 1864; F. Mueller in Walp. Ann. Syst. Bot. 7: 357. 1868; Scheffer, Natuurk. Tijdschr. Nederl.-Indie 31: 354. 1870, ibid. 32: 405. 1873; Vesque in Monogr. Phanerog. 8: 606. 1893; Heyne, Nutt. Pl. Indonesie. ed. Bp 1: 1084. 1950. C. Mie Ridley, Kew Bull. 1938: 121. 1938; Masamune, Enu Phanerog. Born. 475. 1942. Type: Dutch SE. Borneo Glisanian’, Bite eae Motley 54 (holotype, k). Shrub or tree 2-12 meters tall. Terminal bud 5-12(-16) mm. long; twigs flattened, 1.5-2.5(-3.5) mm. across. Petiole 0.4-1.5(-1.9) cm. long; lamina oblong to oval, 3.8-12.5(-15) by 1.9-4.6(-7.3) cm., usually drying chestnut above, latex canals on upper surface not prominent, 6 to 11 veins/5 mm. Inflorescence with 3 to 7 (to 13) flowers; axis 0.3-2.5 cm. long, lowest internode (2—-)4-11 mm. long. Fruit + spherical, 1.2-1.9 cm. long; outer layer 1-2 mm. thick; stone to 1.6 cm. long, walls ca. 0.7 mm. thick. DistripuTION. Bangka, Belitung, and southeastern Borneo (Map 18). SELECTED SPECIMENS SEEN. Sumatra and adjacent islands. BancKka: Djebus, Teysmann, HB 3204 (Bo, L, MEL, u); Soengei Selan, Bukit Raja, 140 m., Buinnemeyer 2002 (Bo, L); Kepo bij Toboali, 60 m., Bunnemeijer 2315 (Bo, K, L); Lobok Besar, 20 m., Kostermans & Anta “45 (a, BO, K, L, SING); Pangkaljunang-Blinjoe, Huitema 35 (Bo); Serdang, de Leeuw 5 (Bo); Koela Soengei Redjein, Burger 28 (po); Gunong Padang, 100 m., Kostermans & Anta 1000 (a, BO, K, L, LAE, P, SING); Soengei Liat dessa Bakem, Teysmann 5.n. (BO). BELITUNG: Tandjong Pandan, Teysmann s.n. (Bo); Kampong Began- tang, de Veer s.n., May 1949 (Bo). Borneo and adjacent islands. KALIMANTAN. Timur: W. Kectes. Kelindjau R., near Melan, Kostermans 9591 (a, BO, kK, SING), bij Benoewa toewa, 20 m., Endert 1585 (a, Bo, kK). Selatan: G. Pamatton, Korthals s.n. (L); km. 22 Bangarmasin to Martapura, 10 m., Dransfield & Hamlali 4318 (po); Martapoera, G. Padamaran, bb 1211 (Bo), G. Koepang, bb 1212 (po). Java. Barat: cult. Hort. Bogor. sub VJ C 128 (Bo, k, us). Ecotocy. Locally abundant, rather small tree of sandy soils, sometimes in padangs; 20-140 m. alt. Flowering September, October, and December; fruiting June, September, and November. 358 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Map 18. Distribution of Calophyllum lanigerum var. lanigerum (circles), C. lanigerum var. austrocoriaceum (squares), C. dasypodum agian localized; star, unlocalized), and C. coriaceum (triangles) in Malesia GERMINATION AND YOUNG PLANTS. Saplings (e.g., Tevysmann, HB 3260) have leaves with very short (ca. 3 mm.) stalks and blades that are broadly rounded to subcordate at the base. LocaL uses. The wood is used in house and ship building. 45b. Calophyllum lanigerum Miq. var. austrocoriaceum (T. C. Whitmore) P. F. Stevens, comb. et stat. nov. C. austrocoriaceum T. C. Whitmore, Gard. Bull. Singapore 26: 269. 1973, Tree Fl. Malaya 2: 174. 1973; H. Keng, Gard. Bull. Singapore 28: 244 1976. Type: Malaya, Johore, summit Gunong Panti Barat, 1684 feet [513 m.], 6 May 1968, FRI 7846 coll. Cockburn (holotype, KEP; isotypes, A, SING). Calophyllum sp. 10, M. R. Henderson & Wyatt-Smith, Gard. Bull. Singa- pore 15: 313. pl. 8. 1966. 1980] STEVENS, CALOPHYLLUM 359 Tree (2.1-)4.5-21 meters tall. Terminal bud 0.9-3 cm. long; twigs rather strongly flattened, 3-5 mm. across. Petiole 1-3 cm. long; lamina narrowly ovate to oblong, 8.5-20 by 3.3-6.4 cm., usually drying umber to olivaceous above (often chestnut brown in Kalimantan), in Bornean specimens latex canals on upper surface often as prominent as veins, (5 to) 7 to 15 veins/5 mm. Inflorescences with 7 to 21 flowers; axis 3-7 cm. long, lowest internode 0.4-2 cm. long. Fruit spherical to ellipsoid or obovoid, (1.8-)2.2-2.9 by (1.7-)2-2.4 cm.; outer layer (1.8-)2.2-3.5 cm. thick; stone (1.1-)1.7-2.1 cm. long, walls 0.8-1.3 mm. thick. DistriBUTION. Southern Malaya to the Lingga Archipelago, northwestern Borneo (Map 18) SELECTED SPECIMENS SEEN. Malaya. Jonore: ridge between G. Panti E. and G. Panti W., 427 m., FRI 13819 (a, K, KEP, SING); G. Blumut, 914 m., FRI 7512 (A, KEP, SING); Kluang F.R., 457 m., KEP 98038 (a, K, KEP, SING); Mersing, Jemaluang F.R., KEP 71889 (k, KEP, sING). Singapore: MacRitchie Reservoir area, 30 m., Burkill 1877 (A, K, LAE, sINc); Gardens Jungle, ‘“‘low’’ alt., SFN Singkhep, Sei Manggoe, 15 m., bb 3936 (30); M. Tandai, Teysmann s.n. (so). Borneo. Sarawak. Ist Division: Bako Natl. Park, Lintang Path, 100 m., S 19718 (K, L, SAN, SAR, SING), Sungei Seloa, 60 m., S 7664 (SAN, SING); G. Matang, 732 m., S 12503 (Bo, - L, SAN, SAR, SING); near G. Siol, Omar 373 (k, sinc); Unjam F.R., 9 m., S 7709 (sar, SING); Batu 6 F.R. [Stapok], FA 827 coll. a 8 July 1940 en 3rd Division: Batang Igan, Sungei Tutus logging camp, S 30169 (k, SAR). KALIMANTAN. Barat: Soengei Kenepai, Hallier 2167 (so, k, L); Danau Lamadgian [Seriang], er PB 3463 (FI); Poeloe Madjang, Jaheri s.n., 1893 (Bo), Teysmann, HB 8042 (so); Pontianak, Dedi 864 (Bo); Pasir Tondjong, 2m., bb 9779 (Bo); N. of Mempawah, Pasir Pandjang, 1 m., Polak 724 (so, L). Ecotocy. Varied. Both in Johore, Malaya, and in the First Division, Sarawak, in colline forest—‘‘occasionally one of the tallest [trees] in the forest’’ (SFN 10723, G. Blumut, Johore); at lower elevations in peat swamps (S 30169, Sarawak); heath forest (e.g., S 12340, Sarawak); sandy soil near sea (Polak 724, Kalimantan); or general mixed dipterocarp lowland rainforest (Johore, Singapore); to 950 m. alt. Flowering July, October, and November (Borneo); fruiting January to May (Borneo and Malaya), July (Singapore), and August (Sarawak). GERMINATION AND YOUNG PLANT. The radicle breaks the stone wall immediately adjacent to the ‘‘plug,’’ which is apparently not pushed out during germina- tion. The seedling has two pairs of leaves separated by an internode of ca. 5 mm. Subsequent growth seems to be slow, with internodes less than 1 cm. long (SFN 39452). The young plant is erect, and the terminal bud is func- tional (pers. obs., Singapore). The leaves of saplings are long and narrow (11.8-50 by 1.7-5.4 cm Loca usE. The wood is coarse grained and is used for planks. The epithet austrocoriaceum (‘‘southern coriaceous’’) was coined because 360 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Calophyllum austrocoriaceum was thought to be related to C. coriaceum and was found to the south of it. Calophyllum lanigerum can be recognized by its plump, usually densely tomentose terminal bud; by its coriaceous, more or less oblong lamina, which has clear, not very dense venation on at least one surface; and by its sharply wrinkled, more or less spherical fruit with a thick outer layer that detaches cleanly from the stone. The spherical, thick-walled stone often has an obscure triradiate marking at the apex and a pluglike area at the base. The epithet lanigerum (‘‘wool-bearing’’) emphasizes the well-developed tomentose indu- mentum characteristic of this species. The description of the bark of Calophyllum lanigerum is almost wholly based on jason of (and field notes on specimens of) C. lanigerum var. austrocoriaceum. Both varieties have yellow latex. e type specimens a Calophyllum lanigerum and C. austrocoriaceum are very different and at first sight appear to belong to different species. However, they are connected by intermediates, although the evaluation of the variation pattern is complicated since specimens in flower are not known from Malaya or Singapore. Specimens of C. austrocoriaceum from high altitudes in Malaya have less dense venation and less well developed indumen- tum than the specimens from Borneo, and the lamina is more or less acute, rather than rounded, at the base; specimens of C. austrocoriaceum from Borneo, and those collected at lower altitudes in Malaya and Singapore, are more like C. Janigerum in these respects. Specimens of C. austrocoriaceum from northwestern Borneo (e.g., 503, Sarawak) have fruits the same size as those of C. lanigerum, and there is also general agreement in details of flower, fruit structure, and indumentum distribution on the inflorescence (dense, tomentose indumentum restricted to the lowest internode). However, the inflorescences of specimens of C. austrocoriaceum from Singapore have indumentum for their entire length; there is perhaps a tendency toward a similar condition in Dedi 864 (C. austrocoriaceum; Pontianak, Kalimantan). Infructescences of specimens of C. austrocoriaceum from Johore are glabrous of the range of C. austrocoriaceum are needed to understand this variation. Hair structure of specimens from all areas and altitudes is similar (Ficure 18, n-q): the base of the hair consists of often thin-walled short cells, with the walls not birefringent in polarized light; the upper part of the hair has elongated cells (often two) with strongly thickened walls that are birefringent under polarized light. The upper surface of the lamina of several specimens of C. austrocoriaceum from Kalimantan dries chestnut brown, as in C. lanigerum, rather than the lighter brown color characteristic of the other specimens of C. austrocoriaceum In view of the underlying similarity between Calophyllum lanigerum and C. austrocoriaceum and the noncorrelation of much of the variation within the complex as a whole, C. austrocoriaceum is reduced to varietal rank under C. lanigerum. Calophyllum lanigerum var. austrocoriaceum can be distinguished from var. lanigerum by the larger size of all its parts (except 1980] STEVENS, CALOPHYLLUM 361 the pedicels and the flowers) and by the frequently lighter brown color of the dried lamina. There is little variation within Calophyllum lanigerum var. lanigerum. Specimens from mainland Borneo tend to have larger leaves than those from Bangka and Belitung; the type specimen of C. frutescens is a specimen of the former type. Some of the variation within var. austrocoriaceum has been discussed above. A few specimens from Malaya (Grubb s.n., 25 Sept. 1970; Shah & Ahmad 2923) have twigs that dry yellowish and leaf blades that dry greenish, but in other respects these specimens are unremarkable. Specimens of Calophyllum lanigerum var. austrocoriaceum from low alti- tudes in Johore and Singapore were cited under C. incrassatum by Henderson & Wyatt-Smith (op. cit., p. 202), albeit with hesitation. However, they have the densely tomentose terminal bud, the distinctive hairs, the clear and rather distant venation, and the yellow latex of the former taxon. 46. Calophyllum castaneum P. F. Stevens, sp. nov. Ficure 17, k. A speciebus aliis Calophylli in gemma terminali magna 2.1-3 cm. longa indumento tomentoso castaneo vel badio praedita, folia magna basi cucullata venulis lateralibus densis 11 ad 22 per 5 mm., et infructescentibus ramosis pedicellis brevibus 2-3.5 mm. longis, differt. Tree 27-38 meters tall, d.b.h. to 58 cm.; trunk without buttresses; outer bark at first yellow-brown, smooth except for long cracks, becoming brown, fissured, scaly, when freshly exposed dark brown, the inner surface orange- brown; under bark dark brown, orange-brown, or reddish; inner bark red to pale rose; latex yellow, opaque, viscous or not. Twigs flattened, 3.5-5.5 mm. across, more or less 4-angled, drying yellowish or grayish brown, color often obscured by dense indumentum, persistently tomentose, the hairs bay to chestnut, to 1 mm. long; axillary innovation (?)without basal scars; internodes 4-6 cm. long; uppermost pair of axillary buds rounded, ca. 2 mm. long, more or less spreading; terminal bud plump, 2.1-3 cm. long, with bay to chestnut, tomentose indumentum (hairs, Ficure 18, r), underdeveloped internode 3-8 mm. long. Petiole 1-2 cm. long, shallowly concave above, convex below, persistently tomentose; lamina elliptic to oblong, (6-)10-25 by (3.5-)4.3-9.5 cm., short-acuminate at apex, rounded to cuneate and more or less cucullate at base, slightly undulate and slightly and narrowly recurved at margin, marginal thickening ca. 0.35 mm. wide, coriaceous, drying bay to umber above and umber below, eomentose on midrib on both surfaces and over entire lower surface, the midrib above narrowing rather abruptly near base, raised, or narrowing gradually, flat at first and becoming raised toward apex, 0.25-0.4 mm. wide at midpoint, below raised, prominent, strongly striate, the venation subapparent on both surfaces, slightly raised, 11 to 22 veins/5 mm., angle of divergence 70-75°. Infructes- cences from foliate axils, with scars of ca. 15 flowers, branched, the branches 3-flowered, to 7.5 mm. long, the axis ca. 2.5 cm. long, sparsely tomentose, especially toward base, lowest internode 3-6 mm. long; bracts not known; pedicels 2-3.5 mm. long, + glabrous. Flower not known. Immature fruit 362 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 ca. 1.5 by | cm., subacute at apex, drying vinaceous-brown, wrinkled; outer layer ca. | mm. thick; stone and spongy layer unknown. Type: Sarawak, 3rd Division, Kapit, Sungei Mengiong, 1500 feet [457 m.], 7 Oct. 1969, S 29755 coll. Suib (holotype, sar). DistrisuTION. Northwestern Borneo (Map 17). ADDITIONAL SPECIMENS SEEN. Borneo. SARAWAK. Ist Division: Semengoh Ar- boretum (Tree 4671), ca. 50 m., Stevens et al. 133 (a); Semengoh F.R., 30 ., Stevens et al. 291 (a). KALManran, Barat: Damenar Tangkawang, de Vries (27) HLB 909,68-60 & -72 (both L); Sambas, Perigi Limoes, 300 m., bb 7044 (so). Sine loco [probably Pontianak or Sintang], de Vries & Teijsmann s.n. HLB 903,343-214, -216, & -217 (all L); Teijsmann s.n. HLB 903,343-215 (L), de Vries s.n. (BO, U). Ecotocy. Lowland and colline mixed dipterocarp forest, 30-457 m. alt. Fruiting in October. Calophyllum castaneum is a distinctive species, even when sterile. It is recognizable by its large leaf blades that have dense venation and are cucullate at the base; its large terminal bud at least 2 cm. long; and its persistent, dark brown, tomentose indumentum on terminal bud, twig, and leaf. The very short pedicels are unusual in a species of Calophyllum that has fruits a moderate size and are a noteworthy feature of C. castaneum. The descriptions of the infructescence and the immature fruit are taken from the type specimen, which is a unicate. The chestnut color of the indumen- tum suggested the specific epithet, castan Calophyllum cucullatum, from the Philippine Islands, also has a large leaf blade that is cucullate at the base and dries a rich brown color. However, C. cucullatum has a terminal bud less than 8 mm. long, very different hairs, a much smaller inflorescence, long and slender pedicels, and a fruit only ca. 1 cm. long. Calophyllum cucullatum is not closely related to C. castaneum. There is a perplexing group of specimens from Sarawak (e.g., S 16599, 3rd Division), Brunei (BRUN 3058), and adjacent Sabah (e.g., SAN 36872, Beaufort) (Map 17) that in many ways resembles Calophyllum castaneum. The specimens have castaneous indumentum with hairs similar to those of C. castaneum in that the two apical cells are much enlarged and are birefringent; however, the cells are not as much enlarged as they are in that species (Ficure 18, m). The leaves have similarly dense venation, and the lamina base is rounded and slightly cucullate. The pedicels seem to be short, and the fruit moderate in size. However, the specimens are much more gracile in appearance, the twigs are less than 3 mm. across, and the terminal bud is less than 1.5 cm. long. Although these specimens are possibly related to C. castaneum (and perhaps to C. rufigemmatum), their identification awaits better collections. Meijer (Bot. Bull. Herb. Forest Dept. Sabah 7: 15. 1967) used the name C. rotundatum for these specimens; this is a nomen nudum. 47. Calophyllum coriaceum Symington ex M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 323. pi. 10. 1956; Wyatt-Smith, Malayan 1980] STEVENS, CALOPHYLLUM 363 Forest Rec. 17: 113. 1952, nomen; Kochummen, Malayan Forest Rec. ed. 2. 17: 216. 1965; T. C. Whitmore, Tree Fl. Malaya 2: 177. 1973, pro parte. Type: Malaya, Pahang, Cameron’s Highlands, Robinson’s Falls, 12 July 1928, KEP 13627 coll. Mead (holotype, sinc; isotype, KEP). C. ?spectabile auct., non Willd.; Henderson, Jour. Malay Branch Roy. Asiatic Soc. 5: 243. 1927. C. near griffithii auct., non King; Henderson, Gard. Bull. Straits Settl. 4: 93. 1927. Tree 4-30 meters tall, d.b.h. to 58 cm.; trunk without buttresses; outer bark yellowish, slightly fissured; under bark deep red-brown; inner bark pink; latex white, sticky or rather fluid, or yellow, thick (smelling like soup stock—KEP 51818). Twigs slightly flattened, 3-4.5 mm. across, + 4-angled, drying brown to blackish, reddish farinose to subtomentose when young; axillary innovations lacking basal scars; internodes 2-5 cm. long; uppermost pair of axillary buds + rounded, to 1.5 mm. long, suberect; terminal bud plump, 1-1.8(-2.7) cm. long, with brown, subcrustaceous to puberulo-tomentose indumentum (hairs, Ficure 19, a—c), underdeveloped internode to 4 mm. long. Petiole 1-3.1 cm long, broadly and shallowly concave above, convex below, glabrous when mature; lamina elliptic to oblong, 8-18.5 by 4.6-8.3 cm., broadly rounded to shallowly retuse and finally short-acute to obtuse at apex, rounded and finally short-acute to acute or cuneate at base, not undulate and slightly to strongly recurved at margin, marginal thickening to 0.8 mm. wide, very coriaceous, drying olivaceous-honey to umber above and honey to umber below, puberulent on midrib on both surfaces when young, glabrescent or not, the midrib above narrowing gradually from or rather quickly near base, + level at first, becoming raised, (0.35-)0.5-0.7 mm. wide at midpoint, center 70-80°. Inflorescences terminal and/or from adjacent foliate axils, with many flowers, branched, the axis at least 6 cm. long, farinose-puberulent, lowest internode ca. 2 cm. long; bracts unknown; pedicels 0.7-2.2 cm. long, puberu- lent, in fruit to 2 mm. thick. Flower (?)hermaphroditic; tepals 8 Ae also 9), the outer pair elliptic-ovate, ca. 10 by 6.5 mm., + puberulent on back, the next pair broadly elliptic, ca. 14 by 10.5 mm., ae crete toward base on back, the inner ones probably ca. 14 by 17 mm.; stamens ca. 370, the filaments ca. 7 mm. long, the anthers oblong, 0.9-1.3 mm. long, he t apex; ovary ca. 2.5 mm. long, the stigma peltate, ca. 1 mm. across, 3-radiate. Fruit ellipsoid, 3-4.5 by 2-2.3 cm., subcuneate to apiculate at apex, drying vinaceous- fe ike with deep, founded wrinkles; outer layer detaching cleanly from stone, 1.5-3 mm. thick, compact except for air spaces developing under skin; ee ellipsoid, 2.4-2.7 by 1.7-1.8 cm., rounded at apex, the walls 1-1.3 mm. thick, smooth, unmarked; spongy layer thick at first. DistriBuTION. Malay Peninsula, the Main Range (Map 18). 364 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 SELECTED SPECIMENS SEEN. Malaya. SELANGor: G. Nuang F.R., Ulu Langat, KEP 51818 (go, KEP, SING). PAHANG: Fraser’s Hill, 1219 m., KEP 87303 (a, BO, K, KEP, SING); Cameron’s Highlands, 1402 m., SFN 23653 (A, BO, K, KEP, SING); Sungei Parang, Jalan Kuda, KEP 27192 (xP): Berenchang, KEP 34015 (K, KEP); trail to G. Jesar, 1463 m., FRI 15459 (KEP, SAN, SAR); Sungei Insar, KEP 25174 (kep). Eco.ocy. Montane rain forest, 1219-1463 m. alt. Flowering in July (flower scented); fruiting May and November (fruit green, eaten by birds (KEP 51818 Calophyllum coriaceum can be recognized by its rather large, very coriaceous lamina with more or less broadly recurved margins, profuse, terminal inflores- cence, and ellipsoid fruit that has a stone with walls at least | mm. thick. The very coriaceous leaf blades suggested the epithet coriaceum Calophyllum coriaceum does not seem particularly closely related to any other species. It is vegetatively similar to C. scriblitifolium; this is discussed further under the latter species. Whitmore (1973) believed that C. austrocoria- ceum was felated to C. coriaceum. Calophyllum austrocoriaceum, however, has axillary inflorescences and hairs of a very different structure; it is here reduced to varietal rank under C. lanigerum. Whitmore (loc. cit.) reported Calophyllum coriaceum from Gunong Padang, Trengganu. The specimens on which this report was based (e.g., FRI 12646) are referred here to C. wallichianum var. incrassatum, although their very coriaceous leaf blades are somewhat atypical. The inflorescences are axillary, and the subspherical fruits are closely wrinkled—both characters of C. wallichianum var. incrassatum. 48. caer dasypodum Miq. Fl. Indiae Batavae 1(2): 511. 1859, nomen m for C. lanceolatum Teijsm. inn.; F. Mueller in Walp. Ann. Sat Bot. 7: 357. 1868; Backer & Bakh. f. Fl. Java 1: 386. 1963, excl. syn.; C. teysmannii Zoll. ex Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 275. 1862, non Mig. 1861, excl. descr. et spec. cit., nomen novum for C. lanceolatum Teijsm. & Binn.; C. lanceolatum Teijsm. & Binn. Nat. Tijdschr. Nederl.-Indié 4: 398. 1853, non Blume, 1825; Kreyenb. Bull. Soc. Imp. Nat. Moscou 27: 259. 1854; Teijsm. & Binn. Nederl. Kruidk. Arch. 3: 395. 1855; F. Mueller in Walp. Ann. Syst. Bot. 7: 357. 1868. Type: Java, Duizend gebergte, van Gesker s.n. (L, U). C. hasskarlii Teijsm. & Binn. ex Planchon & Triana, Ann. Sci. Nat. Bot. V. 15: 273. 1862; Vesque in C. DC. Monogr. Phanerog. 8: 581. 1893, Ficure 19, Hairs a terminal bud, unless otherwise mentioned). a-—c, Calophyllum coriaceum: a, b, Stevens et al. 17; c, KEP 78303, persistent ee only. d-g, C. dasypodum: d, e, Kostermans 6882; f, g, Marche B 371. C. canum: h, S 23960; i, A 859; j, Clemens 31859; k, KEP 99519; I. SAN &3502; m, » Kostermans ‘10369: n, 0, Soepadmo 39; p, S 23968, from axillary bud. 4 , Calophyllum sp. 53 (S 24675). Scale = 60 pm. (in q, r, scale = 120 ny 366 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 pro parte; Koord. & Valeton,. Meded. s’Lands Plant. 61(Bijd. we, Java 9): 384. 1903; Koord.-Schum. Syst. Verzeich. 1(Fam. 187): 1912; Koord. Exkursionsfl. Java 2: 618. 1912; Heyne, Nutt. Pl. ees ed. 3. 1: 1082. 1950. Tyee: — in sylvis, Zollinger 3457, pro parte (holotype, G; isotypes, BM, G, C. pulcherrimum auct., non Wail. ex Choisy; Choisy in Zoll. Syst. Verzeich. 2: 149. 1854, Pl. Javan. 9. 1858. C. tomentosum auct., non Wight; Mig. Pl. Jungh. 291. 1854, Fl. nerd Batavae 1(2): 514. 1859; Planchon & Triana, Ann. Sci. Nat. Bot 15: 294. 1862 Tree 20-30 meters tall, d.b.h. to 50 cm.; trunk without buttresses; outer bark yellow-brown with short, boat-shaped fissures, or dark gray, deeply fissured, the inner surface orange-straw; under bark dark red; inner bark pale pinkish; latex clear yellow, sticky. Twigs slightly flattened, 1.8-3 mm. across, strongly 4-angled, soon becoming subterete, drying blackish brown, subpersistently grayish to brownish puberu- lent to subtomentose; axillary innovations lacking basal scars; internodes (0.2-)1-2.5 cm. long; uppermost pair of axillary buds plump, 2-5 mm. long, suberect; terminal bud plump, 6-8 mm. long, with brown, puberulent indumen- tum (hairs, Ficure 19, d-g), underdeveloped internode 1-4 mm. long. Petiole 0.7-1.1 cm. long, concave above, convex and + persistently puberulent below; lamina ovate to elliptic, 5.5-12.5 by 1.5-3.4 cm., acute to acuminate at apex, cuneate to acute at base, strongly and distantly undulate and flat to slightly recurved at margin, coriaceous, drying umber and shiny above, umber to sepia below, glabrescent or persistently puberulent on midrib below (also above), the midrib above gradually narrowed from base, margins usually somewhat raised but + sulcate at first, 0.2-0.8 mm. across at midpoint, becoming raised in top 1/3, below raised, striate, the venation + apparent on both surfaces, slightly raised, 8 to 15 veins/5 mm., angle of divergence 60-75°. Inflorescences from foliate axils along stem, with 7 to 11 eaten unbranched, the axis 3-5.5 cm. long, puberulent, lowest internode 1-5 m long; bracts + ovate, ca. 2.5 mm. long; pedicels 0.41.3 cm. long, Stier eae Flower (?)hermaphroditic; tepals 4 or 6, or 8 in terminal flowers, sometimes glabrous, the two outer + ovate, ca. 4.5 by 3 mm., the inner ones cel larger; stamens 30 to 55, to 70 in terminal flowers, the filaments to 2 m long, the anthers suboblong, 0.6-0.9 mm. long, slightly retuse at apex; ovary ca. | mm. long, the style ca. 1.5 mm. long, the stigma peltate, ca. 0.5 m across, slightly lobed. Fruit spherical, 1.4-1.7 cm. long and across, nied at apex, drying vinaceous-brown, smooth; outer layer detaching cleanly from stone, 2-3 mm. thick, compact; stone spherical, |.1-1.5 by 1-1.4 cm., rounded at apex, the walls 0.5-0.6 mm. thick, smooth, unmarked or with obscure triradiate apical lines, with a lighter-colored area (?plug) up to 5 mm. across at base; spongy layer thin. DistRIBUTION. Southern Sumatra, western Java, southern Borneo, (?)Phil- ippine Islands (Map 18). SELECTED SPECIMENS SEEN. Sumatra. SELATAN: Lematang Oeloe, 150 m., 1980] STEVENS, CALOPHYLLUM 367 Lambach 1258 (Bo, v (s.n.)); Sepoetik, Soewikis, 25 m., bb 2851 (a, BO). Borneo and adjacent islands. Kacimantan. Timur: Sebatik Is., 10 m., Koster- mans 9154 (a, BO, K, NY, P, SING); Nunukan Is., N. part, Kostermans 8958 (po, L); Lor Haur, W. of Samarinda, 40 m . deastemans 6882 (A, B, BO, AE, NY, P, SING). Selatan: Pleihari, “Kintap, 25 m., bb 2748 (Bo); (?)Doesson, Korthals s.n. (Bo, L, uv). Philippine Islands. CALaMIAN. Busuanga: Malbato, Marche 371 (7) (a, P). Java. Barat: Djasinga, Tjurug, Meijer 2894 (Bo); Salak, Zollinger 1711 ( (‘‘fragm. ex herb. P’’)); Preanger, Soekaboemi, Palaboenratoe, Koorders 2890 (Bo); Bogor, Palace Gardens, Kostermans s.n., Aug. 1953 (A, BO, BRI, K, NY, P, SING). Ecotocy. Sandy soil (Kalimantan), 10-150 m. alt. Flowering March, August, and October (flower scented); fruiting March, May, and September. Calophyllum dasypodum can be recognized by its plump terminal bud; its rather stout twigs with puberulent-subtomentose indumentum; its rather narrow, ovate-elliptic lamina with a deeply undulate margin, a depressed midrib, and a shiny upper surface when dry; its puberulent inflorescence axis; and its spherical, cherry-sized fruits. The epithet dasypodum means “‘hairy foot’”’ and alludes to the conspicuous indumentum on the inflorescence. The only specimen of Calophyllum dasypodum from the Philippine Islands, Marche 381, is sterile. It has dried a color similar to that of the other specimens, but its leaf blades are dull, and although the indumentum is subtomentose, the hairs are typical for the species (Ficure 19, f, g). Marche 381 may also approach C. pisiferum. Van Gesker s.n. agrees quite well with the description of Calophyllum lanceolatum Teijsm. & Binn. and is possibly the specimen on which the original description was based. The way in which the name Calophyllum teysmannii Zoll. ex Planchon & Triana was published makes it, perhaps inadvertently, a new, but superflu- ous, name for C. lanceolatum Teijsm. & Binn. Planchon and Triana placed C. lanceolatum in the synonymy of their C. teysmannii, and in the discussion after the species noted that it was almost certainly a synonym of C. teysmannii, However, both their description and the specimens that they cited are of another taxon, the Javanese form of C. calaba. The name C. teysmannii Zoll. ex Planchon & Triana was published shortly after C. teysmannii Miq.; for further discussion, see C. teysmannii. The confusion between Calophyllum dasypodum and the Javanese form of C. calaba, which resulted in Backer and Bakhuisen’s (Joc. cit.) inclu- sion of the two in one species, apparently also occurs in C. hasskarlii, a synonym of C. dasypodum. In many herbaria Zollinger 3457, the type number of C. hasskarlii, is a mixed collection, with part being the Javanese form of C. calaba (see Vesque, op. cit., p. 582); the two species are not at all closely related. Calophyllum calaba in Malesia lacks the well-developed indumentum of C. dasypodum, its lamina usually has much more dense venation, and its twigs are thinner and are often strongly flattened. The fruits of C. calaba are spherical to ellipsoid and are about two thirds the length of those of 368 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 C. dasypodum, the thinner outer layer becomes disorganized by air spaces, and the stone walls are about a third as thick. 49. se scriblitifolium M.R. Henderson & Wyatt-Smith, Gard. Bull. Singapo : 326. pl. 12. 1956; J. Anderson, Gard. Bull. Singapore 20: “134. 1963: Smythies, Common Sarawak Trees, 64. 1965; Kochum- men, Malayan Forest Rec. ed. 2. 17: 217. 1965; T. C. Whitmore, Tree Fl. Malaya 2: 192. 1973; J. Anderson, Trees Peat Swamp Sarawak, 88. pl. 27D. 1973. Type: Malaya, Perak, Utan Melitang Forest Reserve, 12 Oct. 1949, KEP 69001 coll. Shaari (holotype, KEP). C. wallichianum auct., non Planchon & Triana; Merr. Bibl. Enum. Bornean Pl. 394. 1921, pro par C. revolutum Wyatt- Smith, Malayan Forest Rec. 17: 43. 1952. Nomen. Tree 24-44 meters tall, d.b.h. to 96 cm.; trunk without buttresses or stilt roots, but knee roots sometimes present (Sarawak); outer bark reddish brown or brown and soft, flaky, to gray, fissured and scaling, to chocolate, the inner surface (bronze) brown; under bark orange-brown; inner bark red to pink; latex white to creamy white, slightly sticky to sticky, turning gray-cream on exposure Twigs (slightly) flattened, 3.5-6.5 mm. across, + 4- or 8-angled when young, drying blackish to dark brown (rarely grayish), farinose to puberulent at least when young; axillary innovations perhaps with scars within 5 mm. o base; internodes (0.3-)0.7-3 cm. long; uppermost pair of axillary buds ana ounded, to 1.5 mm. long, spreading; terminal bud plump, 1.2-2.2(-2.9) m. long, with grayish to brown, crustaceous to subtomentose indumentum ee Ficure 20, d, e), underdeveloped internode to 4.5 mm. long. Petiole 1.3-2.8 cm. long, broadly concave above, convex below, + persistently puberulent; lamina oblong to elliptic, 7.8-19 by 1.8-7.6 cm., rounded to retuse (mucronate near midrib) at apex, cuneate to acute at base, slightly undulate but strongly and broadly recurved at margin, very coriaceous, drying olivaceous to honey color above and umber-sepia to fulvous below, usually persistently puberulent on midrib on both surfaces, the midrib above abruptly to gradually narrowed at base, depressed at first, becoming + raised, 0.3-0.5 mm. wide at midpoint, below strongly raised, rounded to striate, the venation apparent to subobscure on both surfaces, 9 to 16 veins/5 mm., angle of divergence (60-)80-85°. Inflorescences from foliate axils along twigs, with 9 to 11 flowers, unbranched, the axis 3.5-5.5 cm. long, densely puberulent, lowest internode Ficure 20. Hairs (from terminal bud, unless otherwise mentioned). a-c, Calophyllum woodii: a, Stevens et al. 160; b, SAN 35504; c, bb 34418. d, e, h, C. scriblitifolium: d, e, S 16170, from stem (d, from above); h, Native collector 1276. f, g, . lowii: f, Grashoff 709, common hair type of species, , van Rossum 44. i, C. glaucescens (Haviland 2058). j-l, C. Cee ales 413 k, SAN 52956; 1, s 29914. m, C. depressinervosum (KEP 98808). complanatum (Stevens et al. 407). o-q, Calophyllum sp. 54 (bb 19334), gre from stem. Scale = 60 pm. (in p, q, scale = 120 pm.). 370 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 1.8-2.8 cm. long; bracts unknown; pedicels 0.6-1.2 cm. long, densely puberu- lent. Flower (?)hermaphroditic; tepals 4, the outer pair ovate, 6-7 by 5.5-6 mm., puberulent on back, the inner pair obovate, ca. 12.5 by 6 mm., puberulent in strip down back; stamens ca. 200 plus, the filaments to 6 mm. long, the anthers oblong, 0.9-1.5 mm. long, retuse at apex; ovary ca. 2.5 mm. long, style and stigma unknown. Fruit ellipsoid, ca. 3.4 by 2 cm., rounded at apex, drying blackish, smooth or with inconspicuous longitudinal wrinkles; outer layer detaching cleanly from stone, 1.3-1.5 mm. thick, compact; stone ellipsoid, ca. 2.7 by 1.6 cm., rounded at apex, the walls ca. 0.5 mm. thick, smooth, with faint reticulations but otherwise unmarked; spongy layer un- known Distripution. Malay Peninsula, Sumatra, and Sarawak (Map 16). ong Pueh F.R., S 6419 (sar); Buntal, anon., March 1912 (sa os Bs Division: Loba Kobang South Protected Forest, S 2740 (KEP, SAR); Naman F.R., Sungei Assan, 6 m., S 7361/10 (sar, sinc). Sine loco, native vlleaay 1276 (?) (A, BO, P, UC). Ecoxocy. Rather large tree of peat swamps, rarely also in kerangas forest in Sarawak; below 10 m. alt. Flowering and fruiting in October. GERMINATION AND YOUNG PLANT. The radicle probably breaks through the stone to one side of the base. The seedling has but a single pair of leaves. The first flush of growth generally consists of two pairs of leaves: first an internode 5(-10) mm. long, then a pair of scale leaves, then an internode ca. 3 cm. long, and finally a pair of fully expanded leaves (13 of the 15 seedlings seen). Scale leaves are rarely absent (2 of the 15 seedlings seen). The terminal bud is functional, and the young plant is erect (Stevens et al. 147A). Calophyllum scriblitifolium is best recognized by its coriaceous leaf blades with their strongly and broadly revolute margins; its axillary, grayish puberulent inflorescences; its flowers, which have four tepals, the outer pair of which is grayish puberulent on the back; and its ellipsoid, almost smooth- -drying fruit that is rounded at the apex and the base. Fresh fruits of Calophyllum scriblitifolium are about 5.7 by 2.2 cm., with the outer layer ca. 5 mm. thick, to judge from fallen fruits seen in Sarawak. The leaves, with their strongly recurved margins, seem to have suggested the specific epithet (scriblitifolium means ‘‘tartlike leaves’’). alophyllum scriblitifolium is perhaps most closely related to C. canum, which at least in Borneo frequently grows in swamps, and which also has white latex, flowers with four tepals, and a similar distribution of indumentum on inflorescence axis and flower. However, the vegetative and fruit characters 1980] STEVENS, CALOPHYLLUM aval given above readily separate the two, and the hairs of the two differ in structure (Ficure 20, d, e; cf. 19, h-p). Calophyllum scriblitifolium is vegeta- tively similar to C. coriaceum, another species with white latex. However, C. coriaceum has fruits that are pointed-ellipsoid and dry sharply wrinkled, with stone walls at least 1 mm. thick; the inflorescences are terminal. Calophyllum coriaceum is a tree of montane forest, and C. scriblitifolium one of peat swamps; the two are not closely related. More collections of Calophyllum scriblitifolium are needed. The variation in hair type in Sarawak is not understood (Ficure 20, h). 50. Calophyllum canum Hooker f. Fl. Brit. India 1: 271. 1874; King, Jour. Asiatic Soc. Bengal, II. 59: 177. 1890; Vesque in C. DC. Monogr. Phanerog. 8: 573. 1893; Curtis, Jour. Straits Branch Roy. Asiatic Soc. 25: 78. 1894; Ridley, Fl. Malay Penin. 1: 185. 1922, pro parte; I. H. Burkill & M. R. Henderson, Gard. Bull. Straits Settl. 3: 347. 1925; Heyne, Nutt. Pl. Indonesié. ed. 3. 1: 1082. 1950; M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 306. 1956; J. Anderson, Gard. Bull. Singapore 20: 153. 1963; Pukol & Ashton, Checklist Brunei Trees, 93. 1964; Smythies, Common Sarawak Trees, 59. 1965; Kochum- men, Malayan Forest Rec. ed. 2. 17: 219. 1965; Burkill, Dict. Econ. Prod. Malay Penin. ed. 2. 1: 412. 1966; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 7: 16. 1967, pro parte; T. C. Whitmore, Tree Fl. Malaya 2: 176. 1973; Corner, Gard. Bull. Singapore Suppl. 1: 104. 1978. Type: Malaya, Malacca, 26 March 1865/1866, Maingay 1645 (Kew dist. 175) (holotype, k). C. borneense Vesque, Epharmosis 2: tt. 28, 29. 1889, in C. DC. Monogr. Phanerog. 8: 580. 1893; Merr. Bibl. Enum. Born. Pl. 393. 1921; Masamune, Enum. Phanerog. Born. 475. 1942; Keith, N. Borneo Forest Rec. ed. 2. 2: 313. 1952. Type: Borneo Cttibae) eae Beccari, PB 2101 (holotype, Pp; isotypes, a (frag.), FI, C. spectabile auct., non Willd.; Ciris ied “Straits Branch Roy. Asiatic C. retusum auct., non Choisy; J. Anderson, Gard. Bull. Singapore 20: 154. Fe quoad syn. C. lowii auct., non Hooker f.; J. Anderson, Trees Peat Swamp Sarawak, SD. 1973, excl. ill. Tree 7.5—36 meters tall, d.b.h. to 77 cm.; trunk without buttresses or spurs; outer bark yellowish or light brown to almost black, with rather close lines of lenticels, or finely cracked, scaling or not, hoop marked, the inner surface dirty brown to orange-brown; under bark reddish to orange, or blackish and red mottled; inner bark pale reddish; latex white, not sticky, in Malaya also reported to be clear yellow-brown and sticky, in Borneo opaque to clear yellow, sticky or becoming sticky. Twigs slightly to strongly flattened, 2-3(-4.5) mm. across, obscurely to strongly 4-angled, drying blackish to dark brown, grayish to brownish farinose- puberulent when young; axillary innovations lacking basal scars; internodes 1-9 cm. long; uppermost pair of axillary buds rounded, to | mm. long, spreading ate JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 or pointed, ca. 3(-8) mm. long, erect; terminal bud plump to narrowly conical, (0.3-)1-2.2 cm. long, with short, grayish brown to rufous, compact indumentum (hairs, Ficure 19, h-p), underdeveloped internode 1|.5-5(—-11) mm. long. Petiole (3.3-)9-22 by (1.2-)3.5-7.5 cm., acuminate (rarely acute) at apex, rounded to acute at base, strongly to moderately undulate and slightly recurved at margin, coriaceous, drying brown-vinaceous to sabelline above and umber to fulvous below, often nitid, entire lamina subconcave or not, glabrous to transiently farinose to puberulent on midrib above and especially below, the midrib above narrowing gradually from base, depressed, margins raised, becoming + raised, or narrowed abruptly near base (Borneo only), depressed at first, soon becoming raised, (0.15-)0.25-0.6 mm. wide at midpoint, below raised, + striate, the venation above and below + apparent, raised, 10 to 18 (to 21) veins/5 mm., angle of divergence 70-80°. Inflorescences from foliate axils usually near ends of twigs (rarely terminal—eastern Borneo), with 7 to numerous flowers, usually unbranched (or flowers flabellately arranged, branches to | cm. long, with up to 5 flowers), the axis 3-7(—12.5) cm. long, grayish- to rufous-puberulent, lowest internode (0.5—)1-3.5 (-4.8) cm. long; bracts + ovate, to 1.2 by 0.6 cm., deciduous; pedicels 0.5-1.5 cm. long, puberulent. Flower (?)hermaphroditic; tepals 4 (rarely 5 to 8), the outer pair elliptic to broadly ovate, (4.5-)5-6.5 by (2.5-)3.5-5 mm., puberulent on back, the inner ones elliptic to ovate, 5.5-8.5 by 3-5(-6) mm., usually puberulent in strip down back (rarely entirely puberulent or puberulent only near base); stamens (180 to) 350 to 600 (to 900 plus), the filaments to 3.5 mm. long, the anthers elliptic to oblong, 0.4-1 mm. long, slightly retuse at apex (very rarely with hairs (Borneo)); ovary 1.2-1.5 mm. long, the style (1.5—)2.8-4.5 mm. long, the stigma peltate, 0.5—0.7 mm. across, 2- to 4-radiate. Fruit spherical to ovoid, 1.5-2.2 by 1.4-1.8 cm., usually stoutly and sharply apiculate, drying brown, rather closely and sharply wrinkled; outer layer not detaching cleanly from stone, 0.6-1.4 mm. thick, compact; stone + spherical, 1.2-1.7 by 1.1-1.5 cm., rounded at apex, the walls 0.3-0.4(-0.8) mm. thick, smooth, unmarked; spongy layer (?)thin. DistripuTIoN. The typical form in Malaya, Sumatra, and northwestern Borneo; the variant only in Borneo (Map 19). SELECTED SPECIMENS SEEN. (Specimens from Borneo that closely match the type specimen of C. borneense denoted by ‘‘b’’; specimens that are + typical C. canum denoted by ‘‘c.’’) Malaya. Kepau: Gunong Boungsu F.R., 610 m., FRI 6956 (kK, KEP, SING); Bukit Enggang F.R., KEP 81345 (kep). PINANG: Government Hill, 305 m., Curtis 1543 (k, sinc); road to the Spout, 240 m., Curtis = 1153 (sinc); Waterfall Garden, 30 m., SFN 3695 (sinc). Perak: Larut, G. Boobo Range, 240-457 m., King’s collector 7704 (c, uc); Larut, less than 30 m., King’s collector 5420 (£, FI, P, SING); Batu Gaja, Wells s.n., Feb. 1918 (kep); Pondok Tanjong F.R., KEP 63467 (ker); Tualang F.R., KEP 63269 (KEP). SELANGOR: Weld Hill F.R., 60 m., KEP 99519 (a, CANB, K, KEP, LAE, SAN, SAR, SING); F.R.I. [Forest Research Institute], Kepong, KEP 99629 (A, KEP, SAN, SAR, SING); Ginting Simpah ridge, 914 m., KEP 70994 (xkeEpP). 1980] STEVENS, CALOPHYLLUM 373 Map 19. Distribution of Calophyllum canum in Malesia. Necri SEMBILAN: Sungei Menyala F.R., anon. (KEP); Senawang F.R., KEP 609 (BO, K, KEP, SING). MALacca: Batang Malaka, Goodenough 1791 (SING). Kuantan, Pekan Road, KEP 43188 (xep); Bukit Setongkol, KEP 883 (KEP, sinc); Pekan, Haviland s.n., April 1890 (sinc); Ulu Peroh, KEP 11213 (KeEp); Bentong, Kemesul F.R., 45 m., KEP 78718 (ker); Temerloh F.R., KEP 5464 (KEP, SING); Balok F.R., KEP 97919 (ker, sinc). JoHore: Sungei Sedili, SFN 36891 (A, KEP, SING); Ma’ Okil F.R., KEP 71284 (Kep); Kluang F.R., KEP m., KEP 80557 (kep). Sumatra and adjacent islands. Riau: Lingga Arch., Pulau Singkep, Koeloe Laboeh, 9 m., bb 5370 (Bo); Pakanbaru, Tenajan R., low alt., Soepadmo 39 (a, BO, C, E, KEP, L, LAE, SING). SELATAN: Rawas, 8 4 i . AK. Ist Division: Lundu, Hewitt 47 (c) (sar); Kuching, Haviland 27 (c) (sinc); Semengoh F.R., 20 m., Stevens et al. 290 (c) (A); Sabal Tapang, 105 m., Stevens et al. 167 (b) (a); G. Matang, 290 m., Stevens et al. 257 (b) (a); G. Santubong, 54 m., Stevens et al. 295 (b) (a); Stapok F.R., 5 m., Stevens et al. 154 (c) (A). 3rd Division: Lepah P.F., Anderson 10/2 (c) (sar); Batang Igan, Sungei Tutus logging camp, S 30255 (c) (sar). 4th Division: Sungei Kelaput, ca. 5 m. S. of Pulau Bruit, S 28 coll. Anderson, 13 Nov. 1952 (c) (KEP, SAR); Sambas Road near Poak, C. Hose s.n., 7 Dec. 1911 (c) (BM). Brunei: Anduki F.R., sea cia (c) (KEP); Temburong, Labu F.R., 15 m SAN 17433 (c) (A, Bo, SAN, SAR, SING). SABAH. Sipitang: Sipitang, KEP 80046 (c) (KEP). pee Mesapol F.R., SAN 43268 (c) (kK, L, SAN); 374 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 1.6 km. NE. of Beaufort township, 105 m., SAN 15056 (A, BO, BRI, K, KEP, L, SAN, SING); Beaufort Hill, San 58588 (san). Ranau: Kinabalu Natl. Park, 457 m., SAN 54034 (k, L, SAN); Penibukan, 1219 m., Clemens 31859 (a, L, ny). Labuk & Sugut: Mile 87!/2, Telupid Road, Hap Seng logging area, 120 m., SAN 83502 (a). Sandakan: Sibuga, Kebun China, SAN 37454 (a, K, KEP, L, SAN, SAR, SING); Sepilok F.R., A 859 (a, Bo, K, KEP, L, SING); Sungei Mayan, Om., Stevens et al. 375 (A). KALIMANTAN. Timur: E. Kutei, Belajan R. near a Bleh, 50 m., Kostermans 10369 (b) (A, BO, CANB, K, KEP, L, LAE, P, nc); W. Kotei, Long Bleh, 30 m., bb 16065 (c) (A, BO, L); Keloem mpang, bb 16927 (Bo, L); Boven omer Liroeng Poedoeng, 50 m., bb 20620 (?) (A, Bo). Tengah: Koeala Kapeoas, Goehoeng, bb 2171 (?c) (Bo, L); Sampit, Karoeng, 10 m., bb 9924 (Bo); Tamoelian, bb 2477 (Bo). EcoLocy. Well-drained mixed dipterocarp forest in Malaya, the variant in Borneo in similar habitats; typical form in Borneo and Sumatra usually in peat swamps; to 1220 m. alt. Flowering April to June, September, and December (advanced bud in January); flower scented. Fruiting May, August, and October (submature fruit December and January); fruit greenish. YOouNG PLANT. The young plant is erect, and the terminal bud is functional (Bornean form). Loca uses. It is used in tuba fishing (species of Derris Lour. probably the poison) in Sarawak. The wood is good for masts and spars (Malaya, Sabah). The typical form of Calophyllum canum can be recognized, even when sterile, by its plump terminal bud and its distinctive leaf blades, which usually dry shiny and brown-vinaceous on the upper surface and have a strongly undulate margin; the midrib is more or less depressed and narrows gradually from the base. The flowers of the typical form usually have four tepals, and there are usually very numerous (350 plus) stamens. The dried fruit or ‘‘gray’’; it is very appropriate since the indumentum often makes the whole inflorescence look grayish. The delimitation of Calophyllum canum has been somewhat troublesome. The species is most closely related to C. lowii; the differences separating the two are discussed under the latter. The typical form of C. canum shows relatively little variation, although in Borneo it seems to prefer peat swamps and in Malaya it usually grows in better-drained ground. However, although of flower and fruit, they differ somewhat vegetatively. These Bornean specimens have a leaf blade that usually dries fulvous above, the margin 54). More surprising variation occurs in three specimens from western 1980] STEVENS, CALOPHYLLUM 375 Sabah that have hairs on the anthers (SAN 15056 (a few only), SAN 54034, Clemens 31859). The single flower of Clemens 31859 examined has ca. 180 stamens, the terminal bud is only ca. 0.3 mm. long, and the hairs are much branched (Ficure 19, j); the two flowers of SAN 54034 dissected have 220 and over 450 stamens, respectively, and the hairs are also strongly branched. There is no indication of a relationship between these specimens and C molle, C. gracillimum, or C. rubiginosum, the other species with hairy anthers. Collections such as S$ 23913, 23960, and 25043 growing in colline forest in the 3rd Division of Sarawak have the facies of C. canum, but somewhat different hairs (Ficure 19, h). They are probably to be placed in C. wallichian- um, the variation of which in Borneo is poorly understood. The specimen from which Calophyllum borneense was described is similar to the Bornean variant of C. canum, although its leaf blade has dried very flat, the inflorescence is usually only five-flowered, and the flowers have four to eight tepals. Further collections referable to C. borneense sensu stricto were made in Sarawak (see especially Stevens et al. 257). In field characters these specimens were very similar to the form of C. canum growing near Sandakan, Sabah, and the two have ovoid fruits and similar flowers. However, in Stevens et al. 257 (C. borneense, Gunong Matang, Sarawak) the submature fruits are ca. 1.9 by 1.5 cm., with the outer layer ca. 2.5 mm. thick and the stone walls ca. 0.8 mm. thick. In Stevens et al. 375 (near Sandakan, Sabah; submature fruits at about the same stage of development) the fruits are ca. 3.2 by 2.3 cm., the outer layer being ca. 5.5 mm. and the stone walls only ca. 0.4 mm. (all measurements from material preserved in alcohol). Although further studies of the whole C. canum complex in Borneo are needed, C. borneense seems best reduced to synonymy under C. canum. Calophyllum canum is close to C. lowii but can be distinguished by the characters given in TaBLeE 10. 51. Calophyllum lowii Hooker f. Trans. Linn. Soc. 33: 163. 1860; Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 271. 1862 (‘‘C. lowei’’); F. Mueller in Walp. Ann. Syst. Bot. 7: 357. 1868; Vesque in C. DC. Monogr. Phanerog. 8: 604. 1893; Merr., Bibl. Enum. Born. Pl. 393. 1921; Masamune, Enum. Phanerog. Born. 476. 1942. Type: in ora septentrionali insulae Borneo [?Sarawak], Low s.n. (holotype, k; isotypes, K, P (frag.)). C. palustre Ridley, Kew Bull. 1938: 121. 1938; Masamune, Enum. Phanerog. B . 1942. Type: Dutch SE. Borneo [Kalimantan], Bangarmassing, Motley 1129 (holotype, k). C. wallichianum auct., non Planchon & Triana; Merr. Bibl. Enum. Born. Pl. 394. 1921, pro parte; Heyne, Nutt. Pl. Indonesié. ed. 3. 1: 1086. 0. Tree to 20 meters tall, d.b.h. to 50 cm.; trunk and bark unknown; latex yellow, clear. Twigs flattened, 3-5.5 mm. across, + 4-angled when young, or with rounded edges, drying blackish, shortly gray-brown-—puberulent when young; axillary 376 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 TaBLe 10. Some differences between Calophyllum canum and C. lowii. C. canum C. lowti Twics LAMINA Bracts INFLORESCENCES STAMENS Fruit STONE WALLS Hairs 2-3(-4.5) mm. across Often nitid above, 10 to 18 (to 21) veins /5 mm., margin strongly undulate Conspicuous in young inflorescence, to 1.2 ‘cm. long Rarely branched or flabellate; flowers often rather congested (180 to) 350 plus, the anthers 0.4—1 mm. long Apex sharply pointed; surface densely and sharply wrinkled 0.3-0.4(-0.8) mm. thick Apical cells rarely 3-5.5 mm. across Often + dull above, 10 to 15 veins /5 mm., margin somewhat undulate Inconspicuous Usually branched, flabellate; flowers not congested Fewer than 210 (to 310), the anthers 1-2 mm. long Apex rounded; surface at most shallowly and distantly striate Less than 0.2 mm. thick Apical cells thin-walled thin-walled innovations lacking basal scars; internodes 2-7.5 cm. long; uppermost pair of axillary buds pointed, to 3 mm. long, spreading; terminal bud plump, (0.8-)1-2.2 cm. long, with grayish brown, crustaceous indumentum (hairs, Ficure 20, f, g), underdeveloped internode 0.4-1.5 cm. long. Petiole 1.4—3 cm. long, shallowly and broadly concave above, convex below, drying blackish, glabrous when mature; lamina narrowly ovate to elliptic, 11-25 by 1.5—8.5 cm., or ovate, ca. 18 by 7.2 cm., gradually acuminate at apex, acute at base, distantly undulate and narrowly or not recurved at margin, coriaceous, drying olivaceous above and umber to hazel below, puberulent on midrib on both surfaces, or glabrous, the midrib gradually narrowed from base, depressed at first with edges + raised, becoming raised at ca. !/3 length of leaf, drying blackish, 0.2-0.5 mm. across at midpoint, raised and + striate below, the venation on both surfaces + apparent (subobscure), raised, 8 to 15 veins/5 mm., angle of divergence 70-80°. Inflorescences from upper foliate axils (very rarely terminal—bb 2635), with 11 to numerous flowers, often lowest flowers flabellately arranged, usually with branches up to 5 cm. long and with 7 flowers, the axis 5-15 cm. long, puberulent, lowest internode 1.5-7.5 cm. long; bracts ovate, to 6.5 mm. long, soon deciduous; pedicels 0.9-2.5 mm. long, puberulent, rather slender, incrassate in fruit, to 2.5 mm. across. Flower (?)hermaphroditic; tepals 4 (to 7), the outer two 1980] STEVENS, CALOPHYLLUM art ovate to broadly elliptic, 6-6.5 by 4.5 mm., puberulent on back, the inner ones suboblong to broadly elliptic, 9.5-10.5 by 4.5-8.5 mm. (rarely suborbicu- lar, ca. 9 mm. long and across), puberulent in strip toward base of back; stamens 130 to 210 (to 310), the filaments to 5 mm. long, the anthers oblong, 1-2 mm. long, retuse at apex; ovary ca. 2 mm. long, the style ca. 3 mm. long, the stigma peltate, ca. 1.2 mm. across, 3-radiate. Fruit spherical to ellipsoid, 2-2.5 by 1.8-2 cm., rounded at apex, or apiculate, wee brown, finely striate; outer layer not detaching cleanly from stone, 0.9-1 . thick, compact, rather hard; stone + ellipsoid, 1.8-2.3 by 1.6-1.8 cm., ain at apex, the walls less than 0.2 mm. thick, smooth, unmarked; spongy layer thin. DisTRIBUTION. Sumatra to Borneo (Map 16). SELECTED SPECIMENS SEEN. Sumatra and adjacent islands. Riau: Indragirische bovenlanden, Kwala Belilas, 60 m., bb 27643 (Bo, L), Pagaroembei (Tjenako- Rivier), 8 m., bb 26108 (Bo, L, sinc), Belimbing, 6 m., bb 28464 (Bo, L, Mo), Keritang, 5 m., 5b 28691 (Bo, ny). Dyambi: Djambi, Gergai, 5 m., bb 11328 (Bo). SELATAN: Bangoeasin, Koeboestreken, 20 m., Grashoff 709 (Bo, L); Palembang, Grashoff 907A (L). Bancka: Lobok Besar, 5 m., bb 34046 (a, Bo, L); Bikang, bb 15095 (Bo); B[e]linjoe, Grashoff 40 (Bo, L). BELITUNG: Tandjong Pandan, Bantan, 30 m., bb 4/06 (Bo). Borneo. Sarawak. Ist Division: near G. Siol, Omar ie (K, SING); near Kuching, Haviland & Hose 3343 (BM, BO, K, L, SAR, UC). KALIMANTAN. Selatan: ond. afd. Pleihari, Pematang Landjar, bb 2129 (Bo, ty Tengah: Sampit, bij Seramau, bb 653 (Bo), Mandawei, 10 m., bb 2635 (Bo); ond. afd. Beneden Djak, Sei. Teroesan, bb 209! (Bo, , P. Kladan, 5 m., bb 12251 (Bo); Koeala Kapoeas, Sungei Mangkoetoep, Dachlan 2197 (Bo), P. Telo, bb 2174 (po); Beneden Merlan, Sungei Kemdeinan- gan, I m., bb 14417 (Bo). EcoLocy. Lowland rainforest, frequently periodically inundated, below 60 m. alt. Flowering October to December and March; fruiting in February. LocaL uses. The wood is used in house building. The latex boiled with coconut milk is used as a remedy for itching and skin infection (Heyne, loc. cit.). Calophyllum lowii is apparently related to C. canum; for the differences distinguishing the two see TasLe 10. Calophyllum lowii is larger in all parts (but has fewer stamens) than C. canum; its fruit is rounded at the apex, dries almost smooth, and has a thin-walled stone; its pedicel is notably incrassate in fruit; and its leaf does not often dry the brown-vinaceous color common in C. canum. The epithet commemorates H. Low, the collector of the type specimen. A number of sterile specimens cited under C. lowii (e.g., those from the Indragiri a (Sumatra)) are included here only provisionally. Planchon and Triana’s description of Calophyllum lowei (sic) appears to be independent of that of Hooker, published the preceding year. Planchon and Triana cited the specimens that they saw as ‘‘Low in herb. Hook. [the holotype of C. lowii Hooker f.] et inde in herb. Planch.’’; fruiting material 378 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 that they thought was another species of Calophyllum is referable to C. soulattri. The type specimen of Calophyllum palustre is a good match with that of C. lowii; both have profuse inflorescences and large leaf blades with a rather broad, blackish drying midrib. 52. Calophyllum woodii P. F. Stevens, sp. nov. Ficure 17, I-n. C. molle auct., non King; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 7: 15. 1967. A speciebus aliis Calophylli in indumento tomentoso gemmis, ramulis, costis laminarum, axibus inflorescentiarum, pedicellis et tepalis exterioribus praedito, ramulis in siccitate saepe pallide coloratis, lamina elliptica vel oblonga 9-22.5 cm. longa venulis lateralibus (6 ad) 8 ad 16 (ad 20) per 5 mm., et fructu subsphaerico strato exteriore 2.5-3 mm. crasso e putaminé haud munde secedenti et putamine parietibus 0.4—-0.6(-0.9) mm. crassis, differt. Tree 21-30 meters tall, d.b.h. to 80 cm.; trunk without buttresses; outer bark yellowish brown to brown, shallowly fissured or with lines of lenticels, becoming strongly fissured, the inner surface orange-red or brown-orange; under bark dark red or greenish straw; inner bark brown-red or pale red; latex brownish, opaque or pellucid, not viscous (rarely yellow (Kostermans 6618) or viscous (Stevens et al. 16 Twigs strongly flattened, 2.3-3.5 mm. across, rounded to + 4-angled, drying pale brown or yellowish to grayish, subpersistently brown-tomentose, hairs to 0.8 mm. long, puberulent; axillary innovations lacking basal scars; internodes 1-4 cm. long; uppermost pair of axillary buds rounded, 1-2. mm. long, + spreading; terminal bud plump to subconical, 0.6—1.2(-1.7) cm. long, with brown, tomentose indumentum (hairs, Ficure 20, a—c), underdevel- oped internode to 3(—6) mm. long. Fenole - 1.9 cm. long, concave to V-shaped above, convex below, with sul lamina elliptic to oblong, 9-22.5 by 3.4-6.3(-8) cm., gubrounded to acuminate at apex, broadly cuneate to subrounded (slightly cucullate) at base, broadly undulate and slightly recurved at margin, coriaceous, drying sepia to umber above and below, often shiny, usually tomentose on midrib on both surfaces, the midrib above abruptly to gradually narrowed at base, raised, 0.25—0.35 mm. wide at midpoint, below raised, slightly striate, the venation subobscure to apparent above and apparent below, raised, (6 to) 8 to 16 (to 20) veins/5 mm., angle of divergence 70—80°. Inflorescences from foliate axils, with 7 to 11 flowers, unbranched, short-tomentose, the axis 2-5.5 cm. long, lowest internode 0.4-1.7 cm. long; bracts ovate, to 5 mm. long, tomentose below, deciduous; pedicels 0.7-2.3 cm. long, slender, short-tomentose. Flower (?)hermaphroditic; tepals 4 (rarely 5), the outer pair ovate, ca. 5.5 by 4.5 mm., tomentose on back, the inner pair subelliptic, ca. 8.5 by 5.5 mm., with strip of tomentum down back; stamens 190 to 270, the filaments to 5.5 mm. long, the anthers elliptic, 0.8-1 mm. long, rounded at apex; ovary ca. 2.5 mm. long, the style ca. 3.5 mm. long, the stigma peltate, ca. 1.4 mm. across, 3-radiate. Fruit 1980] STEVENS CALOPHYLLUM 379 subspherical to ellipsoid, 1.3-2 by 1.2-1.7 cm., rounded at apex, drying brown, wrinkled; outer layer generally detaching cleanly from stone, 2.5-3.3 mm thick, compact, but with air spaces developing under skin; stone ellipsoid to subspherical, 0.7-1.3 by 0.65-1.1 cm., rounded to mucronulate at apex, the walls 0.4-0.6(-0.9) mm. thick, smooth, unmarked; spongy layer initially thick. Type: North Borneo, Sandakan, mile 8, Labuk road, outside Sibuga F.R.., 1 Sept. 1964, SAN 42949 coll. Burgess (holotype, L; isotypes, K, SAR). DisTRiBUTION. Borneo, mostly in the eastern part (Map 17). ADDITIONAL SPECIMENS SEEN. Borneo. SARAWAK. Ist Division: Sabal Tapang, 140 m., Stevens et al. 160 (a, sar); Semengoh F.R., 30 m.; Stevens et al. 124 (a). 3rd Division: Bukit Raya, Btg. Rejang, Ashton 6004 (a). SABAH. Sipitang: Ulu Mendalong, 10 km. SSE. of Malaman, 533 m., SAN 16725 (A, BO, BRI, oe L); Ulu oe 13 km. SSE. of Malaman, 838 m., SAN 16693 (A, BO, KEP, L, SAN, SING). Labuk & Sugut: Mile 42, Labuk Road, SAN 71708 (SAN). Seen ’ Mile 8 ee (tree originating from Sepilok), SAN 35504 (kK, KEP, L, SAN, SAR,-SING); Sepilok F.R., 30 m., SAN 19728 (KEP, SAR), Stevens et all. 331 (A), 335 (a); Plantation boundary, Gum Gum, mile 17, 12 m., SAN 81224 (san); Arboretum, mile 14, SAN 72843 (san), 82500 (a). Tawau: ca. 5 km. from Brantian on Tawau road, 45 m., Stevens et al. 434 (a). KALIMANTAN. Timur: Salembatoe, 100 m., bd 11293 (BO). Samarinda: Loa Djanan, W. of Samarinda, Kostermans 6618 (A, BO, L, P, Mahakam R., Tandjong Bangko, 20 m., Kostermans 7229 (Bo, L); Balikpapan, Sungei Wain, 20 m., bb 34418 (Bo, 1); W. Koetai, Moejoer, 25 m., bb 16764 (A, BO), Kahalu, 25 m., bb 28354 (pisH, Bo, P). Selatan: Pleihari, bb 2033 (BO). Ecotocy. Well-drained lowland to colline mixed dipterocarp forest, some- times in swamps; 30-838 m. alt. Flowering April and September (flower smells like syringa (SAN 35504)); fruiting June, September, October, and December (fruit greenish to pale cream). GERMINATION AND YOUNG PLANT. Young plants ca. 30 cm. tall are erect and have functional terminal buds. Calophyllum woodii can be recognized by the tomentose indumentum on twigs, terminal buds, midrib of the lamina, inflorescence, pedicels, and tepals; the hairs appear to be rough when viewed under high magnification since almost every cell has a small protrusion. The lamina is elliptic to oblong, with fine, moderately dense, and rather widely ascending venation. The inflorescences are axillary and unbranched, and the flowers usually have four tepals. The fruit is generally subspherical and dries wrinkled; the outer layer is 2.5-3.3 mm. thick, and the stone usually has walls less than 0.6 mm. thick. The specific epithet commemorates G. H. S. Wood, who made some of the earliest collections of this species in Sabah. Calophyllum molle has been confused with C. woodii, but in the former the hairs are smooth, the stamens are more numerous, and the anthers, 380 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 ovary, and fruit are hairy. Calophyllum woodiiis closely related to Calophyllum sp. 53; the differences between the two are discussed under the latter. The specimens of Calophyllum woodii from Sarawak and adjacent western Sabah have a narrower terminal bud than those from eastern Sabah, and the base of the lamina is not cucullate to slightly so. However, all specimens agree in general facies and in hair type. Stevens et al. 160 (Sarawak) had sticky latex, whereas in other specimens seen in both Sarawak and Sabah the latex was free flowing, and the fruits had stone walls about 0.9 mm. thick—somewhat thicker than those of other specimens. 53. Calophyllum sp. Ficure 17, j. Tree to 25 meters tall, d.b.h. to 80 cm.; trunk without buttresses or spurs; outer bark yellowish brown, with lines of close-set lenticels, not clearly hoop marked, the inner surface dull yellow-gray or greenish slate and orange-brown mottled; under bark mid-brown or dark reddish; inner bark red; latex clear yellow, sticky, becoming cloudy. Twigs flattened, 3.3-4 mm. across, obscurely 2- or 4-angled, soon becoming rounded, drying yellowish brown or grayish brown, subpersistently brown- tomentose, hairs to 1 mm. long; axillary innovations lacking basal scars; internodes 1.5-6(-10) cm. long; uppermost pair of axillary buds rounded, 1-2(-3.5) mm. long, + spreading; terminal bud obtusely conical, 6.5-9 mm. long, brown-tomentose (hairs, Ficure 19, q, r), underdeveloped internode absent. Petiole 0.8-1.2 cm. long, broadly and deeply concave above, convex and tomentose below; lamina elliptic to oblong, 12.5-20 by 4.6-6.8 cm., broadly rounded and finally mucronate or slightly retuse at apex, broadly rounded and slightly cucullate at base, slightly and distantly undulate and recurved at margin, coriaceous, drying khaki above and sabelline below, persistently tomentose on midrib below, the midrib above narrowing quickly at or near base, raised, center + sulcate, 0.3-0.4 mm. wide at midpoint, below strongly raised, striate, the venation + apparent above and apparent below, raised, 11 to 13 veins/5 mm., angle of divergence 75—85°. Inflorescences terminal and from upper foliate axils, with ca. 15 flowers, flabellate, with 3-flowered branches to 1.2 cm. long, the axis 2.8-5.2 cm. long, densely tomentose, basal internode 1.3-2 cm. long; bracts unknown; pedicels 1-1.5 cm. long, tomentose. Flower (?)hermaphroditic; tepals (?)4, the outer pair ovate-elliptic, ca. 6 by 5 mm., tomentose on back, the inner pair + obovate, ca. 10 by 5 mm., with strip of tomentum down back; stamens (?)numerous, the filaments ca. 5 mm. long, the anthers elliptic-obovate, ca. 0.5 mm. long, strongly retuse at apex; ovary ca. 2 mm. long, style and stigma unknown. Immature fruit spherical, ca. 8 mm. long and across, rounded at apex, drying brownish, smooth; outer layer ca. 1.8 mm. thick. DistripuTion. Northwestern Borneo (Map 17). SPECIMENS SEEN. Borneo. Sarawak. Ist Division: Semengoh Arboretum, Stevens et al. 129 (a); Semengoh F.R., 30 m., Stevens et al. 289 (a), S 24675 (A, L, SAR, SING). 1980] STEVENS, CALOPHYLLUM 381 Eco.tocy. Lowland mixed dipterocarp forest; ca. 30 m. alt. GERMINATION AND YOUNG PLANT. The seedling has two pairs of leaves separated by an internode less than 7 mm. long. Subsequently produced internodes are longer, the terminal bud is functional, and the plant is erect (Stevens eta Calophyllum sp. 53 is a distinctive taxon recognizable by its oblong lamina, which is broadly rounded both toward the apex and at the more or less cucullate base, and which dries a khaki color on the upper surface; the venation is fine but distinct. The inflorescence is apparently at least sometimes terminal, and there is tomentose indumentum on the back of the outer pair of tepals. The short branches of the hairs (Figure 19, q, r) are visible at a magnification of x 30; the hairs appear rough. Although the leaves of Calophyllum sp. 53 dry in a fashion similar to those of C. woodii, and both species have rough hairs, the latter has smaller leaf blades that are more or less acute at the apex, an inflorescence that is not (?ever) terminal, and a plump terminal bud. However, the two taxa are probably more closely related to one another than to other species of Calophyllum. The leaf blades of the young plant are obovate in shape, acuminate at the apex, and narrowly cuneate at the base 54. Calophyllum sp. Figure 17, a. ?Tree; trunk and bark unknown. Twigs slightly flattened, ca. 5 mm. across, rounded or with 4 raised lines, drying brown to blackish, persistently tomentose; axillary innovations lacking basal scars; internodes |-4 cm. long; uppermost pair of axillary buds rounded, ca. 0.7 mm. long, spreading, inconspicuous; terminal bud plump, ca. 1.5 cm. long, tomentose (hairs, Ficure 20, o-q), underdeveloped internode absent. Petiole 1.6-3 cm. long, narrowly concave above, convex below, short-tomen- tose; lamina subobovate to subelliptic or suboblong, 11.5-19.5 by 4-7 cm., obtuse to short-acuminate at apex, cuneate to subrounded at base, strongly and distantly undulate and slightly recurved at margin, coriaceous, drying greenish olivaceous and shiny above and umber-sabelline below, persistently tomentose on midrib above and on entire lower surface, the midrib above gradually narrowed from base, raised, 0.25-0.35 mm. wide at midpoint, below raised, + striate, the venation apparent on both surfaces, raised, 8 to 14 veins /5 mm., angle of divergence 70-75°. Infructescences terminal and from adjacent Gee axils, with scars of ca. 11 flowers, (?)branched, the axis to 6.5 cm. long, short-tomentose, lowest internode ca. 2.2 cm. long; bracts unknown; pedicels 1.5-2 cm. long, tomentose. Flower unknown. Immature fruit ca. 1.2 cm. long, furfuraceous. DisTRIBUTION. Sumatra (Map 17). SPECIMENS SEEN. Sumatra. Utara: Sibolga, 200 m., bb 19334 (a, Bo); Provincia di Padang, a Sungei bulu (+ 0 m.), Beccari PS 953 (FI, K, L, MEL 382 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Calophyllum sp. 54 can be readily recognized by the more or less tomentose indumentum that persists on all parts of the plant except the upper surface of the lamina and the ovary, by its terminal inflorescences, and by its furfuraceous fruits. Calophyllum sp. 54 is perhaps superficially similar to C. molle and C. rubiginosum, but both of these species differ in their tomentose ovaries and their leaves that do not dry so bicolored; C. molle also has axillary inflores- cences. Calophyllum woodii and Calophyllum sp. 53, two Bornean species with tomentose indumentum, both have hairs with numerous papillae and short branches, and leaves that do not dry bicolored and have denser venation; C. woodii has axillary inflorescences. 55. Calophyllum cucullatum Merr. Philip. Jour. Sci. 17: 288. 1920, Enum. Philip. Fl. Pl. 1: 78. 1923. Type: Philippine Islands, Mindanao, Surigao Province, 25 April 1909, BS 34518 coll. Ramos & Pascasio (isotypes, A, BM, BO, K, NY, P, US). Small tree; trunk and bark unknown. Twigs slightly flattened, 2.2-2.6 mm. across, with rounded edges, drying black, at first brown-tomentose, hairs to 2 mm. long; axillary innovations lacking basal scars; internodes 8.5-11.5 cm. long; uppermost pair of axillary buds rounded, ca. 1.5 mm. long, suberect; terminal bud 4.5-6.5 mm. long, brown-tomentose, long hairs obscuring shape (hairs, Ficure 22, a, b), under- developed internode absent. Petiole I|-1.5 cm. long, narrowly concave above, convex below, drying black, subpersistently tomentose below; lamina ovate to suboblong, 9.5-20.5 by 4.8-8.8 cm., (short-)acuminate at apex, rounded and cucullate at base, not undulate and slightly recurved at margin, coriaceous, drying umber above and below, with sparse, subpersistent hairs toward base on midrib on both surfaces, the midrib above abruptly narrowed near base, raised, ca. 0.2 mm. wide at midpoint, below raised, rounded, slightly striate, the venation above usually subobscure, below clear, slightly raised, 7 to 10 veins/5 mm., angle of divergence 65-70°. Infructescences from foliate axils, flabellate, with scars of ca. 15 flowers, with 3-flowered branches to 1 cm. long, the axis 3-5.5 cm. long, sparsely tomentose toward base, lowest internode 0.5-3 cm. long; bracts narrowly elliptic, ca. 5.5 mm. long, tomentose beneath, subpersistent; pedicels 1.1-2.3 cm. long, glabrous. Flower unknown. Fruit ellipsoid, ca. 1 by 0.8 cm., apiculate, drying grayish, smooth; outer layer not detaching cleanly from stone, very thin—when pressed less than 0.3 mm. across, with large air spaces developing; stone ellipsoid, ca. 9 b 7 mm., rounded at apex, the walls ca. 0.3 mm. thick, at base fibrous and ca. 0.5 mm. thick, smooth, (?)unmarked; spongy layer thin. DistriBuTION. The Philippine Islands, Mindanao, known only from the type collection (Map 20). EcoLocy. Thickets along streams at an iron deposit (Merrill, 1920, Joc. cit.); fruiting in April. Calophyllum cucullatum is a distinctive species that can be recognized 130 | | -———_—___— ——10 | y N wf pe | ee > | “0 an a ae ty A iene’ GA Qe . ~~ 4 eee oi ie Py Pall Map 20. Distribution of Calophyllum pulcherrimum (squares), Calophyllum sp. 61 (star in solid circle), C. complanatum (half-circles), C. oliganthum (open stars), C. cucullatum (solid star), and C. pelewense (triangle) in Malesia. WNTIAHdOTVO ‘SNAAALS [0861 €8e 384 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 by its tomentose indumentum; relatively short terminal bud; large, ovate-oblong leaf blades that are cucullate at the base; branched inflorescence; and small, ellipsoid fruits that have a thin outer layer disorganized by air spaces. The epithet alludes to the cucullate (hooded) base of the lamina. Merrill (1920, loc. cit.) compared Calophyllum cucullatum with C. soulattri, but noted that C. cucullatum differed in its cucullate lamina base. Calophyllum cucullatum also differs in drying a much darker color, in its short terminal bud, and in its fruits, which have a thin outer layer; the outer layer of the fruit of C. soulattri is characteristically at least | mm. thick. 56. Calophyllum glaucescens Ridley, Kew Bull. 1938: 120. 1938; Masamune, Enum. Phanerog. Born. 475. 1942. Type: Sarawak, Kuching, Dec. 1892, Haviland 2058 (holotype, k; isotypes, L, p (frag.), SING). Small tree ca. 7 meters tall; outer bark smooth, gray, hoop marked; inner bark light brown, finely laminated; latex yellow. Twigs slightly flattened, 1.5-2 mm. across, with 4 obscure raised lines, drying yellowish to whitish, glabrous; axillary innovations apparently with basal scars; internodes 2.5—5 cm. long; uppermost pair of axillary buds rounded, less than 0.5 mm. long, inconspicuous; terminal bud rather plump, I-1.8 mm. long, with brown, puberulent indumentum (hairs, Ficure 20, i), under- developed internode inconspicuous. Petiole 1-1.5 cm. long, broadly concave above, convex below, drying black; lamina elliptic to suboblong, 7-14.6 by 3.3-6.6 cm., acuminate at apex, acute to broadly cuneate at base, not undulate but narrowly recurved at margin, coriaceous, drying umber above and livid-vinaceous below, glabrous, the midrib above narrowing gradually from base, flat, adjacent lamina raised, 0.4-0.7 mm. wide at midpoint, below raised, rounded, the venation on both surfaces subobscure, ca. 25 veins/5 mm., angle of divergence 75-80°. Inflorescences from foliate axils along stem, with ca. 7 flowers, unbranched, the axis 1-1.5 cm. long, glabrous, lowest internode ca. 0.5 mm. long; bracts broadly ovate, ca. 2 mm. long, brown- puberulent beneath, persistent until anthesis; pedicels 6-9 mm. long, glabrous. Flower known only from advanced bud, (?)hermaphroditic; tepals 4, the outer two obovate, ca. 5.5 by 4 mm., subcucullate, the inner two + obovate, ca. 6 by 4.5 mm.; stamens 50 to 70, the filaments to 2.7 mm. long, the anthers ellipsoid, 0.5-0.7 mm. long, rounded to subretuse at apex; ovary ca. 1.3 mm. long, the style ca. 2 mm. long, the stigma slightly expanded, 0.3 mm. across. Fruit unknown. DisTRIBUTION. Borneo (western Sarawak), local (Map 23). ADDITIONAL SPECIMEN SEEN. Borneo. Sarawak. Ist Division: Sempadi F.R., 9m., S 6012 (sar). Eco.ocy. Small tree on dry podzol, 9m. alt. ($6012). Flowering in December. Calophyllum glaucescens is a distinctive, albeit poorly known, species recognizable by its very short terminal bud, pale-drying twigs, and lamina with dense venation. The midrib on the upper surface of the lamina is 1980] STEVENS, CALOPHYLLUM 385 surrounded by raised blade, and the lower surface of the lamina dries a livid-vinaceous color. The epithet alludes to the rather glaucous underside of the leaf. Calophyllum glaucescens is somewhat similar to C. cucullatum, but the latter species has larger leaf blades at least 9.5 cm. long with a sharply raised midrib on the upper surface, a rounded, cucullate base, less dense venation, and a lower surface that is not pruinose. Calophyllum cucullatum also has a longer (4.5-6.4 mm.) terminal bud, tomentose indumentum, and inflorescences up to 5 cm. long. Although better material is needed for confirmation, C. glaucescens appears to have scars at the bases of the axillary innovations; such scars are lacking in C. cucullatum. 57. Calophyllum nodosum Vesque, Epharmosis 2: ft. 10, 11. 1889, in C. DC. onogr. Phanerog. 8: 559. 1893; Merr. Bibl. Enum. Born. Pl. 393. 1921; Masamune, Enum. Phanerog. Born. 476. 1942; M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 337. 1956; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 7: 14. 1967; T. C. Whitmore, Tree Fl. Malaya 2: 171. 1972. Type: Sarawak [Ist Division], Gunong Poe, July 1866, Beccari PB 2440 (holotype, pP; isotypes, 4 (frag.), FI, K). IGURE 21, e. C. microphyllum T. Anderson in Hooker f. Fl. Brit. India 1: 272. 1874, non Planchon & Triana (1862) nec Scheffer (1873); King, Jour. Asiatic Soc. Bengal, II. 59: 173. 1890; Ridley, Jour. Straits Branch Roy. Asiatic Soc. 35: 8. 1901; C. kingianum Engler in Engler & Prantl, Nat. Pflanzen- fam. 3(6): 221. 1893 (7 Nov.), nomen novum; C. parvifolium Vesque in C. DC. Monogr. Phanerog. 8: 604. 1893 (Dec.), non Choisy (1823), nomen novum; Ridley, Fl. Malay Penin. 1: 182. 1922. Type: Malaya, Malacca, Mt. Ophir, Aug. 1867-68, Maingay 2576 (Kew dist. 165) (lectotype, kK). C. depressinervosum auct., non M. R. Henderson & Wyatt-Smith; Pukol & Ashton, Checklist Brunei Trees, 93. 1964; Smythies, Common Sarawak Trees, 99. 1965. C. pulcherrimum auct., non Wall. ex Choisy & C. rupicola auct., non Ridley; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 7: 14. 1967. Shrub 15 cm. tall to tree to 25(-36) meters tall, d.b.h. to 42 cm.; trunk without spurs or buttresses; outer bark yellowish (brown), with lenticels or boat-shaped fissures, very brittle in old trees, the inner surface black to dull orange-brown; under bark mid to deep red; inner bark reddish; latex clear (rarely cloudy) yellow, sticky. Twigs slightly flattened, 0.5-1.5 mm. across, 4-angled to 4-alate, drying brown, glabrous or sparsely and transiently puberulent; axillary innovations with (sub)basal pair of scars, nearly always with additional pair within 5 mm. of base; internodes 0.3-2.5 cm. long; uppermost pair of axillary buds rounded, to 0.5 mm. long, inconspicuous; terminal bud rather plump, 0.5-1 mm. long, with short, grayish to brownish, subadpressed indumentum (hairs, Ficure 20, j-l), often not functional, underdeveloped internode to 3.5 mm. long. Petiole 1-5.5 mm. long, deeply concave above, convex below, glabrous; 386 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Ficure 21. a, b, Calophyllum stipitatum (FRI 3656), fruit, x 1.5: om outside; b, longitudinal section. c, C. complanatum (SAN 53254), habit x 0.5. d, C. pulcherrimum (Chew 1365), fruit, transverse section, x |. nodosum ‘Ss 19726), fruit, transverse section, pee ae ae On pa ieee (KEP 104281), fruit, longitudinal section, x 3. g-i, C. roseocostatum. g, h, fruit (S 14941), x 0.75: g, longitudinal section; h, from outside. 1, terminal bud (S 26299), x 6 lamina obovate or elliptic to subcuneiform or narrowly elliptic, (0.7—-)1.4-6.5 (-10) by (0.15-)0.4-2.5(-3.6) cm., rounded to shortly and bluntly acuminate at apex, narrowly cuneate at base, usually minutely and abruptly rounded at very base, slightly undulate and not recurved at margin, thinly coriaceous, drying umber to near grayish olive above and umber to near sabelline or olive below, glabrous, the midrib above gradually narrowed from base, + 1980] STEVENS, CALOPHYLLUM 387 level to sulcate, surrounding lamina slightly raised, 0.5-1.5(-2) mm. wide at midpoint, often disappearing ca. 2 mm. short of apex, below level to slightly depressed, inconspicuous, striate, the venation subapparent to subob- scure above and below, slightly raised (latex canals sometimes clearly impressed), 6 to 13 (to 15) veins/5 mm., angle of divergence 50-70(-75)°. Inflorescences from foliate axils along stem, with 1 to 9 (to 13) flowers, unbranched, the axis to 3.5 cm. long, glabrous (rarely puberulent) at base, lowest internode less than 2 mm. long; bracts ovate, ca. 1.7 mm. long, soon deciduous; pedicels 0.5-2.2 cm. long (-3 cm. long in fruit), glabrous, slender. Flower (?)hermaphroditic; tepals 4 (to 8), the two outer ovate (rarely sub- triangular), 3-3.5 by 2.2-3 mm., sometimes puberulent on back at apex, the inner two elliptic to obovate, 3.5-5.5 by 2.3-3.7 mm. (5 by 2.2 mm), any others often much smaller; stamens 27 to 50 (to 60), the filaments to 3 mm. long, the anthers (elliptic-)oblong, 0.5-1 mm. long, slightly retuse at apex; ovary 1.2-1.5 mm. long, the style 2-2.5 mm. long, the stigma peltate, ca. 0.4 mm. across, + 3-radiate to infundibular. Fruit spherical, (4-)5.5-10 mm. long and across, rounded to apiculate at apex, drying grayish brown to yellowish brown, smooth or shallowly wrinkled; outer layer not detaching cleanly from stone, ca. 0.3(-0.7) mm. thick, with large air spaces developing; stone ellipsoid to subspherical, (3.5-)5-7.5 by (3-)5-6.5 mm., rounded at apex, the walls less than 0.15 mm. thick, smooth, unmarked; spongy layer thin. DistripuTion. Malay Peninsula, local; Borneo, widespread in the northwest (Map 21 Map 21. coe Paha of Calophyllum nodosum (squares; half-circle, report- ed locality) in Malesi 388 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61] SELECTED SPECIMENS SEEN (‘‘f’’ denotes Bornean specimens with fruits). Malaya. Jonore: G. Mering and G. Ledang, Ridley 3222 (pm, sinc); G. Ledang (Mt. Ophir), 975 m., FRI 1235] (x, kep). Borneo. Sarawak. Ist Division: A 532 coll. Mashor, 14 Aug. 1948 (ku); Bako Natl. Park, Lintang Path, 120 m., Sinclair & bin Tassim 10317 (aA, B, BM, E, K, M, NY, SAR, SING); Mt. Matang, 1021 m., Clemens 22353 (Ny); G. Pueh F.R., 30 m., S 7530 (sar); Bungoh Range, Bau, 792 m., S 29022 (a, sar); Sabal Tapang, 190 m., Stevens et al. 183 (a). 3rd Division: Ulu Sama, Batang Rajang, SFN 36057 (a, k, KEP, L, LAE, SING); Hose Mts., Ulu Temalad, Mujong, 305 m., S 17236 (a, K, KEP, SAN, SAR, SING); Bintulu, Merurong Plateau, 853 m., S 8714 (sar); Kapit, Melinau, Bukit Pantu, 732 m., S 25720 (KEP, L, SAR, SING); Balingian, Ulu Sungei Arip, Bukit Iju, 180 m., S 23786 (a, Bo, K, KEP, SAN, SAR, SING); Usun Apau Plateau, R. Julai, 965 m., S 38/1] (sar, sinc). 4th Division: Baram distr., Kelabit Highland, 1067 m., S 35343 (k, sar); Lambir proposed Natl. Park, Miri, S 24064 (a, K, KEP, SAN, SAR). 5th Division: Malignan Range, 1494 m., S' 33035 (sar). Brunet: Bukit Sagan, 549 m., BRUN 3110 (L, sar); Batu Ketam, Ulu Ingei, 255 m., BRUN 5608 (L, sar); Bukit Patoi, 240 m., Ashton s.n. Aug. 1958 (sar); Berakas F.R., 15 m., S 7822 (Bo, BRI, K, L, NY, SAR, SING): Andalau F.R., 21 m., Sinclair & bin Tassim 10448 (A, E, K, KEP, SAR, SING); G. Pagon Periok, 1860 m., BRUN 2389 AS SAR); Bados, KEP 35589 (KEP); between Danau and Tutong, van Niel 3736 (L, mo); Sungei Singap, KEP 34570 (KEP); Kuala Belait, S 10652 (sar). SaBan. Sipitang: Menggalong F, Sibubu R., SAN 21829 (KEP, SAN, SAR); Mesapol, Teck Gua SAN 50620 (SAN). Beaufort: Bukit Sunngau, Weston, SAN 55658 | (f) (L, SAN, SAR, SING); Beaufort Hill, SAN 35227 (san). Kota Belud: Mt. Templer F.R., S. of Sungei Talupit, 610 m., SAN 76266 (k, SAN, SAR). Tenom: Kapulu, way to Rendum, 1097 m., SAN 72058 (f) (kK, san); Mandalom F.R., SAN 31948 (san). Keningau: Mile 9, Kampong Nalayan, SAN 5846] (f) (kK, L, san); Mile 10, Rashna road, Nabawan, 305 m., SA N 83839 (f) (a, kK). Tambunan: Trusmadi F.R., above Ulu Koingaran R., 1514 m., SAN 41779 (k, L). Ranau: Kinabalu Natl. Park, Tohubang near Kampong Kiau, 918 m., SAN 51428 (san). Labuk & Sugut: Jambongan Is., 3 m., SAN 3899 (so, k); Bukit Gambaran near Telupid, 457 m., SAN 51525 (kK, san, sar); Bukit Tangkunan, 425 m., Stevens et al. 408 (a). Sandakan: Sungei Paitan W.C., SAN 23977 (f) (xk, KEP, L, SAN, SAR, SING); Mt. Walker F.R., 30-60 m., SAN 53171 (xk, KEP, L, SAN, SAR, SING); Kiabau Hill, 150 m., SAN 4381/4 (san); Sungei Mayan, 40 m., Stevens et al. 38] (a). Kinabatangan: Brassey Range, 810 m., Stevens et al. 481 (A); Lamag, Bukit Kuamat, SAWN 70729 (san); S. slope of G. Lotung Inarat, 1036 m., SAN 83312 (a). Lahad Datu: Pulau Sakar, Look Magulang, m., SAN 15524 (f) (k, L, SAN, SAR); Bakapit, Silabukan Falls, 457 m., SAN 52956 (kK, L, SAN); Mt. Silam, 457 m., SAN 22711 (san). KALIMANTAN. Timur: Nonoekan, 3 m., bb 26213 (?) (Bo, L); W. Kutei, Belajan R., Mt. Palimasan near Tabang, Kostermans 12913 (f) (Bo, CANB, K, L, P, SING); bij L. Petah, 800 m., Endert 3209 (f) (Bo, L). Barat: between S. & G. Kenepai, Hallier 1948 (a, Bo, kK, Ii Bengkajang, G. Bawang, 1400 m., bb 9667 (Bo); Palo, Becking 23 (f) (B Eco_ocy. Common in kerangas forest on acid and often sandy soil, sometimes in waterlogged places; 13-1860 m. alt. Unusually plastic in form: ‘‘procumbent shrub 2 ft. tall rooting along branches’’ (Purseglove 5604); ‘‘cushion plant’’ 20-40 cm. tall (S 4447); ‘‘shrub 6”-1 ft. high’? (Sinclair 10317)—the last 1980] STEVENS, CALOPHYLLUM 389 specimen, at least, is fertile. Near Sandakan, Sabah, to 42 cm. d.b.h.; other individuals nearby somewhat smaller. Flowering February to June; somewhat less frequently July to December. Fruiting January to November (June and December on Malay Peninsula); fruit yellowish to grayish. GERMINATION AND YOUNG PLANT. The radicle breaks the stone near the base of the seed. The seedling has four to six (or seven) pairs of scale leaves and (one or) two pairs of expanded leaves, all separated by well-developed internodes; the internode between the cotyledons and the first pair of scale leaves is ca. 5(-11) mm. long. The terminal bud is at least initially functional, and young plants ca. 30 cm. tall are erect (Stevens et al. 381A Calophyllum nodosum is distinguished by its small leaf blades with the midrib above surrounded by raised lamina and below level or slightly depressed; its axillary innovations with basal scars; and its small fruits with a thin outer layer and a thin stone wall. The epithet nodosum alludes to the rather closely set nodes, which are especially prominent after the leaves have fallen off. Calophyllum nodosum is difficult to separate from C. pulcherrimum when not in fruit, and it appears also to be related to C. depressinervosum; for the differences separating these species see TasLe 11 and the discussion after C. pulcherrimum. Calophyllum nodosum is also related to C. complana- tum (q.V.). Calophyllum nodosum is extremely variable in leaf size, shape, and—to a certain extent—color on drying. Specimens from the 4th Division of Sarawak (Miri: S 24064) and from Batu Ketam and Bukit Patoi in Brunei have a narrowly elliptic to obovate lamina 2.5—4.4(-7.5) by 0.15-0.75(-1) cm. and are at first sight different from the other specimens. S 24064 is in fruit; the fruits are somewhat smaller (4 mm. long) than normal. In all other features the narrow-leaved specimens agree well with the broader-leaved ones. They appear to represent one extreme of the considerable variation in leaf shape and size shown by C. nodosum, and they grow in the same habitat; it does not seem worthwhile to give them formal recognition. The terminal buds in mature plants of Calophyllum nodosum are sometimes functional; those of the young plants seen were functional. Specimens of Calophyllum nodosum from Borneo agree well with those from southern Malaya, despite the considerable distance between the two areas. Ridley (1901, loc. cit.; as C. microphyllum) suggested that C. nodosum may also occur on Gunong Panti, Johore. The apparent absence of C. nodosum from Sumatra is perhaps surprising in view — the abundance and rather wide altitudinal range of the species in Borne Calophyllum microphyllum T. Anderson is “lccloepiliea by Maingay 2576 at Kew. Anderson also cited specimens collected by Lobb and Griffith. 58. Calophyllum depressinervosum M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 335. pl. 17. 1956; C. depressinerve Wyatt-Smith, Malayan Forest Rec. 17: 113. 1952, nomen; Kochummen, Malayan Forest Rec. ed. 2. 17: 220. 1965; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 7: 15. 1967; Corner & Watanabi, Illus. Guide Trop. Pl. 390 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 181. fig. 1969; T. C. Whitmore, Tree Fl. Malaya 2: 179. 1973; Corner, Gard. Bull. Singapore Suppl. 1: 104. 1978. Type: Malaya, Pinang, Moniot’s Road, 1000 feet [305 m.], May 1886, Curtis 830 (holotype, SING; 1SOtypes, K, SING). Ficure 21, f. Calophyllum sp. near floribundum Hooker f.; Curtis, Jour. Straits Branch Roy. Asiatic Soc. 35: 79. 1894. Tree 4-36 meters tall, d.b.h. to 86 cm.; trunk without buttresses or spurs; outer bark brown to yellowish, with shallow, rather numerous, short, boat- shaped fissures, not hoop marked, the inner surface (dark) brown (blackish); under bark reddish to red-brown; inner bark dark red to pink; latex yellow, clear, or becoming cloudy, or cloudy, always sticky. Branches + pendulous. Twigs slightly flattened, 1-1.7(-2.5) mm. across, obscurely 4-angled, soon becoming rounded, drying brown, glabrous; axillary innovations with | to 3 pairs of scars within 3 mm. of base; internodes (0.5-)1-6 cm. long; uppermost pair of axillary buds rounded, ca. 0.5 mm. long, spreading, not conspicuous; terminal bud plump to conical, 1.5-2.2 mm. long, with short, subadpressed, brownish indumentum (hairs, Ficure 20, m), usually not functional, under- developed internode absent. Petiole 3.5-8 mm. long, (deeply) concave above, convex below, glabrous; lamina ovate to obovate or subelliptic, 3.2-10 by 1.2-4.1 cm., strongly acuminate at apex, acumen ca. | cm. long, cuneate to acute at base, slightly and distantly undulate and not recurved to slightly so at margin, thinly coriaceous (rarely coriaceous), drying umber to olivaceous above and cinnamon to sabelline below, glabrous, the midrib above gradually narrowed from base, slightly raised (adjacent leaf raised), 0.1-0.25(-0.35) mm. wide at midpoint, below flat to depressed, striate (rarely somewhat raised), the venation on both surfaces subobscure (subapparent), raised, latex canals frequently (and rarely veins) impressed above, 4 to 10 veins/5 mm., angle of divergence 55-70°. Inflorescences from foliate axils along stem, with 7 to 13 flowers; unbranched, the axis 0.5-5 cm. long, glabrous, lowest internode to 2 mm. long; bracts unknown; pedicels (0.25—-)0.4-1.3 cm. long, glabrous, slender, to 1 mm. thick in fruit. Flower (?)hermaphroditic; tepals 4, the outer two broadly ovate, ca. 3 mm. long and across, the inner two obovate, ca. 5.5 by 2.5 mm.; stamens ca. 40, the filaments to 3 mm. long, the anthers subelliptic, 0.4-0.7 mm. long, rounded to retuse at apex; ovary ca. 1.2 mm. long, the style ca. 2 mm. long, the stigma peltate, ca. 04. mm. across, (?)lobed. Fruit spherical, 0.8-1.1 cm. long and across, apiculate, drying orange-brown, smooth or broadly wrinkled, wrinkles rounded; outer layer not detaching cleanly from stone, 0.2-0.7 mm. thick, with large air spaces developing; stone ellipsoid, 6-8 by 5-6.5 mm., rounded at apex, the walls less than 0.1 mm. thick, smooth, unmarked; spongy layer thin. DistripuTion. Cambodia and Thailand to Borneo, excluding Java (Map 22). SELECTED SPECIMENS SEEN. Cambodia: Pursat, entre Anlong Krauch et Veal Veng, 500 m., Martin 1797 (p). Thailand: Chantabuli, anon., 13 Aug. 1933 (BkF). Malaya. Kepan: Gunong Jerai F.R., KEP 7745 (x, KEP, sinc); Bukit Lada Acheh, KEP 7570 (ker); Gunong Inas F.R., FRI 4632 (ker); Perangin Tas_e 11. Differences between Calophyllum pulcherrimum, C. nodosum, and C. depressinervosum. C. pulcherrimum C. nodosum C. depressinervosum LaMINA LENGTH (cm.) APEX ACUMINATE VENATION DENSITY (veins /5 mm.) LATEX CANALS IMPRESSED WHEN DRY VEINS VISIBLE ON LOWER SURFACE CINNAMON-SABELLINE BELOW WHEN DRY Fruit LONGER THAN 1.4 cM. OUTER LAYER AT LEAST | MM. THICK STONE WALLS AT LEAST 0.3 MM. THICK SEED AT LEAST SOMETIMES POLYEMBRYONIC YOUNG PLANT CA. 20 CM. TALL, ERECT (2.7-)4-8(-11) 7 to 13 (to 17) +(and sometimes latex canals) (0.7-)1.4-6.5(-10) —(+ 6 to 13 (to 15) —(+ +(and latex canals) a) + +? 3.2-10 + 4 to 10 +(=) + + neat WOATIAHdOTVO ‘SNAARLS [0861 16€ 392 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 Map 22. Distribution of Calophyllum depressinervosum in Southeast Asia and Malesia F.R.,90m., KEP 79284 (ker); Enggang F.R., 37 m., KEP 74991 (ker). PINANG: Tiger sae Road, 457 m., Haniff 265 (c, UC). Perak: road from Kampong Ayer to G. Bubu massif, 396 m., FRI 13898 (a, K, KEP, SAR, SING); Kirita, anne Saiong F.R. KEP 65947 (KEP); Ulu Kenderong, KEP 11038 (kep). SELANGOR: Kepong, Bukit Lagong F.R., 150 m., KEP 98701 (A, K, KEP, SAR, SING); Kuang, KEP 22059 (keEp, sinc); Bangi F.R., Kajang, KEP 14578 (x, KEP, SING); Sungei Buloh F.R., KEP 14926 (kep). NEGRI SEMBILAN: Kuala Pilah, Kepis F.R., KEP 62954 (a, Bo, K, KEP, sING); Hutan Simpon Paseh, KEP 100176 (KEP). KELANTAN: Ulu Temiang F.R., KEP 104281 (a, K, KEP, L, SAN, SING); G. Rabong, 366 m., Soepadmo & Mahmud 1198 (a, K, KEP, KLU, L 3km.N. of Kuala Trengganu, FRI 20289 (kep). PAHANG: Jengka F.R., Temerloh, KEP 98594 (Kk, KEP, SAN, SAR); Mentakab, KEP 7922 (ker); G. Tahan, Wong & Wyatt-Smith 49 (ker). Jonore: Mawai, Corner s.n., 7 Oct. 1934 (KEP, po Kluang F.R., 30 m., KEP 69922 (KeEp); Mersing, G. Arong F.R., 30 m., 84703 (KEP); “‘Labis, ‘Segamat, Mambai F.R., KEP 69862 (KEP). Sumatra a adjacent islands. Riau: Lingga, P. Singhap, oetan darat Njakar (Bakong), 150 m., bb 2043 (?) (Bo, L); Ngina, Ri/J-8 (Bo). Barat: Loeboek Sikaping, Tandjoeng Boengi, 370 m., bb 6502 (Bo). Utara: Sibolga, P. Poene, 10 m., bb 3783 (Bo); Baroes, Pangkalan Tapoes, 25 m., bb 29524 (po, L); Morsala 1980] STEVENS, CALOPHYLLUM 393 Is., Silabua, 10 m., Kostermans’ collector 13, Jan. 1951 aCe BO, SING). BANGKA: W. Bangka, Penganah, 30 m., bb 26138 (Bo, sinc). Borneo. Sarawak. Ist Division: Sadong, Sabal F.R., S 13454 (a, Bo, K, L, SAN, Fay 2nd Division: Bukit Ubah Ribu, Ulu Sungei Kaup, 610 m., S 33782 (k, KEP, SAN, SAR). Brunel. Temburong, mile 11/4 Bangar-Batu Apas road, 9 m., SAN 17093 (A, BO, K, KEP, L, SAN, SING). SABAH. Sipitang: Mile 8, Sipitang road, SAN 35125 (KEP, L, SAN, SAR, SING); Mesapol F.R., SA N 73347 (san). Papar: Mandahan Hill, 225 m., SAN 78001 (san). Sandakan: Sepilok Extension, sea level, SAN 24912 (K, KEP, L, SAN, SING). Tawao: Brantian, 30 m., Stevens et al. 435 (A); Brassey Range, 660 m., Stevens, sight record. KALIMANTAN. Tengah: Midden Doeson, Lemoedajak, 60 m., bb 11596 (so). Barat: Moeara Kajang, 10 m., bb 7465 (Bo) Ecotocy. Mixed dipterocarp forest, on ridges and other well-drained places, sometimes in forest approaching kerangas forest (Sarawak) or swamp forest (Johore—Corner, loc. cit.; the single tree I saw in a swamp was on an eminence ca. 1.5 m. above surrounding swamp); to 762 m. alt. In Malaya flowering January, May, November, and December; fruiting January to March and May, November, and December. In Borneo fruiting March, April (most), and June. Ripe fruit yellowish. GERMINATION AND YOUNG PLANTS. Vigorously growing young plants up to 1.2 meters tall with functional terminal buds have been seen (Sarawak, Sabal Tapang) Calophyllum depressinervosum can be most readily recognized by its moderate-sized to small leaf blades that are strongly acuminate at the apex, that usually have obscure, rather distant venation when dry, and that often have clearly impressed latex canals. The lower surface of the dried leaf blades is often a yellowish brown, at least in specimens that are not too old. The flowers have four tepals. The fruits are small (about 1 cm. long), and the stone has thin walls. The epithet depressinervosum really refers to the latex canals, which are depressed when the leaf is dry (note that the veins are rarely depressed!). Calophyllum depressinervosum is similar to C. nodosum and, to a lesser extent, to C. pulcherrimum; some differences between the species are listed in TABLE 11. Calophyllum depressinervosum also has less dense venation than does C. nodosum: four to ten veins/5 mm. vs. six to thirteen (to fifteen) veins /5 mm. Leaf blades and fruits of specimens of C. nodosum from Sarawak (e.g., S 17236, 21136, 21243) dry a similar color to those of C. depressinervosum, but the species are not otherwise similar. The above-cited specimens of Calophyllum depressinervosum from Cambo- dia and Thailand are sterile, but there is no doubt as to their identity. The specimens from Kalimantan and some from elsewhere in Borneo (e.g., SAN 17093, from Brunei) are robust, with twigs up to 2.5 mm. in thickness, and the leaf blades are rather coriaceous and tend to dry boat shaped. SAN 17093 has fruits characteristic of C. depressinervosum, albeit borne on rather short pedicels. The specimens from Kalimantan are sterile. 394 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 59, Calophyllum complanatum P. F. Stevens, sp. nov. Ficure 21, c. A speciebus aliis Calophylli in lamina lingulata vel oblonga basi cordata costa supra lamina elevata circumdata infra non nisi vel leviter elevata, et inflorescentia axillari internodio infimo 0-1 mm. longo, differt. Tree ca. 9 meters tall, d.b.h. ca. 5 cm.; outer bark yellow-green. Twigs strongly flattened, 1.2-2.6 mm. across, strongly 4-angled, drying brown, glabrous; axillary innovations with pair of basal scars and often another pair within 5 mm. of base; internodes 2-4 cm. long; uppermost pair of axillary buds rounded, less than 0.5 mm. long, inconspicuous; terminal bud plump, 1.5-3 mm. long, with short, brown, tomentose indumentum (hairs, Ficure 20, n), underdeveloped internode absent. Petiole 1-2 mm. long, shallowly concave above, convex below, glabrous; lamina lingulate to suboblong, 9.5-14.8 by 1.5-4.5 cm., subacuminate to acute at apex, cordate at base, slightly undulate and recurved at margin, thinly coriaceous, drying grayish sepia above and sabelline below, glabrous at maturity, the midrib above abruptly narrowed near base, raised, surrounding lamina also raised, 0.1—0.25 mm. wide at midpoint, below raised, angled to substriate, the edges subde- pressed, the venation + apparent above, slightly raised, below subobscure, one canals slightly impressed above and strongly impressed below, 5 to 7 veins/5 mm., angle of divergence ca. 80°. Inflorescences from foliate axils, with ca. 7 flowers, unbranched, the axis 1.2-3 cm. long, glabrous, slender, lowest internode to 1 mm. long; bracts not known; pedicels 0.7-2.2 cm. long, glabrous. Flower (?)hermaphroditic; tepals 4, the outer pair ovate- aaa ca. y 2 mm., the inner pair + obovate, 5-6 by 3-4 mm.; stamens ca. 1, the filaments to 3 mm. long, very slender, the anthers elliptic, 0. ie ie S mm. long, retuse at apex; ovary ca. 0.7 mm. long, the style ca. 2.7 m long, the stigma slightly expanded, ca. 0.3 mm. across. Immature a (?)subspherical, ca. 5 mm. long; outer layer ca. 0.2 mm. thick; stone with walls ca. 0.2 mm. thick. Type: Sabah, Kinabatangan, Karamuak, Lamag, east of Sg. [Sungei] Meliau, 45 m., 17 June 1965, SAN 53254 coll. Teorodop (holotype, k; isotype, SAN). DistrisuTION. Northeastern Borneo (Map 20). ADDITIONAL SPECIMEN SEEN. Borneo. SABAH. Labuk & Sugut: Mile 86, Bukit Tangkunan, 425 m., Stevens et al. 407 (a). Eco.ocy. On soils derived from ultramafic rock, 45-425 m. alt. Flowering in June. Calophyllum complanatum can readily be distinguished from the other species of the genus even when sterile: its axillary innovations have basal scars, and its leaf blades are cordate at the base, with the midrib on the upper surface flat and surrounded by raised lamina. The twigs are notably flattened, hence the specific epithet. Calophyllum complanatum is probably most closely related to C. nodosum, which also has innovations with basal scars, the upper surface of the lamina 1980] STEVENS, CALOPHYLLUM 395 with the midrib broadly raised, inflorescences with short basal internodes, and flowers with four tepals. Calophyllum nodosum differs most obviously from C. complanatum in its smaller leaf blades that are never cordate at the base. The two species grow within a few meters of each other on Bukit Tangkunan (for C. nodosum see Stevens et al. 408); young plants of C. complanatum up to 3 meters tall were quite common but showed little variation. The fruits of Calophyllum complanatum are known only from the isotype at Sandakan. These fruits are small and have been distorted during pressing, but they have a well-defined stone wall ca. 0.2 mm. thick. It is therefore likely that the mature fruits of C. complanatum are less than 1 cm. long. 60. Calophyllum eee Wall. ex Choisy, Mém. Guttif. Inde, 41. 1849, Mem. Soc. Phys. Hist. Nat. Geneve 12: 421. 1851; Wall. Catal. 4848. 1831, nomen; ee & Triana, Ann. Sci. Nat. Bot. IV. 15: 274 1862; T. Anderson in Hooker f. Fl. Brit. India 1: 271. 1874, pro parte; Kurz, Jour. Asiatic Soc. Bengal, II. 39: 64. 1870; Pierre, Fl. Forest. Cochinch. 1: pl. 104B. 1885, pro parte; King, Jour. Asiatic Soc. Bengal, II. 59: 174. 1890, pro parte; Vesque in C. DC. Monogr. Phanerog. 8: 570. 1893, pro parte; Ridley, Jour. Straits Branch Roy. Asiatic Soc. 33: 47. 1900, Fl. Malay Penin. 1: 182. 1922, pro parte; Heyne, Nutt. Pl. Indonesié. ed. 3. 1: 1085. 1950; M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 334. 1956; Gilliland, Common Malay PI. 37. 1958; Kochummen, Malayan Forest Rec. ed. 2. 17: 220. 1965; Tree Fl. Malaya 2: 181. 1973; H. Keng, Gard. Bull. Singapore 28: 244. 1976; Corner, Gard. Bull. Singapore Suppl. 1: 104. 1978. Type: Singapore, Sept. 1822, Wallich dist. 4848, pro parte (holotype, Gc; isotypes, BM, FE, FI, G, K, M, MEL, NY, P, W). Ficure 21, d. C. plicipes Miq. Fl. Indiae Batavae Suppl. 1(3): 499. 1861; Kurz, Nat. Tijdschr. Nederl.-Indié 27: 192. 1864; F. Mueller in Walp. Ann. Syst. Bot. 7: 357. 1868. Type: eae prope Djeboes, Teysmann, HB 3209 (holotype, u; isotypes, B C. mesuaefolium Wall. Catal. 4850. — Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 293. 1862. Nom C. retusum auct., non Choisy; King, Jour. Asiatic Soc. Bengal, II. 59: 176. 1890, pro minore parte. C. pulcherrimum Wall. var. gracile (Miq.) Boerl. Catal. Horto Bogor. 2: 82. 1901, quoad spec. cit C. foetidum Ridley, Jour. Straits Branch Roy. Asiatic Soc. 54: 18. 1910, pro minore parte. Tree 10—23(-30) meters tall, d.b.h. to 40 cm.; trunk rarely with buttresses to 1 meter tall; outer bark yellowish brown, often mottled, with vertical lines of lenticels, or cracked (rarely slightly scaly), or with boat-shaped fissures, the inner surface red-brown to reddish black; under bark green-brown, red, or blackish, outer bark often detaching only with difficulty; inner bark pinkish 396 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 to mid-red; latex usually clear yellow (rarely cloudy yellow or brown-yellow), sticky. Branches often pendulous Twigs somewhat flattened, 0.7-1.2 mm. across, strongly 4-angled, drying brown or dark brown, glabrous; axillary innovations with | (or 2) pair(s) of basal scars; internodes |-4.5 cm. long; uppermost pair of axillary buds rounded, less than 1 mm. long, spreading, inconspicuous; terminal bud conical (plump), 0.7-1.3 mm. long, with subadpressed, brownish indumentum (hairs, Ficure 22, j), usually not functional, underdeveloped internode absent (—3.5 mm. long). Petiole 3-11 mm. long, deeply concave above, convex below, gla- brous; lamina elliptic to suboblong (rarely obovate), (2.7—)4—8(-11) by (0.8-) 1.6-3.4(-4.2) cm., bluntly acuminate (rarely acute to subrounded) at apex, acute to attenuate toward base, very base minutely rounded, slightly undulate and not recurved at margin, coriaceous, drying umber to sepia above and umber to sabelline below (often with thin, grayish, waxy covering, especially on the upper surface), glabrous, the midrib above gradually narrowed from already narrow base, raised, surrounding leaf raised, 0.15—0.3(-0.55) mm. wide at midpoint, below flat to depressed, striate, the venation subobscure to apparent above and below, slightly raised, 7 to 13 (to 17) veins/5 mm., angle of divergence 65-70(-80)°. Inflorescences from foliate axils along twigs, with 7 to 11 (to 17) flowers, unbranched, the axis 1.5-4.5 cm. long, glabrous, lowest internode less than 2.5 mm. long; bracts ovate to obovate, ca. 2 mm. long, soon deciduous; pedicels 0.4-3 cm. long, glabrous, slender. Flower (?)hermaphroditic; tepals 4 (rarely 8), the two outer ovate to elliptic, 3.5-5.5 by 2.5-4 mm., very rarely with few hairs near apex on back, the inner ones + obovate to elliptic, 6-9 by 2.5-3.5 mm., rarely glabrous; stamens (25 to) 65 to 140, the filaments to 5 mm. long, connate for up to 1 mm., the anthers oblong, 0.6-1.2 mm. long, + retuse at apex; ovary |.3-1.5 mm. long, the style 2-3.2 mm. long, the stigma peltate, 0.4-0.8 mm. across, 3-lobed. Fruit ovoid to spherical, 1.5-1.7 by 1.2-1.8 cm., when young ovoid, sharply pointed at apex, smooth, when older apiculate or not, drying brown, promi- nently wrinkled; outer layer detaching cleanly from stone, 1—-2.2 mm. across, compact; stone subspherical, 1-1.3 by 0.95-1.3 cm., rounded at apex, the walls 0.3-0.5 mm. thick, smooth, unmarked; spongy layer thin. DistripuTION. Southern Malaya to western Borneo, excluding Java (Map ae 22. Hairs (from terminal bud). a, b, Cee eieailanien (BS 34518). c, d, C. oliganthum (BS 34520). e, f, C. andersonti: e, S 24679; f, Haviland = 1812. g, h, C. pelewense (Kanehira 386). 1, Calophyllum sp. 61 (S 4732). j, C. pulcherrimum (Shah & Shukor 2395). k, C. roseocostatum (S 26299). 1, C. stipitatum (FRI 3656). m, C. confertum (SAN 17146). n-p, C. praetermissum (Haviland & Hose ie Se ara in only one plane p, apex and base of hair ca. 500 pm. long). q, C. flavo-ramulum (FRI 7521). r-t, C. celebicum (Cel. /II-213), the a thin- Rone apical cells fall off, leaving flat tops. Scale = 60 398 JOURNAL OF THE ARNOLD ARBORETUM {voc. 61 SELECTED SPECIMENS SEEN (‘‘f’? denotes Bornean specimens with fruits). Malaya. Necri SEMBILAN: Pengkalen Kempas, K EP 4213 (x, sina); Pasir Panjang, KEP 4202 (k, sinc); Port Dickson, Cape Rachado, 120 m., KEP 71340 (keEpP); Sungei Menyala F.R., 45 m., KEP 64091 (ker). Macacca: Pulu Besar, Maingay 1065 (Kew dist. 169, pro parte) (x); Sungei Udang, Goodenough 1738 (sinc, w); Merlimau, Derry 156 (sinc); Bukit Sabukor, Derry 104 (pm). PaHano: Sungei Rompin, Bukit Leuting, KEP 3230 (x, sinc); E. edge of Chini F.R., 150 m., FRI 14560 (a, K, KEP, SING); Baloh F.R., KEP 6842 (x); Lesong F.R., ridge N. of Sungei Jekatih, 180 m., FRI 20006 (KEP, sar); Menchali F.R., KEP 29655 (ker); Temerloh, 643/4 miles Maran Road, 180 m., KEP 77964 (KEP). Jonore: Sungei Kayu, SFN 31990 (k, KEP, sinc); G. Ledang (Mt. Ophir), Ulu Sungei Belemang, FRI 12327 (a, k, KEP, SING); Banang F.R., 270 m., FRI 2122 (a, K, KEP, SAN, SING); 10th mile Mersing—Endau Road, FRI 2783 (kK, KEP, sING); Tg. Penawar, Johore Coast, 15 m., FRI 7624 (k, KEP, SING); Pulau Tinggi, KEP 70936 (kep); Batu oe Soga F.R., KEP 79519 (KEP); G. Arong F.R., KEP 94884 (kep, sinc); G. Pulai F.R., 455 m., FRI 17553 (KEP); G. Lambak microwave station, Renggam F.R., FRI 17755 (k, KEP, SAR, SING); Panchor, Tg. Sewat Rogei, KEP 62814 (kep); Kota Tinggi, 115 chains in from E. coast, 60 m., KEP 118195 (a, Kk, KEP, SING). Singapore: Seletar Reservoir, Mandai Road, Shah & Shukor 2370 (A, BRI, C, KEP, KLU, sING); Tanjong Gul, Sinclair 10762 (k, FI, G, K, SING); Fairy Point, Changi, Ridley s.n., Nov. 1894 (sinc); MacRitchie Reservoir, SEN rate: (B, K, M, NY, SING); Bukit Timah F.R., SFN 34440 (a, Bo, K, LA NG); Tampenis, Ridley 4639A (Fi, sinc); Garden Jungle, Ridley 13305 aa K, oe. Keranji, Ridley 260 (sm, sinc). Sumatra and adjacent islands. Riau: Riouw, Tandjong Pinang, Kelong, P. Ngalim Besar, 12 m., bb 11444 (so); Lingga, P. Singkep, Tandjong Batang, 15 m., bb 4019 (Bo); Marok Toewa bij Soengei Air Tawar, bb 1687 (Bo). Dsamsi: Djambi, Sei Oeron, 43 m., bb 12277 (Bo). SELATAN: Banjoeasin en Koeboestreken, 15 m., 134 E | P 1036 (Bo, L); Koemering Oeloe, 80 m., Grashoff 565 (Bo); Kaju Agung, Kostermans 14089 (L). BANGKA: Lobok Besar, G. Pading, 20 m., Kostermans & Anta 964 (A, BO, K, KEP, LAE, NY, SING); G. Mangkol, 50 m., Kostermans & Anta 716 (A, BO, K, LAE, NY, P, SING); prope ane Teysmann, HB 3214, pro parte (po); G. Maras, 2 Kostermans & A 41 (A, BO, CANB, K, KEP, LAE, P, SING); Belinjoe, 28 m., Grashoff 72 es. “ ae Majang, bb 7302 (Bo); Soengei Liat, Tey smann, HB 7565 (c, Fl, MEL), G. Rebo 5 m., Bunnemeijer 1886 (Bo, L); Plangas, Teysmann, HB 3251 (so); S. Bangka, Mindik, 25 m., bb 10689 (Bo), Toboali, bb 1941 (Bo, L). BeLtituNc: Tandjong Padan, Bantan, : bb 7377 (Bo); N. von Manggor, Ham 18 (Bo). He and adjacent islands. SarAwAK. Ist Division: Mt. Santubong, S 21510 (f) (a, Bo, K, KEP, P, SAN, SAR, SING); Matang, Beccari PB 1647 (f) (FI); ae Natl. Park, summit of Bukit Gondol, 270 m., S 20924 (a, kK, SAN, SaR, sING); Serian, G. Penrissen, S 16306 (f) (A, BO, K, SAN, SAR, SING); G. Buri, 490 m., S 36950 (f) (sar). Maranga on Tandjung Plateau, 200 m., Kostermans 12542 (Bo, K, L, P, SING); E. Koetai, Godong Lengah (Bengalon), 15 m., bb 7952 (Bo); Loa Djanan, W. of Samarinda, Kostermans 9868 (A, Bo, % KEP, L, P, SING); Sg. Tiram complex, SW. of Samarinda, Kostermans 6056 (A, BO, K, L). Selatan: Bangar- massing, Motley 561 (kK, P); Kintap, Pleihari, Dachlan 2049 (Bo, L); Martapoera, Kalaan, 400 m., bb 12040 (Bo), G. Melati, Dachlan 2156 (so); G. Pamatton, 1980] STEVENS, CALOPHYLLUM 399 Korthals s.n. (f) (L). Tengah: Karrau R., S. Miiller s.n. (L); Poeroek Tjahoe, M [oear]a Djaan, 100 m., bb 10485 (Bo), M[oear]a Laoeng, 80 m., bb 10099 Kelapeh, 200 m., bb 1033 (po). Barat: Karimata Arch., Poelau- Seroetoe, Mondi{h] 170 (f) (BO, K, L, NY, SING, U); Pontianak, Dedi 865 (Bo); Melawi Tjatit, B. Gontoek, 180 m., bb 27014 (f) (A, BISH, BO, K, NY, P, SING); B. Tengkoejoeng, 450 m., bb 25115 (so, 1, mo); Afd. Beneden Matan, S. Kendawangan, 10 m., bb 14428 (Bo, L). Jav va. Barat: cult. in hort. Bogor. (seed from Bangka) sub VI C 47 (Bo, BRI, G, K, LAE, MO, NY, P, SING, US). Ecotocy. Mixed dipterocarp forest, to 300(-590) m. alt. Sometimes on sandy soil; on beach; on dry land with patches of low scrub; in swamp forest. Flowering all year, mostly June to September (flower scented, sometimes foetid (Ridley 13305)); fruiting October to June (fruit dull green). Fruit eaten by fruit bat (Pteropus edulis) in Malaya oe 1930); however, this report may properly refer to fruits of C. tetrapte Anthony (1974) reported several ik on a Citeeree pulcherrimum in Singapore. A psyllid made the leaf revolute with undulations and crisped edges; the midrib became abaxially curved, and the petiole twisted. A possibly undescribed species of the midge genus Bruggmanniella caused fleshy spheroid galls, which were often attached to the midrib on the abaxial surface of the leaf; another midge gall caused swellings of the stem, petiole, and midrib. A moth larva formed an elongated swelling of the lamina roughly parallel to the midrib. GERMINATION AND YOUNG PLANT. The radicle emerges after the stone wall to one side of the base has been broken. The seedling has usually three, but sometimes up to five, pairs of scale leaves and two pairs of normal leaves, all separated by well-developed internodes; the first pair of scale leaves is borne 0.5-1.8 cm. from the cotyledonary node. The terminal bud may produce the first flush of leaves; scale leaves are not produced at the beginning of this flush. However, in the forest even the first flush is often produced by an axillary bud (the terminal bud being damaged?), subsequent flushes seem always to be produced by axillary buds, and there are scale leaves at the bases of the innovations. The young plant is strongly arched, but as the plant gets older the lower part of the stem straightens. (Stevens et al. 47; seedlings from Stevens et al. 333 grown at Kuching but unfortunately later eaten by a dog. Details of the young plant sent by P. J. Martin.) Locat uses. The wood is good for poles, fishing stakes, masts, and similar objects, and is also used in construction and for ships’ knees. The fruit is sour but edible. Specimens of Calophyllum pulcherrimum can be recognized by their usually nonfunctional terminal buds, axillary innovations with basal scars, and leaf blades with the midrib flat to depressed below and broadly raised and continuous with the leaf surface above. The fruit is ovoid, at least 1.5 cm. long, and strongly wrinkled when dry; the outer layer of the fruit is 1-2.2 mm. thick, and the stone walls are 0.3-0.5 mm. thick. The epithet pulcherrimum means ‘‘most beautiful,’’ a not inappropriate name for the often very floriferous specimens of this species. 400 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 Sterile or flowering specimens of Calophyllum pulcherrimum are very distinguished fairly easily. The differences between the three species are summarized in TaBLE 11, but work is needed to confirm the possible differences in seedling and in the growth of the young plant. All three species were found growing by a logging road at Sabal Tapang (Sarawak), and in undisturbed forest there trees of C. pulcherrimum and C. depressinervosum were growing within 30 meters of each other on a ridge. Calophyllum pulcherrimum has often been confused with C. floribundum (= C. tetrapterum var. tetrapterum), and although Henderson and Wyatt-Smith (loc. cit.) clearly separated the two, the confusion still seems to persist, to judge by determinations on specimens. The two species can be separated very easily, even in the absence of flowers and fruits. Calophyllum tetrapterum has a functional terminal bud, its axillary innovations usually lack basal scars, less than 1.3 cm. long and lack the relatively thick outer layer and stone walls of those of C. pulcherrimum. Although the leaf margin of C. tetrapterum is often markedly paler than the rest of the blade when dry, this is not always the case, and the margin of the blade may be a similarly pale color in some specimens of C. pulcherrimum. The confusion between the two species is of long standing. There are specimens of Calophyllum floribundum sensu stricto in some herbaria mounted with the label of Wallich dist. 4845, the type number of C. pulcherrimum. One of the three numbers collected by Maingay and cited as C. floribundum in the original description of that species is a specimen of C. pulcherrimum (see the discussion after C. tetrapterum). Some synonyms of C. tetrapterum have been cited under C. pulcherrimum, and Boerlage (loc. cit.) reduced distributed (cult. Hort. Bogor. sub V/ C 47). Finally, on one sheet of Pierre 3645 at Paris (C. tetrapterum var. tetrapterum, collected in Cambodia), there of this species on Pierre 3645 probably came from Maingay Pierre (/oc. cit.) used to illustrate the fruits of his C. pulcherrimum (properly C. tetrapterum var. tetrapterum). 61. Calophyllum sp. Tree ca. 4.5 meters tall; trunk and bark unknown. Twigs slightly flattened, 1.2-1.8 mm. across, rounded to obscurely 4-angled, drying brown, sparsely puberulent when very young; axillary innovations with basal scars; internodes 0.8-3.2 cm. long, first-developed internode often 1980] STEVENS, CALOPHYLLUM 401 twice as long as others; uppermost pair of axillary buds rounded, ca. 0.6 mm. long, suberect, sometimes subglabrous; terminal bud plump, 1-1.5 mm. long, with subadpressed, brown indumentum (hairs, Ficure 22, 1), underde- veloped internode to 1 mm. long. Petiole 3-4 mm. long, concave above and convex below, puberulent at edge when very young; lamina obovate, 2-3.7 by 0.9-2.3 cm., rounded to shallowly retuse at apex, cuneate at base, slightly undulate and not recurved at margin, coriaceous, drying umber above and umber or sepia below, glabrous even when young, the midrib above abruptly narrowed at base, flat at first, becoming slightly raised, 0.15-0.25 mm. wide at midpoint, below slightly raised, striate, inconspicuous, the venation apparent above and below, raised, 5 to ns/5 mm., angle of divergence 45-55°. Inflorescences from foliate axils, ies Pio i flowers: unbranched, the axis 4-6 cm. long, glabrous, lowest internode and first pair of flowers not developed, next internode 1.8-3.5 cm. long, others 0.8-1.5 cm. long; bracts foliaceous, to 2.6 by 2 cm., persistent; pedicels ca. 8 mm. long, glabrous. Flower (?)hermaphroditic, known only in bud; tepals 8; stamens ca. 120, the anthers elliptic, ca. 0.3 mm. long; stigma peltate. Fruit unknown. DistriBuTION. Known only from one collection made in the 4th Division, Sarawak (Map 20 SPECIMEN SEEN. Sarawak: Baram, G. Api, 1219 m., 12 July 1961, S 4732 coll. Anderson (K, SAR). Eco.tocy. Exposed submontane forest on limestone hill, 1219 m. alt. Calophyllum sp. 61 is a distinctive taxon. It has a small terminal bud, axillary (?and terminal) innovations with basal scars, and small leaf blades in which the midrib is flat on the upper surface and more or less raised on the lower. The axillary inflorescences, with their basal scars, foliaceous, persistent bracts, and long, first fully developed internodes are striking. However, until the fruits are known, it seems premature to name this taxon. 62. reales pelewense P. F. Stevens in Fosberg & Sachet, Smithson. Contr. Bot. 45: 12. 1980. Type: Pelew Islands, July—Aug. 1929, Kanehira 386 nee A; isotypes, BISH, NY). Ficure 24, f. C. cholobtaches auct., non Lauterb.; Kanehira, Bot. Mag. Tokyo 45: 329. 1931, FI. Micronesica, 233. fig. 105. 1933, Jour. Dept. Agr. Kyushu Imp. Univ. 4: 370. 1935. Tree; trunk and bark not known. Twigs flattened, 1.5-2 mm. across, strongly 4-angled to subalate, drying brown, glabrous; axillary innovations with basal scars; internodes (0.5—)1-3 (-5.5) cm. long; uppermost pair of axillary buds rounded, ca. 0.5 mm. long, spreading; terminal bud plump, 1I-2.5 mm. long, with short, gray, adpressed indumentum (hairs, Ficure 22, g, h), underdeveloped internode 1-2.5 mm long. Petiole 0.9-1.5 cm. long, shallowly and narrowly to broadly concave above, convex below, glabrous; lamina elliptic to ovate (rarely siete! 3.5-10.2 by 2.1-4.9 cm., acuminate (rarely + rounded) at apex, acute 402 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 attenuate at base, undulate and slightly recurved at margin, coriaceous, drying umber on both surfaces, glabrous, the midrib above gradually narrowed from base, not depressed or slightly so, the edges not raised or slightly so, 0.35-0.6 mm. wide at midpoint, below raised, + striate, the venation subapparent on both surfaces, raised, 5 to 10 veins/5 mm., angle of divergence 55-70°. Inflorescences from foliate axils (rarely terminal), with 9 to 17 flowers, unbranched, the axis 1.3-8 cm. long, puberulent at base, lowest internode to 2 mm. long; bracts subobovate, ca. 3.5 mm. long, soon deciduous; pedicels 1-2.5 cm. long, glabrous. Flower (?)hermaphroditic; tepals 8 to 11, the outer pair broadly ovate, ca. 4 mm. long and across, the inner ones obovate, 5.5-6 by 3.5-4.5 mm., glabrous; stamens 125 to 230, the filaments ca. 3 mm. long, the anthers suboblong, 0.8-1 mm. long, retuse at apex; ovary ca. 2 mm. long, the style ca. 2.2 mm. long, the stigma peltate, 0.7-1.2 mm. across, + 3-radiate. Fruit probably cllipsosd, ca. 1.5 by | cm., rounded at apex, drying vinaceous-brown, wrinkled; outer layer (?)not deischine cleanly from stone, ca. 0.5 mm. thick, with air spaces developing; stone with walls ca. 0.2 mm. thick; spongy layer unknown. DistripuTion. The Caroline Islands, Palau group (Map 20). ADDITIONAL SPECIMENS SEEN. Caroline Islands: Korror, Raymondus 334 (7) (wrsL); Palau, Garusamao, Takamatsu 1556 (BisH, us); Amiliki [Aimiriik], Hosokawa 7216 (BIsH); Aimiriik, Kanehira 1984 (k, p, Ny, us); Almatan [Armitin], 8 m., Takamatsu 1525 (sisu); Airai, Kanehira 2380 (ny); Ngerikill R., Fisher 25 (us), 55 (us); sine loco, Ledermann 14489 (x). Eco.ocy. River banks at landward edge of mangroves. Flowering in March (flower scented), submature fruit in March, young fruit July and August. Calophyllum pelewense can be recognized by its very short terminal bud; axillary innovations with basal scars; axillary inflorescences with a short basal internode, usually elliptic to ovate leaf blades with rather prominent, distant venation, flowers with eight to eleven tepals; and fruits of moderate size (ca. 1.5 cm. long). The epithet pelewense is derived from Pelew (Palau) Islands, where this species grows. Specimens of Calophyllum pelewense have been labeled C. cholobtaches (= C. soulattri), but the two species are not at all close. Calophyllum soulattri has a longer terminal bud, a greenish-drying lamina, a branched inflorescence, flowers with four tepals, and a spherical fruit with a thick, compact outer layer. 63. Calophyllum pyriforme P. F. Stevens, sp. nov. Figure 23, a-c. A speciebus aliis Calophylli quibus fructibus stipitatis habent in lamina longiore magnis tenuiter coriacea apice longe acuminata pagina infra indumento subpersistenti praedita et fructu apice rotundato, non acuto, differt. Tree 4.5-7.5 meters tall, d.b.h. to 9 cm.; trunk without buttresses; outer bark brown, yellowish, or gray-black, sometimes with shallow fissures, exfoliating in thin scales; inner bark red-brown or yellowish white; latex yellow. 1980] STEVENS, CALOPHYLLUM 403 GURE 23. a-c, Calophyllum pyriforme: a, habit (SAN 64779), x 0.5; b, terminal bud (SAN 21748), x 0.75; c, fruit (SAN 19092), longitudinal section, x 1. d-f, Calophyllum sp. 65 (S 23854). d, e, fruit, x 1.25: d, from outside; e, longitudinal section. f, terminal bud, x 6. Twigs slightly flattened, 2-2.5 mm. across, slightly 4-angled, drying brown, at first tomentose, glabrescent; axillary innovations with basal scars, or scars up to 1.8 cm. from base; internodes (1-)1.5-7(-8) cm. long; uppermost pair of axillary buds rounded, to 2 mm. long, + spreading, not conspicuous; terminal bud conical, 4.5-11 mm. long, with short, tomentose, brown indumen- tum (hairs, Ficure 27, c), underdeveloped internode absent. Petiole 1-1.7 cm. long, narrowly channeled above, convex below, subpersistently tomentose; lamina elliptic to suboblong, 11.5-23 by (3.7-)5-8.8 cm., abruptly acuminate at apex, acumen to 1.5 cm. long, acute at base, undulate and sharply but 404 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 narrowly recurved at margin, thinly coriaceous, drying umber to olivaceous above and honey to pale olivaceous below, tomentose on midrib above and over entire lower surface, the midrib above abruptly narrowed near base, raised, sulcate at first, 0.1-0.2 mm. wide at midpoint, below raised, substriate, the venation apparent to subobscure on both surfaces, raised, 6 to 9 (to 11) veins/5 mm., angle of divergence 70-80°. Inflorescences from foliate axils (very rarely terminal), with 5 to 9 flowers, unbranched, the axis 1.2-3.5(-7.5 in fruit) cm. long, subpersistently tomentose, lowest internode Gbeeni to) 3-8 mm. long; bracts narrowly ovate, ca. 4 mm. long, deciduous; pedicels 0.8-1.2(-2.2) cm. long, subpersistently tomentose. Flower (?)her- maphroditic; tepals 8, the outer pair broadly ovate to elliptic, 5-6 by 4-4.5 mm., short-tomentose on back, the next pair elliptic, 8.5-9.5 by 4-6 mm., the inner ones narrowly elliptic, 8-10 by ca. 3 mm. (rarely ca. 7.5 by 4 mm.); stamens 190 to 265, the filaments to 5.7 mm. long, connate for up to 2 mm., the anthers elliptic-oblong, 0.5-0.8 mm. long, shallowly retuse at apex; ovary ca. 2 mm. long, the style 4-5 mm. long, the stigma peltate, ca. 0.6 mm. across, 4-radiate. Fruit obovoid, 1.7-3 by 1.3-1.6 cm., apiculate, stipe fibrous and 6-10 by 5-7 mm., drying dark brown, sharply wrinkled; outer layer not detaching cleanly from stone, 0.6-1 mm. thick, compact, fibrous; stone ellipsoid, 1.6-1.75 by 1.1-1.3 cm., rounded at apex, the walls less than 0.1 mm. thick, smooth, (?)unmarked; spongy layer thin, Tyre: North Borneo [Sabah], Bettotan, near Sandakan, 10 Aug. 1927, SFN 19092 coll. Boden Kloss (holotype, L; isotypes, K, LAE, NY, SING, UC). DistripuTion. Eastern Borneo (Map 23). Map 23. Distribution of Calophyllum flavo-ramulum (erect triangles), stipitatum (stars in solid circles), C. glaucescens (half-circles), pein ee sp. 65 (solid stars), and C. pyriforme (inverted triangles) in Malesia. Inset: C. elegans (solid circles), C. roseocostatum (open circle). 1980] STEVENS, CALOPHYLLUM 405 ADDITIONAL SPECIMENS SEEN. Borneo. SABAH. Labuk & Sugut: Mile 80 Labuk Road, 150 m., SAN 38780 (san). Kinabatangan: timber camp at Daramakud, SAN 16863 (A, BO, KEP, L, SING); Brassey Range, 620 m., Stevens et al. 475A (a); G. Rara F.R., ca. 32 km. NW. of Luasong camp, 366 m., SAN 75782 (K, L, SAN, SING); ca. 48 km. NW. of Luasong camp, 488 m., SAN 75654 (A, L, SAN, SING); Old Road, Luasong, 27 m., SAN 64779 (xk, L, sAN). Lahad SAN 65829 (K, L, SAN). KALIMANTAN, Timur: Berau, near Telukbajur, Koster- mans 21748 (Bo, CANB, G, L, P, SAR, SING). EcoLocy. Along river banks or in sagen inundated forest, also hilly country, sometimes on sandy soil; to 620 m. alt. Flowering March and April; fruiting August and September (fruit pale a. Calophyllum pyriforme is perhaps the most distinctive species in the group of taxa that have stipitate fruits with a fibrous outer layer and a thin-walled stone; these species all have scars at or near the bases of the axillary innovations (see TaBLe 12). It can be recognized by its rather large leaf blades that are long-acuminate at the apex and that have subpersistent, tomentose indumentum on the lower surface; by the tomentose indumentum on the inflorescence axis, pedicels, and the backs of the outer tepals; and by its fruits, which are rounded at the apex. The hairs are distinctive, with the thickening of the walls so poorly developed that the cross walls are invisible (FicurE 27, b). The pear-shaped fruits suggested the specific epithet. Calophyllum pyriforme has been confused with C. rubiginosum; however, the two species are only superficially similar and are not closely related. Calophyllum pyriforme has twigs that dry brown, not yellow; inflorescences that are nearly always axillary, rather than terminal; and anthers that are glabrous, not hairy. The pyriform fruits of C. pyriforme are not furfuraceous, while the ellipsoid fruits of C. rubiginosum are. Some specimens appear to have rather regular variation in internode length along the main stem. Internodes are periodically shorter; the leaves at the nodes above these short internodes are soon deciduous. These shorter internodes are probably the first of the flush. The presence of scars at the start of axillary innovations is not as constant as it is in other members of this group. 64. Calophyllum roseocostatum P. F. Stevens, sp. nov. Ficure 21, g-i. A speciebus aliis Calophylli quibus fructibus stipitatis habent in ramulis tenuis 1.3-2.2 mm. in transversis, internodio terminali innovationis aliis subaequanti, lamina in siccitate nitida costa (rubro-)brunnea cetera paginae olivacea, et fructu apice saepe rotundato, differt. Tree 14-27 meters tall, d.b.h. to 70 cm.; trunk without buttresses (spurred); outer bark yellow-brown to grayish, with deep, distant, boat-shaped fissures, the inner surface orange-red or blackish; under bark dark red to chestnut; inner bark dark red; latex yellow, opaque, not viscous. 406 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Twigs somewhat flattened, 1.3-2.2 mm. across, 4-angled, drying brown, scars within 5 mm. of base; internodes 1-5 cm. long; uppermost pair of axillary buds rounded, less than 1 mm. long, subspreading; terminal bud narrowly conical to + plump, 2.5-4 mm. long, with compact, brown indumen- tum (hairs, Figure 22, k; cf. 22, 1), underdeveloped internode to 3 mm. long. Petiole 1-1.6 cm. long, slender, concave above and convex below, glabrous; lamina elliptic to oblong, 5.7-11.1 by 2.3-3.1 cm., acuminate at apex, narrowly cuneate to acute at base, undulate and slightly recurved at margin, coriaceous, drying olivaceous and shiny above, greenish olivaceous-buff below, glabrous, the midrib gradually narrowed from base, slightly raised, center + sulcate, 0.25-0.4 mm. wide at midpoint, below raised, striate, the venation apparent on both surfaces, raised, 11 to 15 veins/5 mm., angle of divergence 65-75°. Infructescences from foliate axils, with scars of 5 to 7 flowers, not branched, the axis 0.7-2 cm. long, puberulent, lowest internode to 2 mm. long; bracts not known; pedicels 1-1.5 cm. long, puberulent (glabrous toward apex). Flowers not known; tepals (from scars) 8. Fruit obovoid, 2.6-4.2 by 1.7-2 cm., rounded to obtuse at apex, stipe fibrous and 0.6-1.2 cm. by ca mm., drying brown, wrinkled; outer layer not detaching cleanly from stone, ca. 1.5 mm. thick, compact, but air spaces developing under skin; stone ellipsoid, ca. 2 by 1.3 cm., rounded at apex, the walls less than 0.1 mm. thick, smooth, unmarked; enonsy layer becoming thin. Type: Sarawak, Kuching, Semengoh F.R., 90 m., Sept. 1961, § 1494] coll. Galau (holotype, k; isotypes, L, SAR). DistripuTion. Northwestern Borneo, known only near Kuching (Map 23). ADDITIONAL SPECIMENS SEEN. Borneo. SARAWAK. Ist Division: 12th Mile, Penrissen pe Semengoh F.R., S 26299 (a, K, L, SAN, SAR, SING), S 25388 (K, L, SAR, SING), 60 m., S 14 760 (K, SAR); outside Arboretum, S 3354] (a, SAN, SAR), ee et al. 136 (A). Eco.tocy. Well-drained mixed dipterocarp forest, ca. 60 m. alt. Fruiting April, September, and October Calophyllum roseocostatum can be recognized by its slender twigs, the scars of the bases of the axillary innovations, its short terminal buds, and its medium-sized, elliptic to oblong lamina with the midrib on the upper surface drying reddish brown and almost level. The fruits are stipitate. The epithet alludes to the color of the midrib on the upper surface of the dried lamina. Calophyllum roseocostatum is a member of the group of taxa centered around C. pyriforme; the differences separating the taxa in the group are given in Taste 12. Further collections may change the rank of the taxa recognized here; it is perhaps a little surprising that C. roseocostatum is at least locally quite common around Kuching, but has not yet been found elsewhere. TasLe 12. Differences between Calophyllum pyriforme and its relatives. C. pyriforme — C. stipitatum C. roseocostatum Calophyllum sp. 65 C. elegans Hairs BRANCHED, ALL CELLS UNTHICKENED + = _ = _ GENERAL INTERNODE LENGTH (cm.) 1.5-7 1,2-3.5 1-5 1-3.5 1.5-3 TERMINAL INTERNODE LENGTH (cm.) 1.5-7 0.2-0.6 1-5 1-3.5 (0.3-)0.5-1 TWIG THICKNESS (mm.) 2-2.5 2-2.5 1.3-2.2 2-2.3 1.6-2.5 TOMENTUM SUBPERSISTENT ON STEM AND LOWER SURFACE OF LEAF BLADE + = =a = = LEAF BLADE LENGTH (cm. 11.5-23 5.4-10.5 5.7-11.1 6-9,.2 (3-)4.6-8.5 MIDRIB NARROWING GRADUALLY FROM BASE = + + + + MIpRIB SHARPLY RAISED — — — + + VENATION LESS PROMINENT ON LOWER THAN ON UPPER SURFACE — _ _ + —+ TEXTURE Thinly Coriaceous Coriaceous Coriaceous Coriaceous coriaceous APEX ong- Acuminate Acuminate Acuminate acuminate subacuminate FRUIT ROUNDED AT APEX + + +(—) aa (?)- GEOGRAPHIC DISTRIBUTION . Saba Malaya SW. Sarawak NE. Sarawak SW. Sarawak E. Kalimantan (?)Brunei WN TIAHdOTVD ‘SNAHAALS [0861 LOP 408 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 65. Calophyllum sp. Ficure 23, d-f. Tree ca. 21 meters tall, d.b.h. ca. 48 cm.; trunk and bark unknown. Twigs slightly flattened, 2-2.5 mm. across, strongly 4-angled, drying brown, glabrous when mature; axillary innovations with basal scars; internodes 1-3.5 cm. long; uppermost pair of axillary buds rounded, ca. | mm. long, + spreading; terminal bud plump, 2-3 mm. long, with brown, subadpressed indumentum (hairs, cf. Figure 22, k), underdeveloped internode ca. 2 mm. long. Petiole 6-10 mm. long, deeply concave above, convex below, glabrous; lamina obovate to elliptic, 6-11 by 2.4-4 cm., acuminate at apex, cuneate to acute at base, strongly undulate but only slightly recurved at margin, coriaceous, drying with + persistent, grayish, waxy covering, olivaceous above and sabelline below, midrib and margin fulvous on both surfaces, glabrous at maturity, the midrib above narrowing gradually from base, raised, 0.2-0.3 mm. wide at midpoint, below raised, striate, the venation subapparent, clearer above than below, raised, 12 to I5 veins/5 mm., angle of divergence ca. 70°. Infructescences from foliate axils, with scars of 5 to 9 flowers, unbranched, the axis 2.3-4 cm. long, glabrous, lowest internode to 0.5 mm. long; bracts unknown; pedicels |.6-2 cm. long, glabrous. Flower unknown. Fruit obovoid- ellipsoid, 2.6-3.5 by 0.7-1.3 cm., sharply pointed at apex, stipe 1-1.3 cm. by ca. 8 mm., drying tawny, wrinkled; outer layer not detaching from stone, ca. | mm. thick, compact, fibrous; stone ca. 1.15 by ca. | cm., rounded at apex, the walls ca. 0.2 mm. thick, smooth, unmarked; spongy layer initially thick. DistrRiBUTION. Known only from Sarawak (Map 23). REPRESENTATIVE SPECIMENS SEEN. Borneo. Sarawak. 3rd Division: Bukit Raya, Kapit, Sibu, 180 m., S 23854 (4, Bo, K, KEP, L, SAN, SAR, SING). 4th Division: Bukit Mersing, Ashton 6005 (a). Eco.ocy. Ridge forest, 180 m. alt. Submature fruit in October. Calophyllum sp. 65 can be recognized by its axillary innovations with basal scars, its small terminal bud and relatively long underdeveloped internode, and its lamina, which has a grayish covering, a fulvous midrib, and venation that tends to be more prominent on the upper than on the lower surface. The fruits are stipitate and have a fibrous outer layer and a thin-walled stone. Calophyllum sp. 65 is closely related to such species as C. stipitatum and C. pyriforme; the differences distinguishing it from these taxa are given in TABLE 12. 66. Calophyllum stipitatum P. F. Stevens, sp. nov. Ficure 21, a, b. speciebus aliis Calophylli quibus fructibus stipitatis et innovationibus axillaribus cicatricibus basalibus habent in lamina mediocra in siccitate infra olivacea costa supra plus minusve depressa, et fructu apice rotundato, differt. 1980] STEVENS, CALOPHYLLUM 409 Tree ca. 18 meters tall, d.b.h. to 38 cm.; trunk probably without buttresses; outer bark + smooth, with broad, shallow, distant, boat-shaped fissures; latex yellow, clear, greenish near cambium. Twigs flattened, 2-2.5 mm. across, 4-angled or not, drying blackish, at maturity smooth, glabrous; axillary innovations with one or two pairs of basal scars; internodes 1.2-3.5 cm. long, terminal internode 2-6 mm. long; uppermost pair of axillary buds rounded, to 1 mm. long, spreading; terminal bud broadly conical, 2.3-3 mm. long, with short, tomentose, brown indumen- tum (hairs, Ficure 22, 1), underdeveloped internode to 2 mm. long. Petiole 0.7-1.5 cm. long, broadly concave above and convex below, glabrous; lamina elliptic to obovate, 5.4-10.5 by 2.3-4 cm., acuminate at apex, narrowly acute to long-attenua te at base, margin rather distantly undulate and slightly recurved, coriaceous, drying with grayish covering, sepia above and olivaceous below, the margins pale, glabrous at maturity, the midrib above gradually narrowed from base, + depressed, edges slightly raised, 0.2-0.35 mm. wide at midpoint, below raised, striate, the venation subobscure to apparent on both surfaces, raised, latex canals on lower surface subraised to clearly impressed, 7 to 10 veins/5 mm., angle of divergence 65-70°. Infructescences arising from foliate axils, with scars of 9 to 13 flowers, unbranched, the axis 2.7-4 cm. long, sometimes puberulent at base, lowest internode less than 1 mm. long; bracts not known; pedicels 0.8-1.4 cm. long, glabrous. Flower not known. Fruit (submature) obovoid, ca. 1.8 by 0.9 cm., subpersistently apiculate, stipe fibrous and ca. 8 by 4 mm., drying dark brown with apical part smooth and stipe closely wrinkled; outer layer not detaching cleanly from stone, ca. 0.5 mm. thick, compact, fibrous; stone ellipsoid, ca. 9 by 7.5 mm., rounded at apex, the walls ca. 0.1 mm. thick, smooth, (?)unmarked; spongy layer thick in young fruit. Type: Malaya, Pahang (southeast), Aur forest, reserve at Sungei Aur, flood plain Sungei Aur, 11 May 1967, FRI 3656 coll. Whitmore (holotype, k; iso- types, A, KEP, L, SAR, SING). DistrisuTIoN. The Malay Peninsula, probably also northwestern Borneo, local (Map 23). ADDITIONAL SPECIMENS SEEN: Borneo. Brunei: Andalau, Ashton s.n. (SAR); Kuala Belait, S 10681 (sar). Eco.Locy. Lowland rainforest. Submature fruit in May. Calophyllum stipitatum is clearly related to the other species from Borneo that have stipitate fruits, scars at the bases of the axillary innovations, and inflorescences with very short basal internodes (C. elegans, C. pyriforme, species are listed in Taste 12. The epithet stipitatum alludes to the stipitate ruits. Although the specimens from Brunei cited above as Calophyllum stipitatum have several pairs of scars at the bases of the axillary innovations, they agree with that species, rather than with Calophyllum sp. 65, in vegetative 410 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 characters. However, the identity of these specimens can be confirmed only by the collection of fruiting material. 67. Calophyllum elegans Ridley, Kew Bull. 1938: 118. 1938; Masamune, Enum. Phanerog. Born. 475. 1941; Smythies, Common Sarawak Trees, 59. 1965. Type: Sarawak, near Kuching, 8 Jan. 1892, Haviland 968 coll. Garai (holotype, kK; isotypes, BM, L, P, SAR). Tree 6-22 meters tall, d.b.h. to 20 cm.; trunk without buttresses; outer bark brown to yellowish brown, with broad, shallow, boat-shaped fissures, not hoop marked, the inner surface deep red to reddish brown; under bark deep red to chestnut brown; inner bark pale red; latex opaque to clear yellow, sticky. Twigs slightly flattened, 1.6-2.5 mm. across, + 4-angled when young, later with 6 + raised lines, drying brown to blackish, sparsely puberulent when young; axillary innovations with pair of (sub)basal scars, terminal innovations with ca. 3 pairs of near basal scars; internodes 1.5-3 cm. long; uppermost internode of innovation usually (0.3-)0.5-1 cm. long; uppermost pair of axillary buds rounded, to 1.5 mm. long, spreading; terminal bud plump, 3-3.5 mm. long, abruptly narrowed at apex, with short, subadpressed, brown indumentum (hairs, Ficure 25, c, d), underdeveloped internode to 2 mm. long. Petiole (0.4-)0.8-1.5 cm. long, deeply concave above, convex below; lamina elliptic to oblong or subobovate, (3-)4.6-8.5 by (1.3-)2.7-4.7 cm., retuse to obtuse (subacuminate) at apex, cuneate to acute at base, slightly undulate and recurved raised, rounded toward apex, striate toward base, the venation rather apparent above, apparent to subobscure below, raised, 6 to 11 veins/5 mm., angle of divergence 70-80°. Inflorescences from foliate axils, with 9 to 11 flowers, unbranched, the axis 1.2-3 cm. long, glabrous, lowest internode lacking; basal bracts broadly ovate, ca. 1.7 mm. long, coriaceous, persistent, upper bracts ovate-elliptic, to 3 mm. long, chartaceous, subpersistent; pedicels 7-9 mm. long, glabrous. Flower (?)hermaphroditic; tepals 8, glabrous, the outer pair ovate to elliptic, 3-3.5 by 2.2-3 mm., + keeled, the next pair subobovate, ca. 5 by 2.2 mm., the innermost ones oblong-elliptic, to 3.5 by 1.4 mm; stamens ca. 95, the filaments to 3.5 mm. long, the anthers elliptic-oblong, 0.6-1 mm. long, retuse at apex; ovary ca. 1.8 mm. long, the style ca. 1.8 mm. long, the stigma infundibular, ca. 0.3 mm. across. Immature fruit ellipsoid-obovoid, shape distorted by drying, ca. 2.2 by 0.8 cm., sharply apiculate, stipe ca. 8 mm. long. DistriBuTION. Western Sarawak, local (Map 23). SELECTED SPECIMENS SEEN. Borneo. Sarawak. Ist Division: Semengoh Arbore- tum, 15 m., S 5383 (A, BO, K, KEP, L, P, SAN, SAR, SING); G. Matang, 580 m., Stevens et al. 274 (a); Sempadi F.R., 15 m., S 6095 (sar); Sabal Tapang, 190 m., Stevens et al. 185 (a); Stapok F. R., 10 m., Stevens et al. 153 (a). 1980] STEVENS, CALOPHYLLUM 411 Ecotocy. Edge of swamp forest, mixed dipterocarp forest on sandy clay soil, and kerangas forest; 10-580 m. alt. Flowering January and February (flower scented). Calophyllum elegans is characterized by its rather small, thinly coriaceous a Tib o pp internode of each innovation is nearly always markedly ae than the others, the terminal pair of leaves being smaller than the rest (figures in parentheses in the description above). Although only immature fruits are known, they are clearly of the stipitate type. The specific epithet elegans (‘‘elegant,’’ ‘‘fine’’) is appropriate for the neat-looking specimens of this species. Calophyllum elegans is related to the groups of species centered on C. pyriforme. For the differences separating it from other taxa in this group, see TABLE 12. seer Calophyllum elegans i is superficially similar to some of the forms lack basal scars; the terminal bud is not acutely pointed; the lowest internode of the inflorescence is more or less well developed, and if the inflorescence bracts are subpersistent, they are never small and coriaceous; the fruit is subspherical and lacks a stipe. 68. Calophyllum oliganthum Merr. Philip. Jour. Sci. 17: 289. 1920, Enum. Philip. Fl. Pl. 3: 80. 1923. Type: Philippine Islands, Mindanao, Surigao del Norte Province, Bucas Grande Island, 10 June 1919, BS 35110 coll. Ramos & Pascasio (isotypes, A, BM, BO, K, P, US). Tree 1-5 meters tall, d.b.h. to 10 cm.; trunk and bark unknown; branches pendulous. Twigs slightly flattened, 0.6-1.3 mm. across, not angled, drying blackish when young (grayish when older), glabrous or + transiently puberulent; axillary innovations (?)always lacking basal scars; internodes 1.5—7 cm. long; uppermost pair of axillary buds rounded, 0.5-1 mm. long, suberect, not conspicuous; terminal bud conical, (1.5-)2-3 mm. long, with adpressed indumentum (hairs, Ficure 22, c, d), underdeveloped internode absent (to 2 mm. long). Petiole (0.4-)0.6-1.1 cm. long, shallowly concave above, convex below, usually becoming glabrous; lamina elliptic to ovate (rarely obovate), (2-)3-8 by (0.7-)1.2-3.5 cm., acuminate at apex, (narrowly) cuneate at base, not undulate to slightly so and recurved at margin, coriaceous, drying sepia to sabelline-sepia above, umber to fulvous below, glabrous or with few hairs on midrib below when very young, the midrib above narrowing gradually from base, flat, obscure, surrounding lamina + raised, 0.2-0.3 mm. wide at midpoint, below slightly raised to raised, rounded, the venation subobscure above, + apparent 412 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 and raised below, 8 to 15 veins/5 mm., angle of divergence 60-70°. Inflores- cences from foliate axils along stem, with 3 to 5 (to 7) flowers, unbranched, the axis |-2.5(-3.7) cm. long, sometimes sparsely and transiently subtomentose, at least toward base, lowest internode 0.3-1(-1.7) cm. long; bracts ovate, ca. 1.5 mm. long, caducous; pedicels (0.6-)1-1.8 cm. long, glabrous. Flower (?)hermaphroditic; tepals 4 to 8, the outer two suborbicular, ca. 4 mm. long and across, the next two broadly elliptic, ca. 6 by 4-4.5 mm., others, if present, probably much smaller; stamens 45 to 80, the filaments to 3.5 mm. long, the anthers suboblong, 0.5-1 mm. long, retuse at apex; ovary ca. | mm. long, the style ca. 4.5 mm. long, very slender, the stigma ca. 0.4 mm. across, 2- or 3-radiate. Fruit ellipsoid, 1-1.3 by ca. 0.8 cm., rounded at apex, drying dark brown, + smooth; outer layer not detaching cleanly from stone, less than 0.5 mm. thick, air spaces developing; stone ellipsoid, 8-11 by 5-7 mm., rounded at apex, the walls 0.15—0.3 mm. thick, smooth, unmarked; spongy layer thin. DistripuTIon. The Philippine Islands, Mindanao (Map 20). SELECTED SPECIMENS SEEN. Philippine Islands. Minpanao. Surigao del Norte: Dinagat Is., BS 83812 (Ny); sine loco, BS 34520 (a, kK, P, US), Eco.ocy. Small tree, low alt. On Dinagat Island growing on iron deposit. Flowering in June; fruiting April and June. Calophyllum oliganthum can be recognized by its small, probably nonfunc- tional terminal buds, its lamina with the midrib on the upper surface flat and the adjacent blade slightly raised, and its axillary inflorescences with a well-developed basal internode. The ellipsoid fruit has a thin outer layer and a thin-walled stone. The epithet oliganthum (‘‘few flowers’’) was coined because the inflorescences have few flowers. Calophyllum oliganthum is superficially similar to C. nodosum and C. depressinervosum. All three species have more or less profuse branching ue to a weak or nonfunctional terminal bud, leaves with a rather similar midrib on the upper surface, flowers with usually four tepals and few, small stamens, and fruits of similar structure. However, in C. oliganthum the axillary innovations lack basal scars, the midrib on the lower surface of the lamina is raised, rather than depressed, the inflorescence is only three to five flowered and lacks flowers close to the base, and the fruit is ellipsoid, rather than spherical; polyembryony has not been observed. Venation density and promi- nence are more like those of C. nodosum, but the lamina is larger and dries darker, and the internodes are much longer. 69. Calophyllum garcinioides P. F. Stevens, sp. nov. Ficure 24, a-e. C. cuneatum auct., non M. R. Henderson & Wyatt-Smith; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 7: 15. 1967. A speciebus aliis Calophylli in lamina coriacea canalibus laticibus venulis lateralibus oblique transgressentibus, innovationibus axillaribus cicatricibus basalibus ornatis, floribus cum 12 ad 16 tepalis, et fructu subsphaerico 1.6—2 cm. longo putamine parietibus minoribus 0.2 mm. crassis, differt. 1980] STEVENS, CALOPHYLLUM 413 Ficure 24. a-e, Calophyllum garcinioides. a, b, habit, x 0.5: a, SAN 71859; b, SAN 31788. c, d, fruit (SAN 62033), x 0.75: c, longitudinal section; d, from outside. e, terminal bud (SAN 26052), x 3. f, C. pelewense (Kanehira 386), terminal bud and axillary inflorescence with basal scar (arrow), x 6. Tree (1.8-)9-25(-39) meters tall, d.b.h. to 63 cm.; trunk without buttresses; outer bark usually yellowish brown (brown or gray), fissured, the inner surface brown to greenish brown; under bark red to reddish brown; inner bark red; latex yellow, transparent, sticky (rarely white). Twigs flattened, 1.5-3.9 mm. across, at first slightly 2-, 4-, or 6-angled, soon becoming rounded, drying blackish brown, glabrous or sparsely farinose- puberulent; axillary innovations with scars 1.5-7 mm. from base; internodes 0.5-3 cm. long; uppermost pair of axillary buds rounded, ca. 1 mm. long, spreading; terminal bud plump, 2.5-6 mm. long, with adpressed, brown to grayish indumentum (hairs, Ficure 25, e, f; cf. 22, k), underdeveloped internode to 3(-6) mm. long. Petiole 3.5-7 mm. long, broadly and shallowly concave above and convex below, glabrous; lamina obovate or trapeziform to elliptic 414 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 (rarely ovate), 2.5-9.3 by 1.4-5 cm., usually subacuminate to acute (rarely retuse) at apex, narrowly cuneate to rounded at base, distantly and shallowly recurved and at most slightly recurved at margin, coriaceous, drying umber or sabelline above and umber to sepia below, glabrous, the midrib above narrowing gradually or abruptly at base, usually subdepressed (+ raised), 0.2—-0.5(-0.7) mm. wide at midpoint, below raised, obscurely angled to striate, the venation apparent to obscure on both surfaces, raised, beneath usually with latex canals ascending obliquely over veins, 7 to 15 veins/5 mm., angle of divergence 50-75°. Inflorescences terminal and from adjacent foliate axils, with 7 to 17 flowers, branched when terminal (rarely so when axillary), the axis 3-9.5 cm. long, glabrous or puberulent toward base, lowest internode to 3(-11) mm. long; bracts usually soon deciduous (very rarely foliaceous, to 3 cm. long, persistent—Clemens 32480); pedicels 0.7—2 cm. long, glabrous. Flower (?)hermaphroditic; tepals 12 to 16, the outer pair broadly ovate to elliptic, 9.5-10.5 by 5.5-9 mm., the inner ones obovate to elliptic, 11-13 by 5-10 mm.; stamens 240 to 330, the filaments to 6 mm. long, connate for up to 0.6 mm., the anthers oblong, 1.1-1.5 mm. long, rounded to slightly retuse at apex; ovary 2-3 mm. long, the style ca. 4 mm. long, the stigma peltate, 1.1-1.4 mm. across, 3- or 4-lobed. Fruit subspherical, 1.6-2 by 1.1-1.4 cm., rounded to strongly apiculate at apex, drying brown, sharply wrinkled; outer layer detaching cleanly from stone, ca. 1 mm. thick, compact; stone subspherical, 1.5-1.9 by 1-1.3 cm., rounded at apex, the walls less than 0.2 mm. thick, smooth, unmarked; spongy layer thin. Type: North Borneo [Sabah], Ranau, above Mosilau Camp, 15 Aug. 1963, SAN 36197 coll. Mikil (holotype, san; isotypes, A, K, KEP, L, SAR, SING). DistripuTion. Northeastern Borneo (Map 24). ee 39 SELECTED SPECIMENS SEEN (‘‘w’’—white latex; ‘‘y’’—yellow latex). Borneo and adjacent islands. Sarawak. 4th Division: Baram, G. Mulu, 1554 m., Anderson & Keng K 3 (a, BO, K, L, SAN, SAR, SING), 1696 m., S 4259 (k, L, SAN, SAR, SING), 1524 m., Chew 383 (a, L, sinc), 1600-2200 m., Hotta 14731 (L). 5th Division: Lawas, G. Murut, Ba Belalan, 2149 m., S 26502 (L, sar, SING), 1829 m., S 26372 (L, SAN, SAR). BruNEI: G. Pagon Periok, 1859 m BRUN 2386 (x, L, SAR). SABAH. Tambunan: Trusmadi, 2134 m., SAN 31788 (K, L, SAN). Ranau: Kinabalu Natl. Park & vicinity, 2438 m., KEP 71640 Ficure 25. Hairs (from terminal bud, unless otherwise mentioned). a, b, Calophyllum parkeri (Parker 2313), from outside of basal scale on inflores- cence. c, d, C. elegans (S 5383). e, f, C. garcinioides (Clemens 34390, from axillary bud). g-i, C. sclerophyllum: g, Kostermans 8020; h, 5 E-1 P-649; i, A 398, from above. j, C. havilandii (Haviland & Hose = 3345). k, t, C. cordato-oblongum (CP 3823) (t, base of hair ca. 450 wm. long); — | teysmannii var. inophylloide: 1, Kostermans 12792, hair ca. 600 wm. , FRI 5873; n, Kostermans 12734; 0, bb 31797. p, q, C. hosei (Native ee 889) (q, an uncommon type). r, C. grandiflorum isiewmitns 23822). s, C. calcicola ane 13837). Scale = 60 um. (in 1, m, scale = 120 ym.; inr = 1980] STEVENS, CALOPHYLLUM 415 416 JOURNAL OF THE ARNOLD ARBORETUM [voc. 61 Map 24. Distribution of Calophyllum parkeri (triangle), C. fraseri (circles), C. enervosum (half-circles), C. praetermissum (s solid stars), C. confertum (open stars), and C. garcinioides (squares) in Southeast Asia and Malesia. Inset: Calophyllum sp. 73. (kEP), 1219 m., SAN 16232 (Bo, BRI, KEP, L, SING), 1829 m., 37989 (k, KEP, SAN, SAR, SING), 1219 m., 38068 (?) (y) (A, KEP, SAN, SAR), 2438 m., 38325 (y) (K, L, SAN), 71849 (y) (K, L, SAN, SING), 1524 m., RSNB 4308 (w) (k, L, SAN, SING), 1829 m., 4731 (w) (CANB, K, L, SAN, SING), Clemens 32442 (a, , BO, G, K, L, M, NY, UC), 1676 m., 33992 (a, BO, G, L, NY, UC), Marai Parai, 1524 m., Clemens 32480 (probable) (A, BO, L, M, NY, UC), 1524 m., 33194 (probable) (A, BO, NY). Kinabatangan: Lamag, Karamuak, Mt. Tavai, 990 Mis, SAN 51744 (san, pro parte). Eco.ocy. Locally common in lower montane and montane forest, sometimes in soil derived from ultramafic rock; 990-2438 m. alt. Flowering March (Sarawak) and July to September (Sabah); fruiting February to April and June to August (fruit greenish). GERMINATION AND YOUNG PLANT. The young plant is erect, and the terminal bud is functional. Young plants about 0.5 meters tall have axillary innovations with basal scars. 1980] STEVENS, CALOPHYLLUM 417 Although Calophyllum garcinioides is at first sight like other small- and coriaceous-leaved species of Calophyllum found on mountains, it can readily be distinguished from these other species by the scars at the bases of the axillary innovations, the obliquely ascending latex canals on the lower surface of the lamina, the axillary and terminal inflorescences that usually both have a very short basal internode, the flowers with 12 to 16 tepals, and the subspherical fruit with a thin-walled stone. The characteristic latex canals on the underside of the lamina are not always readily visible on dried specimens; however, they have been observed on specimens from all mountains where it grows. They look like the latex canals of Garcinia L.; hence, the specific epithet. Similar inflorescences, leaves, and axillary innovations occur in some specimens of the poorly known Calophyllum fraseri, from Malaya (q.Vv.). However, that species always has axillary inflorescences, flowers with at most eight tepals, and fruits about twice as large with stone walls ca. | mm. thick. There is considerable variation in leaf shape within Calophyllum garcinioides, but the extremes are connected by intermediates. Specimens taken fr trees reported to have white latex (all collected by the expedition of the Royal Society, London) tend to have thicker leaf blades that are more obtuse at the apex, while specimens from trees reported to have yellow latex have thinner leaf blades that are more acute at the apex. However, SAN 29128, with yellow latex, has a lamina that is rounded at the apex. Specimens from Mt. Kinabalu that are cited above with a question mark have relatively small, thin leaf blades that are more or less acuminate at the apex and axillary innovations that lack basal scars. SAN 60644, which has larger leaf blades that are subacuminate at the apex and scars ca. 7 mm. from the base of the axillary innovations, is intermediate between these and more normal specimens. Further variation is shown by the specimens collected by the Clemenses from Marai Parai, which have leaf blades that are strongly retuse at the apex and rounded at the base, while the specimen of Clemens 32480 at a has persistent, foliaceous bracts. The single specimen seen from Mt. Tavai, in Kinabatangan, is sterile. 70. Calophyllum fraseri M. R. Henderson & Wyatt- -Smith, Gard. Bull. Singapore 15: 344. pls. 24, 25. 1956, T. C. Whitmore, Tree Fl. Malaya 2: 182. 1973, pro parte. Tyre: Malaya, Pahang-Selangor border, Fraser’s Hill, 28 Sept. 1923, SFN 11282 coll. Nur (holotype, sinc; isotype, KEP). C. ?praineanum auct., non King; King, Jour. Fed. Malay States Mus. 7: 39. 1916. C. canum auct., non Hooker f.; Henderson, Gard. Bull. Straits Settl. 4: O35 1927. C. symingtonianum M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 338. 1956, pro minore parte; T. C. Whitmore, Tree Fl. Malaya 2: 192, 1973, pro parte. 418 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Tree 15-36 meters tall, d.b.h. to 58 cm.; trunk without buttresses or stilt roots; outer bark dirty deep gray to yellow-brown, dimpled or with shallow fissures at wide intervals; inner bark pale yellow-cream to pink; latex milky (Maxwell’s Hill) or clear yellow, resinous (G. Mengkuang, Kelantan); wood yellow. Twigs rather strongly flattened, 1.8-2.5 mm. across, not 4-angled to slightly so, soon becoming rounded, drying blackish to dark brown, puberulent when young; axillary innovations with or apparently lacking basal scars; internodes 1.5—3(-4.5) cm. long; uppermost pair of axillary buds rounded to subacute, to 1.5 mm. long, adpressed to terminal bud or not; terminal bud plump, 4,5-7 mm. long, with short, subadpressed, ferrugineous indumentum (hairs, cf. Ficure 20, m), underdeveloped internode to 2.5 mm. long. Petiole 0.7-1.2 m. long, usually shallowly concave above, convex below, glabrous when mature; lamina ovate to elliptic, 5-12.8 by 1.8-4.8 cm., + acuminate at apex, acute to cuneate at base, broadly undulate but not recurved or slightly so at margin, coriaceous, drying cinnamon-drab above and sabelline below, glabrous when mature, the midrib above narrowing rather gradually from base, at least margins raised, 0.2-0.6 mm. wide at midpoint, below strongly raised, striate, the venation above and below + apparent, raised, at least sometimes crossed by obliquely ascending latex canals, (7 to) 9 to 14 veins /5 mm., angle of divergence 60-70(-80)°. Inflorescences from foliate axils, sometimes two per axil, with up to 11 flowers, unbranched, the axis 1-4 cm. long, puberulent, especially toward base, or glabrous, lowest internode to 2.5 mm. long; bracts unknown; pedicels 0.5-1.8 cm. long, glabrous, to 2.5 mm. thick in fruit. Flower (?)hermaphroditic; tepals 4 or 8, the outer pair ovate, 4-7 by 3.5-S mm., the next pair ovate to suboblong, 5-8 by 3-4. m., inner ones, where present, suboblong, ca. 9 by 4 mm.; stamens (?)numerous, the filaments to 3.5 mm. long, the anthers oblong, ca. 1.6 mm. long, retuse at apex; ovary ca. 1.5 mm. long, the style ca. 3.5 mm. long, the stigma peltate, 0.8-1.4 mm. across, (?)3-radiate. Fruit ellipsoid, 2.6-3.3 by 2.3-5.2 cm., apiculate or not, drying vinaceous-brown, deeply reticulate or shallowly corrugate; outer layer detaching + cleanly from stone, 1.5-2.5 mm. across, compact; stone ellipsoid, 2.1-3 by 1.6-2.15 cm., rounded at apex, the walls 0.8-1.1 mm. thick, to 3 mm. thick at base, smooth, unmarked; spongy layer initially thick. DistripuTION. Malaya (Map 24). SELECTED SPECIMENS SEEN. Malaya. Kepau: Kedah Peak, 914 m., Robinson & Boden Kloss 6039 (sinc); Gunong Inas F.R., 914m., FRI 9262 (KEP). PAHANG: Fraser’s Hill, 1219 m., F.M.S. Mus. 11576 (56; KEP), path to Pine Tree Hill from Weld Hill, KEP 28955 (xep); oe ridge above Sungei Tanglin to G. Mengkuang, 1067 m., KEP 79614 (ker); Gunong Berumbun, Ridley 13910 (prob.) (k, sinc); Sungei Tkan, KEP 27634 (prot. ) (sING); ee Percival, KEP 35944 (KEP); Maxwell’s Hill, bend 57, 762 KEP 76333 (Kk, KEP); ridge to G. Tapis, 914 m., FRI 10974 (a, kK, KEP, ae Siar G. Rabong, 1295 m., FRI 20685 (KEP, SAN). TRENGGANU: G. Padang, 1219 m., SFN 31878 (KEP, sinc); Ulu Brang, camp 3, FRI 1755 (kep). 1980] STEVENS, CALOPHYLLUM 419 Eco.ocy. Colline or montane rainforest, 760-1295 m. alt. Flowering March and September; submature fruit September and October. Bilabiate, purselike galls probably caused by coccids (see Calophyllum teysmannii) occur fairly commonly (KEP 27634, 35944, 66601; Robinson & Boden Kloss 6126; Ridley 13910). Calophyllum ae is characterized by its elliptic leaf blades that are more or less acu t the base and acuminate at the apex and that have rather clear peice on both surfaces, and by its relatively large (ca. 3 cm. long) fruit that has a compact outer layer and a stone with walls ca. 1 mm. thick. The specific epithet commemorates L. J. Fraser, a trader and mule train operator, who disappeared in 1916 on what was subsequently called Fraser’s Hill. Calophyllum fraseri is perhaps related to C. garcinioides, from northeastern Borneo. Leaves of the two species dry similarly and have latex canals ascending obliquely to the venation; both species also have axillary innovations with scars at the bottom (but see below). However, C. garcinioides has a smaller (less than 2 cm. long) fruit with a thin-walled (less than 0.2 mm.) stone and larger flowers with 12 to 16 tepals. The base of the lamina in C. garcinioides is usually more or less rounded or cuneate; in C. fraseri it is acute. Calophyllum fraseri is perhaps also close to the poorly known C. parkeri (q. Despite recent collections made from Kelantan and Trengganu, oaiechvilun fraseri is a very poorly known species, and its variation is not yet understood. Specimens from Fraser’s Hill (the type locality) have a small (less than 1.2 cm. long) inflorescence axis that is puberulent toward the base and has a clear basal internode; the flowers have four tepals. Only one specimen with flowers has been collected elsewhere; this specimen (FRI 85, from Kelantan) has an inflorescence axis at least 2 cm. long that lacks a clear basal internode, and flowers with eight tepals. Axillary innovations with basal scars have been seen only in specimens from Kedah Peak, Kelantan, Trengganu, and perhaps from Cameron’s Highlands; much of the material is too poor for confirmation of the presence or absence of these scars, although they appear to be absent on material from Fraser’s Hill. Specimens with these scars have more coriaceous leaf blades than the other specimens, and obliquely ascending latex canals are visible on the lower surface. Fruits are known only from Trengganu (G. Padang), and Pahang (Maxwell’s Hill and G. Tapis) (i.e., not from the type locality). Despite this confusing situation, the following points may be noted. First, the specimens of Calophyllum fraseri that are most similar to C. garcinioides in vegetative characters can be clearly distinguished from that species by characters of both flower and fruit. Second, because fruits are not known from the type locality of C. fraseri, it remains to be established whether or not C. fraseri from Fraser’s Hill is conspecific with the other material cited. Henderson and Wyatt-Smith (op. cit.) thought that their C. cuneatum was close to C. fraseri; C. cuneatum (= C. calaba var. cuneatum) also has flowers with four tepals, but differs most obviously in its very much shorter petioles. Third, the specimens cited that have bilabiate coccid galls are mostly 420 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 sterile; I do not know of specimens with both galls and scars at the bases of the axillary innovations. Some specimens determined as C. fraseri by T. C. Whitmore have been removed to C. teysmannii var. inophylloide (q.v.), which has similar galls. The latter taxon is quite variable, and it is possible that some specimens still included in C. fraseri will also have to be removed to C. teysmannii. 71. Calophyllum parkeri Fischer, Kew Bull. 1926: 455. 1926. Type: Burma, Tavoy, Nwalabo, 1050 m., 2 Dec. 1924, Parker 2313 (holotype, k; isotype, A). Small tree; outer bark cracked into irregular plates, otherwise unknown. Twigs slightly flattened, 2.5-3.2 mm. across, rounded, drying blackish, glabrous; axillary innovations unknown; internodes 0.5-2.5 cm. long; upper- most pair of axillary buds and terminal buds unknown (hairs, Ficure 25, a, b). Petiole 5-8 mm. long, broadly and shallowly concave above, convex below, glabrous; lamina elliptic, 5.2-8.5 by 1.6-3.4 cm., cuneate at apex, rounded to subcuneate at base, slightly undulate and somewhat recurved at margin, coriaceous, drying umber above and below, glabrous, the midrib above gradually narrowed from base, flat or slightly raised, 0.3-0.6 mm. wide at midpoint, below raised, striate to subangled, the venation subapparent on both surfaces, slightly raised, 6 to 10 veins/5 mm., angle of divergence 50--60°. Inflorescences axillary and terminal, with up to 13 flowers, unbranched (terminal inflorescences sometimes branched), the axis 3-4 cm. long, glabrous, lowest internode to | mm. long; bracts unknown; pedicels to 1.3 cm. long, glabrous. Flower (?)hermaphroditic; tepals 12 (rarely 13), the outer pair orbicular, 3-3.5 mm. long and across, the next pair eee 6.5-7 b 5.5-6.5 mm., the inner ones elliptic to obovate, 9-12 by 4.5-7.5 mm.; stamens ca. 310, the filaments to 7.5 mm. long, connate for up to "1 mm., the anthers oblong, 1-1.2 mm. long, retuse to rounded at apex; ovary 2-2.5 mm. long, the style ca. 5.5 mm. long, the stigma peltate-infundibular, ca. 1.5 mm. across, not radiate. Fruit unknown. DistriBUTION. Burma (Map 24); known only from the type specimen. Calophyllum parkeri is an imperfectly known species that can be recognized by its rounded twigs; moderate-sized, elliptic leaf blades that are cuneate at the apex and subrounded at the base and have a more or less flat midrib on the upper surface; axillary and terminal inflorescences (the former, at least, have a very short or nonexistent basal internode); and 12-tepaled flowers. It is quite possible that the axillary innovations have basal scars. The epithet commemorates the collector, R. N. Parker. If the axillary inflorescences and axillary innovations of Calophyllum parkeri have basal scars, it may be related to the specimens of C. fraseri, from the eastern part of Malaya, and to C. garcinioides, from Borneo. Calophyllum fraseri apparently has four- to eight-tepaled flowers and axillary inflorescences. Although C. garcinioides has both axillary and terminal inflorescences and flowers with at least twelve tepals, the tepals of the outermost pair are 1980] STEVENS, CALOPHYLLUM 42] about three times the size of those of C. parkeri; it also has obliquely ascending latex canals on the lower surface of the lamina. 72. Calophyllum flavo-ramulum M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 310. pls. IC, 6. 1956; Kochummen, Malayan Forest Rec. ed. 2. 17: 219. 1965; T. C. Whitmore, Tree Fl. Malaya 2: 181. 1973. Type: Malaya, Trengganu, Kemaman, Bukit Kajang, 500 feet [150 m.], 8 Nov. 1935, SFN 30321 coll. Corner (holotype, sinc; isotypes, a, BO, K, KEP, LAE). e 30-38 meters tall, d.b.h. to 88 cm.; trunk without buttresses; outer eer Sein to greenish brown, strongly fiscuned flaking, ca. 8 mm. thick, in transverse section with orange lines demarcating brown inne: the under surface + bright orange; under bark pale straw; inner bark reddish; latex clear yellow, sticky. Twigs + flattened, 2-3 mm. across, rounded or obscurely 4-angled, drying dark to blackish brown when young, yellowish when older, subpersistently reddish brown-—puberulent or not; axillary innovations with basal scars; internodes 1.5-4.5 cm. long; uppermost pair of axillary buds rounded, up to 1.5 mm. long; terminal bud plump, 2.5-6 mm. long, with reddish brown- tomentose to grayish puberulent indumentum (hairs, Ficure 22, q), underde- ele eee absent. Petiole 0.6-1.4 cm. long, shallowly (deeply) concave x below, ec ueuee puberulent or not; lamina oblong ON a a a6 by 1.7-4.8 cm., + gradually to abruptly acuminate at apex, cuneate to attenuate at base, are and slightly recurved at margin, entire lamina + concave, coriaceous, drying brown-vinaceous to umber above and umber to fulvous below, often persistently puberulent on midrib above and especially below, the midrib above usually narrowing gradually from rather narrow base, raised, usually sulcate at first, 0.15-0.35 mm. wide at midpoint, below raised, striate, the venation + apparent above and below, raised, 10 to 19 veins/5 mm., angle of divergence 70-80°. Inflorescences terminal and from adjacent foliate axils, with 7 to 15 (axillary inflorescences) to man (terminal inflorescences) flowers, unbranched (flabellate), or with 3-flowered branches to | cm. long, the axis 2.4-7 cm. long, reddish puberulo-tomentose (rarely subglabrous except at base), lowest internode of axillary inflorescences to 3 mm. long; bracts not seen, early deciduous; pedicels 0.5-1.5(-2.2) cm. long, puberulent (rarely glabrous), slender at first, in fruit to 3 mm. thick. Flower (?)hermaphroditic; tepals 8, the outer two broadly ovate, ca. 5 by 3.5-4.5 mm., + puberulent on back, the next pair obovate, 6.5-10 by 3.5-7 mm., sometimes puberulent on back near base, the inner ones obovate, 7.5-11 by 3-5 mm., stamens 100 to 135, the filaments to 3 mm. long, the anthers oblong-elliptic, 0.5-0.7 mm. long, rounded at apex; ovary ca. 1.5 mm. long, the style ca. 3 mm. long, the stigma peltate, ca. 0.5 mm. across, + 3-lobed. Fruit ellipsoid to subspherical, 2.5-3.5 by 2-2.6 cm., rounded to sharply pointed at apex, drying vinaceous-brown, with shallow, fairly dense, rounded wrinkles, or almost smooth (Sumatra); outer layer detaching + cleanly from stone, |.5-2.3 mm. thick, compact; stone ellipsoid, 2—2.6 by 1.75-2.2 cm., 422 JOURNAL OF THE ARNOLD ARBORETUM [vo. 61 rounded at apex, the walls 0.9-1.2 mm. thick, thinner to one side of base, smooth, unmarked, perhaps with basal plug ca. 5 mm. across; spongy layer thin. DistripuTion. The eastern part of the Malay Peninsula, the Lingga Archipela- go, perhaps in northeastern Sumatra (Map 23). SELECTED SPECIMENS SEEN. Malaya. TRENGGANU: Kuala Trengganu, 45 m KEP 76064 (ker); Dungun, Karteh Ulu Chukai F.R., 150 m., KEP 93882 (KEP); Jerengau State Land, KEP 76661 (KEP); Bukit Bauk F. R., 150 m., KEP 93973 (ker); Kemaman, Bukit Kajang, 150 m., SFN 30565 (kee, SING). PaHanc: Kuantan, Bukit Goh F.R., KEP 77978 (xk, ker); G. Tahan, Wong & Wyatt-Smith 34 (2) (KEP); Sungei Kelui, G. Tapis, FRI 10083 (?) (KEP); Baloh F.R., KEP 40216 (kep). JoHore: Kluang F.R., 90 m., FRI 7521 (a KEP, SING); Mersing F.R., 15 m., KEP 77944 (xep); Jemaluang F.R., KEP 73552 (KEP); Panti F.R., 366 m., KEP 99237 (kep); Lenggor F.R., 35 m., Stevens et al. 57 (A). Sumatra and adjacent islands. Riau: Lingga, P. Singkep, Marok Toewa bij Soengei Soelit, bb 1708 (= Amat 22) (Bo, L), S. Poelak, 25 m., bb 4031 (= Amat 6) (Bo). Utara: Laboehan Batoe, Rattosan, Basau, 50 m., bb 8978 (2) (Bo). Ecotocy. Lowland or colline mixed dipterocarp forest, 15-427 m. alt. Flowering January, April, July, and August; fruiting February, May, and July (fruit mauvish black— KEP 77978). Some inflorescences of SFN 30321 are galled, having been converted into cauliflowerlike structures ca. 1 cm. across. Calophyllum flavo-ramulum can be distinguished from the other large- leaved species of Calophyllum by its short terminal bud, axillary innovations with basal scars, leaf blades that dry boat shaped, and terminal inflorescences. (Although C. flavo-ramulum was described as having axillary inflorescences, some inflorescences, even on the type specimen, are terminal.) The older twigs of C. flavo-ramulum dry a yellowish color, hence the specific epithet. Calophyllum flavo-ramulum has been confused with C. dasypodum, but the two species are not close. The terminal bud of C. dasypodum is longer, its older twigs do not dry yellow, its leaves do not dry boat shaped, its inflorescences are axillary, and its flowers usually have four tepals. The specimens cited from the Lingga Archipelago are very similar to those from the Malay Peninsula, although they have somewhat smaller flowers and ellipsoid fruits that dry almost completely smooth, with only faint striations. The fruits of the Malayan specimens are variable in shape, but dry wrinkled. In the ellipsoid fruits of KEP 99237, the outer layer is much thickened (7 mm.) at the base. 73. Calophyllum sp. Tree ca. 20 meters tall, d.b.h. to 32 cm.; trunk without buttresses; outer bark pale gray to eae smooth except for diamond-shaped fissures; latex golden, clear, very s Twigs slightly nee ca. 2 mm. across, not angled, drying pale brownish, 1980] STEVENS, CALOPHYLLUM 423 shiny, glabrous; axillary innovations with several pairs of basal scars; inter- nodes 2-3.5 cm. long, uppermost pair of axillary buds rounded, ca. 0.7 mm. long, inconspicuous; terminal bud plump, 2-5 mm. long, consisting of several pairs of glabrous, imbricate scales up to 2.5 by 4 mm., underdeveloped internode absent. Petiole 0.7-1.3 cm. long, shallowly concave above, convex below, drying blackish, glabrous; lamina elliptic to suboblong, 7—-13.5 by 3.4—5 cm., acuminate at apex, acute at base, slightly recurved and undulate at margin, entire lamina + concave, coriaceous, drying + vinaceous-brown above and subchestnut-brown below, glabrous, the midrib above gradually narrowed from base, raised, 0.25-0.5 mm. across at midpoint, below rather narrow, raised, rounded, the venation on both surfaces apparent, raised, 9 to 14 veins /5 mm., angle of divergence 75-80°. Inflorescences, flowers, and fruits not known. DistriBUTION. The Malay Peninsula, Trengganu (Map 24). SPECIMENS SEEN. Malaya. Trenccanu: Dungun, Bukit Bauk F.R., 30 m., KEP 53353 (kEP), 77983 (KEP). Calophyllum sp. 73 is a remarkable taxon that is easily recognized, even when sterile, by its terminal buds, which are composed of a number of decussate, glabrous, imbricate perulae. se buds are unique in the genus, and fertile material of this taxon is much needed. KEP 53353 was cited as Calophyllum EE by Henderson and Wyatt-Smith (1956), but in that species a pair of indumentum-covered perulae completely enclose the rest of the bud. 74. Calophyllum enervosum M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 352. p/. 32. 1956; Kochummen, Malayan Forest Rec. ed. 2. 17: 221. 1965; T. C. Whitmore, Tree Fl. Malaya 2: 180. 1973. Tyre: Malaya, Johore, Mersing, Jemaluang F.R., 30 m., 20 April 1954, KEP 71894 coll. Wyatt-Smith (holotype, KEP; isotypes, BO, K, SING). Tree 24-33 meters tall, d.b.h. to 50 cm.; trunk without buttresses; outer bark greenish olive to yellowish, smooth to rough and cracking very finely, hoop marked, the inner surface dull brownish orange; under bark dull red- dish; inner bark reddish, laminated; latex clear he very sticky, report- ed to turn cloudy (Henderson & Wyatt-Smith, loc. cit.). Twigs slightly flattened, 2-2.7 mm. across, Gute 4-angled, soon becoming rounded, drying blackish when young, later whitish to brown, + brown-tomentose when young; axillary innovations with pair of scars 2-5 mm. from base; internodes 1-3 cm. long; uppermost pair of axillary buds rounded, less than 1 mm. long, inconspicuous; terminal bud plump, 2-4 mm. long, with ferrugineous, puberulo-tomentose indumentum (hairs, Ficure 27, a), underdeveloped internode absent. Petiole (0.3-)0.5-1 cm. long, flat to broadly and shallowly concave above, convex below, glabrous; lamina elliptic to obovate, 3-9 by 1.4-4.8 cm., rounded to shortly and rather obscurely acuminate at apex, narrowly cuneate to attenuate and finally decurrent at base, not or broadly and rather distantly undulate and not or slightly recurved 424 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 at margin, marginal thickening ca. 0.3 mm. wide, coriaceous, drying umber above and below, transiently tomentose on midrib below when young, otherwise glabrous, the midrib above narrowing gradually from sulcate base, becoming flat to slightly raised, 0.15—0.3 mm. wide at midpoint, below raised, striate, the venation + obscure above and below, slightly raised or not, 7 to 10 veins/5 mm., angle of divergence 65-75°. Inflorescences terminal and from foliate axils along twigs, with 7 to 11 flowers, usually unbranched (sometimes with 3-flowered branches to 4 mm. long), the axis 1.5-2.5 cm. long, + persistently short-tomentose, lowest internode 1-2 mm. long; bracts ovate, ca. 4 mm. long, subpersistent, short-tomentose below; pedicels in fruit 5-10 mm. long, persistently subtomentose or not. Flowers known only in very young bud. Submature fruit ellipsoid to obovoid, 1.4-1.7 by 1.1-1.2 cm., + rounded at apex, drying vinaceous-brown, 7 and sharply wrinkled; outer layer detaching cleanly from stone, 1-1.8 mm. thick, compact, except by 0.8-0.95 cm., rounded at apex, the walls ca. 0.15 mm. thick, smneuths unmarked; spongy layer thick at first. DistripuTion. The Malay Peninsula (Johore) and the Lingga Archipelago (Map 24) ADDITIONAL SPECIMENS SEEN. Malaya. Jonore: Armas KEP 5823 (x); ae Serah, Mersing F.R., 30 m. plus, KEP 92104 (kep); Jemaluang F.R., m., KEP 72907 (KEP), Stevens et al. 72 (a); G. Panti E., Jumali & eae 194 Galea: ?sapling) (Bo). Sumatra and adjacent islands. Riau: Lingga Arch., P. Singkhep, Sungei Roengkil, 10 m., bb 4082 (Bo); Tandjong Ba- tang, 8 m., bb 4016 (Bo). Eco.tocy. Mixed dipterocarp forest, 8-40 m. alt. Locally common, at least on Lingga Archipelago. Submature fruit April and May GERMINATION AND YOUNG PLANT. The young plant is erect, and the terminal bud is functional. When young the leaf blades are bright red with a broad green stripe ca. 5 mm. wide down the middle. Calophyllum enervosum is an easily recognized species because of its rather small, coriaceous leaf blades that dry nearly flat with almost invisible venation and that are narrowly cuneate to decurrent at the base. The rather small, deeply wrinkled fruit with its relatively thick outer layer is also distinctive. The almost invisible venation of the dried leaves suggested the specific epithet. The specimens cited above from the Lingga Archipelago are good matches with those from Malaya, although their indumentum is somewhat longer and their leaf blades are a little more pointed at the apex. Fruits are known from both areas and are identical; they have not been described before. To be concluded Erratum—On page 43 of Volume 61 of the Arnold Arboretum, January, | 1 should read as follows: FIGURE , number 1, of the Journal 980, the caption to FIGURE Vang, discoverer of Metasequoia glyptostroboides, right, with his teacher, Professor _T.N. Liou, seated, and S. Y. Hu, left, July, 1975. Journal of the Arnold Arboretum April, 1980 CONTENTS OF VOLUME 61, NUMBER 2 © A Revision of the Old World Species of Calophyllum Hetitenaey. POPPE TEVENS eae eae ee ee ae Ae ae ST ARONOMIG. PHSTORY os tsiccn ater sive a ten eae vie aaa arenes ese =e YAIATON IN THE GENUS AND Its USE IN CLASSIFICATION See eo as 121 F Sen. sic sashes ponrer ate Pass Vs aa RE REI ee 144 Boca DistriBution, AND-ECOLOGY.. 28> s. “ion on. ie 148 AICKNOWEEDGMENTS test sre See ana a 166 SOTERA TIGL REARMENT con cals en eens Seo nc urn ge ee geri nesta cee 167 Prati i fe een ee ie a ome a oe OS ee ea GOERS Volume 61, Number 1, including pages 1-116, was issued March 21, 1980. JOURNAL or te. ARNOLD ARBORETUM HARVARD UNIVERSITY VOLUME 61 NUMBER 3 en ISSN 0004-2625 Journal of the Arnold Arboretum Published quarterly in January, April, July, and October by the Arnold Arboretum, Harvard Universit ity. Subscription price $25.00 per year. Subscriptions and remittances should be sent to Ms. E. B. ‘Schmidt, Arnold Arboretum, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. Claims will not be accepted after six months from the date of issue. POSTMASTER: send address changes to Ms. B. Schmidt, Arnold Arboretum, 22 Divinity Avenue, Cambridge, pee 02138, U.S.A. Volumes 1-51, reprinted, and some back numbers of volumes 52-56 are available from the Kraus Repent Corporation, Route 100, Millwood, New York 10546, U.S.A EDITORIAL COMMITTEE S. A. Spongberg, Editor E. B. Schmidt, Managing Editor P. S. Ashton K. S. Bawa P. F. Stevens C. E. Wood, Jr. Printed at Edwards Brothers, Inc., Ann Arbor, Michigan ER: Th L + Act: OV it single scohome (magnified nearly 160 times) of a Fijian endemic, can ie ‘ewcocarpum A. C. Smith, form the basis of this year cover, W Ss ap rin As in recent years, the designs, based on holotype material in ihe herbarium of the Ainold A rboretum, were draw. n by Karen Stoutsenberger Calophyllum L. Se is a wide-ranging genus of tropical forest trees in both the New and Old Worlds, and some species are of considerable economic importance as sources of valuable Laas Its use on eon of this volume of the Journal of the Arnold Arboretum is appropriate since the April and Jul numbers are devoted 2 P. F. Stevens’s revision of the Old World representatives of this genus.—S. A. Second-class postage paid at Boston, Massachusetts, and additional offices. JOURNAL OF THE ARNOLD ARBORETUM VoLumE 61 JuLty 1980 NuMBER 3 A REVISION OF THE OLD WORLD SPECIES OF CALOPHYLLUM (GUTTIFERAE)* P. F. STEVENS 75. Calophyllum confertum P. F. Stevens, sp. nov. FicureE 26, f, g. A speciebus aliis Calophylli ere quibus internodia terminali innova- tionis breviore aliis habent in ramulo in siccitate minute reticulato, pagina superiore laminae in siccitate ee pagina inferiore pallidiore et venulis lateralibus 3 usque ad 6 per 5 mm. (in speciebus aliis 5 usque ad 19 per mm.), differt Tree ca. 36 meters tall; trunk and bark not known. Twigs slightly flattened, 1.8-2.2 mm. across, 4-angled, drying brown to yellowish, surface minutely reticulate, glabrous; axillary innovations lacking basal scars; internodes mostly 1-3 cm. long; uppermost internode of innovation up to 7 mm. long; uppermost pair of axillary buds rounded, less than 0.5 mm. long, inconspicuous; terminal bud plump to narrowly conical, 1.3-2.5 mm. long, with adpressed, brown indumentum (hairs, Ficure 22, m), under- developed internode absent. Petiole 0.4-1 cm. long, deeply concave above and convex below, glabrous; lamina obovate, (3.2-)5-8.5(-18) by (1.1-)2.3-4 (-7.1) cm., rounded to subacute at apex, narrowly cuneate at base, slightly undulate and somewhat recurved at margin, coriaceous, drying cinnamon above and sepia below, glabrous at maturity, the midrib above ee narrowed near base, peep depressed at first, becoming subelevated, conspicuous, 0.15-.3 mm. wide at midpoint, below raised, somewhat striate, the venation er above and subobscure below, slightly raised, 3 to 6 veins/5 mm., angle of divergence 40-50°. Inflorescences from foliate axils near ends of twigs, with scars of 5 to 7 flowers, unbranched, the axis 3-5 cm. long, glabrous, lowest internode 3.3-3.7 cm. long; bracts not known; pedicels 0.7—-2 cm. by ca. 1.8 mm., glabrous. Flower not known. Fruit spher- ical, 1.5-1.8 cm. long and across, rounded at apex, drying with shallow and rounded wrinkles, sharply wrinkled when young, pale brown; outer layer detaching cleanly from stone, 1.7-2 mm. thick, compact; stone spherical, *Continued from Volume 61, page 424. © President and Fellows of Harvard College, Journal of the Arnold Arboretum, 61: 425-690. ae 1980. 426 JOURNAL OF THE ARNOLD ARBORETUM [ VoL. 6 _ Ficure 26. a, Calophyllum praetermissum (Haviland & Hose 3342), terminal bud, x 3. b, C. aerarium (Cel. /II-354), terminal bud, x 3. c, C. andersonii (Haviland = 1812), terminal bud, x 6. d, e, C. teysmannii var. bursiculum (S 17230): d, terminal bud, x 4; e, habit, x 0.5. f, g, C. confertum (SAN 17146): f, habit, x 0.5; g, fruit, transverse section, x |. 1980] STEVENS, CALOPHYLLUM 427 ca. 1.2 cm. long and across, rounded at apex, the walls less than 0.2 mm. thick, smooth, unmarked; spongy layer (?)thin. Type: Brunei, Temburong, summit of Bukit Patoi, 255 m., 5 April 1957, SAN 17146 coll. Smythies, Wood, & Ashton (holotype, a; isotypes, Bo, K, KEP, L, SAN (Several), SING). DistripuTion. Northeastern Sarawak, Brunei (Map 24). ADDITIONAL SPECIMENS SEEN. Borneo. Sarawak. 3rd Division: Ulu Arip, Balingian, Ashton 6002 (a). 4th Division: Nyabau Catchment area, Ashton 6001 (4). Brunet: Andalau F.R., 60 m., Ashton s.n., July 1959 (sar); Bukit Biang, Ashton s.n., July 1959 (sar). Sine loco: Brunig Li 50 (sar). EcoLocy. Forest, 60-255 m. alt. Specimens of Calophyllum confertum can readily be recognized, even when sterile, by the minutely reticulate twigs, leaves aggregated at the ends of the twigs, and obovate leaf blades with distant, steeply ascending venation. The epithet confertum means ‘‘crowded,”’ an allusion to the congestion of the leaves at the end of each innovation. Calophyllum obliquinervium also has innovations with short terminal inter- nodes, but its leaf has denser venation, and the midrib on the lower surface of the blade is less prominent than in C. confertum and often does not reach the apex. It differs further in having terminal inflorescences, and fruits with the outer layer less than 0.5 mm. thick and not compact. Calophyllum confertum appears to be most closely related to the imperfectly known C praetermissum; for the differences between the two, see the latter. All specimens of Calophyllum confertum except the type are sterile. However, the other specimens have the characteristic reticulation of the twig (except sometimes in young growth), the short terminal internode, and the distant venation of the type specimen. Although the leaf blades are somewhat larger than those of the type (measurements in parentheses above), the specimens may have been taken from young trees. 76. Calophyllum praetermissum P. F. Stevens, sp. nov. Ficure 26, a. A speciebus aliis Calophylli in foliis apicibus innovationibus quaternis agegregatis, ramulis in siccitate griseo-brunneis vel albescentibus, lamina mediocra margine haud undulata, et inflorescentia axillaris internodio basali 2-7 cm. longo, differt. Tree 10-17 meters tall, d.b.h. to 35 cm.; trunk without spurs or buttresses; outer bark dark to pale brown, tending to have shallow cracks, scaly; inner bark pink; latex opaque, sparse, barely sticky (S 2761). Twigs slightly flattened, 1.2-2.3 mm. across, transiently to rather persistently 4- or 6-angled, drying gray-brown to whitish, glabrous or sparsely short-tomen- tose; axillary innovations lacking basal scars; basal internodes of innovation to 5.5 cm. long, intermediate internodes 0.7-2.5 cm. long, terminal internodes 0.1-1 cm. long; uppermost pair of axillary buds inconspicuous, less than 428 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 0.5 mm. long; terminal bud conical, 2-3 mm. long, with short, tomentose, brown indumentum (hairs, Ficure 22, n—p), underdeveloped internode absent. Petiole 4-7 mm. long, broadly concave above, convex below, drying blackish, when young with few hairs on edges; lamina obovate, 3.2-7.4(-12) by 1.5-3.6(-4) cm., rounded to retuse at apex, acute to cuneate at base, not undulate and slightly to clearly and narrowly recurved at margin, coriaceous, drying dark brick above and sepia below, glabrous, the midrib above rather abruptly narrowed at base, somewhat depressed at first, becoming slightly raised, 0.1-0.2 mm. wide at midpoint, below slightly raised, striate, the venation on both surfaces + apparent, raised, 5 to 9 veins/S mm., angle of divergence 50-65°. Inflorescences from uppermost foliate axils, with 5S to 9 flowers, unbranched, sometimes flabellate, the axis 4.5-9.5 cm. long, glabrous or very sparsely puberulent toward base when young, lowest internode 2-7 cm. long; bracts ovate, ca. | mm. long, subtomentose below, not persistent; pedicels 1-2.2 cm. long, glabrous. Flower known only from bud, (?)hermaphroditic, tepals 4, the outer pair broadly ovate to orbicular, ca. 5 by 4.5 mm.; stamens 80 to 110, the anthers oblong, 1.5-2.2 mm. long, slightly retuse at apex, ovary ca. 1.5 mm. long, glabrous, the style ca. 3 mm. long, the stigma peltate, ca. 0.6 mm. across. Immature fruit subspherical, ca. 8 by 7 mm., drying finely wrinkled, brown; outer layer ca. 1.3 mm. thick. Type: Sarawak, near Kuching, 24 Nov. 1894, Haviland & Hose 3342 (holotype, BM; isotype, Kk). DistripuTion. Northwestern Borneo (Map 24). ADDITIONAL SPECIMENS SEEN. Borneo. Sarawak. Ist Division: near Kuching, Bartlett s.n., anno 1893 (pm); Selang F.R., S 7360 (?) (sar). 3rd Division: Batang Igan, Sungei Tutus logging camp, S 30526 (sar); Loba Kabang South Protected Forest, S 2761 (sar). KALIMANTAN. Selatan: Amoentai, Rantau Kodjang, 10 m., bb 7787 (po). Barat: Palo, 5 m., Becking 41 (Bo). Eco.tocy. Swamp forest, low. alt. Late bud in November. Calophyllum praetermissum differs most obviously from C. confertum in its twigs, which do not dry finely reticulate, and in its leaf blades, which have denser venation and dry darker on the upper surface than on the lower. Although the two taxa have a similar distinctive inflorescence type and leaf arrangement, they have different hairs and ecology. The epithet praetermissum Sarl refers to the superficial similarity of this species and C. confertu At a Sight sterile specimens of Calophyllum praetermissum are somewhat like those of C. andersonii. However, in the latter species only the older twigs dry a pale gray-brown color, so there is less contrast between dark petiole and light twigs; the lamina is concave toward the base rather than broadly convex as in C. praetermissum. Neither the inflorescence nor the leaf arrangement of C. andersonii is like that of C. praetermissum S 7360 is a fragmentary specimen and is included here with hesitation. 1980] STEVENS, CALOPHYLLUM 429 77. Calophyllum andersonii P. F. Stevens, sp. nov. Figure 26, c. ee Ridley, Kew Bull. 1938: 120. 1938, typo excepto; J. Anderson, Bull. Singapore 20: 154. 1963, Trees Peat Swamp Forest Sarawak, = i Zia 1972. A speciebus aliis Calophylli in gemma terminalis corpulenta, lamina mediocra obovata apice plus minusve retusa venulis lateralibus subdistantibus validisque, tepalis 4, et fructu ovoideo usque ad 1.6 cm. longo apice acuto strato exteriore circa 1.5 mm. crasso et putamine parietibus circa 0.8 mm. crassis, differt Tree 15-23(-30) meters tall, d.b.h. to 20 cm.; trunk with buttresses to 0.9 meters tall, or stilt roots to 1.2 meters tall; outer bark dark brown to pale yellowish gray, narrowly fissured, the inner surface dull orange-brown; under bark dull orange- elie inner bark reddish to dark brown; latex sparse, whitish, (?)sticky or no Twigs slightly Aeaed 1.3-3 mm. across, rounded, striate when old, drying brown when young, when older grayish to yellow-brown, glabrous or brown- farinose when young; axillary innovations lacking basal scars; internodes 0.5-4(-5.5) cm. long; uppermost pair of axillary buds rounded, to 1.5 mm. long, erect; terminal bud plump (narrowly conical), 3-6 mm. long, with gray-brown, subcrustaceous to puberulent indumentum (hairs, Ficure 22, e, f; some also moruloid), underdeveloped internode to 2.5 mm. long. Petiole 0.7-2.1 cm. long, broadly concave above and convex below, glabrous, drying blackish; lamina obovate, 2.6-10(-13) by 1.8-5.5(-6.7) cm., retuse to rounded (rarely cuneate) at apex, acute to cuneate at base, slightly and distantly undulate and somewhat recurved at margin (with submarginal vein 0.4 mm. from margin, or vein obscured by thickening), coriaceous, drying sabelline to greenish olivaceous above and sepia to fulvous below, when young farinose-puberulent on midrib below, the midrib above narrowing gradually rom base, subdepressed at first, becoming raised, 0.1-0.2 mm. wide at midpoint, below raised, rounded to subangled, the Nis subobscure to apparent above and apparent below, raised, 4 to 9 (to 12) veins/5 mm., angle of divergence 45-70°. Inflorescences from foliate axils along twigs, with 7 to 11 flowers, unbranched, the axis 1.3-2.7 cm. long, puberulent at least near base, lowest internode 2-9 mm. long; bracts elliptic, to 3 mm. long, deciduous; pedicels 0.5-1.3 cm. long, frequently farinose when young, slender. Flower aie iad oe 4, the outer pair broadly ovate, 3.5-4.5 by ca. 4 mm., the inner pair broadly obovate, ca. 6 by mm.; stamens 70 to 85, the filaments to 2.5 mm. long, the anthers alo 0.8-1.2 mm. long, + retuse at apex; ovary ca. 1.3 mm. long, the style ca. 1.5 mm. long, the stigma subpeltate, 0.55-0.7 mm. across, not clearly radiate. Fruit ovoid, 1.4-1.6 by ca. | cm., acute at apex, drying vinaceous-brown, closely and rather shallowly wrinkled; outer layer detaching cleanly from stone, |.2-1.8 mm. thick, compact; stone ellipsoid, 8.5-10 by 6.5-7 mm., acute to obtuse at apex, the walls 0.7-0.8 mm. thick, smooth, unmarked; spongy layer (?)thin. Tyre: Sarawak, 3rd Division, Sibu [Loba Kabang South Protected Forest] , 430 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 20 Feb. 1954, S$ 480 coll. Ahmady (holotype, sar; probable isotypes ‘‘coll. Anderson,’” KEP, SAR, SING). DistripuTion. Northwestern Borneo (Map 33). ADDITIONAL SPECIMENS SEEN. Borneo. Sarawak. Ist Division: ca. 3 km. from Kuching, Haviland 1812 (BM, L, SAR, SING), = 1812 (BM, BO, GH, L); Stapok F.R., S 24679 (a, L, SAR, SING), 30590 (sar); (?)G. Pueh F.R., S 7523 (sar); Sabal Tapang, 150 m., Stevens et al. 178 (a). 3rd Division: Rantau Panjang, S 2739 (sar); Loba Kabang South Protected Forest, S 2698 (sar); Sungei Tutus logging camp, S 30543 (sar). Brunet: H. Simpai Berakas, S 2003 (sar). KALIMANTAN. Barat: Palo, 5 m., Becking 76 (Bo). Ecotocy. Usually in peat swamp (alan forest dominated by Shorea albida Symington), also in mixed dipterocarp forest on low hills at Sabal Tapang; 5-150 m. alt. Flowering August, September, and December (flower scented); submature fruit in February. Calophyllum andersonii can be recognized by its short, plump terminal buds, its twigs, which dry a grayish- or yellowish-brown when older, and its obovate leaf blades with retuse apices and relatively distant venation. The flowers have four tepals, and the small fruits have a well-developed outer layer and stone walls about 0.8 mm. thick. The epithet commemorates J. A. R. Anderson, well known for his work on the peat swamps of northwestern Borneo where this species usually grows. Calophyllum andersonii is somewhat like C. teysmannii var. inophylloide ; the latter has similar bark, but the latex is yellow. However, living leaves of C. andersonii are notably less rigid than those of C. feysmannii var. inophylloide, the midrib is not raised (seen also in the dried leaf), and the older twigs dry striate and pale yellowish brown, in contrast to the blackish petioles. Calophyllum teysmannii var. inophylloide also has larger fruits that are rounded at the apex and have a relatively much thinner stony layer. The filaments of C. andersonii appear to be papillate toward the apex, a character that has not been noticed in C. teysmannii, but more material is needed to confirm this difference. The paratype of Calophyllum fragrans, Haviland 1812, is to be referred to C. andersonii. However, the two species are not at all close, C. fragrans (= C. hosei) having, for example, axillary innovations with basal scars, leaf blades with much denser venation, and ellipsoid fruits. S 5935 (Brunei, Andalau F.R., 45 m.) may be a specimen of Calophyllum andersonii, but it has leaf blades with very broadly recurved margins and a terminal bud 6.5-9 mm. long on an underdeveloped internode 0.4—1.5 cm long. There are two types of fruit associated with the specimen, although neither has been seen attached to a shoot. One is somewhat similar to that of C. andersonii, although a little larger (to 1.8 by 1.4 cm.) and more deeply wrinkled. The other is spherical, ca. 2 cm. long, and has an outer layer about 0.5 mm. thick; it is perhaps similar to that of C. alboramulum (see that species; KEP 80089 is also similar to $ 5935), 1980] STEVENS, CALOPHYLLUM 431 78. Calophyllum teysmannii Mig. Fl. Indiae Batavae Suppl. 1(3): 499. Dec. 61. Type: Sumatra, prope Paja-kombo, Teysmann, HB 650 (holotype, U; isotypes, BO, L, P). Tree 3-40 meters tall, d.b.h. to 95 cm.: trunk very often but not always with spurs or small buttresses to 70 cm. tall: outer bark brown to gray-brown, fissured and scaling, not hoop marked, the inner surface dirty orange-brown to blackish; under bark dark orange-brown to deep red; inner bark red; latex yellow, clear to opaque, sticky to fluid: sapwood pink to yellow; heartwood hard. Twigs slightly to definitely flattened, (1-)1.5-3.5(-5) mm. across, not or 2-, 4-, or 6-angled, often with transverse lines at nodes, drying brown to blackish, transiently puberulent to subpersistently tomentose; axillary innova- tions lacking basal scars; internodes 0.5-3(-5) em. long; uppermost pair of axillary buds rounded, to 1.5 mm. long, erect to spreading; terminal bud plump to conical, (2-)3.5-9.5(-12.5) mm. long, with subcrustose grayish to tomentose, erect to adpressed, brown indumentum (hairs, Figures 25, l-o: 27, ci; some also moruloid), underdeveloped internode to 5 mm. long. Petiole 0.35—-2.2(-3.5) cm. long, broadly and shallowly concave above, convex below, sometimes subpersistently tomentose; lamina obovate (elliptic, oblong, cunei- form, subovate, or suborbicular), (2-)3.3-13.8 by (1.3-)2.1-6.6 cm., retuse (rounded or acute) at apex, acute to cuneate (rarely rounded) at base, flat to slightly undulate and sharply recurved at margin, marginal thickening absent to 1.2 mm. wide, coriaceous to very coriaceous, drying bay to umber (pale khaki) above and umber (fulvous) below, usually + transiently puberulent on midrib on both surfaces, or subpersistently tomentose—also on margin when young, the midrib above often narrowing + abruptly near or gradually from base, raised (adjacent blade obscurely raised) (+ depressed), 0.1-0.8 mm. wide at midpoint, below strongly raised, striate to angled, the venation above subobscure to apparent, below + apparent, raised, (4 to) 6 to 1 (to 21) veins/S mm., angle of divergence 50-80°. Inflorescences from foliate axils near apex and along twigs, with 3 to 11 flowers, unbranched (rarely 3-flowered branches to 1.2 cm. long, and/or flabellate), the axis (0.5—)1.3-6 cm. long, puberulent (tomentose) especially toward base, lowest internode (0.1-)0.5-3.4 cm. long; bracts unknown; pedicels 0.5-3.5(-4.3) cm. long, glabrous (rarely puberulent). Flower (?)hermaphroditic; tepals 4 to 8, the outer pair ovate to suborbicular, (4—-)5.5-7.5 by (3.5-)5—6(-6.5) mm., strongly concave, the next pair broadly elliptic, 5.5-10 by 4-7 mm. (rarely 7 by 6 mm.), the inner ones, if any, obovate to lingulate, 6.510 by 4-6.5 mm. (rarely 10 by 3 mm.); stamens 65 to 145, the filaments to 4.5 mm. long, the anthers oblong, 1.2-2 mm. long, slightly retuse to truncate at apex; ovary 1-2 mm. long, the style 2-3 mm. long, the stigma peltate, 0.5-1.2 mm. across, 2- or 3-radiate. Fruit spherical to ellipsoid, 1.7-3.7 by 1.4-3.2 cm., rounded (rarely pointed) at apex, drying dark vinaceous-brown, faintly striate to smooth; outer layer rarely detaching cleanly from stone, ( 1-)1.5-3 mm. thick, compact and rather hard; stone spherical to ellipsoid, 1.4—3 by 1.1-2.6 cm., rounded at apex, the walls 0.1-0.4(-0.6) mm. thick, smooth, unmarked: spongy layer thin; cotyledons of dried embryo separating easily. 432 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Key to the Varieties of Calophyllum teysmannii — _ Terminal bud usually + conical, initially enclosed by petiole bases of uppermost pair of leaves; stem often with horizontal lines at node; lamina drying + flat; inflorescences borne in upper leaf axils; neal (0.5-) 1-3.5(-4.3) cm. long. 2. Twig with prominent, V-shaped lines at node. . 78b. var. bursiculum. 2. Twig with at most obscure horizontal lines atnode. ....-.---: nae ee oe eee Fee Ae ee ee 8a. var. teysmannii. _ Terminal bud often plump, rarely enclosed by petiole bases of uppermost pair of leaves; stem practically never with horizontal lines at nodes; lamina often drying concave; inflorescences often borne in leaf axils along stem; pedicels 0.5-1.8(-3.5) cm. long. ........-.-- 78c. var. inophylloide. —_— 78a. Calophyllum teysmannii Mig. var. teysmannii C. teysmannii Mig.; F. Mueller in Walp. Ann. Syst. Bot. 7: 357. 1868; H. Keng, Gard. Bull. Singapore 28: 245. 1975; C. miquelii Vesque in C. DC. Monogr. Phanerog. 8: 607. 1893, nomen novum for C. teysmannii Miq. C. intramarginale M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 342. pl. 22. 1956. Type: Malaya, Trengganu, Gunong Padang, 4000 feet [1218 m.], June 1937, SFN 31900 coll. Moysey & Kiah (holotype, SING; isotypes, A, K, KEP). C. inophylloide King var. singapurense M. R. Henderson & Wyatt- -Smith, Gard. Bull. Singapore 15: 316. p/. 9. 1956; Kochummen, Malayan Forest Rec. ed. 2. 17: 215. 1965; a Common Sarawak Trees, 61. 1965, excl. spec. cit.; T. C. Whitmore, Gard. Bull. Singapore 26: 270, 1973, Tree Fl. Malaya 2: 186. 1973; Corner, Gard. Bull. Singapore Suppl. 1: 104. 1978. Type: Singapore, MacRitchie Reservoir, 10 Nov. 1936, SFN 32518 coll. Corner (holotype, sING; isotypes, K, KEP). C. inophylloide auct., non King; Ridley, Jour. Straits Branch Roy. Asiatic Soc. 34: 47. 1900, Fl. Malay Penin. 1: 186. 1922, pro parte. C. rhizophorum auct., non Teijsm. & Binn.; Meijer, Bot. Bull. Forest Dept. Sabah 7: 15. 1967. Figure 27. Hairs ae terminal bud, unless otherwise noted): a, Calophyl- lum enervosum (KEP 71894), from abaxial surface of bract, most hairs completely unthickened. b, C. pyriforme (SAN 75654). cH, C. ena c- ae var. teysmannil: c, d, S 11249, apex and base of hair ca. 1200 p of slightly birefringent hair ca. 390 pm. long; i, bb 27611, apex of hair ca. 1000 wm. long. j-p, C. ferrugineum, expanded apical cells ee iagieseey j-m, var. eM: j, k, KEP 105163, apex and base of hair ca. 720 ym. long; 1, m, SFN 28196, apex and base of hair ca. 660 wm. lo an n, var. are te (FRI 2594). 0, p, var. orientale (BRUN 5697), hairs to 1450 wm. long: o, from stem. tr, C. macrocarpum (S 15898), rare hair type from inside 4 s, t, C. aerarium (Cel. /I-354), Scale = 120 wm. (in c, d, scale = 240 p 434 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 61 C. ?cymosum auct., non Miq.; Koord.-Schum. Syst. Verzeich. 2: 39. 1910, quoad Koorders 10336. Calophyllum sp. 44 M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore > 350. pl. 30. 1956; T. C. Whitmore, Tree Fl. Malaya 2: 173. 1973 Corner, Gard. Bull. Singapore Suppl. 1: 105. 1978. Tree 3-36 meters tall; sometimes with loop roots; latex from cut bark yellow, fluid, clear, becoming opaque when rubbe Twigs (1—)1.5-2.5 mm. across, with obscure ped lines at nodes; terminal bud conical or + plump, (2-)3-8 mm. long, with sericeous to tomentose indumentum. Lamina usually obovate (rarely cuneiform, suboblong, subor- bicular, or subovate), (2.6—-)3.4-9 by (1.3-)2.1-4.6 cm., slightly recurved at margin, marginal thickening 0.4—0.8(-1.2) mm. wide, drying + flat, the midrib flat to raised, 0.15—0.3(-0.4) mm. wide at midpoint. Inflorescences from upper foliate axils, sometimes two together, lowest internode (0.7—)1.5—3.4 cm. long; pedicels (0.5—)1-3.5 cm. long. Distripution. Northeastern and southeastern Malay Peninsula to Borneo (Map 25). SELECTED SPECIMENS SEEN. Malaya. KEeLantan: G. Rabong, 671 m., FRJ 20622 (KEP); Sungei Perias near K. May, 305 m., FRI 4114 (k, KEP). TRENGGANU: along Sungei Pelong, FRIJ 14854 (k, KEP, sinc); Ulu Brang, near K. Lallang, 305 m., FRI 12593 (k, KEP, sinc); G. Padang, 1219 m., FRI 12644 (a, KEP, SAN, sina): Ulu Sungei Loh, 716 m., FRI 10840 (a, k, KEP, SING); Kuala Trengganu—Sengau, KEP 81266 (KeEp); Jerengau State Land, KEP 79853 (KEP); Bukit Bauk F.R., 120 m., KEP 77980 (ker); Dungun, Bukit Besi, KEP eed (KEP); Rasau Kerlih FR. 310 m., KEP 67753 (a, BO, K, KEP, SING); Kemam Bukit Kajang, Ulu Bendong, 150 m., Corner s.n., 3 Nov. 1935 (?) aes PaHANG: G. Tahan, Wong & Wyatt-Smith 168 (ker); Taman Negara near Map 25. Distribution of Calophyllum a ene var. teysmannili (circles) and C. teysmannii var. bursiculum (star) in Malesi 1980] STEVENS, CALOPHYLLUM 435 Kuala Tahan, 210 m., FR/J 20136 (kep). Jonore: G. Arong F.R., 30 m., KEP 54250 (Bo, Kk, KEP, SING); Kluang F.R., 305 m., KEP 98026 (k, KEP, sinG); Jemaluang F.R., KEP 73551 (ker); G. Blumut, 518 m., FRI 8839 (KEP); G. Panti E., 305-457 m., Shah & Ahmad 2917 (a, KEP, LAE, SING, uc); Kota Tinggi, PantiF.R., 210 m., KEP 70184 (ker, sinc); Kg. Hubong, Endau, Kadim & Noor 321 (k, L, sinc); Jason Bay, Sungei Semadan, Corner s.n., 30 April and 16 June 1934 (kep, sinc); Tanjong Jurat, Lake & Kelsall s.n., anno 1892 (sinc); Mawai, Corner s.n., 21 May 1934 (sinc). Singapore: Seletar Reservoir, Mandai ae Shah & Shukor 2398 (a, c, sinc); Botanic Gardens, SFN 28662 (A, BO, K, KEP, LAE, SING); MacRitchie Reservoir, SFN 32518 (kep); Stag Mount, Ridley s.n., anno 1909 (k); Bukit Timah, Baker 5339 (Gc, KEP, Nsw). Sumatra. Riau: Siak-Tanggana, Koorders 10336 (?) (Bo). Dsampi: Djambi, Simpang, 45 m., bb 13162 (Bo). Barat: Taram, E. of Pajakumbuh, 500-1000 m., Meijer 6914 (L). Borneo. Sarawak. Ist Division: Kuching, Stapok Road, Brooke 9413 (Gc, L). 4th Division: Baram, sag F.R., S 8289 (kK, L, SAN, SAR). BRUNEI: Bukit Puau, 24 m., Ashton , Aug. 1958 (sar). Saban. Beaufort: Lumut, on railway line, mile 21/2, SAN 58410 (L, SAN). Papar: Kimanis F.R., 6 m KEP 80288 (ker); Bongawan F.R., A 397 (a, K, KEP, L, SING). KALIMANTAN. Timur: Boelongan, Binai, Rutten 17 a Tengah: Bonjoet, bij Boentok, Obi 1880 (= bb 578) (Bo, L). Eco.ocy. Peat swamps, ‘‘secondary forest on mangroves’’ (SAN 58410), flat-lying mixed dipterocarp forest, kerangas vegetation, and ridges in lower montane rain forest; to 1220 m. alt. Flowing February, June to August, November, and December (flower scented); fruiting March to May, and July (inside of fruit magenta—FRI 8839). Lotong (wolverines) eat fruit (Corner n., 8 Feb. 1935) Bilabiate, purselike galls (Ficure 5, h) occur on plants of this taxon throughout its range (e.g., Malaya, Shah & Shukor 2398; Sumatra, Meijer 7037; Sarawak, Brooke 9413). These galls seem to be most common on young plants. Caused by a coccid (Lecanodiaspididae ~ Amorphococcus sp.), they develop abaxially on the midrib and shorten the internodes (Anthony, 1974). Anthony also reports psyllid galls that may affect each half of the leaf separately, thus causing it to become revolute. Elongate, slitlike galls on the lower and sometimes also on the upper surface of the lamina, generally found along the midrib or near the margin, are also known (e.g., KEP 66957, Malaya: SFN 32518, Singapore). Teysmann, HB 650, from Sumatra, has a row of raised, pustular swellings on either side of the midrib on the upper surface of the lamina. GERMINATION AND YOUNG PLANT. The radicle breaks the stone wall ca. 5 mm. to one side of the base. The seedling usually has two pairs of leaves separated by an internode less than 8 mm. long. (Some seedlings apparently have up to four pairs of leaves separated by internodes of ca. | cm. or more; the lowest two pairs of leaves drop off (Stevens et al. 108).) Subsequent growth is erect, the terminal bud is functional, and the internodes produced are well developed (more than | cm. long). In the young plant at least the terminal bud has tomentose indumentum. Leaves produced by young plants are oval-elliptic, less than 12 cm. long, and usually rounded (to subacute) at the apex. (KEP 77881, 99236; Stevens et al. 108.) 436 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 78b. Calophyllum teysmannii Miq. var. bursiculum P. F. Stevens, var. nov. Ficure 26, d, e. A varietatibus aliis Calophylli teysmannii in ramulis teretibus ad nodos lineis elevatis V-formatis basibus foliorum conflatentibus ornatis et lamina elliptica mediocri plana percrassa, differt. Tree 15-21 meters tall; latex unknown. Twigs 3.5—4 mm. across, with prominent, V-shaped lines at nodes, otherwise terete; terminal bud conical, 6-9 mm. long, sericeous. Lamina elliptic to suboblong (rarely suborbicular), 7.2-10.3 by 3.6-5.4 cm. (rarely ca. 4.2 by 4 cm.), slightly recurved at margin, marginal thickening not obvious, drying flat, the midrib slightly depressed, 0.15—0.55 mm. wide at midpoint. Inflores- cences from upper foliate axils, lowest internode 2—2.5 cm. long; pedicels in young fruit 2.4—4.3 cm. long Type: Sarawak, 3rd Division, Hose Mountains, Mujong, Ulu Telamud, 310 m., 25 March 1964, § 17230 coll. Ashton (holotype, k; isotypes, A, BO, L, SAN, SAR, SING). DistTRIBUTION. Sarawak, known only from the 3rd Division (Map 25). ADDITIONAL SPECIMEN SEEN. Borneo. SARAWAK. 3rd Division: Tatat Memuas, Ulu Tiau, Mujong, Balleh, 180 m., S 2/225 (k, L, Sar, SING) Ecotocy. Heath forest on Kakus sandstone plateau, 180 m. alt. (S 2/225); pole forest on dacite knoll, 310 m. alt. (S 17230). Elongate, slitlike galls ca. 1 cm. long occur on the lower surface of the lamina (S 17230). The name bursiculum (‘‘little purse’’) alludes to the way in which the terminal bud is enclosed by the petiole and lamina bases. 78c. Calophyllum teysmannii Miq. var. inophylloide (King) P. F. Stevens, comb. nov C. inophylloide ae Jour. Asiatic Soc. Bengal, II. 59: 178. 1890; Vesque in C. DC nogr. Phanerog. 8: 549. 1893; Ridley, Fl. Malay Penin. 1: 186. 1922, oe parte; C. inophylloide King var. inophylloide M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 315. 1956; Pukol & Ashton, Checklist Soe. 93. 1964; Kochummen, Malayan Forest Rec. ed. 2. 17: 213. 1965; T. C. Whitmore, Tree Fl. Malaya 2: 183. 1973. Type: Malaya, Perak, ce Padang district, Aug. 1885, 300-500 feet [90-150 m.], King’s collector [Kunstler] S112 (isotypes, kK, Uc). C. marginatum Wall. Catal. 4845. 1831; Choisy, Mém. Soc. Phys. Hist. at. Geneve 12: 424. 1849; Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 293. 1862; T. Anderson in Hooker f. Fl. Brit. India 1: 276. 1874; Vesque in C. DC. Monogr. Phanerog. 8: 608. 1893. Nomen C. borneénse auct., non Vesque; Pukol & Ashton, Checklist Brunei Trees, 93. 1964, pro parte. C. fraseri auct., non M. R. Henderson & Wyatt-Smith; T. C. Whitmore, Tree Fl. Malaya 2: 182. fig. 3. 1973, pro parte. 1980] STEVENS, CALOPHYLLUM 437 C. cymosum auct., non Miq.; Koord.-Schum. Syst. Verzeich. 2: 39. 1910, quoad Koorders 10335. Tree 18-40 meters tall; latex from cut bark yellow, clear, sticky, rather fluid, becoming opaque when rubbed. Twigs 2-5 mm. across, rarely with raised lines joining opposing leaf bases; terminal bud plump to conical, with grayish subcrustaceous to brown-tomen- tose indumentum. Lamina obovate to elliptic or oblong, (3.3-)5—13.8 by (2.2-) 2.6-8 cm., somewhat recurved at margin to strongly so, marginal thickening to 0.2-0.8 mm. wide, tending to dry + boat shaped, the midrib depressed to + raised, 0.15-0.7 mm. wide at midpoint. Inflorescences usually from foliate axils along stem, lowest internode (0.1-)0.3-1(-2) cm. long; pedicels 0.5-1.8 (-3.5) cm. long. DistripuTion. Malay Peninsula to Borneo (Map 26). SELECTED SPECIMENS SEEN (letters refer to discussion in text). Malaya. Kepau: Kedah Selatan Kulim, G. Inas, 305 m., KEP 105249 (kep). Pinanc: Telok Pahang, KEP 66369 (ker); Pantai Acheh F.R., KEP 27793 (kep); Pulau Penang, Curtis s.n., anno 1894 (sinc). Perak: near Tapah, Ridley 3014 (x); Larut, KEP 15 (KEP, SING); i F.R. (near Gri), KEP 31086 (ker); Changkat Jong F.R., 30 m., FRI 5873 (a, k, KEP); Taipeng, Selama, KEP 273 (KEP); es Kinta, KEP "39464 i Bruas F.R., 60 m., KEP 69406 (kep); Chikus KEP 3071 (ker); Pondok Tandjong, ‘KEP 65558 (KEP); Batang Padang 7 ae 15m., KEP 65053 (kep, sinc); Piah F.R., 853 m., FRI 9051 (f) (KEP). SELANGOR: Bukit Lagong F.R., KEP 13281 (kep, SING); Ulu Gombak F.R., nae 24703 (KEP, SING), 768 m., KEP 94716 (f) (A, K, KEP, SAN, SAR, SING); 1 r Gap, 800-900 m., Ando ‘et al. 88 (f) (KEP); Kanching, KEP 30106 pe NEGri SEMBILAN: Kuala Pilah, Serting, KEP 62956 (KEP, sING). TRENGGANU: along Sungei Pelong, FRI 14856 (k, KEP, sING); Dungun, Jerengau State Land, KEP 81407 (kEP); Kemaman, Ulu Bendong, Bukit Kajang, Corner s.n., Map 26. Distribution of Calophyllum eased var. inophylloide (circles) and C. celebicum (stars in solid circles) in Malesi 438 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 1935 (sinc); Kuala Trengganu, proposed Belara F.R., 60 m., KEP 76066 (KEP); Bukit Bauk F.R., 150 m., KEP 104709 (xkeEp); 28th Mile Jalan Trengganu, 30 m., KEP 78551 (kep); Mandi Angin exped., S. watershed of Sungei Loh, 792 m., FRI 12034 (f) (a, kep, sinc); Sungei Trengganu near Kuala Panchor, 610 m., FRI 20561 (?f, pro parte) (KEP). G. Padang exped., Ulu Brang, 1006 m., FRI 17776 (f) (A, KEP, SING). PAHANG: Sungei Kelui, G. Tapis, 518 m., FRI 10052 (ker); Pekan Road, Kuantan, KEP 43/02 (ker); Ulu Kali, 914 m., FRI 15643 (f) (kep); Fraser’s Hill, Bukit Jeriau, 1036 m., FRI 20412 (f) (KEP, SING). JoHoreE: Bukit Jelakoi, Kluang F.R., 225 m., KEP 76290 (Bo K, KEP, SING, US); Labis F.R., FR/J 3843 (KEP); Ginting Simpah, 610 m., KEP 71248 (kep); G. Pulai F.R., 610 m., FRI 9955 (ker). Sumatra and adjacent islands. Riau: Lingga, Sei. Soeling, Ri/I-3 (po); Indrag. Bovenlanden, Kwala Belilas, Buwalda 6754 (Bo, L); ae Langkat, Aloer Goetra, 50 m., bb 16592 (a, Bo). Barat: bij Padang, 80 m., Koorders 10335 (so); Bangkinang, Katoer, 300 m., bb 23395 (Bo, : Mo); Sidjoendjoeng, Moearo, bb 9098 (Bo). yee Sarawak. Ist Division: Semengoh F.R., Arboretum, 60 m., S 32979 (k, SAN, SAR, SING); G. Matang, 305 m., Stevens et al. 231 (a), 241 (c) (A); Sabal Tapang, 120 m., Stevens, sight record; G. Penrissen, 914 m., S 16397 (c) (A, BO, K, L, SAN, SAR, SING). 2nd Division: Simanggang, Kampong Mentu, 305 m., S 6527 (c) (K, L, SAR, SING). 3rd aan a Bukit Raya, Kapit, Pelagus, 270 m., S 14359 (kK, L, SAR, SING); Hose , Bukit Kajang Carapa, 900 m., $ 19090 (A, BO, L, SAR, SING). 4th ee “Miri NE. Lambir Hills, Tukau formation, 120 m., S 16756 (c) (K, L, sar). Brunet: G. Pagon Periok, 1829m., BRUN 2442(?) (san): Sagan Hill, 365 m., S 18711 (c) (sar); Temburong, Kuala Sekurop, 457 m., BRUN 733 (pri, K, L, SAR, SING); Kuala Belait, Andalau F.R., 60 m., SAN 17476 (c) (A, BO, K, KEP, L, P, SING); Berakas F.R., 30 m., S 2004 (c) (KEP, SAR, SING); Ulu Sugei, 60 m., S 10/3 (K, SAN, SAR, SING); Kuala Belalong, 610 m., Ashton s.n., Nov. 1959 (sar). SaBan. Beaufort: Beaufort Hill, 20 m., SAN 44588 (kK, L, SAN, SAR, SING). Ranau: Penibukan, 2134- 2438 ils SAN 75844 (k) (SAN), Colombon Basin, 1524 m., Clemens 40003 (k) (A, Bo, L, NY, Uc). Labuk & Sugut: Bukit Malawali, Labuk Road, SAN 51770 (san); Mt. Meliau near Kiabau, 487 m., SAN 51505 (c) (SAN); Bidu-Bidu hills above Kiabau, SAN 43803 (c) (SAN); near Sungei Binuang, NW. of Sapi, 457 m., SAN 44001 (?) (san); Bukit Tangkunan, Mile 86 Labuk Road, 425 m., Stevens et al. 410 (2c) (A), 75 m., Stevens et al. 414 (a). Kinabatangan: Lamag, Karamuak, E. of Sungei Meliau, 60 m., SAN 53255 (c) (SAN), Bukit Tavai, 150 m., SAN 53312 (san). Lahad Datu: Silam Mining Camp, 610 m., SAN 37481 (SAN, SAR). Tawau: Brassey Range, 770 m., Stevens 2 ee — (A). KALIMANTAN. Timur: Tidoengsche Landen, bb 18240 (A, BO, L); Telowek Daoen, Sungei Kasei, 75 m., bb 12195 (po); W. Kutei, Mt. ee near Tabang on Belajan R., 190 m., Kostermans 13144 (c) (Bo, kK, L, SING); 500 m., Kostermans 12792 (c) (Bo, L), 600 m., Kostermans 12878 (Bo, CANB, L, p), near Mt. Kemoel, 1200 m., Endert 3458 (k) (po, 1); Long Bleh, L). Barat: Melawi Tja atit, B. es 180 m., bb 27004 (Bo, K, L, SING); Landak, Ngabang, 40 m., bb 6402 (Bo); Mempawa, Toho, 300 m., bb 12388 (BO). Ecotocy. Usually well-drained lowland to colline mixed dipterocarp forest, 1980] STEVENS, CALOPHYLLUM 439 also in Trengganu (Malaya) in freshwater swamp, in Sarawak and Brunei in kerangas vegetation, in Kalimantan on (waterlogged) acid white sands, in Sabah on ultramafic rock; 4-1400(-2438?) m. alt. Flowering April, June to August, and September to November; fruiting December to February, April, August, and September (fruit greenish). Bilabiate, purselike coccid galls are known from Malaya (e.g., KEP 15, Stevens et al. 30) and from Kalimantan (Kostermans 12840). Elongate, slitlike, pustular galls on the lower surface of the lamina are known from Sarawak (Stevens et al, 275, Ficure 5, d). GERMINATION AND SEEDLING. As for Calophyllum teysmannii var. teysmannii. The leaves of the young plant are often larger (up to 22 by 7.5 cm.) and are shortly, but markedly, acuminate at the apex. (KEP 62957, 76055, 79305: FRI 23084; Stevens et al. 30, 137, 231, 410.) Loca use. The wood is rather hard and is used in construction. The epithet inophylloide (‘‘like [the leaves of] Calophyllum inophyllum’’) was chosen because the leaves of the type of C. inophylloide are similar to those of C. inophyllum. Although Calophyllum teysmannii is a variable taxon, it can generally be readily recognized by its moderate-sized, obovate (to oblong) leaf blades that are retuse at the apex and usually distinctly thickened at the margin. The axillary inflorescences are few flowered, and the flowers have four to eight tepals. The fruit is of moderate to large size and is more or less smooth when dry; the outer layer is thick ((1-)1.5-4 mm. across) and compact, and the stone walls are 0. 1—0.4(-0.6) mm. thick. The hairs are usually branched at the base and are moderately birefringent under polarized light, or they may be almost moruloid but still obviously branched. The seedlings nearly always have two pairs of leaves separated by an internode less than 8 mm. long. Young plants of vars. inophylloide and teysmannii may have characteristic bilabiate, purselike, coccid galls, while adult plants of all three varieties are known to have slitlike galls. The outer bark is brown, fissured, and scaly; stilt roots or spurs occur frequently. The specific epithet commemorates the indefatigable collector, J. E. Teijsmann. Specimens of saplings of Calophyllum teysmannii from Malaya have been confused with sterile specimens of C. subhorizontale. Although the two have similar terminal buds, the venation density of the leaves of C. teysmannii saplings is greater than that of C. subhorizontale, the twigs of the former do not dry the greenish color of the latter, and the thickened margin of the leaf common in C. teysmannii is not found in C. subhorizontale. Some of the specimens of Calophyllum teysmannii with very coriaceous leaf blades may be confused with C. sclerophyllum. The two species have similar fruits, and C. sclerophyllum also has hard heartwood and well-developed stilt roots. In the low-lying forest of southern Johore, C. sclerophyllum and C. teysmannii var. teysmannii (the former described as Calophyllum sp. 44 by Henderson & Wyatt-Smith, Joc. cit.) grow together, but they are quite distinct. In Sarawak both the local representative of C. teysmannii var. 440 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 inophylloide and its thick-leaved variant are quite distinct from the C. sclerophyllum pede awhousl in this case there is no ecological overlap. Calophyllum scl t obviously from C. teysmanniiin having the lower surface oe ne lamina often punctate, the terminal bud never enclosed by the bases of the petioles, the seedling with but a single pair of leaves and with subsequently produced leaves alternate, and the young plant without purselike coccid galls. Variation within Calophyllum teysmannii The variation within Calophyllum teysmannii is mostly in terminal bud shape and indumentum; leaf size, texture, and indumentum; inflorescence position; pedicel length; and fruit size. There is a certain amount of variation in the young plant and also in bark characters. Even when dealing with the Malayan plants, Henderson and Wyatt-Smith had trouble pices at which rank to recognize C. inophylloide var. inophylloide (= C. teysmannii var. inophylloide) and C. inophylloide var. singapurense (= C. teysmannii var. teysmannii). This problem has become exacerbated by the wides altitudi- nal, ecological, and geographic range of the species as here circumscribed. Much more collecting is still needed, especially from higher altitudes in both Malaya and Sabah, and also of the large, very coriaceous-leaved form that occurs throughout Borneo. Although there are a number of differences between Calophyllum teysmannii var. teysmannii (and var. bursiculum) and C. teysmannii var. inophylloide, these differences intergrade. The conical terminal bud of C. teysmannil var. teysmannii is often enclosed by the upright petioles of the terminal pair of leaves, whereas the plump bud of var. inophylloide is not often so enclosed since the petioles are spreading. Young plants of C. teysmannii var. inophylloide have transverse lines across the stem at the nodes, and each innovation seems to consist of only a single pair of leaves. The former, and probably also the latter, are characteristic of adult plants of the other two varieties, but C. teysmannii var. inophylloide probably produces more pairs of leaves each innovation. In Malaya and Singapore, the latex of the two varieties differs: in C. teysmannii var. teysmannii it is clear and fluid, turning opaque when rubbed; in var. inophylloide it is clear but sticky. There are perhaps a few exceptions. The latex of KEP 76055 (Trengganu; var. inophylloide) was reported to be nonsticky; however, it was also slow to appear, while the nonsticky latex of var. teysmannii flowed freely. In Sarawak, on the other hand, the slash characters and, to a certain extent, the leaf characters (e.g., Stevens et al. 273, 275) of C. teysmannii var. inophylloide are those of var. teysmannii in Malaya; unfortunately, I did not see any trees of var. teysmannii in Sarawak. It therefore seems most appropriate to reduce Calophyllum inophylloide toa variety of C. teysmannii. On Bukit Jelakoi, at the Kluang Forest Reserve, Johore, Malaya, the two varieties may grow quite near one another (KEP 76290, 225 m., var. inophylloide; KEP 76291, 210 m., var. teysmannii); this may also happen in northeastern Malaya. Calophyllum teysmannii var. bursiculum is described because, although 1980] STEVENS, CALOPHYLLUM 441 it has the bud, hair type, and inflorescence characteristic of var. teysmannii, it can be readily distinguished by its very coriaceous, short-petioled leaves that lack obvious marginal thickening. Its distinctive twigs, with their V-shaped lines at the nodes, also distinguish var. bursiculum from both other varieties. There is s relatively little variation within Calophyllum teysmannii var. Johore; in all characters of seedling and bark slash they were identical with C. teysmannii var. teysmannii, Some specimens tend to have a rather narrow lamina with a length : breadth ratio of 2-3 (that of fertile specimens of C. teysmannii var. teysmannii is usually less than 2). The fruit has an outer layer 1-1.3 (vs. more than 1.5) mm. thick. These are slight differences; Calophyllum sp. 44 is a local variant of var. teysmannii not distinct enough for formal recognition. Two specimens from Trengganu (FRI 12593, 14854) have obovate to cuneiform leaf blades and ellipsoid fruits that are more or less pointed at the apex; the outer layer of the fruit is only ca. | mm. thick and detaches cleanly from the stone. The subtomentose indumentum on the twigs, midrib, and base of the inflorescence is better developed than is usual in adult plants of Calophyllum teysmannii var. teysmannii. Although these specimens are referred to this variety, the characters listed above suggest that they may be hybrids with C. ferrugineum var. oblongifolium, which grows in the same area. Another specimen collected in the mountains of Trengganu, FRI 12644, looks similar to the form of Calophyllum teysmannii var. inophyl- loide growing on the same mountain (Gunong Padang, see below). However, it has the terminal bud and transversely lined nodes of var. teysmannii, and it is included in this taxon. Calophyllum teysmannii var. inophylloide is a heterogeneous taxon as delimited here. Whitmore (1973, Joc. cit.) included in his concept of C. fraseri specimens that seem to represent a local high-altitude form of C. teysmannii var. inophylloide (‘‘f’’ in the list above). They sometimes have the stilt roots of the latter taxon, while C. fraseri lacks them. Stones and germinating seeds of FRI 23084 (from Fraser’s Hill) are identical to those of C. teysmannii, and hairs of specimens of this type, although very small, can perhaps be compared to the branched bases of the low-altitude form. These specimens have rather narrow, long-petiolate leaves ee the blades lacking the obvious thickened margin of C. teysmannii. FRI 19869 (Pahang, 792 m.) has bilabiate coccid galls. Specimens collected from The Gap (Selangor) seem to show a transition between this form and more typical C. teysmannii var. inophylloide, as suggested by the series of specimens collected by Ando et al. (88 (the form), 85, 151, 125 (close to var. inophylloide)). There are a number of specimens from Borneo with large, very coriaceous leaf blades that are strongly recurved at the margin and that have a more or less depressed midrib (‘‘c’’ in the above list). These robust specimens are at first sight very different from the rest of Calophyllum teysmannii var. inophylloide. However, specimens like Kostermans 13144 have a more 442 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 or less raised midrib, while others such as Kostermans 12743 and S§ 1115 (the latter is in fruit) have lines across the stems at the nodes. Indumentum is quite variable: Kostermans 12743 and Stevens et al. 241 have short, tomentose indumentum on the bud, stem, midrib, and leaf margin; SAN 17476 and Kostermans 13144 have short, grayish, almost furfuraceous indumentum on the terminal bud and little obvious indumentum elsewhere. The shape of the terminal bud varies from conical to plump. It would be unwise to recognize these large, coriaceous-leaved specimens formally, but it should be emphasized that this variation is not understood. On Gunong Matang (Sarawak) two forms, ordinary var. inophylloide and a very coriaceous-leaved form with subtomentose indumentum, grow on the same ridge; on Mt. Palimasan (Kalimantan) two similar forms, as well as a very coriaceous-leaved form that lacks tomentose indumentum, grow in the same general area. In northern Borneo there are some specimens that dry rather similar to the local form of Calophyllum biflorum. One of these is Stevens et al. 125 (from Sarawak), which has the bark and slash characters of C. teysmannii var. inophylloide. Another, S 1013 (from Brunei, has leaves drying somewhat like those of C. ferrugineum var. orientale (close to C. biflorum, see below), as well as the inconspicuous tomentose indumentum of that species. However, the hairs have many expanded, rather thin-walled apical cells in comparison to the two to four thick-walled cells found in C. ferrugineum var. orientale. The field characters of S 10/3 are those of C. teysmannii. On the lower slopes of Mt. Kinabalu, Sabah, and on Mt. Kemoel, Kaliman- tan, acres is perhaps another form of Calophyllum teysmannii var. inophylloide in the above list). In the young plants the terminal buds are protected by the petiole bases (Stevens et al. 495), and the hair, bark, and slash characters are compatible with those of this variety. However, the only specimen with flowers (Clemens 40003) has puberulo-tomentose indumentum on the inflores- cence axis and pedicels, and the flowers have 175 to 325 stamens. Fruits are unknown. More collections of this form are needed to establish its status; it has not been incorporated in the description. Synonymy and Nomenclature There is no doubt about the synonymy of the names listed under Calophyllum teysmannii var. teysmannii. The type specimen of C. feysmannii has rather small, obovate leaf blades with a prominent thickened band at the margin, and inflorescences with few rather large, long-pediceled flowers. Calophyllum intramarginale was described from similar specimens that were growing at lower altitudes, but that otherwise closely match the type specimen of C. teysmannil. The publication of Calophyllum teysmannii Miq. just antedates that of C. teysmannii Zoll. ex Planchon & Triana. That part of the supplement to Miquel’s Flora Indiae Batavae in which the name C. teysmannii Miq. is validly published came out in December, 1861 (the name is also mentioned on p. 193 of the supplement published in June, 1861 (Stafleu, 1967)). However, the portion of Planchon and Triana’s ‘‘Mémoire sur la famille des Guttiferes 1980] STEVENS, CALOPHYLLUM 443 in which the name C. teysmannii Zoll. ex Planchon & Triana was published appeared early in 1862.'* Calophyllum teysmannii Zoll. ex Planchon & Triana is itself a superfluous name for C. dasypodum (q.v.). alophyllum marginatum, a manuscript name of Wallich, has been something of a mystery to earlier authors. Vesque (/oc. cit.) said that the name made him think of C. pulcherrimum (properly C. tetrapterum), the leaves of which often have a pale margin. However, the specimen of Wallich dist. 4845 at Kew was taken from a young plant of C. teysmannii var. inophylloide and has a thickened leaf margin, lines across the twigs at the nodes, and purselike 79. Calophyllum aerarium P. F. Stevens, sp. nov. Ficure 26, b. A speciebus aliis Calophylli in gemma terminali 0.6-1.5 cm. longa, lamina oblonga vel obovata apice rotundata in siccitate aerata, inflorescentiis axillari- bus puberulentibus, et fructu circa 2 cm. longo in siccitate arcte rugoso strato exteriore valido circa 1.3 mm. crasso putamine parietibus tenuissimis, i Tree ca. 30 meters tall, d.b.h. ca. 50 cm.; trunk and bark unknown; latex yellow. Twigs flattened, 2.5-4 mm. across, + 4-angled when young, often with obscure horizontal line at nodes, drying brown, transiently brown-farinose; axillary innovations lacking basal scars, internodes (1—)2-8.5 cm. long; upper- most pair of axillary buds rounded to pointed, 1-3 mm. long, spreading, conspicuous; terminal bud narrowly conical, 0.6-1.5 cm. long, with brown, + furfuraceous indumentum (hairs, Ficure 27, s, t), underdeveloped inter- node 1-3 mm. long. Petiole 1-3 cm. long, shallowly concave above, con- vex below, drying blackish, transiently farinose; lamina obovate or subob- long to elliptic, 6.5-14 by 3.5-7 cm., broadly rounded at apex, whether or not apiculate, acute at base, rather deeply but distantly undulate and strongly recurved at margin, coriaceous, drying umber above and sabelline-sepia below, + transiently farinose on midrib on both surfaces, the midrib above narrowing gradually from base, + flat at first, center sulcate, becoming raised (surrounding lamina obscurely raised), 0.2-0.4 mm. wide at midpoint, below raised, angled (rounded toward base), venation + apparent above and below, raised, 6 to 10 veins/5 mm., angle of divergence 65-76°. Infructescences from upper- most foliate axils, with ca. 5 flowers, eam: the axis 2-3.5 cm. long, puberulent, lowest internode 0.4-2.2 cm. long; bracts unknown; pedicels 4-10 mm. long, + puberulent. Only oes flowers known, (?)hermaphroditic; type asap of Quiina decaisneana Planchon & Triana, soe in the *s article (p. 315), is cited as ‘“‘Guyana (Melinon, serres du Muséum, 186 )? Ina a, article in the same volume (H. F. Hance, ‘““Symbolae ad floram Sinicam,’’ pp. 220-230), specimens are cited as having been Spee in Canton as late as ae Te) (p. 229), while on page 220 Hance wrote ‘“‘Whampoa sinensium Scrip 1861. H. F. H.’’ This makes a publication date of eae 1862 for PIEnChO and Tia s article very probable. (I am extremely grateful to Dr. H. Heine, Paris, for drawing this evidence to my attention.) 444 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 tepals 6 (?always), the outer pair broadly ovate, ca. 6 by 7.5 mm., puberulent on back, the inner ones to 13 by 6.5 mm.; stamens numerous, the filaments to 4.5 mm. long, the anthers oblong, ca. 1.2 mm. long, retuse at apex; ovary ca. 2.5 mm. long, style and stigma unknown. Fruit + spherical, ca. 2.1 by 1.9 cm., apiculate, young fruit ovoid and sharply apiculate, drying brown, closely, sharply, and rather shallowly wrinkled; outer layer not detaching cleanly from stone, ca. 1.3 mm. thick, compact and very tough; stone spherical, ca. 1.6 cm. long and across, rounded at apex, the walls less than 0.05 mm. thick, smooth, (?)unmarked; spongy layer thin. Type: Celebes, Malili, Oesoe, 100 m., 2 May 1921, Cel. /II-354 coll. Waturandang (holotype, Bo). DistriBUTION. Celebes (Map 27). ADDITIONAL SPECIMENS SEEN. Celebes. SuLawesi: Malili, Oesoe, 100 m., Cel. /II-354 coll. Reppie (Bo), Cel. /II-354 coll. anon. (a, L); Pankadjene, Teysmann, HB 12086 (po); Baleh-angien, Teysmann, HB 12496 (Bo). Ecovocy. Fruiting in February. Crateriform, pustular galls occur just in from the margin on the lower surface of the lamina in Cel. /I/-354 coll. Waturandang. Calophyllum aerarium can be recognized by its long, narrow, terminal bud; its long petioles; its fairly large, obovate to elliptic lamina that is broadly rounded at the apex (whether or not the very apex is apiculate); its axillary, puberulent inflorescences; and its closely wrinkled fruit with a well-developed, compact, very strong outer layer and a very thin stone wall. The leaves Map 27. Distribution of Calophyllum sclerophyllum (circles, localized speci- mens; triangles, unlocalized specimens) and C. aerarium (stars in solid circles) in Malesia. 1980] STEVENS, CALOPHYLLUM 445 dry a color close to bronze, hence the specific epithet aerarium (‘‘made of copper or bronze’’). Calophyllum aerarium is superficially like C. alboramulum, but the latter has uppermost axillary buds that are suberect, leaf blades that are acuminate at the apex, twigs that dry a paler color, and terminal inflorescences. It also approaches C. sclerophyllum, but can be distinguished by its larger flowers and hairs, longer terminal buds, and thinner leaves drying a different color. 80. Calophyllum celebicum P. F. Stevens, sp. nov. FicureE 28, a—c. A speciebus aliis Calophylli in ramulis valde 4-angulatis, lamina suboblonga plus minusve valde coriacea spissescenti marginali 0.5—-0.8 mm. lati, floribus cum 8 tepalis, fructu strato exteriore valido 1-1.2 mm. crasso, putamine parietibus tenuissimis, differt. Tree 20-30 meters tall, d.b.h. to 50 cm.; trunk and outer bark not known; latex yellow to yellowish pink. Twigs slightly flattened, 3.5-5.5 mm. across, strongly and persistently 4-angled to 4-alate, drying blackish brown to blackish, transiently brown-fari- nose; axillary innovations lacking basal scars; internodes 1-3.5 mm. long; uppermost pair of axillary buds subrounded, ca. 1(-2) mm. long, + spreading, inconspicuous; terminal bud plump, 7-9 mm. long, with crustaceous, gray to brown indumentum (hairs, Ficure 22, r-t; some also moruloid), underdevel- oped internode 1-3 mm. long. Petiole 0.8-2 cm. long, broadly and shallowly concave above and convex below, glabrous, drying black; lamina oblong or elliptic to obovate, (7.5-)10-20.2 by (3.7-)4.6-9.6 cm., rounded (rarely obtuse or shallowly retuse) at apex, cuneate to broadly rounded at base, slightly and distantly undulate and somewhat recurved at margin, marginal thickening 0.5-0.8 mm. wide, very coriaceous, drying sepia above and umber below, transiently to subpersistently farinose on midrib below, the midrib above usually abruptly narrowed at (rarely narrowing gradually from) base, depressed, the margins raised or not, 0.35—-0.6 mm. wide at midpoint, becoming long; bracts ovate, to 5 mm. long, deciduous; pedicels ca. 1 cm. long, + glabrous, in fruit 1.8-2.6 cm. by up to 3.5 mm. Flower known only in bud, (?)hermaphroditic; tepals 8, inner ones glabrous; stamens ca. 150, the anthers oblong, |-1.6 mm. long; stigma peltate, ca. 0.7 mm. across. Fruit ovoid to ellipsoid, 2-2.3 by 1.6-1.7 cm., rounded to acute at apex, drying brown, finely striate; outer layer not detaching cleanly from stone, 1-1.2 mm. thick, compact, tough; stone ca. 1.7 by 1.3 cm., rounded at apex, the walls less than 0.1 mm. thick, smooth, (?)unmarked; spongy layer thin. Type: Celebes, Ond. Malili, Oesoe, 200 m., 27 Jan. 1934, Cel. /1-213 coll. Waturandang (holotype, a; isotype, Bo). 446 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 N ne iG a us Figure 28. a-c, Calophyllum celebicum (Cel. /I-213). a, terminal bud, x .b, c, fruit, x 1: b, from outside; c, transverse section. = C. ceriferum. d, f, Poilane 6172: d, terminal bud, x 6; f, habit, x 0.5. e, Robinson 1748, young infructescence, x 0.5. g, h, C. macrocarpum, fruit, en in section, x 0.5: g, FRI 10926; h, S 15898. i, C. a aes (Kostermans 14022), terminal bud, x 6. j-l, C. havilandii (SAN 17449). j, terminal ps x 6. k, 7 it, but in indumentum distribution and type it is similar to other Specinieae eee to C. ferrugineum; the t taxa are fae same, and the correct name for the combined taxon is C. ferrugine ee follooane Vesque (see above) and misled by the appearance of Wallich dist. 4846, authors working on the flora of the Malay Peninsula have recently used the name ‘“‘Calophyllum retusum’”’ for the taxon described below as C. sundaicum. Calophyllum sundaicum has a shorter inflorescence and less conspicuous indumentum than are found in C. ferrugineum, and the venation on the upper surface of the leaf blade is the same density as that on the lower The synonymy given above for Calophyllum ferrugineum and its varieties presents no problems. The name obdlongifolium must replace the more appropriate neriifolium for the most northerly variant of C. ferrugineum. The name C. ferrugineum itself is here lectotypified by the sheet of Ridley 10842 at Singapore; Ridley (Joc. cit.) also cited Ridley 4799. 85. Calophyllum sundaicum P. F. Stevens, sp. nov. Ficure 28, m-o. C. retusum auct., non Wall. ex Choisy; M. R. Henderson’& Wyatt-Smith, Gard. Bull. Singapore 15: 331. pl. 15. 1956; J. Anderson, Gard. Bull. Singapore 20: 154. 1963; Kochummen, Malayan Forest Rec. ed. 2. 17: 218. 1965; Smythies, Common Sarawak Trees, 61. 1965, excl. spec. cit.; T. C. Whitmore, Tree Fl. Malaya 2: 188. 1973; H. Keng, Gard. Bull. Singapore 28: 255. 1976; Corner, Gard. Bull. Singapore Suppl. 1: 104. 1978. A speciebus aliis Calophylli in foliis mediocris ellipticis vel oblongis apice plerumque retusis in siccitate marginibus planis nervis lateralibus densis (11 usque ad 18 per 5 mm.), et fructu ellipsoideo strato exteriore rugoso e putamine munde secedenti pagina interiore nitida plus minusve striata, differt. Tree 22-28 meters tall, d.b.h. to 65 cm.; trunk without buttresses (with loop roots); outer bark brown to yellowish, shallowly and closely fissured, or with lenticels in lines, or cracked, the inner surface dark brown to black; under bark orange-brown to dark straw; inner bark pale to dark red; latex yellow, clear, very sticky. Twigs flattened, (1.5-)2-3 mm. across, 4-angled, drying dark brown to blackish, brown-puberulent when young; axillary innovations lacking basal scars; internodes 0.7-4 cm. long; uppermost pair of axillary buds subrounded, to 3 mm. long, erect; terminal bud plump, 5-10 mm. long, with brown to grayish, puberulent to subtomentose indumentum (hairs, Ficure 29, c-e), underdeveloped internode to 3 mm. long. Petiole 0.8-2 cm. long, concave to V-shaped above and convex below, + puberulent when young; lamina elliptic to oblong, (3.2—)4.5—10(-14.5) by 2.5-5.5 cm., + retuse (rounded to subacute) at apex, cuneate to rounded at base, not undulate to slightly so 468 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 and recurved at margin, coriaceous, drying bay to castaneous or dark brick above and + umber below, puberulent on midrib on both surfaces, the midrib above usually narrowing near base, + depressed at first, becoming raised, or raised from base, 0.15-0.3 mm. wide at midpoint, below raised, striate to subangled (edges depressed), the venation subobscure on both surfaces, slightly raised, 11 to 18 veins/5 mm., angle of divergence (50-)70-80°. Inflo- rescences from foliate axils (rarely terminal—hb 4951), with 7 to 11 flowers, unbranched, terminal flowers tending to be 5 together, others in fours, the axis 0.9-2.2(-4) cm. long, puberulent toward base, lowest internode (0.15-) 0.4-1.8 cm. long; bracts ovate, 1.5-2.7 mm. long, not persistent; pedicels ca. 10 mm. long, glabrous to puberulent. Flower (?)hermaphroditic; tepals 4, the outer pair ovate, ca. 6 by 4 mm., rather fleshy, the inner pair + obovate, 7-7.5 by 4.5-6 mm.; stamens 100 to 170, the filaments to 4 mm. long, the anthers oblong, 1.2-2 mm. long, retuse at apex; ovary ca. 1.5 mm. long, the style ca. 3.2 mm. long, the stigma peltate, 1.2—-1.6 mm. across, 3-lobed. Fruit ellipsoid, 1.5-2.1 by 1-1.4 cm., rounded at apex, drying brown, pruinose, wrinkled; outer layer detaching cleanly from stone, 0.5-1.3 mm. thick, compact, inner surface shining and striate; stone ellipsoid, 1.3-1.8 by 1-1.2 cm., rounded at apex, the walls 0.15-0.2 mm. thick, smooth, unmarked; spongy layer thin. Type: Singapore, Seletar Forest, behind Nee Soon vegetable gardens, 16 June 1951, SFN 39252 coll. Sinclair (holotype, sinc; isotypes, K, KEP, L), DistripuTion. Southern Malay Peninsula to western Borneo, excluding Java (Map 30 SELECTED SPECIMENS SEEN. Malaya. SELANGor: Sungai Tinggi, Kuala Selangor, SFN 34087 (a, K, KEP, L, LAE, SING); Klang, Telok F.R., KEP 23230 (k, sING); Selatan, KEP 96602 (ker); Kuala Langat, KEP 53951 (ker); Olar Simpit, KEP 43692 (kep). PAHANG: Pekan, State Land, Batu 2 Jalan Kuantan, KEP 69601 (Ker); Menchali F.R., KEP 67623 (ker); Kuantan, Pekan—Ninen road, KEP 77956 (kep). Jonore: Pontian, Api-Api, KEP 75603 (Bo, kK, KEP, L, SING), Pengkalan Raja, SFN 36637 (sinc); Pontian Besar, KEP 70261 te Kiri Mudek [Sungei Benut], KEP 73053 (ker); Ayer Hitam, KEP 7020 eee sinc); 8th Mile, Kota Tinggi-Mawai road, SFN 29050 (Bo, K, KEP, SING); miles Muar-Pt. Sulong road, KEP 74107 (kep). Singapore: Jurong, SFN pe (A, K, KEP, L, LAE, SING); [11/2 miles Mandai Road, SFN 40268 (a, sina); Botanic Gardens, SGN 1633 Cenc. (sING). Sumatra and adjacent islands. Riau: Karimoen, Telok Sabuk, , bb 5309 (Bo), Pangka, 10 m., bb 7360 (Bo), Tandjong Poendoer, IS m., bb 6306 (Bo); Indragirische Bovenlanden, . Gelang, 4m., bb 29116 (a, Bo, K, L, NY, sinc); Bengkalis, Poeloe Rangsang, Panglong 222, 3 m., Beguin 505 (Bo, L), Telok Oekis, 3 m., bb 12853 (Bo), P. Mendal, Keloemang, bb 12473 See Selatpandjang, Kampar-monding, 5 m., 5b 22059 (Bo, L); Laboean Batoe, Laboean Bilik, 5-10 m., bb 4951 (Bo); Soengei Ranah, 5-10 m., Prunier 286 (Bo). Utara: Bila, 40 m., Lérzing 14227 (so). Borneo, Sarawak. Ist Division: Setapok F.R., 30 m., S 4817 (k, L, SAR, SING); Unjam F.R., 0 m., S 14464 (sar). 3rd Division: Daro F.R., Si 126 coll. Anderson, Sept. 1953 (sar); Loba Kabang South Protected Forest, KEP 79329 (sar); Pulau Burit, Sungei Kelepu, 3 m., S 8039 (?) (sar). 4th 1980] STEVENS, CALOPHYLLUM 469 Map 30. Distribution of Calophyllum sundaicum in Malesia. Inset: C. havilandii. Division: Miri, Sungei Dalan F.R., 10 m., S 1422 (KEP, sar). KALIMANTAN. Barat: Soemban, 10 m., bb 18386 (a, Bo, L); Batoe Lajang, 4 m., bb 9675 (?) (Bo). Eco.tocy. Locally common in peat swamps, to 40 m. alt. In Sarawak sometimes in kerangas forest (S 1422), Flowering June and September to December; fruiting March and May (fruit greenish). GERMINATION AND YOUNG PLANT. The radicle probably breaks through the stone to one side of the base. The seedling has two pairs of leaves separated by an internode ca. 5 mm.(—1 cm.) long. Subsequently produced internodes are longer, the terminal bud is functional, and the young plant is erect. (Stevens et al. 41.) Calophyllum sundaicum can be recognized by its puberulent indumentum and by its medium-sized, elliptic to oblong leaf blades that are rounded to retuse at the apex, that have rather dense venation above and below, and that have almost flat margins even when dry. The inflorescence axis is fairly short, and the flowers have four tepals and tend to be in groups of four. The fruit is sharply wrinkled, with the outer layer detaching cleanly from the stone; the inner surface of the outer layer is shiny. The places where C. sundaicum grows, although now separated by water, were once part of Sundaland, hence the specific epithet. The characters that separate Calophyllum sundaicum from related species are summarized in TasLe 13. The nomenclatural problems surrounding the name C. retusum, which has been used for this taxon, are discussed under C. ferrugineum. 470 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 The inflorescence of Calophyllum sundaicum is characteristically short and congested, with the axis less than 2.5 cm. and the lowest internode often less than | cm. However, in specimens such as KEP 75603 (Malaya), and Si 126 and S 4817 (Sarawak), the inflorescences are longer and more like those characteristic of C. ferrugineum, the basal internode being more than 1 cm. long. In the herbarium of the Botanic Gardens, Singapore, there is a specimen of Calophyllum sundaicum that may have been collected in Malacca. The specimen bears the number ‘‘463’’ and has a local name written in Arabic (‘‘kayo bintangor batu’’); the writing is similar to that on some specimens of C. pisiferum, also probably collected in Malacca. 86. Calophyllum costulatum M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 344. pl. 26. 1956; T. C. Whitmore, Tree Fl. Malaya 2: 173. 1973; H. Keng, Gard. Bull. Singapore 38: 244. 1976. Type: Malaya, Johore, Tanjong Bunga, June 1894, Ridley 6332 (holotype, SING; 1SOtypes, BM, W). Tree 5-30 meters tall, d.b.h. to 18 cm.; trunk without buttresses or spurs; outer bark brown to yellowish brown, with vertical rows of flat to pustular lenticels, hoop marked, the inner surface yellow to bright orange; under bark dull reddish; inner bark pale red to pink; latex yellow, slightly opaque or (?)clear, very sticky. Twigs flattened, 2-3 mm. across, 4-angled, drying blackish to yellowish when young, when older yellowish, + persistently (pale) brown-tomentose; axillary innovations lacking basal scars; internodes 0.5—3.5 cm. long, lowest internode of axillary innovation often notably longer than others, uppermost pair of axillary buds rounded, 2-2.5 mm. long, erect; terminal bud plump, 3.5-5.5 mm. long, with brown, tomentose indumentum (hairs, Ficure 29, f-i), underdeveloped internode |-2.5 mm. long. Petiole 2.5-6.5 mm. long, broadly and rather deeply concave above, convex below, + persistently short-tomentose; lamina elliptic to oblong, 2.4-7(-11) by 1.6-3.1(-4.2) cm., retuse at apex, broadly rounded at base, not undulate to slightly so and recurved at margin, coriaceous, drying umber to sepia above (margins much paler) and fulvous to sabelline below, + persistently tomentose on midrib below and often above and on margins, the midrib above narrowing quickly or rather quickly near base, raised, 0.12-2 mm. wide at midpoint, below strongly raised, striate to rounded, venation apparent above, apparent to subobscure below, raised, 10 to 16 veins/5 mm., angle of divergence 70-80°. Inflorescences from foliate axils (very rarely terminal), with 5 to 11 flowers, terminal flowers often in fives and sevens, others in fours, unbranched, the axis 3.8-9.5 cm. long, tomentose, lowest internode 1.7-5.2 cm. long; bracts unknown; pedicels I-1.7 cm. long, tomentose. Flower (?)hermaphrodi- tic; tepals 4, the outer pair suborbicular, ca. 4 mm. long and across, backs tomentose-canescent, the inner ones broadly eee to elliptic, 6-7 by ca 6 mm.; stamens 210 to 260, the filaments to 4.2 mm. long, the anthers broadly elliptic, 0.35-0.5 mm. long, retuse at apex; ovary ca. 1.2 mm. long, the 1980] STEVENS, CALOPHYLLUM 471 style ca. 3.5 mm. long, the stigma peltate, ca. 0.8 mm. across, 2-lobed. Fruit spherical, ca. 1.4 cm. long and across, rounded at apex, drying brown, m cm. across, rounded at apex, the walls ca. 0.25 mm. thick, smooth, unmarked; spongy layer thin. DistripuTIon. Malaya and Singapore (Map 36). ADDITIONAL SPECIMENS SEEN. Malaya. Perak: G. Besout F.R., Slim R., 90 m., FRI 1767 (kK, KEP, L, SAN, SING). JoHorE: Kpg. Hubong, Endau, Kadim & pe oe (A, L, LAE, SING); Panti F.R., Kota Tinggi, 564 m., KEP 94367 (Ca ; G. Arong F. R., 15 m., Stevens et al. 84 (2) (a). Singapore: Bee on Langlassé 123 (<, P). Eco.tocy. Swamps, flat land, or hillsides, primary and secondary forest; 15-564 m. alt. Flowering June to August (flower scented); fruiting in December (fruit green). Calophyllum costulatum can be recognized by the dense, often whitish, tomentose indumentum on the twig, terminal bud, midrib, inflorescence axis, epithet costulatum, the diminutive of costa (midrib), refers to the narrow midrib on the upper surface of the lamina Calophyllum costulatum is clearly related to the C. ferrugineum complex; the characters by which it differs are given in TaBLe 13. At least sometimes C. costulatum has a discolored, pale leaf margin like C. biflorum. In the long, lowest internode of the inflorescence, it is like C. ferrugineum, and in general fruit structure it agrees with all the members of the complex, although the dried fruit is more finely wrinkled than it is in the other taxa. The hairs of C. costulatum are slightly birefringent, and the apical cells, although often very large, are unthickened and empty (hence the whitish color of the indumentum); they are perhaps most similar to the hairs of C. ferrugineum vars. ferrugineum and oblongifolium. The lamina of Stevens et al. 84 dried similar to that of Calophyllum biflorum, and the scars of the flowers on the infructescence do not appear to be in fours; itis included in C. costulat nly with hesitation. A more problematic specimen is Soepadmo 53 (Sumatra, Upper Riauw, Pakanbaru, Tonajan River, in Swamp; A, B, BO, C, L, NY, SING), which has the very coriaceous lamina and the indumentum distribution of C. costulatum, but the lamina is cuneate to acute at the base, the venation is obscure on both surfaces, and the midrib on the lower surface is coarsely striate. There does not seem to be any tendency for the scars of the flowers on the infructescence to be in fours. Both specimens have hairs like those of more typical specimens. On some duplicates of Ridley 6332, the collection date is given as May 7. Whether this is an error or not is unknown 472 JOURNAL OF THE ARNOLD ARBORETUM [voL. 6] 87. ar seta biflorum M. R. Henderson & Wyatt-Smith, Gard. Bull. apore 15: 349. p/. 29. 1956; Kochummen, Malayan Forest Rec. = 2. 17: 217. 1965; T. C. Whitmore, Tree Fl. Malaya 2: 176, 1973; H. Keng, Gard. Bull. Singapore 28: 244. 1976. Type: Malaya, Johore, Gunong Arong Forest Reserve, 18 Nov. 1948, KEP 35963 coll. bin Talib (holotype, KEP; isotype, K). C. aren auct., non Wall. ex Choisy; Smythies, Common Sarawak Trees, 1965, quoad spec. cit. a tae King var. singapurense auct., non M. R. Henderson & Wyatt-Smith; Smythies, Common Sarawak Trees, 61. p/. 20. 1965, et quoad spec. cit. Tree 6-36 meters tall, d.b.h. to 95 cm.; trunk without buttresses (very rarely with spurs); outer bark yellowish and brownish mottled, with distant, shallow, boat-shaped fissures or lines of lenticels and with areas of darker, rougher bark, or brownish, sometimes scaly, often hoop marked, the inner surface mottled orange and dirty brown, to deep brown; under bark reddish; inner bark red, laminated; latex white, often curdy, + sticky (yellow, clear or opaque). Twigs flattened, 1.2-3(-3.5) mm. across, + 4-angled, or rounded, drying (rarely pale) brown or dark brown, transiently farinose to subpersistently tomentose; axillary innovations lacking basal scars; internodes 1-6.5(-7.5) cm. long; uppermost pair of axillary buds + rounded, 0.5-3 mm. long, erect; terminal bud plump to conical, 2.5-9.5 cm. long, usually with brown, tomentose (subcrustaceous, grayish) indumentum (hairs, Ficure 29, j-l), underdeveloped internode to 3(-7) mm. long. Petiole 0.5-1.8 cm. long, broadly concave above, convex below, glabrescent or subpersistently tomentose; lamina oblong to subelliptic or obovate, 3.5-12.5 by 1.7-4.5 cm., retuse to rounded (rarely subacute) at apex, acute to cuneate at base, slightly to strongly and distantly undulate and barely recurved at margin, coriaceous to very coriaceous, drying grayish sepia to cinnamon above and hazel to sabelline below, with subpersis- tent, brown, farinose to tomentose indumentum on midrib on both surfaces, or soon glabrescent, the midrib above usually narrowing rather quickly near or gradually from base, raised, 0.15—0.35 mm. wide at midpoint, below raised, striate, the venation apparent (rarely subobscure) above and below, + raised, 7 to IS (to 18) veins/5 mm., angle of divergence 55-75°. Inflorescences from foliate axils near ends of twigs, with 5 to 21 flowers, sometimes flabellate, and/or with 3-flowered branches to 1.5 cm. long, the axis 0.8-6.5 cm. long, + tomentose on lowest internode, lowest internode (0.2-)0.6-3.2 cm. long; bracts narrowly ovate or elliptic, 2-3 mm. long, deciduous; pedicels 0.8-3.2 cm. long, glabrous, pronouncedly incrassate in fruit or not. Flower (?)her- maphroditic; tepals 4, glabrous, the outer pair ovate to broadly elliptic, 4.5—6 by 3-4.5 mm., the inner pair obovate, ca. 8 by 4 mm.; stamens 60 to 225, the filaments to 3 mm. long, the anthers oblong, ca. 1.2 mm. long, retuse at apex; ovary 1.5-2 mm. long, the style ca. 2.5 mm. long, the stigma peltate, ca. 0.5 mm. across, obscurely 3-radiate. Fruit ellipsoid to spherical or ovoid, 1.1-2.3 by 1.1-2.2 cm., truncate to rounded at apex, drying brownish, sharply 1980] STEVENS, CALOPHYLLUM 473 wrinkled (obscurely striate); outer layer detaching cleanly from stone, 0.5-3 mm. thick, + compact, air spaces sometimes developing, the inner surface striate, dull or shiny; stone subspherical to ellipsoid or ovoid, 1-1.8 by 0.8-1.35 m., rounded to obtusely pointed at apex, the walls 0.1-0.3(-0.65) mm. thick, smooth, unmarked; spongy layer thin. DistriBuTION. Southern Malay Peninsula, Singapore, Borneo (Map 31). SELECTED SPECIMENS SEEN. Malaya. JoHore: Mersing, 30 m., KEP 76288 (a, BO, K, KEP, SING, US), Endau Road, 30 m., KEP 76281 (a, K, KEP, SING), Ulu Endau, near summit of Bukit Kendek, 396 m., KEP 105019 (robust) (A, K, KEP, SING); Kluang, Renggam F.R., 427 m., KEP 98951 (A, K, KEP, SAN, SING); Kluang F.R., Bukit Jelakoi, 210 m., KEP 76296 a KEP, SING); Kota Tinggi, Panti Fr. R., 90 m., KEP 73055 (k, KEP, sING); G. Arong F.R., 45 m., KEP 94883 (KEP). Singapore: Sungei Hantu, opposite S penis m., 39532 (A, E, K, KEP, SING); ‘‘Woodlands,”’ Albero s.n., March 1866 (FI). Borneo. SaRAWAK. Ist Division: Semengoh F.R., Omar 387 (ic, sinc); Bako Natl. Park, along Lintang Path, S 24865 (k, L, SAR); Sempadi F.R., S 61/47 (sar); Matang W. ridge, 305 m., S 1524] (a, Bo, K, L, SAN, SAR, SING); Serian, G. Penrissen, S' 16303 (a, BO, K, L, SAN, SAR, SING); Sabal Tapang, 150 m., Stevens et al. 177 (aA). 3rd Division: Kapit, Yong, Ulu Bediri, 750 m., S 36381 (sar), Ulu Sungei Kapit, Bukit Goran, 750 m., S 36187 (sar), ridge between Sungei Balang and Sg. Balleh, 518 m., S 28260 (A, L, SAR, SING); Usun Apau Plateau, R. Jalan, 965 m., S 3802 (SAR, SING), between R. Biak and Sut, 721 m., S 2938 (sar, SING, us). 4th Division: Miri, Sungei Miri, S 12944 (Bo, xk, L, SAN, SAR, SING); Baram, Ulu Melinau, 180 m., S 1049 coll. Brunig, Aug. 1958 (sar); Bintulu, Nyabau F.R., S 14487 (k, L, SAN, SAR, SING); Bukit Lambir, Map 31. Distribution of Calophyllum biflorum in Malesia. 474 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 ew S 26780 (A, E, K, SAR, SING). 5th Division: Trusan, Bukit Tudal, 975 a 8742 (sar); Lawas, oe Range, 1067 m., . 18704 © SAR). BRUNEI: ee F.R., 15 m., S 7811 (a, BO, K, KEP, L, SAR, SING); R. Ingei, 30 m., BRUN 131 (aA, K, KEP, L, ae Kuala Belait, S$ 1944 (sar); Andalau F.R., BRUN 251 (po, K, KEP, L, SAR). SABAH. Sipitang: W. side of G. Lumaku, 16 km. SSE. of Malaman, 1372 m., SAN 16718 (2) (BRI, KEP, L, SAN, SING). Ranau: Poring turnoff, 640 m., Stevens et al. 558 (a). Labuk & Sugut: Bukit Tangkunan, 180 m., SAN 61260 (san). Tawau: Tawau R. F.R., 50 m., SAN 19449 (), below summit Mt. Lucia, 1067 m., SAN 22691 (Bo, KEP, SAN, SAR); Mt. Wullersdorf, 610 m., SAN 61428 (san); Brassey Range, 770 m., Stevens et al. 477 (a). Kinabatangan: Bukit Tingka, 365 m., SAN 23268 (?) (KEP); Lamag, S. side of G. Lotung Inarat, 1524 m., SAN 83235 (2) (A). KALIMANTAN. Timur: Bulungan, Sg. Sebakis region, Kostermans 9247 (Bo, kK, L, P); Ti doengsche Landen, 16 m., bb 18276 (a, Bo, L); Salimbatoe, Sungei Pingping, 150 m., bb 11178 (Bo); W. Koetai, Mt. Palimasan near Tabang on Belajan R., 800 m., Kostermans 12959 (Bo, L); Keloempang, 50 m., bb 16939 (a, BO, L); Boven Mahakam, Taliba (G. Leng, Bov. Pari), 600 m., bb 26556 (BISH, BO, K, L, NY, P, SING). Tengah: Beneden Dajak, Tewai Baroe, 40 m., bb $170 (v0): Poerokijahoe, ecg ee 25 m., bb — (2), BO). Barat: foet van G. Kenepai, Hallier 1633 (a, Bo, BRI, K, L, SING); Melawi as B. Kemoenting, 225 m., bb 29651 (?) ee L, SING); aie Dawak, 50 m bb 12401 (xo). Ecotocy. Usually in well-drained mixed dipterocarp forest, to 1067(—1372) m. alt. In Borneo frequently in kerangas vegetation, once in peat swamp (S 12944, Sarawak); also in soil derived from ultramafic rock. Flowering May, October, and November; fruiting January to March, May, June, and October (fruit pale green). GERMINATION AND YOUNG PLANT. The radicle almost certainly breaks the stone to one side of the base. The seedling has two pairs of leaves separated by an internode less than | cm. long. Subsequently produced internodes are successively longer, the terminal bud is functional, and the young plant is erect. The leaves of the seedling are 1.1-1.9 cm. wide, and successively produced ones are notably narrower (initially only 0.6-1.3 cm.). Axillary innovations of the young plant may have basal scars. (KEP 76296; Stevens et al. 50, 134A, 255, 406A.) Loca uses. The wood is used to make planks (Kuching). The latex of S 3802 (3rd Division, Sarawak) was noted as being poisonous, but there may have been confusion between this plant and a member of the Anacardia- ceae (the local name of S 3802 is given as “‘betaho’’). Calophyllum biflorum can be recognized by its usually oblong lamina that as a retuse to subacute apex and venation of equal density on both sides; it often dries with the margin a paler color than the rest of the blade. Its flowers have four tepals, and its fruits are broadly wrinkled when dry. The outer layer of the fruit detaches cleanly from the stone, and the inner surface of the outer layer is striate and usually shiny. Indumentum, other than on the terminal bud, the very base of the inflorescence, and the young parts 1980] STEVENS, CALOPHYLLUM 475 ou the plant, is usually inconspicuous. The specific epithet biflorum means “two flowered’’; Henderson and Wyatt- “Smith incorrectly thought that this species had two-flowered inflorescences. There is considerable variation within Calophyllum biflorum in Borneo, even in a single locality. On Gunong Matang (near Kuching, Sarawak) at least two forms of the species grow together (Stevens et al. , a very coriaceous-leaved form, and Stevens et al. 237, with less coriaceous leaves— more like the Malayan form). Some specimens from Borneo have yellow latex: Stevens et al. 134, from near Kuching, had yellow latex, yet is very close vegetatively to the Malayan form; some of the specimens from the 3rd Division of Sarawak are also reported to have yellow latex. Some Bornean specimens may lack the paler leaf margin otherwise common in the species, and brown outer bark seems to be notably more common in Borneo than in Malaya. In Johore trees of C. biflorum often have broad bands or patches of brown, scaly bark from which latex has exuded and dried a more or less pellucid greenish white color; this may be caused by damage to the trunk. I did not observe this phenomenon in trees of C. biflorum from Sarawak and Sabah, although there was black dammar at the base of the trunk of Stevens et al. 128 (Sarawak). Branched inflorescences are not present in Malayan specimens of C. biflorum, although they appear to occur quite commonly in Sarawak and Brunei, and at least sometimes in Sabah (SAN 83235). Most Bornean specimens have leaf blades that are more coriaceous and terminal buds that are longer than those of the Malayan specimens. However, the difference is not absolute; KEP 105019 (Malaya) is very similar to many Bornean specimens. Despite this considerable variation, it would be imprudent to recognize infraspecific taxa in Calophyllum biflorum. The pattern of variation in Borneo is not well understood; not only do distinct forms seem to grow in close proximity (Gunong Matang, Sarawak: see above), but the knowledge of variation in individual characters is imperfect. Thus, in Borneo inflorescence type is known only from a few specimens, and despite the abundance of C. biflorum in Johore, Malaya, flowers have still not been collected there. Although there is considerable variation in hair type (Ficure 29, j-l), the hairs are similar on some Bornean and Malayan specimens. The specimens cited above from the 3rd Division of Sarawak are included in Calophyllum biflorum only with hesitation. Some specimens have large truits and/or leaf blades that are more or less acute at the apex, and the margin of the leaf does not dry notably discolored. The striate inner surface of the outer layer of the fruit is dull rather than shiny. S 26780 (4th Division, Sarawak) also has subspherical fruits, the outer layer of which has a dull inner face; this specimen was taken from a large tree reported to have had "The inflorescence of Calophyllum biflorum was originally described as being few it was called two flowered. However, both the young inflorescences and the infructes- censes of the material that Henderson and Wyatt-Smith had at hand have or had five to seven flowers 476 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 thick, low buttresses—an unusual condition in the species. Calophyllum biflorum is most closely related to C. ferrugineum and its relatives; for the differences separating these taxa, see Taste 13. Sterile specimens of C. biflorum from Borneo, especially from the Semengoh Forest Reserve, near Kuching, Sarawak, can be confused with C. teysmannii var. inophylloide. The latter taxon has a more or less obovate (rather than oblong) lamina frequently with a clearly thickened margin; although the margin of C. biflorum is often discolored, the veins are not obscured by thickening as they are in C. teysmannii var. inophylloide. Calophyllum teysmannii var. inophylloide apparently always has yellow latex, and the trunk often has spurs and/or stilt roots; the latex of C. biflorum is often white, and spurs and stilt roots are very uncommon. SAWN 83235 (Sabah) at least superficially approaches C. canum: its leaf blades are more or less pointed at the apex and its latex is reported to be ‘‘white yellow.” 88. Calophyllum hosei Ridley, Kew Bull. 1938: 120. 1938; Masamune, Enum. Phanerog. Born. 475. 1942; J. Anderson, Trees Peat Swamp Sarawak, 86. pl. 27B. 1973. Type: Sarawak [4th Division], Baram District, Jan. 1895, Hose 146 (holotype, kK; isotypes, BM, L, SING). Calophyllum fragrans Ridley, Kew Bull. 1938: 120. ene Hee excepto; asam Enum. Phanerog. Born. 475. 1942. Type: Sar , 2 miles [3 km.] from Kuching, 10 Dec. 1894, Haviland : Hose 3355, oe K; isotypes, A, BM, BO, L, P, SAR, UC, W). Tree 7.5—18(-36) meters tall, d.b.h. to 45 cm.; trunk without buttresses, small knee roots with prominent, corky lenticels reported (Anderson, Joc. cit.); outer bark grayish to brown and yellow, often mottled, with lines of lenticels, often hoop marked, the inner surface dark brown; under bark orange-brown; inner bark red; latex yellow (white), opaque to clear, sticky (watery). Twigs flattened, 1.5-2.5 mm. across, rounded (4-angled), drying blackish brown, glabrous even when young; axillary innovations with pair of basal scars and another pair ca. 1.5 mm. from base; internodes 1-4 cm. long; uppermost pair of axillary buds rounded, ca. | mm. long, spreading; terminal bud plump, |.8-4 mm. long, with short, adpressed, grayish brown indumentum (hairs, Figure 25, p, q; some an hia aeie underdeveloped internode 1-4 mm. long. Petiole (0.5—)0.8-1.3 . long, broadly concave above, convex below, glabrous; lamina oblong to "elliptic, (3-)4-9.5 by (1.7-)2.2-5 cm., retuse (acute-subacuminate) at apex, acute [cuneate] at base, not undulate or shallowly and distantly so and slightly recurved at margin, coriaceous to very coriaceous, drying umber to sabelline-olivaceous above and umber to fulvous-olivaceous below, glabrous, the midrib above gradually narrowed . Inflorescences from foliate axils near ends of twigs, with 7 to 13 1980] STEVENS, CALOPHYLLUM 477 flowers, unbranched, the axis 4-8 cm. long, glabrous, lowest internode to 0.3(-2.2) cm. long, ultimate internode often much shorter than penultimate, or terminal flowers 5 together; bracts not seen, probably quite large; pedicels 1-3 cm. lon labrous, to 2 mm. across at apex in fruit. Flower (?)her- maphroditic; iene 4, the outer pair suborbicular, 4.5-6(-7) by 4.5-6 mm., deeply concave, thick, the inner pair obovate to broadly elliptic, (5.5-)8-11 by (5-)6.5-9 mm., thinner, less concave or not; stamens 90 to 240 [to 370 to 420], the filaments to 5 mm. long, connate for up to ca. 1.5 mm., the anthers oblong, 0.9-1.5 mm. long, shallowly [to deeply] retuse at apex; ovary 1.3-1.7 mm. long, the style 3-3.7 mm. long, the stigma peltate, 0.7-0.9 [-1.8] mm. across, 2- or 3-radiate. Fruit ovoid to ellipsoid, ca. 1.9 by 1.2 cm., apiculate, drying vinaceous-brown, smooth to shallowly wrinkled; outer layer detaching cleanly from stone, I-1.2 mm. thick, compact; stone ellipsoid, ca. 1.6 by 1 cm., + rounded at apex, the walls ca. 0.2 mm. thick, smooth, unmarked; spongy layer thin. DistripuTION, Southeastern Sumatra, Borneo (not in the northeast) (Map SELECTED SPECIMENS SEEN. Sumatra. Djampi: Sampit, 45 m., bb 13128 (Bo). Borneo. Sarawak. Ist Division: Bako Natl. Park, T. Pandan Kechil, 0-30 mee Carrick & Enoch 135 (Kivu, sar); Setapok F.R., 30 m., S 8934 (so, SAR, SING); Sirak Mangrove F.R., FA 1126 coll. Browne, 24 Nov. 1951 (sar); G. Pueh F.R., S 6298 (sar). 2nd Division: Botong Distr., Tj. Keranji, 9 m., S 1596 9 (a, K, SAR, SING). 3rd Division: near Sibu, Anderson & Ding Hou 475 (a, Bo, Cc, K, L, SAR, SING). 4th Division: Marudi F.R., 9-15 m., S 9971 ee Brunel: ‘‘Forest Reserve,’’ FA 602 coll. Igan, 29 March 1935 (KEP). SABAH. Beaufort: Lupak area, 3 m., SAN 15861 (KEP, SAN, SAR); Lumak, 24 m., SAN 5091/4 (san); Bukit Siungau, 30 m., SAN 80634 (kK). KALIMANTAN. Timur: Tidoengsche Landen, Sepoetoek, bb 17832 (?) (Bo, L); Pembiliangan, 8m., bb 18141 (a, Bo). Selatan: Amoentai, Tanupekan, Rantau Kodjang, 10 m., bb 7784 (Bo); Bulungan, Sebakis R. region, Kostermans 9304 (Bo, L, SING). Tengah: Sampit, Lei Pemoelian, bb 2085 (Bo, L); Moera Tewe, Moera Montalat, bb 9997 (po). Barat: Soengei Landak, Teysmann, AB 11381 (Bo, Fi, K, L Ecotocy. In freshwater peat swamps, less commonly in kerangas forest; 3-30 m. alt. Flowering November to January and July to September; fruiting in February. Calophyllum hosei can readily be distinguished from other species of the genus by its small terminal bud generally borne on a pronounced underdevel- oped internode, by the almost complete absence of hairs except on the terminal bud, by the scars at the base of the axillary innovations, and b the usually short basal internode of the inflorescence coupled with the often much-shortened terminal internode. The fruit is smooth, even when almost mature. The specific epithet commemorates the well-known collector, C. ose. The leaves of the saplings of Calophyllum hosei are ovate, acuminate at the apex, and up to 16 by 7 cm 478 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Calophyllum hosei is perhaps superficially similar to C. ferrugineum and its relatives. However, the characters given above readily separate the two: C. ferrugineum and its relatives have a longer terminal bud, much more obvious indumentum, axillary innovations without basal scars, and wrinkled fruits. In several respects the type specimen of Calophyllum hosei (characters enclosed in brackets in the description) differs from the others. It also has a somewhat more coriaceous lamina that is cuneate at the base and has a very prominent, rounded midrib on the lower surface. It is a good match with S 8943 (in bud), which was collected from kerangas forest. However, since all specimens cited agree in the other characters noted above, and there are intermediates in the other characters, only one taxon is recognized. In the original description of Calophyllum fragrans, Ridley cited Haviland 1812, inaddition to the type, Haviland & Hose 3355. However, Ridley annotated the former specimen at Kew as being the type, as a note by H. K. Airy Shaw on the specimen indicates that it almost certainly should be. Although the description of the leaves and the position of the inflorescences of C. fragrans perhaps agree better with Haviland 1812, and O. Stapf’s earlier identification of the species as ‘“‘C. near retusum,’’ which Ridley mentions, is on this specimen, the length of the inflorescence and the field notes were apparently taken from Haviland & Hose 3355. Hence, Ridley seems to have based his description on both specimens, and his typification as given in the Kew Bulletin is followed. Haviland 1812 is here included in C. andersonii. 89. Calophyllum cordato-oblongum Thwaites, Enum. Pl. Zeyl. 407. 1864; Bedd. Fl. Sylvat. 3: xxi. 1871; T. Anderson in Hooker f. Fl. Brit. India 1: 275. 1874; Vesque, Epharmosis 2: tr. 13, 14. 1889; Trimen, Handb. Fl. Ceylon 1: 103. 1893; Vesque in C. DC. Monogr. Phanerog. 8: 562. 1893; Alston in Trimen, Handb. FI. Ceylon (Suppl.) 6: 21. 1931. Tyre: Ceylon [Sri Lanka], Hinidoon Patoo, Sept. 1863, CP 3823 (isotypes, BM, G, GH, K, L, MEL, NY, P, W). Tree ca. 20 meters tall, d.b.h. to 40 cm.; trunk apparently without buttresses; outer bark brown to dark red-brown, (shallowly) fissured, the strips 2-3 cm. wide; inner bark red. Twigs flattened, 2.2-4.5 mm. across, 4-angled, with raised lines decurrent from middle of petiole, drying dark brown, somewhat yellowish when older, sparsely brown-tomentose when young; axillary innovations apparently lacking basal scars; internodes 2—9(-13) cm. long; uppermost pair of axillary buds rounded to pointed, ca. 1.5(-4.5) mm. long, + erect; terminal bud plump, 0.9-1.3 cm. long, with brown, tomentose indumentum (hairs, Ficure 25, k, t), underdeveloped internode to 1 mm. long. Petiole 3-7 mm. long, concave above, convex below, subpersistently puberulent below or not; lamina oblong to elliptic-oblong, 8-17(-25.5) by 4-7(-11.5) cm., rounded at apex, cordate at base, usually rather distantly undulate and slightly recurved at margin, coriaceous, drying sepia above and cinnamon-sepia below, subpersistently dark brown-puberulent on midrib below, the midrib above abruptly narrowed 1980] STEVENS, CALOPHYLLUM 479 at base, depressed for up to half length of leaf, becoming raised, 0.2-0.4 mm. wide at midpoint, below raised, angled, striate toward base, the venation above subobscure to subapparent, below + apparent, raised, (3 or) 4 to 9 veins /5 mm., angle of divergence 60-70°. Inflorescences terminal and from adjacent foliate axils, with 7 to 15 flowers, often flabellate, with 3-flowered branches to 2.5 cm. long, the axis (2-)6-10 cm. long, subpersistently sparsely to densely tomentose, lowest internode (0.5—)1-4 cm. long; bracts unknown: pedicels 0.5-1.3 cm. long, persistently tomentose. Flower (?)hermaphroditic; tepals (6 to) 8, the outer pair orbicular to broadly ovate, 5-7 by 7-9 mm deeply concave, tomentose on back, the inner ones obovate to narrowly elliptic, 1.2-1.5 by 0.5-1.1 cm., outer two sometimes with few hairs toward base on back; stamens ca. 235, the filaments to 7.5 mm. long, the anthers oblong, 0.8-1.5 mm. long, rounded at apex; ovary 2.5-3 mm. long, the style 7-8.5 mm. long, the stigma peltate, ca. 1.5 mm. across, + 3-radiate. Fruit subspherical, 1.7-2.2 by 1.5-2 cm., rounded at apex, drying brown, smooth: outer layer not detaching cleanly from stone, 1.5-2 mm. thick, compact, with few fibers; stone subspherical, to 1.9 cm. long and across, rounded at apex, the walls less than 0.1 mm. thick, smooth, (?)unmarked; spongy layer thin. DistripuTIon. Southwestern Sri Lanka (Map 8). SELECTED SPECIMENS SEEN. Sri Lanka: Kanneliya forest near Hiniduma, Kos- termans 24771 (a, £, us); Galle Distr., Beriliya forest near Elpitiya, Kos- ermans s.n., Aug. 1974 (k). Eco.ocy. Apparently rather uncommon, in rain forest, low elev. Flowering in September; fruiting in May. Calophyllum cordato-oblongum is a distinctive species that can readily be distinguished from the other taxa with cordate lamina bases by its terminal inflorescences with rather persistent, tomentose indumentum: its usually eight-tepaled flowers; and its fruits, which have stones with walls less than 0.1 mm. thick. The epithet refers to the leaf shape: more or less oblong, and cordate at the base. Although Anderson (/oc. cit.) described the flowers as having eight petals (i.e., presumably a total of twelve tepals), neither Vesque (1893, loc. cit.) nor I have found flowers with more than eight tepals. The surface of the fruit appears to be coarsely furfuraceous, but this is probably caused by damage. The relationships of Calophyllum cordato-oblongum are unclear; its hairs are reminiscent of those of C. bracteatum, but the species are otherwise quite dissimilar. 90. Calophyllum venulosum Zoll. Syst. Verzeich. 2: 149, 150. 1854. Type: Java, ex montosis Seribu, Zollinger 993, pro parte (holotype, P). Tree 4-45 meters tall, d.b.h. to 80 cm.; trunk without buttresses (shortly spurred); outer bark grayish to pale brown, yellowish when young (dark 480 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61] brown when older), usually cracked and flaking, newly exposed bark darker in color (deeply fissured) (rarely hoop marked), the inner surface dull brown to yellowish (mottled darker); under bark dull straw to reddish, mottled or not: inner bark dark red or pale brown to pink; latex clear (becoming opaque) yellow (brown-yellow or orange-yellow), sticky; sapwood cream; heartwo od reddish. Twigs flattened, 1.5-4.5 mm. across, 6-angled (very strongly flattened and 2-angled) (with transverse raised lines at nodes), drying yellowish or whitish, transiently brown-subfarinose to subpersistently tomentose, hairs to 0.8 mm. long; axillary innovations lacking basal scars; internodes (0.5—)1-10 cm. long; uppermost pair of axillary buds rounded to pointed, to 3.5 mm. long, spreading; terminal bud plump, 3-13 mm. long, with adpressed grayish to spreading brown-tomentose indumentum (hairs, Ficure 29, o-q, s—u), underdeveloped internode to 3 mm. long. Petiole 1-6 (apparently to 25) mm. long, concave above, convex below, sometimes subpersistently tomentose below; lamina elliptic, ovate, or suboblong to obovate, 3.7-23.5 by 1.2-11.5 cm., acute to rounded (subacuminate) at apex, cordate to auriculate or minutely rounded (narrowly decurrent) at base, deeply and rather distantly undulate but not recurved at margin, the whole lamina + concave, thinly coriaceous to coriaceous, drying bay or honey to brown above and bay or umber to ochraceous below, usually subpersistently farinose to tomentose on midrib below (also above), the midrib above + abruptly narrowed at or near base, at first level or slightly depressed, becoming raised, 0.1-0.4(-0.6) mm. wide at midpoint, below raised, + angled, the venation above subobscure to apparent, below apparent, raised, sometimes clearly branched, 4 to 10 veins /5 mm., angle of divergence 60-80°. Inflorescences from foliate axils (rarely terminal), sometimes two together, with (1 to) 5 (to 11) flowers, very rarely flabellate, unbranched, the axis to 0.5-3.5 cm. long, puberulent to short- tomentose toward base, lowest internode to 0.3-1.7 cm. long; bracts ovate to elliptic, 2.5-5.5 mm. long, caducous; pedicels 0.6-3 cm. long, glabrous, slender, to 4.5 cm. by 2.5(-5) mm. in fruit. Flower (hecmapheodit tepals 4 (very rarely 8), the outer pair ovate, 4-7 by 3.7-5 mm., the inner pair obovate, 7-13 by 4.5-10 mm.; stamens 85 to 205, the filaments to - mm. long, connate for up to 0.7 mm., the anthers oblong, I-1.6(-2.1) mm. long, rounded to subretuse at apex; ovary |.5-3 mm. long, the style 3-5 mm long, the stigma peltate, 0.7-1.3 mm. across, + 3-lobed. Fruit usually subspherical, |.2-2 by 1.1-2 cm. (rarely ellipsoid, ca. 2.6 by 2.1 cm.), minutely apiculate, drying dull brown, smooth; outer layer detaching cleanly from stone, 1-4 mm. thick, compact; stone often subobovoid, 0.95—1.25(-1.5) by 0.9-1.1 cm., rounded at apex, the walls 0.1-0.4 mm. thick, smooth, usually with triradiate or biradiate marking at apex; spongy layer thin. Key to the Varieties of Calophyllum venulosum — . Petiole 1-6 mm. long, clearly distinct from lamina; terminal bud 3-13 Lit I ee Bk eee ee ee oe ee Oa. var. venulosum. . Petiole apparently (0.8-)1-2.5 cm. long, not clearly distinct from lamina; terminal bud 3-5 mm. long... ........... 90b. var. fenuivenium. — 1980] STEVENS, CALOPHYLLUM 481 90a. Calophyllum venulosum Miq. var. venulosum C. venulosum Mig.; Walp. Ann. Syst. Bot. 4: 366. 1857; Miq. Fl. Indiae Batavae 1(2): 511. 1858; Choisy, Pl. Javan. 9. 1858; Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 279. 1862; Vesque, Epharmosis 2: tt. 23, 24. 1889, in C. DC. Monogr. Phanerog. 8: 575. 1893; Koord. & Valeton, Meded. s’Lands Plant. 61(Bijd. Booms. Java 9): 379, 1903: Koord.-Schum. Syst. Verzeich. 1(Fam. 187): 6. 1912; Koord. Exkurs. Java 2: 617. 1912; Merr. Bibl. Enum. Bornean Pl. 394. 1921; Heyne, Nutt. Pl. Indonesié. ed. 3. 1: 1086. 1950; Backer & Bakh. f. Fl. Java 1: 385. 1963. C. javanicum Migq. Pl. Jungh. 3: 292. 1854; Walp. Ann. Bot. Syst. 4: 367. 1857; Mig. Fl. Indiae Batavae 1(2): 510. 1858; Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 294. 1862; Keith, N. Borneo Forest Rec. ed. 2. 2: 315. 1952 (““C. sp. ex aff. javanicum’’); Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 318. 1956, pro parte; Kochummen, Malayan Forest Rec. ed. 2. 17: 214. 1965, pro parte; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 7: 15. 1967; T. C. Whitmore, Tree Fl. Malaya 2: 187. 1973, pro parte. Tyre: Java, m. Gunong-Saribu, van Gesker s.n. (holotype, vu). C. auriculatum Merr. Philip. Jour. Sci. C. 4: 291. 1909, Enum. Philip. Fl. Pl. 3: 78. 1923. Tyre: Philippine Islands, Mindanao, Zamboanga, Sax River, 500 feet [150 m.], 20 Feb. 1905, Williams 2339 (isotypes, 4, k, ny (several), us). C. ee Boerl. & Koord. in Koord.-Schum. Syst. Verzeich 2: 1911. Type: Sumatra [Tapanoeli], Tapos, 20 Feb. 1891, Koorders 10331 (holotype, os C. Ua eagle auct., non Wall. ex Choisy; Baker f. Jour. Bot. London oo 8. 1924. C. griffithii auct., non. T. Anderson; Baker f. ibid. Tree 4-45 meters tall, d.b.h. to 80 cm. Terminal bud 3-13 mm. long. Petiole 1-6 mm. long; lamina elliptic, ovate, or suboblong to subobovate, 3.7-23.5 by 1.2-11.5 cm., usually cordate or auriculate (rarely acute) at base. DistripuTion. Malay Peninsula (somewhat doubtful) and Sumatra to the Philippines (Map 32 SELECTED SPECIMENS SEEN (letters refer to discussion in text). Malaya. Perak: Kroh F.R., 30 m., KEP 71909 (ker). PaHanc: Kemasul F.R., KEP 78679 (KEP). Jouore: Bukit Hantu F.R., 60 m., KEP 79170 (kep); Pontian, KEP 70251 (KEP, SING). Sumatra. SELATAN: Tandjong- Ning, R. Moesoe-oeloe, 1524— 829 m., Forbes 2756 (a) (BM, L, Mo, SING); Mengkoelem [Mengkulem], 457 m., Forbes 3069 (a) (A, BM, FI, GH); Rawas, Grashoff 1122 (e) (Bo, 4 Moesi Veloe, Endert 27 (leaf base rounded to acute, ?e) (Bo); ond. afd. Redjang, bij Loeboek Bindjai marga Sindang Blibi, 150 m., bb 3022 (a) (A, Bo, L), Loeboek Blingbing, 600 m., bb 7903 (a) (Bo). Barat: Pajakombo, Deloe Air, 1000 m., Sjamsoeddin 26 (Bo); Sidjoendjoeng, Moearo, 200 m., bb 2974 (e) (A, BO, 1); Padang Lawas, 390 m., bb 6633 (Bo). Utara: Angkola en Sipirok, Panobasan, 500 m., bb 26114 (Bo, L, Mo, stNc); Nandsiling c. a., Djoeloe, 1000 m., bb 6180 (Bo); Sibolangit, cult. (seed from Sibajak at 1200 m.), 500 m., Lorzing 11684 (po); Langkat, Halaban Keden, 20 m., bb 8486 (c) (Bo). 482 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 61 Borneo, Sarawak. Ist Division: Mattang, Beccari PB 1988 (c) (Fi, kK); Kuching, Semengoh F.R., S 26260 (c) (A, kK, L, SAN, SAR, SING); Sabal Tapang, 120 m., Stevens, sight record. 3rd Division: R. Biak (R. Luar), 586 m., S 2974 (Cc) (L, SAR, SING, US). 4th Division: Mt. Murud, upper Baram, Moulton 99 ©) (SING). SABAH. Sipitang: Ulu Mendalong, ca. 10 km. SSE. of Malaman, 533 m., SAN 16763 (c) (Bo, BRI, KEP, L, SING). Beaufort: ee 8 km. WNW. of Tenom, 270 m., SAN 15112 (c) (a, BO, BRI, K, KEP, L, SING). Kota Belud: Mt. Templer F.R., slope of Mt. Madalon, 610 m., SAN 76223 (?f) (A, SAN, SAR, SING); Tuburan, 305 m., SAN 3337 (c) (Bo, Kk). Tenom: Rayoh F.R., SAN 41445 (c) (Kk, KEP, L, SAN, SAR). Tambunan: Trusmadi F.R., 1524 m., SAN 31444 (L, san). Lahad Datu: Ulu Sungei Segama, 500 m., Stevens, sight record. Tawau: Kalabakan, 10 acre plot, Luasong, 15 m., SAN 59783 (c) (sAN); Brassey Range, 600 m., Stevens et al. 462 (c) (a). KaLaacipan. Timur: Salimbatoe, be Pingping, 150 m., bb 11171 (wo); Kabiran, S. Bengaloen, 100 m., bb 11707 (Bo); W. Koetai, no. 36, near L. Petah, 500 m., Endert a (€) (A, Bo, kK), no. 24, L. Iboet, 150 m., Endert 4787 (e) (a, Bo, K, L); Maranga on Tundjung Plateau, 200 m., Kostermans 12250 (f) (Bo, pos K, L, NY, P, sING); C. Kutei, Belajan R., G. Kelopok near Tabang, Kostermans 10578 (f) (Bo, L); peak of Balikpapan, Beoul, 600 m., oe 7290 (f) (Bo); E. Kutei, Sg. Menubar region, 5 m., Kostermans 5034 (b) (A, BO, K, L, SING); Sangkulirang Distr., Sg. Mandu region, 150 m., Kostermans 13312 (b) (Bo, kK, L). Selatan: path from Djaro Dam to Mt. Serempaka, ca. 20 km. NE. of Muara Uja, 680 m., de Vogel 1071 (f) (L). Barat: Melawi Oeloe, Soengei Semangka, 450 m., bb 29056 (Bo, K, L, NY, SING); Soeka Lanting, Hallier 160 (2b) (so). Philippine Islands. Minpanao. Lanao: Kalambugan, FB — (uc); Lake Lanao, Camp Keithley, Clemens 1019 (d) (r, G). Zamboanga: Sax 150 m., Williams 2339 (d) (a, Kk, Ny, Us). BAsiLAN: Isabela de Basilan, a 884 (d) ce BISH, E, MO, NY). Java. Barat: Batavia, Pasir Tjihideung, . v. Leuwiliang en Buitenzorg, m., Dakkus 180 (between a and b) (Bo, L); Bantam, G. Pangisisan (G. arang), Tjamara, 10-200 m., Koorders 2884 (a) (Bo); Tjioendar, bij Pondok Sean (G. Karang), 1050 m., Koorders 5712 (a) (Bo); Baya [Bajah], Hasskarl s.n. (a) (L, p); cult. in Hort. Bogor. sub numero VI C 135 (between a and b) (no, k, L, P, Us) Ecotocy. Usually in well-drained mixed dipterocarp forest, (5— )200-610 (-1524) m. alt. Sometimes in marshes (Kostermans 5034; 5 m, alt.), quite frequently along streams. Flowering January, February, May, July, and September; fruiting January to May, and August to November (fruit yellowish to brownish, acid in taste (Kostermans 10581)). There are spherical galls ca. 1 mm. across on and near the midrib on both surfaces of the leaf on Dakkus 180; these are caused by midges (Docters van Leeuwen-Reijnvaan & Docters van Leeuwen, 1926). Raised pustules occur near the margin and the midrib on the lower surface of the leaf in Koorders 10331. GERMINATION AND YOUNG PLANT. The seed germinates by breaking the stone to one side of the base (SAN 16763). A single seedling seen had a pair of reduced leaves and a pair of expanded leaves (de Vogel, 1980, pl. 15). Initially the young plant grows slowly, with internodes less than 3 cm. long and only one pair of leaves being produced per flush; after the plant reaches 1980] STEVENS, CALOPHYLLUM 483 ca. 40 cm. in height, growth is more rapid. The plant is erect, and the terminal bud is functional. Locat use. In Java (Bantam) the wood is used for oars (Heyne, op. cit.). 90b. Calophyllum venulosum Zoll. var. tenuivenium (M. R. Henderson & Wyatt-Smith) P. F. Stevens, comb. et stat. nov. C. tenuivenium M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 311. pl. 7. 1956; T. C. Whitmore, Tree Fl. Malaya 2: 193. 1973; Corner, Gard. Bull. Singapore Suppl. 1: 105. 1978. Type: Malaya, Johore, 8th mile Kota Tinggi-Mawai road, 18 Feb. 1937, SFN 32274 coll. Corner (holotype, sING; isotypes, A, BO, K, KEP, LAE, NY, P, SING). Tree ca. 20 meters tall, d.b.h. ca. 30 cm. Terminal bud 3-5 mm. long. Petiole apparently (0.8-)1-2.5 cm. long; lamina elliptic, (2.6-)6-10 by (0.8-)2-4.5 cm., attenuate at base. DistripuTION. Known only from southern Malaya (Johore). SELECTED SPECIMENS SEEN. Malaya. Jonore: Panti F.R., 5 m., Stevens et al. 115 (a); G. Sumalayang, 305 m., Chin 602 (KLU). Ecotocy. Seasonally inundated or colline forest, 5-305 m. alt. Flowering in February; flower scented. GERMINATION AND YOUNG PLANT. Growth of the young plant is initially slow, with the internodes of plants 20 cm. tall being less than I(-2) cm. long; growth is erect, and the terminal bud is functional. The stems very soon become yellowish. (Stevens et al. 116.) The epithet ftenuivenium emphasizes the fine, but distinct and distant, venation on the type specimen. Calophyllum venulosum is usually readily recognizable, despite the variation in its vegetative characters. The twigs dry yellowish white and usually have six raised lines. The leaves often vary considerably in size on a single specimen (Ficure 30, g); the petiole is short; and the blade is usually cordate to minutely auriculate at the base, often has rather distant venation, and frequently dries bicolored—a dark or grayish brown above and often a somewhat orange-brown below. The inflorescence usually has fewer than seven flowers, and the flowers generally have four tepals. The fruit is characteristic—smooth when dry and with a rather thick outer layer that detaches cleanly from the smooth stone; the stone is nearly always triradiately marked. The epithet venulosum refers to the venation, which is often notably apparent and prominent in this species. Rank of Calophyllum tenuivenium Calophyllum tenuivenium and C. venulosum are similar in all characters except for the apparently long-petiolate leaves of the former: Henderson & Wyatt-Smith (Joc. cit.) mentioned the petiole of C. tenuivenium as usually JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 1980] STEVENS, CALOPHYLLUM 485 being 1-2 cm. long. When dry the petiole is transversely rugulose for ca. the basal 4 mm. only; the remainder dries longitudinally striate. In C. venulosum the short petiole dries transversely rugulose, while the midrib on the lower surface of the lamina dries more or less striate. This suggests that the lamina of C. tenuivenium is in fact very narrowly attenuate, with the petiole proper being ca. the basal 4 mm. Very young plants of C. tenuivenium (Stevens et al. 116, collected under an adult tree) have leaves with a short (2.5 mm.) petiole; the lamina is gradually narrowed toward the base and is shortly and abruptly rounded at the very base (Ficure 30, f). The tree under which these young plants were growing had the ‘‘long petioles’? normal for C. tenuivenium, and it is presumed that in successively produced leaves the lamina becomes more and more strongly narrowed toward the base and is eventually not developed there. There is a tendency for the base of the lamina to be cuneate or acute in other specimens of Calophyllum venulosum, although leaves with blades that are minutely auriculate at the base also occur on these specimens. This tendency is shown by some specimens collected by Forbes from Sumatra (Ficure 30, a, b), by Grashoff 1122 and Endert 27 (also from Sumatra), by specimens that are otherwise a good match with the type specimen of C. ijzermannii (see below), and by specimens collected by Endert (e.g., Endert 3451) from West Koetei, Kalimantan. However, in none of these specimens is the petiole more than 7 mm. long, and the petiole and the lamina are always quite clearly separated. Thus, it seems best to reduce Calophyllum tenuivenium to varietal rank under C. venulosum. Not only are the leaves of the young plants of C. tenuivenium similar to those of C. venulosum, but the slow growth of the young plant and the rather distinctive appearance of the bark of the mature tree of C. tenuivenium are both similar to those of C. venulosum seen in Sabah and Sarawak. Young plants of Calophyllum venulosum var. tenuivenium have leaves with blades that are acuminate at the apex and up to 15.6 by 5.4 cm Variation within Calophyllum venulosum var. venulosum The type specimens of Calophyllum javanicum and of C. venulosum are STE and are like a number of specimens from Java and Sumatra (denoted *» in the list of specimens examined). The lamina is small to medium sized (less than 14 by 4 cm.), and thin and acute to more or less rounded Ficure 30. Variation in leaf; extent of variation in some collections shown by smaller outlines inside larger ones. a-g, Calophyllum venulosum d, e, g, var. venulosum: a, b, Forbes 3189; d, S 26277; e, van Gesker SAW. g, SAN 15112 (leaves from same shoot). c, f, var. ftenuivenium: c, SFN 32274, arrows mark extent of narrowly attenuate lamina; f, Stevens et al. 116 (from very young plant). h-o, C. pentapetalum. h, i, k-n, var. cumingii: h, i, BS 26928; k, BS 43260: 1, BS 44802; m, n, BS 30206. j, 0, var. pentapetalum: j, Santos 6178; 0, Cuming 1212. All x 0.67. 486 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 at the apex. The petiole is noticeable (3 mm. or more long), and the terminal somewhat ill-defined group of specimens from Java and southern Kalimantan (‘‘b’”’ in the list) have leaf blades that tend to dry greenish and subnitid. The apex of the lamina is acuminate, and the short petiole is obscured by the cordate base. In Kalimantan such specimens have been collected from trees 4-28 meters tall. The form that is most common in Borneo, and which also occurs in Sumatra (‘‘c’’ in the list), has well-developed, tomentose indumentum on the vegetative parts; the terminal bud is 6-11 mm. long; the twigs are stout (2-4.5 mm. across); the lamina tends to be large, although it is notably variable in size ((3.3-)6-23.5 by (2-)3-11.5 cm.); and the base of the lamina is usually clearly cordate or auriculate. Such specimens have been collected from trees 27—42 meters in height. Specimens from the Philippine Islands (‘‘d’’ in the list) are similar to those in group c above. However, the indumentum is not well developed except on the terminal bud, where it may be tomentose (e.g., Ebala 884) or short and adpressed (e.g., Williams 2339). The inflorescence is sometimes rather congested. Calophyllum auriculatum was described from a specimen of this type. Specimens have been collected from trees 5—18 meters tall. Some specimens from Sumatra and southern Kalimantan (‘‘e’’ in the list) have moderate-sized to large leaves and terminal buds. The lamina is subcoria- ij In both Sumatra and Kalimantan there are apparently independent tendencies for the base of the lamina to become more or less cuneate or acute. Specimens in group e have been taken from trees 12-30 meters tall. The final group of specimens (‘‘f’’ in the list), known only from Kalimantan, also have a subcoriaceous, rather densely veined lamina, but it does not dry dark brown and nitid. The terminal bud is ca. 5 mm. long, and the plant does not have very well-developed indumentum. One specimen (Koster- mans 10581) has mostly ellipsoid fruits up to 2.6 by 2.1 cm. The specimens from Malaya cited above are included here with hesitation. All are sterile. The specimens from Johore have stout twigs, large leaves, and rather short indumentum on the terminal bud, and are perhaps intermediate between Calophyllum venulosum var. venulosum group d and C. subsessile (q.v.). The specimens from Perak and Pahang have a rather small (4.6-10.6 by 1.6-3.6 cm.) lamina, and their indumentum is well developed; their latex is reported to be yellow or milky yellow. At first sight the latter group of specimens is very distinctive, but more collections of C. venulosum and its relatives from the entire Malay Peninsula are needed to assess the significance of the variation discussed above, as well as the status of C. subsessile, Specimens taken from a tree once cultivated at the Botanic Gardens at 1980] STEVENS, CALOPHYLLUM 487 Bogor under the number V/ C 1/33 are atypical in several respects (see also Calophyllum subsessile). The inflorescence has up to eleven flowers and is sometimes branched, the flowers have eight tepals, the fruits dry rather deeply and sharply wrinkled, and the stone lacks triradiate markings. 91. Calophyllum grandiflorum J. J. Sm. Bull. Jard. Bot. Buitenzorg, III. 1: 396. tt. 45, 46. 1920; Backer & Bakh. f. Fl. Java 1: 385. 1965. Type: Java, Soekaboemi, G. Tjimerang, ca. 700 m., Jan. 1914, ten Oever s.n. (lectotype, Bo; isolectotype, L). Tree 15-28 meters tall, d.b.h. to 60 cm.; trunk without buttresses or spurs; outer bark yellowish brown and smooth at first, becoming mid-brown, with vertical lines of lenticellary welts, scaling, scales small, the inner surface yellow-straw; under bark reddish; inner bark dark red; latex opaque yellow, slightly sticky. Twigs flattened, 3-6 mm. across, + 4-angled when young and with prominent raised lines decurrent from each petiole, 2-angled when older (with obscure transverse raised lines at nodes), drying brown (yellowish), glabrous or almost so; axillary innovations lacking basal scars; internodes 4-10 cm. long; upper- most pair of axillary buds acute, up to 7 mm. long, spreading; terminal bud narrowly conical, 1.2—2.2 cm. long, with grayish or brownish, subcrustose indumentum (hairs, Figure 25, r), underdeveloped internode to 2 mm. long. Petiole 2-5 mm. long, concave above, angled below, glabrous or puberulent: lamina oblong to elliptic or lingulate, 15-36 by 6-10 cm., acute at apex, cordate at base, undulate and slightly recurved at margin, coriaceous, drying umber to sabelline above and cinnamon to sabelline below, glabrous or with sparse hairs on midrib below when young, the midrib above narrowing quickly near base, + depressed in bottom 3-S(-7) cm., becoming sharply raised, 0.5-0.8 mm. wide at midpoint, below prominent, angled, the venation above and below subapparent to subobscure, raised, 4 to 9 veins/5 mm., angle of divergence 70-85°. Inflorescences from axils of topmost or adjacent leaves, with 3 to 11 flowers, often flabellate (with 3-flowered branches to 3 cm. long), the axis 3-12 cm. long, + glabrous, lowest internode (1—-)3—7.5 cm. long; bracts reported to be foliaceous, up to 9.25 by 5.2 cm.; pedicels 3-7.5 cm. long, glabrous, in fruit to 3 mm. thick. Flower (?)hermaphroditic; tepals 8, sometimes glabrous, the outer pair reflexed at anthesis, + orbicular, ca. 11.5 mm. long and across, the inner ones obovate to elliptic, 1.7-2 by 1-1.4 cm.; stamens 320 to 370, the filaments to 8 mm. long, the anthers oblong, 1,5-2.5 mm. long, rounded to retuse at apex; ovary ca. 3.5 mm. long, the style 9-12 mm. long, the stigma peltate, 1.5-2.5 mm. across, 3-radiate. Fruit + spherical to ellipsoid, ca. 2 by 1.5 cm., apiculate, with stipe ca. 2.5 by 4 mm., scars of tepals and androecium prominent, drying yellow-brown, finally dark brown, strongly and sharply wrinkled, furfuraceous; outer layer detaching cleanly from stone, ca. 2 mm. thick, compact; stone spherical to ellipsoid, ca. 1.6 by 1.3 cm., rounded at apex, the walls ca. 0.3 mm. thick, smooth, with 2 to 4 pale longitudinal stripes from apex; spongy layer thin. 488 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 DistRiBUTION. Western Java (Map 32). SELECTED SPECIMENS SEEN. Java. Barat: Lengkong, estate ee 0 m., Kostermans 23822 (a, AAU, BO, CANB, G, K, , Bodjong Lopang, Backer 16990 (Bo, L); Bantam, G. Tompok, bij pea 300- 400 m., Backer 7328 (Bo). Ecotocy. Locally common in colline forest, 300-700 m. alt. Flowerin February, September, October, and November (flower scented); fruiting in January (fruit yellowish brown when ripe) GERMINATION AND YOUNG PLANT. The radicle breaks the stone wall i diatel to one side of the base of the stone. The seedling has a single pair of leaves. Subsequent growth is slow, with the internodes less than 3 cm. long until the plant reaches 20-30 cm. in height; after that the internodes produced are up to 20 cm. long. The terminal bud is functional. (Stevens et al. 723.) Calophyllum grandiflorum is one of the most easily recognized species of Calophyllum: its large leaf blade is cordate at the base and has relatively distant venation, its terminal bud is long and thin, and its inflorescence has - “eo | eet e 2 SP ST nl AIO 2 Bi a Map 32. Distribution of Calophyllum venulosum var. venulosum (squares), C. subsessile (triangles), C. grandiflorum (half-circles), C. mukunense (solid stars), . aa (open star), and Calophyllum sp. 95 (star in solid circle) in Males 1980] STEVENS, CALOPHYLLUM 489 a long basal internode and large flowers (up to 5 cm. across when fully opened—hence the appropriate specific epithet). The stipitate, strongly wrin- kled fruit is also striking. Calophyllum grandiflorum is related to C. venulosum: the morphology and bark of the two species are similar, and both have young plants that grow only slowly at first. However, the characteristics mentioned above readily distinguish the two species. The specimen fen Oever s.n. (collected in January, 1914) at Bogor herbarium is made the lectotype of Calophyllum grandiflorum. The locality data for this collection are those given on the label; they are slightly different from those given by J. J. Smith (/oc. cit.). The other specimen that Smith cited, Backer 7328, is sterile. 92. Calophyllum subsessile King in Ridley, Ann. Bot. Gard. Calcutta 5: 142. pl. 171. 1896; Ridley, Fl. Malay Penin. 1: 187. 1922; I. H. Burkill & M. R. Henderson, Gard. Bull. Straits Settl. 3: 347. 1925. Synrypes: Malaya, Perak, Larut, less than 100 feet [30 m.], Nov. 1884, King’s collector [Kunstler] 6877 (gm, k, L, P), less than 100 feet [30 m.], Feb. 1885, King’s collector |Kunstler| 7311 (x). C. javanicum auct., non Miq.; M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 318. 1956, pro parte; Kochummen, Malayan Forest Rec. ed. 2. 17: 214. 1965, pro parte; T. C. Whitmore, Tree Fl. Malaya 29 187.. 1993 pro patie. Tree to 24 meters tall, d.b.h. to 90 cm.; trunk without buttresses, but knee roots reported; outer bark golden, yellow, green, and brown [mottled], with long, irregular fissures, roughly scaly; under surface yellow; inner bark pale pink, clearly laminated; latex clear golden, stick Twigs slightly flattened, 2-3.5 mm. across, with 6 prominently raised lines, drying + shiny, whitish yellow, initially sparsely brown-puberulent; axillary innovations apparently lacking basal scars; earniee 1.5-4 cm. long; upper- most pair of axillary buds rounded, ca. m. long, inconspicuous; terminal bud plump, 4-6 mm. long, with grayish one farinose-puberulent indumentum (hairs, Ficure 29, n), underdeveloped internode absent. Petiole 2-3 mm. long, shallowly concave above, + angled below, glabrous; lamina elliptic to oblong, 6.8-14 by 3.7-6 cm., rounded at apex, cordate at base, slightly undulate and not recurved or slightly so at margin, coriaceous, drying umber above and below, glabrous, the midrib above rather quickly narrowed near base, depressed in bottom quarter, becoming raised, 0.2-0.5 mm. wide at midpoint, below raised, angled, the venation above subobscure, below + apparent, raised, 5 to 8 veins/5 mm., angle of divergence 65-75°. Inflorescences terminal and axillary, with 5 to 13 flowers (flabellate and/or with 3-flowered branches to 1.7 cm. long), the axis 5.5-7 cm. or more long, slightly puberulent toward base or not, lowest internode 2-4 cm. long; bracts subelliptic, ca. 5 mm long, not persistent; pedicels to 1.3 cm. long, glabrous. Flower bud only seen, (?)hermaphroditic; tepals 8, the outer pair suborbicular, to 5 by 4.5 mm.; stamens 155 to 165, the anthers oblong, 1.2-1.7 mm. long, rounded 490 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 to subretuse at apex; ovary ca. | mm. long, the style ca. 1.5 mm. long, the stigma peltate, ca. 0.7 mm. across, 3-lobed. Fruit spherical, ca. 2.5 cm. long and across, rounded at apex, drying brown, with broad, rounded wrinkles; outer layer detaching cleanly from stone, ca. 2 mm. thick, compact; stone broadly obovoid, ca. 2.1 by 1.9 cm., rounded at apex, the walls ca. 0.4 mm. thick, smooth, with triradiate marking at apex; (?)spongy layer thin. DistripuTion. The Malay Peninsula, known only from Perak (Map 32). ADDITIONAL SPECIMEN SEEN. Malaya. Perak: Gelong Raya, 4Ist mile Bruas— Lumut road, KEP 43570 (KeEp). Eco.tocy. Lowland freshwater swamps or seasonally inundated forests. Flowering in November; fruiting in February. Calophyllum subsessile is closely related to C. venulosum, and like that species, it has leaf blades that are more or less rounded at the base and fruits with a thick, compact outer layer that detaches cleanly from the stone; the stone has a triradiate marking at the apex. It can be distinguished from . venulosum by its whitish-drying terminal bud, its more coriaceous leaf blades, its longer inflorescences that are sometimes terminal and branched and that have more numerous flowers, its flowers, which have eight tepals, and its larger fruits (spherical fruits of C. venulosum are less than 2 cm. long). The almost sessile leaves presumably suggested the epithet subsessile. Calophyllum subsessile is provisionally maintained as a distinct species, since the only specimens of C. venulosum that have flowers with eight tepals and inflorescences approaching those of C. subsessile in size are cultivated specimens from Bogor (VJ C 133), which in other characters are unlike C. subsessile. The field notes of KEP 43570 report knee roots; although these have not been observed in C. venulosum, infraspecific variation in the presence or absence of knee roots is quite common in Calophyllum. The distribution of C. subsessile in Malaya (Map 32) is not unexpected for a species with eastern affinities (Keng, 1970). Saplings of Calophyllum subsessile have leaf blades up to 24.5 by 6.5 cm.; the midrib on the upper surface of the blade is raised for its entire length. The fruits of Calophyllum subsessile were reported by King (loc. cit.) to be minutely pubescent; they are in fact glabrous. 93. Calophyllum mukunense P. F. Stevens, sp. nov. A speciebus aliis Calophylli quibus laminis basibus cordatis et fructibus putaminibus signatis habent in inflorescentiis terminalibus et axillaribus (huis cum internodiis basalibus usque ad 2 mm. longis), floribus cum 8 tepalis, et fructu strato exteriore tenui circa 0.5 mm. crasso, differt Tree 18-26 meters tall, d.b.h. to 50 cm.; outer bark yellowish, exfoliating irregularly, the scales elongated, thin, | mm. thick; inner bark pink; latex yellow (Kostermans ). Twigs strongly flattened, 3.2-5 mm. across, rounded to obscurely 4-angled, 1980] STEVENS, CALOPHYLLUM 49] drying yellowish, shiny, glabrous at maturity; axillary innovations apparently lacking basal scars; internodes 1-4.5 cm. long; uppermost pair of axillary buds rounded, to | mm. long, inconspicuous, sometimes glabrous; terminal bud broadly conical, 3.5-4.5 mm. long, with brown, crustaceous to subap- pressed indumentum (hairs, Figure 29, m; also moruloid), underdeveloped internode to 2.5 mm. long. Petiole 4-8 mm. long, broadly and shallowly concave above and convex below, glabrous; lamina elliptic to suboblong, 6.3-23.3 by 2.6-7.3 cm., acute to subacuminate (rarely rounded) at apex, cordate to rounded at base, strongly and distantly undulate and slightly recurved at margin, coriaceous, drying shiny, ochraceous to umber above and umber below, glabrous, the midrib above gradually narrowed from base, depressed, (0.25-)0.5-1.1 mm. wide at midpoint, below raised, becoming flat to slightly raised, angled (striate or rounded toward base), the venation usually subobscure above and subapparent below, raised to flat, 4 to 6 veins/5 mm., angle of divergence 65-80°. Infructescences terminal and from adjacent foliate axils, with scars of 11 to 17 flowers, unbranched (rarely with 3-flowered branches to 4 cm. long), the axis 3.5-7.5 cm. long, stout, puberulent at base, lowest internode in axillary inflorescences to 2 mm. long; bracts not known; pedicels 1.3-3 cm. long, glabrous, slender, in fruit to 5.5 cm. by 4 mm. Flower not known. Fruit ellipsoid to spherical, 1.8—2 by 1.4-1.8 cm., rounded at apex, drying grayish brown, strongly wrinkled; outer layer detaching cleanly from stone, ca. 0.5 mm. thick, compact, inner surface shiny; stone subspherical, 1.3-1.7 by 1.2-1.4 cm., rounded at apex, the walls ca. 0.2 mm. thick, smooth, with 3 longitudinal stripes from apex; spongy layer thin. Tyre: Borneo [Kalimantan], Sg. [Sungei] Mukun near Sangasanga (Sa- marinda), 5 m., 3 Aug. 1952, Kostermans 7725 (holotype, a; isotypes, B, BO, K, L, LAE, P, SING). DisTRiBUTION. Sumatra and southwestern Borneo (Map 32). ADDITIONAL SPECIMENS SEEN. Sumatra. Riau: Selatpandjang, Kampar Monding, 5 m., bb 22058 (Bo). Diamsi: Maera Pidjoean, 89 m., bb 12831 (Bo); Simpang, (Bo); Beneden Djak, Troesas, 1 m., bb 9872 (Bo); Sampit, Sei Kereng Bindjai, Selongau, 3 m., bb 7939 (Bo). Barat: Koeboepadi, 5 m., bb 6364 (Bo). Eco.Locy. At least periodically flooded forest, 1-90 m. alt. Fruiting March (once), and August (once; submature). Calophyllum mukunense can be recognized by its shiny, at most slightly angled twigs, its rather large leaf blades that are rounded to cordate at the base, its terminal and axillary inflorescences with at least eleven flowers, and its fruits that dry wrinkled and that have an outer layer about 0.5 mm. thick and a triradiately marked stone. The axillary inflorescences have a very short basal internode; however, there do not seem to be scars at the bases of the axillary innovations, although this must be confirmed. The epithet mukunense comes from the name of the river near which the type specimen was collected. 492 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 The closest relatives of Calophyllum mukunense are probably C. venulosum, C. subsessile, and C. grandiflorum, all three of which have similarly shaped leaf blades and stones with triradiate markings. However, the other characters mentioned above readily differentiate C. mukunense from these species. Although the pedicels of C. venulosum var. venulosum are sometimes notably incrassate in fruit (e.g., Endert 27, from Sumatra), such specimens agree with C. venulosum in all other respects. 94. Calophyllum calcicola P. F. Stevens, sp. nov. FIGuRE 28, 1. A speciebus aliis Calophylli quibus laminis basibus cordatis habent in innovatione axillari saepe cicatricibus basalibus ornata, lamina mediocra coriacea vel percoriacea nitida, inflorescentiis axillaribus, floribus cum 8 tepalis, et fructu putamine haud signato, differt. Shrub or tree to 10 meters tall, d.b.h. to 20 cm.; outer bark yellowish, rough, cracked. Twigs slightly flattened, 1.5-2 mm. across, with 6 elevated lines, drying brown to yellow when young, later whitish, glabrous to sparsely brown- puberulent when young; axillary innovations often with basal scars; internodes 1-5 cm. long; uppermost pair of axillary buds rounded, ca. 0.4 mm. long, + spreading, inconspicuous; terminal bud plump, 2-3 mm. long, with brown, crustaceous indumentum (hairs, Figure 25, s), underdeveloped internode absent. Petiole 2-4 mm. long, concave above and convex below, glabrous, drying black; lamina obovate to elliptic-oblong, 1.2—8.5 by 1-3.8 cm., rounded to retuse (rarely apiculate) at apex, auriculate to rounded at base, undulate but not recurved at margin, coriaceous to very coriaceous, drying shiny, bay above and umber below, glabrous, the midrib above abruptly narrowed near base, not obvious, flat to slightly raised, ca. 0.15 mm. wide at midpoint, below raised, striate, the venation subobscure above and apparent below, raised, 4 to 6 veins/5 mm., angle of divergence 40-70°. Inflorescences from foliate axils, with 5 flowers, unbranched, the axis (0.2—)2.5-3.5 cm. long, glabrous or sparsely puberulent at base, lowest internode 0.2-1.5 cm. long; bracts ovate, ca. 2 mm. long, deciduous; pedicels 2—2.7 cm. long, glabrous. Flower (?)hermaphroditic; tepals 8 (rarely 7), sometimes glabrous, the outer pair ovate, ca. 5.5 by 4.5 mm., the inner ones ovate to obovate, 6-8 by 4-5.5 mm.; stamens 60 to 75, the filaments to 3.7 mm. long, the anthers oblong, 1.2-1.6 mm. long, + retuse at apex; ovary 1.5-2 mm. long, the style ca. 2.5 mm. long, the stigma peltate, ca. 0.6 mm. across, (?)lobed. Submature fruit ellipsoid, ca. 1.5 by 1-1.4 cm., + rounded at apex, drying brown, smooth; outer layer detaching cleanly from stone, ca. | mm. thick, compact; stone subspherical, ca. 1.3 by 1.2 cm., almost round at apex, the walls ca. 0.8 mm. thick, smooth, unmarked; spongy layer initially thick. Tyre: Borneo [Kalimantan], Berouw, Mt. Has Bungaan, 700 m., 12 Dec. 1957, Kostermans 13837 (holotype, L; isotypes, BO, CANB, K, LAE, P, SING). DistripuTION. Southeastern Borneo (Map 32). 1980] STEVENS, CALOPHYLLUM 493 ADDITIONAL SPECIMENS SEEN. Borneo. KALIMANTAN. Timur: Berouw, Mt. Ilas Bungaan, 700 m., Kostermans 13834 (Bo, K, L, SING); top of Mt. Ilas Mapulu, 800 m., Kostermans 14022 (Bo, K, KEP, L, SING). EcoLocy. On limestone, 700-800 m. alt.; sometimes shrub pendulous on limestone walls (Kostermans 13834). Flowering in September Calophyllum calcicola can be recognized by its short terminal bud; small, very coriaceous, nitid leaf blades that are rounded to auriculate at the base; axillary, five-flowered inflorescences; flowers with eight tepals; and spherical fruits with an unmarked stone. The epithet calcicola means “‘limestone dwell- er,’ a reference to the ecological proclivities of this species. Calophyllum calcicola is perhaps related to the widespread and variable . venulosum. However, it differs in its axillary innovations that often have scars at the Base (such scars are absent in C. venulosum), its coriaceous and nitid leaf blade, its small terminal bud, its flowers that always have seven or eight tepals (in C. venulosum I know of only one cultivated specimen with eight tepals), and its fruit with a thinner outer layer and an unmarked stone. In C. venulosum the outer layer is 1-4 mm. thick, and the stone has longitudinal markings. 95. Calophyllum sp. Tree 8 meters tall, d.b.h. 10 cm.; trunk and bark unknown. Twigs flattened, 2.5-3 mm. across, with 4 + raised lines, drying yellowish, tomentose when young; axillary innovations apparently lacking basal scars; internodes (0.3-)0.7-6 cm. long; uppermost pair of axillary buds rounded, ca. 1 mm. long, erect; terminal bud conical, 7.5-8.5 mm. long, brown-tomen- tose, underdeveloped internode absent. Petiole 2-3.5 mm. long, concave above, convex below, tomentose; lamina elliptic to subobovate or suboblong, 10-20 by 3.8-7.1 cm., cuneate at apex, broadly rounded to shallowly cordate at base, obscurely undulate and slightly recurved at margin, drying near sepia on both surfaces, with grayish covering above, + persistently tomentose over entire lower surface, the midrib above quickly narrowed at base, raised, center strongly sulcate at first, 0.35-0.4 mm. wide at midpoint, below raised, striate, the venation above and below subobscure, raised, 8 to 13 veins/5 mm., angle of divergence 75-80°. Infructescences from foliate axils, with scars of 3 to 5 flowers, unbranched, the axis 2.5—4 mm. long, glabrous when mature, lowest internode I-2.5 mm. long; bracts unknown; pedicels 8-10 by ca. 2.5 mm., glabrous. Flower unknown. Fruit probably spherical, ca. 2 cm. long and across, minutely apiculate, drying vinaceous-brown, smooth; outer layer detaching + cleanly from stone, |.5-2.5 mm. thick, compact, small air spaces under skin; stone subspherical, ca. 1.4 by 1.2 cm., rounded at apex, the walls 0.3-0.4 mm. thick, smooth, perhaps with triradiate marking at apex; spongy layer thin. DistrisuTion. The Philippine Islands (Map 32); known only from a single collection. 494 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 SPECIMEN SEEN. Philippine Islands. MinpAnao. Agusan: Butuan, San Matea Bo., Tungao Bo., along Ojot R., Maasin Line km. 25, 350 m., PNH 41930 (L, PNH, SING). EcoLocy. Colline forest, 350 m. alt. Fruiting in June (fruit green). Loca use. The wood is used in construction. Calophyllum sp. 95 can be recognized by its bluntly conical, tomentose terminal bud; its rather large, sepia-drying leaf blades that are rounded to shallowly cordate at the base and that have subobscure venation; and its spherical fruits with a thick outer layer and a rather thin-walled stone. eer tomentose indumentum persists on the lower surface of the lamina. The duplicate of PNH 41930 at Leiden appears to have furfuraceous fruits, but this is apparently the result of damage. 96. Calophyllum pentapetalum (Blanco) Merr. Sp. Blancoanae, 266. 1918; Tovomita pentapetala Blanco, Fl. Filip. 432. 1837. Type: Philippine Islands, Luzon, Lepanto subprovince, Species Blancoanae 184 coll. la Pena (neotype, a; isoneotypes, BO, F, GH, K, L, MO, NSW, NY, P). Shrub or tree to 7.5 meters tall; trunk and bark unknown. Twigs not flattened to slightly so, |-2.3 mm. across, usually strongly 4-angled when young, drying brown to blackish, initially tomentose; axillary innovations lacking basal scars; internodes 0.5-2(-4) cm. long; uppermost pair of axillary buds rounded, to 0.7 mm. long, spreading; terminal bud plump, I|-4 mm. long, with brown, + tomentose to subadpressed indumentum (hairs, Ficures 29, r, v-x; 31, b, e), underdeveloped internode not apparent. Petiole 0.5—6 mm. long, concave above, convex below, subpersistently tomentose; lamina ovate to oblong or obovate, I-9.5(-15) by (0.5—)0.9-4.5 cm., acute to rounded at apex, cordate to cuneate at base, neither undulate nor recurved at margin, coriaceous, drying sepia to hazel above and sepia to near sabelline below, + deciduously and sparsely puberulent to tomentose on and near midrib below, the midrib above + quickly narrowed at base, flat to slightly raised (surrounding lamina also raised), 0.2-0.4 mm. wide at midpoint, below slightly raised, subangled to striate, the venation above and below subobscure to subapparent, + raised, 8 to 13 veins/5 mm., angle of divergence (20-)40-75°. Inflorescences terminal and/or from adjacent foliate axils, with 7 to 21 flowers, Ficure 31. Hairs (from terminal bud). a, Calophyllum clemensorum (Clemens c, Elmer 13217. d, e, C. ceriferum: d, Robinson 1478; e, Poilane 6207. f, h-k, C. pisiferum. f, SFN 34747 (atypical specimen). h, 1, k, Pierre 3648: h, i, apex and base of hair ca. 510 pm. long; j, oe. 10292. g, Calophyllum sp. 98 (SAN 21074). 1, 1, C. dispar: 1, Achmad 1751; 1, SFN 37715, base of hair ca. 480 wm. long (uncommon type). m ,n,q, C. tetrapterum var. tetrapterum: m, FRI 14014 (atypical specimen); n, Kerr 9175; q, van Rossum 34. 0, p, C. rupicola: o, — termans 22040; p, FRI 12526. s, C. lineare (Martin 1783). Selle = = 120 496 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 sometimes flabellate and/or with branches to 4 cm. long and 9 flowers, the axis 3.5-12 cm. long, + tomentose toward base, lowest internode 1-3 cm. long; ae ovate to elliptic, to 5 mm. long, deciduous (foliaceous, to 3.2 cm. long, subpersistent); pedicels 0.7-2.1 cm. long, glabrous or with sparse hairs. Flower (?)hermaphroditic; tepals 6 to 12 (to 17), the outer pair suborbicular, 2.5—5 by 2.2-4.5(-6) mm., the next pair broadly elliptic to obovate, (4-)6-8 by (2-)4.5-6 mm., the inner ones elliptic to obovate, (4-)6-11 by (2-)3-6 mm., sometimes glabrous; stamens 40 to 145 (to 180), the filaments to 6 mm. long, the anthers suboblong, 0.65—-1.8 mm. long, + retuse at apex; ovary 1.4-2 mm. long, the style 3-4.8 mm. long, the stigma peltate, 0.7-1.7 mm. across, + 3-lobed. Fruit subspherical to ovoid, 8.5-18 by 7.5-10 mm., + rounded at apex, drying purplish brown to gray, pruinose, smooth, sharply wrinkled when young; outer layer not detaching cleanly from stone, 0.2-0.4 mm. thick, air spaces developing, especially under skin; stone subspherical to ellipsoid, 6-12 by 5-7 mm., rounded at apex, the walls 0.1-0.3 mm. thick, smooth, unmarked; spongy layer thin. Key to the Varieties of Calophyllum pentapetalum — . Lamina cordate at base; petiole 0.5—2(-4) mm. long... ........2.. ese) OF face Brack Be: Ok We a ae ee Se ee a 96a. var. ea . Lamina rounded to cuneate at base; petiole at least 2 mm. Zi Tepals 12 to 17; stigma at least | mm. across; lamina shovate to trape- —_ ee ee ee ee ee ee 96c. var. pulgarense. a Tepals (6 to) 8 (to 12); stigma less than | mm. across; eae variable n shape, but rarely obovate or trapeziform. . 96b. var. cumingii. 96a. Calophyllum pentapetalum (Blanco) Merr. var. pentapetalum C. pentapetalum (Blanco) Merr.; Merr. Enum. Philip. Fl. Pl. 3: 80. 1923. Ochrocarpus pentapetalus (Blanco) Fernand.-Vill. Novis. App. 17. 1880. C. amplexicaule Choisy ex Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 281. 1862; Vidal, ics Cuming. Philip. 96. 1885, Rev. Pl. Vasc. Filip. 54. 1886; Vesque, Epharmosis 2: r. 16. 1889, in C. DC. Monogr. Phanerog. 8: 564. 1893; Merr. Philip. Jo < C. 5: 199. 1910. Tyee: Philippine Islands, Luzon, Albay oo Ilocos Norte], Cuming 1212 (holotype, G; isotypes, BM, FI, K, MEL). Shrub or small tree to 5 meters tall, d.b.h. to 15 cm. Petiole 0.5-2 mm. long; lamina ovate or elliptic to oblong, 1.1-5.8(—8) by 0.9-3.3 cm., + cordate at base, angle of divergence of venation 60-75°. Flower with 6 to 12 tepals, outer pair 3.2-5 by 2.9-4.2 mm.; stigma 0.6—0.9 mm. across. DistripuTiIon. Philippine Islands: Palawan and western Luzon (Map 33). SELECTED SPECIMENS SEEN. Philippine Islands. PaLawan: km. 112, Pinagbatuan, PNH 91186 (L, pNH). Luzon. Ilocos Norte: sine loco, Cuming 1841 (k). Ilocos Sur: Candon, Clemens 18675 (Bo, c, sinc, uc, w). La Union: San pn Species Blancoanae 969 (A, BM, BO, F, GH, L, MO, NSW, NY, P, US, W). Beng 1980] STEVENS, CALOPHYLLUM 497 Map 33. Distribution of Calophyllum ceriferum (open stars), C. rupicola (squares), C. rupicola variant (solid stars), C. andersonii (circles), C. clemen- sorum (stars in solid circles), C. pentapetalum var. pulgarense (large triangle), C. pentapetalum var. pentapetalum (inverted small triangles), and C. pentapetalum var. cumingii (erect small triangles) in Southeast Asia—Malesia. sine loco, Loher 67 (k). Pangasinan: sine loco, FB 28413 (a). Zambales: Santa Cruz, Barrio Dinabag, Santos 6176 (L, us Eco.ocy. Dry forests, sometimes with pine; also in secondary forest and in rocky places in littoral forest; low alt. Flowering December, February, and March; fruiting November, December, and March. 96b. Saree pentapetalum (Blanco) Merr. var. cumingii (Planchon & Triana) P. F. Stevens, comb. et stat. nov. C. cumingii Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 259. 1862; Vidal, Phanerog. Cuming. Philip. 96. 1885, Rev. Vasc. Pl. Filip. 54. 1886; Merr. Enum. Philip. Fl. Pl. 3: 78. 1923. Type: Philippine Islands, Luzon, Albay [or gue Cuming 1077 (holotype, G; isotypes, BM, E, FI, G, K, L, MEL, NY, P CG: pseudotacamahaca Blanches & Triana, Ann. Sci. Nat. Bot. IV. 15: 270. 1862; Vidal, Phanerog. Cuming. Philip. 96. 1885, Rev. Pl. Vasc. 498 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Filip. 54. 1886; Vesque, Epharmosis 2: rt. 16-18. 1889, in C. DC. Monogr. Phanerog. 8: 565. 1893. Type: Philippine Islands, Luzon, Albay [or Zambales?], Cuming 1047 (holotype, G; isotypes, BM, E, FI, G, K, L, MEL, MO, NY, P, W). C. buxifolium Vesque, Epharmosis 2: tt. 18, 19. 1889, in C. DC. Monogr. Phanerog. 8: 538. 1893; Merr. Enum. Philip. = 3: 78. 1923. Type: Philippine Islands, anno 1853, Llanos s.n. (holo G). C. tacamahaca auct., non Willd.; Choisy, es Gultif, Inde, 43. 1849, Mem. Soc. Phys. Hist. Nat. Geneve 12: 423. 1851, pro parte. Tree 1-7.5 meters tall, d.b.h. to 10 cm. Petiole 2-6 mm. long; lamina elliptic to obovate, (1—)2-9.5(-15) by (0.5-)1-4.5 cm., cuneate to rounded at base, angle of divergence of venation (20-)40-70°. Flower with 8 or 9 tepals, outer pair 3.5-5.5 by 3-3.5 mm.; stigma 0.4—0.8 mm. across. DistripuTion. Philippine Islands (Map 33). SELECTED SPECIMENS SEEN (for explanation of letter, see discussion). Philippine Islands. CALAMIAN. Culion: sine loco, Merrill 609 (GH, K, NY, US). MINDORO: Golo, Lubang Is., PNH 36738 (cf. a) (K, BISH, L, PNH, SING). Luzon. Hocos Norte: Burgos, BS 26728 (a, k, us); Bangui, BS 27419 (a, Bo). Ilocos Sur: Barrio Lugong, Santa Maria and vicinity, Clemens 17096 (uc). La Union: San Fernandes, 4 m., Sete 174 (uc). Pangasinan: So. Cabalatinawan, Sual, 450 m., 7 30206 (£, uc). Zambales: Mt. Canaynayan, Castillejos, BS 26538 (A, BM, F, L, MO, P, uc, us); Mt. Marayep, 396 m., BS 44802 (a, Ny, we); Subig, Merrill 1768 (us); Sitio Apulul, Barrio Amungan, Iba, Santos 6142 (L, us); Botolan, Merrill 2592 (sinc, us); Anuling, BS 44602 (B, c, Ny, uc); Mt. Tapaloa, 917 m., BS 44718 (1, ny, uc). Quezon: Quinayangan, Vidal 2152 (a) (hi K,.. 1); Guminan, ye — be sING). Guimaras Is.: sine loco, FB 229 (cf. a) (BM, BO, F, K, SING, US). Panay: Iloilo, FB 25433 (A, BO, BISH). Micba io: cna sine cies. FB 25284 (a, P, US). EcoLocy. Habitats similar to those of var. pentapetalum, to 450 (rarely to 917) m. alt. Flowering November to March, and May; fruiting December to March, May, and August (ripe fruit black (BS 44602)). The epithet commemorates H. Cuming, the noted collector of shells, plants, and other items of natural history. 96c. Calophyllum pentapetalum (Blanco) Merr. var. pulgarense (Elmer) P. F. Stevens, comb. et stat. nov. C. pulgarense Elmer, Leafl. Philip. Bot. 5: 1786. 1913 (“‘C. pulgarensis’’), Merr. Enum. Philip. Fl. Pl. 3: 80. 1923. Type: Philippine weet Palawan, Puerto Princesa (Mt. Pulgar), 4250 feet [1295 m.], 1911, Elmer 13217 (neotype, A; isoneotypes, BISH, BM, BO, E, F, FI, - GH, K, L, LY, MO, NSW, NY, P, U, US, W). Tree ca. 3 meters tall. Petiole 2.5-5 mm. long; lamina obovate to trapeziform, 2.1-5.5 by 1.3-3 cm., cuneate at base, angle of divergence of venation 40-55°. Flower with 12 to 17 tepals, outer pair to 5 by 6 mm.; stigma 1.4—1.7 mm. across. 1980] STEVENS, CALOPHYLLUM 499 DistripuTIon. Philippine Islands, known only from Palawan (Map 33). SELECTED SPECIMEN SEEN. Philippine Islands. PaLawan: sine loco, BS 567 (BO, GH, K, SING, NY, US). EcoLocy. Locally abundant in rocky, mossy summit scrub, 1100-1300 m. alt. Flowering in May; fruiting in December. The epithet is taken from the name of the mountain on which the type specimen was collected. Calophyllum pentapetalum can be recognized by its usually rather small and coriaceous leaf blades that often dry sepia-brown and nitid on the upper surface and that have moderately dense venation. The terminal bud is small, with more or less well-developed, tomentose indumentum. The inflorescences are usually terminal, and the flowers are relatively large (often more than 1 cm. across) for the size of the fruit (usually less than 1.5 cm. long). The very thin outer layer of the fruit becomes disorganized by air spaces, and the stone has thin walls. The epithet pentapetalum (‘‘five-petaled’’) was coined because Blanco thought that the specimen he studied had five ‘‘petals.”’ Variation within Calophyllum pentapetalum There is a great deal of variation in Calophyllum pentapetalum; leaf shape, in particular, varies considerably on a single individual (Ficure 30, h-o). Despite the (at first sight) very different appearance of the extreme forms, Vesque already suspected in 1893 that the details of lamina anatomy and indumentum of the three species he recognized were so similar that specific rank would probably not be maintained; however, “‘il faudrait avoir le courage d’ opérer la fusion et de distinguer simplement des varietés.’’ Vesque recognized C. pseudotacamahaca, C. buxifolium (both = C. pentapetalum var. cumingii), and C. amplexicaule (= C. pentapetalum var. pentapetalum). Although most specimens of Calophyllum pentapetalum from low altitudes in the Philippine Islands are easily placed in either var. pentapetalum, which has a lamina that is cordate at the base, or var. cumingii, which has a lamina that is cuneate at the base, there are two specimens that are more or less intermediate: BS 30047 (Luzon, Pangasinan, Mt. San Isidro) and PNH 92450 (Luzon, Laguna, Llavac). A few specimens of C. pentapetalum var. cumingii (‘‘a’’ in the list above) have somewhat denser venation (10 to 13 veins/5 mm.) that diverges from the midrib at only 20-40°; the apex of the leaf is more acute than usual. In leaf shape such specimens approach the type of C. buxifolium. Finally, FB 27842 has long, narrow leaf blades (9-15.5 by 2.5-3.2 cm.), although in other respects it is not unusual. Although specimens referable to Calophyllum buxifolium have very small leaves and at first sight appear to represent yet another variety, C. buxifolium and C. pentapetalum var. cumingii are connected by intermediates. FB 26728, FB 27419, and BS 26838 form a series in leaf size: the first two (from Ilocos Norte Province, Luzon) are placed in C. buxifolium, while FB 26838 at the Arnold Arboretum (Zambales, Luzon) has one shoot somewhat like a comparatively large-leaved C. buxifolium and the other like a fairly small-leaved 500 JOURNAL OF THE ARNOLD ARBORETUM [voL. 6l C. pentapetalum var. cumingii (see also Figure 30, h-o). The type specimen of C. buxifolium, Llanos s.n., has narrower leaves; the lamina was described as 1.6-3.2 by 0.5-1 cm., although on the type specimen at Geneva the largest leaf remaining is only 2.6 by | cm. It is therefore not possible to recognize a taxon based on C. buxifolium. Calophyllum pentapetalum var. pulgarense grows at higher altitudes than do the other varieties (ca. 1300 m. vs. usually less than 450 (rarely to 917) m.) and is known only from Mt. Pulgar, on Palawan Island; not surprising- ly, its leaves are more coriaceous than those of the other two varieties. In these varieties the midrib on the lower surface of the lamina is often demarcated from the adjacent surface by a line that is absent in var. pulgarense. Calophyllum pentapetalum var. pulgarense always has numerous tepals in each flower (vs. usually only eight in the other varieties), and its stigma is very much larger. In all other details, including those of the fruit (in an envelope on the sheet of FB 3873 at Kew), it is similar to the other two varieties. Nomenclature and Synonymy I follow Merrill in his interpretation of Blanco’s name Tovomita pentapetala (Merrill, 1918, Joc. cit.), and the duplicate of Species Blancoanae 184 at the Arnold Arboretum is designated the neotype of the name (Blanco’s description is poor). e type specimen of Calophyllum pseudotacamahaca, Cuming 1047, has especially well-developed, leaflike bracts, but similar bracts occur in C. pentapetalum vars. pentapetalum and cumingii (the sheet of Cuming 1047 at FI is exceptional in that the flowers are single and axillary). Planchon and Triana specifically excluded the fruiting shoot of Cuming 1047 at c from their C. pseudotacamahaca because they thought that the small fruits on this shoot were unlikely to have come from flowers the size of those on the other shoot. However, as Vesque (1893, loc. cit.) suggested, the two are conspecific. (The localities for the specimens collected by Cuming are taken from Merrill, 1915.) Cuming 1077, the type specimen of Calophyllum cumingii, is very similar to the type of C. pseudotacamahaca, although it has smaller flowers. It may be noted that if the taxon here called C. pentapetalum var. cumingii is recognized at the specific level and includes C. pseudotacamahaca in its circumscription, the correct name for this taxon is C. pseudotacamahaca rather than C. cumingii (cf. Merrill, 1923, loc. cit.). Vesque (1893, loc. cit.) reduced C. cumingii to synonymy under C. pseudotacamahaca The specimen of E/mer 13217 at the Arnold Arboretum is designated the neotype of Calophyllum pulgarense; the holotype was destroyed 97. Calophyllum ceriferum Gagnep. ex P. F. Stevens Ficure 28, d-f. C. ceriferum Gagnep. in Humbert, Fl.. Gen. Indo-Chine Suppl. 1: 272. fig. 24, 1-8. 1943; Pham, Cay-Cdé Mién Nam Viét-Nam. ed. 2. 2: 301. fig. 1970. Nomen invalidum. 1980] STEVENS, CALOPHYLLUM 501 A Calophyllo pentapetalo, quo aliter simile est, in indumento minus evoluto, internodio terminali innovationis saepe breviore quam aliis, pedicellis in fructo valde incrassatis usque ad 2.5 mm. in transversis (minus quam | mm. in transversis), et fructibus corrugatis strato exteriore circa | mm. crasso, compacto (minus quam 0.3 mm. crasso, haud compacto) et putamine parietibus circa 0.7 mm. crassis (minus quam 0.3 mm. crassis), differt. Tree 5-12 meters tall; trunk and bark not known. Twigs flattened, 1.2-1.6 mm. thick, 4-angled, drying blackish (with grayish, waxy cover me ae when young; axillary innovations lacking basal scars; interno .4-3.3 cm. long, terminal internode often shorter than others; pene pair of ailany buds rounded, ca. 0.5 mm. long, subspread- ing; terminal bud plump, |.3-2 mm. long, with short-tomentose brown or grayish subadpressed indumentum (hairs, Ficure 31, d, e), underdeveloped internode absent. Petiole 0.25-1.3 cm. long, broadly concave above and convex below, glabrous; lamina obovate (rarely subelliptic or trapeziform), (1.6—)2.3- 5.5 by (0.6-)l-3.5 cm., obtuse to rounded (rarely subacute to retuse) at apex, cuneate to acute at base, neither undulate nor recurved at margin, coriaceous, drying umber above (color obscured by grayish covering) and below, glabrous or transiently puberulent on midrib below, the midrib above gradually narrowed from base, + flat, 0.1-0.25 mm. wide at midpoint, below raised, rounded, narrow, inconspicuous, the venation subapparent on both surfaces, raised, 5 to 8 (to 10) veins /5 mm., angle of divergence 30-55°. Inflorescences terminal and from adjacent foliate axils, with 7 to 15 flowers, flabellate or with 5-flowered branches to 5 cm. long, the axis 3-7 cm. long, puberulent, or with scattered hairs toward base, lowest internode (0.5—)1.5—4 cm. long; bracts not known; pedicels 0.5-3.1 cm. long, glabrous to transiently puberulent, slender, up to 2.5 mm. thick in fruit. Flower (?)hermaphroditic; tepals 8, the outer pair ovate, 5-6 by 3.5-5 mm., the inner ones elliptic to obovate, 5.5-10 by 3.5-6 mm.; stamens 100 to 125, the filaments to 5 mm. long, the anthers oblong, 1-1.5 mm. long, subretuse at apex; ovary 1.7-2 mm long, the style ca. 3 mm. long, the stigma peltate, ca. 1 mm. across, 3-radiate. Submature fruit spherical, ca. 1.7 cm. long and across, acute at apex when young, becoming + rounded, drying pale brown, closely and deeply wrinkled; outer layer not detaching cleanly from stone, ca. 1 mm. thick, compact; stone spherical, ca. 1.5 cm. long and across, rounded at apex, the walls ca. 0.7 mm. thick, smooth, not marked; spongy layer initially thick. Tyre: Annam [Vietnam], Nhatrang, presqu’ile de Nui Hon Heo, 100 m., 3 May 1925, Poilane 6172 (holotype, B; isotypes, 4, K). DistriBuTION. Vietnam (Map 33). ADDITIONAL SPECIMENS SEEN. Vietnam: Nhatrang, presqu’ile de Nui Hon Heo, Poilane 6207 (p), Poilane 6234 (B, BM, K); Nhatrang and vicinity, C. B. Robinson 1478 (Kk, Ny, P); Phanrang, Cana, 200 m., Poilane 9612 (xk, p), 400 m., Poilane 12512 (B, K, P Eco.ocy. Tree sometimes of rather poor form, stony ground, 100-400 m. 502 JOURNAL OF THE ARNOLD ARBORETUM [voL. 6] alt. Flowering April, May, and October (flower scented); submature fruit in October Calophyllum ceriferum can be recognized by its short, plump terminal bud; its small, more or less obovate leaf blades with subdistant, steeply ascending venation; and its more or less obvious grayish, waxy covering on the vegetative parts of the plant. The pedicels are markedly incrassate in fruit, the flowers have eight tepals, and the submature fruit has a compact outer layer about 1 mm. thick and stone walls about 0.7 mm. thick. The epithet ceriferum (‘‘wax bearing’’) refers to the waxy covering of the plant. Calophyllum ceriferum is perhaps close to the variable C. pentapetalum, from the Philippine Islands, but can be distinguished from that species by the characters given in the diagnosis The specimens of Calophyllum ceriferum from Phanrang, as wellas Robinson 1478, from Nhatrang, have small leaves with rather dense venation; the leaves are frequently, but not always, separated by internodes of equal length, and the bud has subadpressed, grayish indumentum. The other specimens have larger leaves with more distant venation, the terminal internode of an innovation is often markedly shorter than the others, and the terminal bud has brown, subtomentose indumentum. However, all specimens have a short, plump, terminal bud, the hairs are similar in structure, there is no absolute difference in leaf size and venation density, the inflorescences and flowers are similar, and the pedicels are long, becoming markedly incrassate in fruit. The grayish (waxy) covering of the leaf alluded to by Gagnepain (Joc. cit.) is not always obvious, but is best developed on Robinson 1478. Formal description of Calophyllum ceriferum is necessary since Gagnepain’s description is invalid, being in French. 98. Calophyllum sp. C. brachyphyllum auct., non Merr.; Meijer, Symp. Ecol. Res. Humid Trop. 347. 1965. Tree 7.5-20 meters tall, d.b.h. to 20 cm.; trunk with rounded buttresses (SAN 51735); outer bark pale yellow to red-brown, thin, shallowly fissured, smooth, or scaly-flaky in oblong scales. Twigs slightly flattened, 1-1.7 mm. across, + strongly 4-angled, drying blackish brown, at least sometimes puberulent when young; axillary innova- tions lacking basal scars; internodes 0.5—4 cm. long; uppermost pair of axillary buds rounded, ca. 0.3 mm. long, erect; terminal bud narrowly conical, 1-2 mm. long, with adpressed, brown indumentum (hairs, Ficure 31, g), under- developed internode absent. Petiole 1.5-5 mm. long, deeply concave above, convex below, glabrescent; lamina suborbicular to elliptic or obovate, 1.1-4.2 by 0.9-2.5 cm., slightly retuse to bluntly pointed at apex, cuneate to shallowly cordate at base, slightly undulate and not recurved at margin, coriaceous, drying umber to fuscous above and bay to umber below, glabrous when mature, the midrib above abruptly narrowed at base, flat to slightly raised, 0.15-0.25 mm. wide at midpoint, below raised, striate, not very conspicuous, 1980] STEVENS, CALOPHYLLUM 503 the venation obscure to apparent above and + apparent below, raised, 7 to 9 veins /5 mm., angle of divergence 40-60°. Inflorescences terminal and/or from adjacent foliate axils, with 5 to 11 flowers, unbranched, the axis 2-3.5 cm. long, sparsely brown-tomentose, especially near base, lowest internode 0.5-1.3 cm. long; bracts foliaceous (?or not), to 7 mm. long, deciduous; pedicels 4-8 mm. long, glabrous. Flower known only in bud, (?)hermaphroditic; tepals 8 (rarely 9), the outer pair suborbicular, ca. 4.5 by 4 mm.; stamens 75 to 105, the anthers oblong, ca. 1.5 mm. long, truncate to retuse at apex; ovary ca. | mm. long, style ca. 1.5 mm. long, stigma ca. 0.5 mm. across. Immature fruit + ellipsoid, ca. 8 by 5 mm.; outer layer probably ca. 0.3 mm. thick; stone walls probably very thin; spongy layer unknown. DistRriBUTION. Northeastern Borneo (Map 35). SPECIMENS SEEN. Borneo. SABAH. Ranau: Bukit Ampuon, 1280 m., SAN 21074 (A, CANB, K, KEP, L, LAE, SAN, SAR). Kinabatangan: Mt. Tavai, Karamuak, 610 m., SAN 51735 (a, san). Lahad Datu: NW. ridge of Mt. Silam, 19 km. WSW. of Lahad Datu, 762 m., SAN 15045 (a, BO, BRI, KEP, L, SING), 853 m., SAN 75168 (SAN). Ecotocy. Hillsides and ridges, in soil derived from ultramafic rock; 610- 1280 m. alt. Flowering March and July. Calophyllum sp. 98 is very similar to small-leaved specimens of C. pentapetalum var. buxifolium, differing mainly in its more narrowly conical terminal buds and in the characteristically dark-drying (umber to fuscous) upper lamina surface. The lowest internode of the axillary innovation is often notably longer than the others. All specimens were collected at moderate elevations in areas of ultramafic rock, and this may in part be responsible for their rather distinctive facies. However, it is best to segregate them as a distinct, but unnamed, taxon close to C. pentapetalum, at least until fruits are known. Calophyllum sp. 98 can be readily distinguished from C. brachyphyllum by its less coriaceous leaves with less dense venation, and by its larger, often terminal inflorescences. The two taxa are not immediately related. Although Calophyllum sp. 98 shows considerable variation in lamina shape, the extremes (suborbicular blade subcordate at the base vs. obovate blade cuneate at the base) are connected by intermediates. 99. Calophyllum clemensorum P. F. Stevens, sp. nov. Ficure 32, c, d. C. rotundifolium auct., non Ridley; Meijer, Symp. Ecol. Res. Humid Trop. 7. 1965. A speciebus aliis Calophylli in lamina basi cordata, venulis lateralibus manifestis, inflorescentia terminali, floribus saepe cum 12 tepalis, et fructu parvo circa 1.1 cm. longo strato exteriore compacto circa 0.2-0.3 mm. crasso e putamine munde secedens, differt. Shrub to tree 3.9-15 meters tall, d.b.h. to 30 cm.; bark not known. 504 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Twigs flattened, 2-2.5 mm. across, usually strongly 4-angled, drying dark brown, sparsely and transiently puberulent; axillary innovations usually with basal scars; internodes |.5-3 cm. long; the upper pair of axillary buds rounded, to 0.7 mm. long, + spreading; terminal bud conical, 2-2.5 mm. long, with grayish indumentum (hairs, Ficure 3l, a), underdeveloped internode to | mm. long. Petiole 1-3 mm. long, narrowly concave above and convex below, glabrescent; lamina orbicular to lingulate, (1.7—)2.7-7.3 by (1.5—)2.4-6.3 cm., obtuse to retuse at apex, cordate at base, slightly undulate and somewhat recurved at margin, coriaceous, drying umber to sepia on both surfaces, glabrous at maturity, the midrib above + quickly narrowed near base, flat to slightly raised, 0.15—0.3 mm. wide at midpoint, below raised, slightly striate, the venation + apparent on both surfaces, raised, 7 to 12 veins/5 mm., angle of divergence 65-75°. Inflorescences terminal (also from adjacent foliate axils), with 9 to 25 flowers (flabellate and/or with branches to 3.5 cm. long and with 7 flowers), the axis 3-4.5 cm. long, glabrous or sparsely puberulent toward base, lowest internode 0.6—1.2(-2) cm. long; bracts oblong to ovate, 2.5-4 mm. long, or foliaceous, to |.5 cm. long, subpersistent; pedicels 0.5—1.5 cm. long, sometimes sparsely puberulent when young. Flower (?)hermaphrodi- tic; tepals (11 or) 12 (or 13) [8 or 9 in SAN 17252], the outer pair broadly ovate, 4-6 by 3.7-5 mm., the inner ones obovate to elliptic, 6.5-12.5 by 2.5-6 mm., sometimes glabrous; stamens 110 to 195, the filaments to 4.5 mm. long, the anthers oblong, (0.9—)1.3-1.8 mm. long, retuse at apex; ovary 1.2-1.7 mm. long, the style ca. 3 mm. long, the stigma subpeltate, ca. 0.7 mm. across. Submature fruit ellipsoid, ca. 1.1 by | cm., rounded at apex, drying vinaceous-brown, sharply wrinkled; outer layer detaching cleanly from stone, 0.2-0.4 mm. thick, compact; stone ellipsoid, ca. 10 by 8 mm., rounded at apex, the walls ca. 0.2 mm. thick, smooth, unmarked; spongy layer (?)thin. Tyree: North Borneo [Sabah], Penibukan, 5000 feet [1524 m.], Il Nov. 1933, J. & M. S. Clemens 50316 (holotype, A; isotypes, G, K, MICH, NY, UC). DistripuTION. Northeastern Borneo (Map 33). ADDITIONAL SPECIMENS SEEN. Borneo. Saban. Ranau: Mt. Kinabalu, Penibukan, head of Dahobong R., 1524 m., Clemens 40705 (a, G, K, Ny, Uc), 1524 m. Clemens 30984 (po), 1219-1524 m., Clemens 31428 (Bo); W. Marai Parai, 1219 m., Clemens 35038 (so, Ny); Pentaturan Basin, 2134 m., Clemens s.n., 27 July 1933 (po). Kinabatangan: G. Tonsuon, 975 m. SAN Li2o2, @) (es, BO, EP); EcoLocy. Lower montane rain forest, 975-2135 m. alt.; near Mt. Kinabalu on sometimes dry ridges. Flowering February, May, July, and October; submature fruit in April. In Clemens 35038 the stem is much swollen in places, apparently being galled, and is up to | cm. across. This swelling may also occur on the midrib of the lamina, which then becomes woody. Calophyllum clemensorum can readily be distinguished from the other Malesian species of Calophyllum with a cordate-based lamina by its strongly 1980] STEVENS, CALOPHYLLUM 505 four-angled twigs, distinct venation, terminal inflorescence, flowers usually with more than eight tepals, and ellipsoid fruits with a thin, compact outer layer that detaches easily from the thin-walled stone. This species is named after the collectors and missionaries J. and M. S. Clemens (for an excel- lent photograph of the couple, see Fl. Males. 1(1): 109. 1950). Although the leaves of Calophyllum clemensorum are superficially like those of C. rotundifolium (see Meijer, loc. cit.; E. D. Merrill also identified Clemens’s specimens as C. rotundifolium), that species has more profusely branched, only slightly angled twigs, leaves with the midrib below more or less depressed and not very distinct, and a terminal inflorescence consisting of only one or two flowers. The flowers of C. rotundifolium are larger than those of C. clemensorum and have very thick outer tepals. Calophyllum clemensorum is superficially similar to C. vergens (C. thwaitesii Planchon & Triana ‘B’), from the mountains of Sri Lanka. However, the latter species has axillary inflorescences, flowers with only eight tepals, a somewhat longer terminal bud, and much larger fruits with a thicker outer ayer. SAN 17252 is the only specimen of Calophyllum clemensorum that has eight (or nine) tepals, and it also has a thicker lamina than the others. It is a poor specimen, however, and further collections from Gunong Tonsuon may show that it belongs to the variable C. teysmannii complex. The above description of the fruit is based on detached fruits on the sheets of Clemens 35038. 100. Calophyllum tetrapterum Mig. Pl. Jungh. 291. 1854. Type: Sumatra, Angkola superior, 1000-3000 pedes [310-925 m.], Junghuhn s.n. (holo- type, U; isotypes, Bo, L). Shrub 1.5 meters tall or tree 3-20 meters tall, d.b.-h. to 40 cm.; trunk lacking buttresses but occasionally with stilt roots; outer bark whitish to yellowish or brown, shallowly fissured or smooth (rarely brown, deeply fissured), usually hoop marked, the inner surface orange(-red) to blackish; under bark reddish or reddish and yellow mottled; inner bark reddish; latex usually clear to opaque yellow, sticky (rarely white to whitish or yellow, fluid, not sticky). Twigs slightly flattened, |.3-2.8(-3.5) mm. across, strongly 4-angled (to + rounded), drying brown to blackish or yellowish, glabrous or + sparsely and transiently brown-pubescent; axillary innovations usually lacking basal scars; internodes 0.5-5 cm. long; uppermost pair of axillary buds rounded, 0.5—1.5 mm. long, spreading; terminal bud plump, |.5-4 mm. long, with short, adpressed, grayish, to short-tomentose, brown indumentum (hairs, Ficure 31, m, n, q; cf. 20, i-l, and 22, k, 1), underdeveloped internode 0.5—2(-3) mm. long. Petiole 0.4-1.4(-2) cm. long, + deeply concave above, convex below, at most transiently puberulent below; lamina elliptic to obovate (oblong), (2.2—)3.5—-14 by (1-)1.6-6.5 cm., acuminate or acute to rounded at apex, cuneate to acute at base, at margin sometimes with band of thickening to 1 mm. wide, or clear submarginal vein, undulate but not recurved or only slightly 506 JOURNAL OF THE ARNOLD ARBORETUM [vo. 61 so, + coriaceous, drying umber to gray-olivaceous above, margin often paler, or color obscured by pruinose covering, and cinnamon to olivaceous below, glabrous or sparsely puberulent to subtomentose on midrib below, the midrib above gradually narrowed from base, usually sharply raised (almost level), 0.15-0.45 mm. wide at midpoint, below raised or slightly raised, somewhat striate, the venation on both surfaces usually apparent, raised, (4 or) 5 to 14 (to 17) veins/5 mm., angle of divergence (50-)65—75°. Inflorescences usu- ally axillary (terminal), with 3 to Il flowers, unbranched, the axis (0.3-)1-4 (-7.5) cm. long, glabrous or short-puberulent to subtomentose toward base, lowest internode 0.4-3.5 cm. long; bracts usually small and fugaceous (folia- ceous, to 2 cm. long, subpersistent); pedicels (0.2-)0.5—-2(-3) cm. long, gla- brous, usually very slender, frequently incrassate and to 2.5 mm. thick in fruit. Flower (?)hermaphroditic; tepals 4 or 8 (rarely 5 to 7, or 10), the outer pair ovate to broadly elliptic, 2.2-5 by 2-4 mm., sometimes papillate or puberulent on back near apex, the inner ones elliptic to lingulate, 3.5-8.5 by 1.5-3.5 mm.; stamens 25 to 105 (to 135), the filaments to 4.5 mm. long, the anthers suboblong, (0.4-)0.7-1.2 mm. long, shallowly retuse at apex; ovary 0.8-1.3 mm. long, the style to 3.5 mm. long, the stigma peltate, ca. 0.4 mm. across, slightly lobed. Fruit ellipsoid to spherical, 6.5-16 by 5-12 m., apiculate or rounded at apex, drying grayish to pale brown, sharply wrinkled when young, at least sometimes smooth when mature; outer layer usually not detaching cleanly from stone, 0.25-0.5(-1.3) mm. thick, with large air spaces developing; stone spherical to ellipsoid, 5.5—11 by 5-10mm., rounded at apex, the walls 0.1—-0.2(-0.35) mm. thick, smooth, unmarked; spongy layer thin. Key to the Varieties of Calophyllum tetrapterum — Lamina with clear submarginal vein, or submarginal band of thickening (0.2-)0.4-1 mm. wide, (4 or) 5 to 7 (to 10) veins/5 mm.; floral bracts SOMetiines SUDPEISISIEN: «644 a6. 6@ eae & oo 8 100b. var. obovale. Lamina lacking distinct submarginal vein, marginal band of thickening less than 0.4 mm. wide, usually more than 7 veins/5 mm.; floral bracts soon deciduou 2. Lamina ih obscure venation on lower surface, 5 to 7 veins/5 mm. Tere Te ee ee eT ee ee c. var. blumutense. 2. Lamina with + clear venation on lower surface, (5 to) 8 or more veins /5 MMs % OG He ee oy a 100a. var. tetrapterum. — 100a. Calophyllum tetrapterum Miq. var. tetrapterum C. tetrapterum Miq.; Walp. Ann. Syst. Bot. 4: 367. 1857; Miq. Fl. Indiae Batavae 1(2): 510. 1857, ibid. Suppl. 3(2): 1861; Planchon & Triana, Ann. Sci. Nat. Bot. [V. 15: 293. 1862; Vesque in C. DC. Monogr. Phanerog. 8: 610. 1893; H. Keng, Gard. Bull. Singapore 28: 245. 1976. C. bancanum Miq. FI. Indiae Batavae, Suppl. 1(3): 499. 1861; Kurz, Nat. Tijdschr. Nederl.-Indié 27: 192. 1864; F. Mueller in Walp. Ann. Syst. Bot. 7: 357. 1868; Scheffer, Nat. Tijdschr. Nederl.-Indie 31: 354. 1870, ibid. 32: 405. 1873. Type: Bangka, Djebus, Teysmann, HB 3214 (holotype, U; lsotypes, BO, K, L). 1980] STEVENS, CALOPHYLLUM 507 C. gracile Miq. Fl. Indiae Batavae, Suppl. 1(3): 498. 1861; F. Mueller 5 var. gracile (Miq.) Boerl. Catal. Horto. Bogor. 2: 82. 1901, excl. spec. cit. Type: Sumatra, prope Paja-Kombo, Teysmann, HB 649 (holotype, U; Isotypes, BO, K (s.n.), L, MEL (s.7.)). Ce floribundum Hooker f. Fl. Brit. India 1: 272. 1874, pro parte; King, Jour. Asiatic Soc. Bengal, II. 59: 175. 1890; Ridley, Fl. Malay Penin. 1: 184. 1922; M. R. Henderson, Gard. Bull. Straits Settl. 4: 224. 1928; M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 332. pi. 16. 1956; Smythies, Common Sarawak Trees, 61. 1965; Kochummen, 2: 180. 1973; Corner, Gard. Bull. Singapore, Suppl. 1: 104. 1878. Type: ne Malacca, | Jan. 1867, Maingay 1660 (Kew dist. 170) (lectotype, C. sa anon King, Jour. Asiatic Soc. Bengal, II. 59: 175. 1890; Vesque in C. DC. Monogr. Phanerog. 8: 550. 1893; Ridley, Fl. Malay Penin. 1: 183. 1922. Syntypes: Malaya, Perak, Larut, less oe 100 feet a. m.], Dec. 1883, King’s collector [Kunstler] 5366 (BM, FI, G, K, P, UC), 800- ae a [240-305 m.], Feb. 1885, King’s collector ane 7243 (BM, FI, G, pene King, Jour. Asiatic Soc. Bengal, Il. 59: 180. 1890; Vesque in C. DC. Monogr. Phanerog. 8: 549. 1893; Ridley, Fl. Malay Penin. 1: 186. 1922. Type: Malaya, Perak, Larut, 300-350 feet [90-105 m.], July 1885, King’s collector [Kunstler] 7763 (isotypes, FI, G, Kk). C. foetidum Ridley, Jour. Straits Branch Roy. Asiatic Soc. 54: 18. 1910, pro maxime parte, Fl. Malay Penin. 1: 186. 1922, pro parte. Type: Sines Gardens Jungle, anno 1904, Ridley 11958 (lectotype, sING; isolectotypes, BM, K). C. lanceola Ridley, Jour. Straits Branch Roy. Asiatic Soc. 82: 170. 1920, Fl. Malay Penin. 1: 182. 1922. Type: Malaya, Kedah, Kedah Peak, 4000 feet [1219 m.], Ridley 5751 (holotype, sinc). C. rupicola Ridley var. elatum T. C. Whitmore, Gard. Bull. Singapore 26: 270. 1970, pro minore parte, Tree Fl. Malaya 2: 169. 1973, pro minore parte. Tyre: Malaya, Kelantan, Ulu Sat F.R., 180 m., 2 Feb. 1970, FRI 2538 coll. Kochummen (holotype, KEP; isotypes, kK, L, SING). C. pulcherrimum auct., non Wall. ex Choisy; T. Anderson in Hooker f. Fl. Brit. India 1: 271. 1874, pro parte; Pierre, Fl. Forest. Cochinch. 1: pl. 104A. 1885, pro parte; Vesque, Epharmosis 2: ¢. 21. 1889, in C. DC. Monogr. Phanerog. 8: 570. 1893, pro parte; Curtis, Jour. Straits Branch Roy. Asiatic Soc. 25: 78. 1894; Pitard in Lecomte, Fl. Gén. Indo-Chine 1(4): 321. 1910; eae Fl. Malay Penin. 1: 182. 1922, pro parte; I. H. Burkill & M. R. Henderson, Gard. Bull. Straits Settl. 3: 347. 1925; Craib, Fl. Siam. Enum. 1: 121. 1931; Gagnep. in Hum bert, Fl. Gén. Indo-Chine, Suppl. 1: 274. 1943; Pham & Nguyen, Cay-Co Mién Nam Viét-Nam, 179. 1960; Pham, Cay-Cd Mién Nam Viét-Nam. ed. 2. 2: 301. fig. 1970, pro parte. C. amoenum auct., non Wall. ex Choisy; Vesque in C. DC. Monogr. Phanerog. 8: 576. 1893, quoad King 548. & ae auct., non T. Anderson; Baker f. Jour. Bot. London 62(Suppl.): C. Serene auct., non Pierre; Craib, Fl. Siam. Enum. 1: 120. 1931; 508 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Gagnep. in Humbert, Fl. Gen. Indo-Chine, Suppl. 1: 274. 1943, pro parte. C. globuliferum Ridley, Kew Bull. 1938: 121. 1938, typo excluso. Calophyllum sp. Craib in Schmidt, Bot. Tidsskr. 32: 328. 1915. Tree 6-30 meters tall, d.b.h. to 38 cm.; outer bark closely fissured; latex clear to opaque yellow (rarely white to whitish (SAN 22940, 71512)), sticky. Twigs slightly flattened, usually strongly 4-angled; axillary innovations nearly always lacking basal scars. Lamina elliptic to obovate (rarely suboblong), 3.7-13.5 by 1.6-5.2 cm., usually acuminate (acute) at apex, usually notably pale at margin, marginal thickening less than 0.4 mm. wide, thinly coriaceous, the venation + clear above and below, (5 to) 8 to 14 (to 17) veins/5 mm. Inflorescences axillary; bracts very fugaceous, usually lost even in very young inflorescences. DistripuTIOoN. Cambodia to Borneo, excluding mainland Java (Map 34). SELECTED SPECIMENS SEEN (‘‘e’’ denotes atypical specimens—see notes at end). Vietnam: Phu Quoc, Pierre 3645 (pr), near Hatien, Poilane 876 (k, p). Cambodia: Kampot, Mt. Bokor, 1000 m., Smitinand & Abbe 6511 (k). Thailand: Krat, Kookrap, 700 m., Kerr 17772 (pm, kK), Kao Saming, Put 567 (kK); Kaw Koh Chang, Klong Munse, Schmidt 528 (c, k); Klong Majum, Schmidt 603A (c, K), Klawng Non Si, 50 m., Kerr 9175 (a, c, E, K, L, Pp); Krabi, Ao Luk, 50 Map 34. Distribution of Calophyllum tetrapterum var. tetrapterum (circles), C. tetrapterum var. blumutense (star in or circle), and C. fetrapterum var. obovale (stars) in Southeast Asia—Malesi 1980] STEVENS, CALOPHYLLUM 509 m., Kerr 18567 (a, BM, C, K, P); Terutas, Satul, 5 m., Kerr 14213 (BM, c, kK, c N itrasirirak 206 (?) (BKF). Andaman and Nicobar Islands. S. ANDAMAN: Tusono- g’s collectors s.n., 23 Jan. 1893 (Bo, k, P, U). Malaya. Kepan: Kedah peak, 792 m., KEP 94422 (a, Bo, K, KEP, NY, SING); Bukit Enggang F.R., 427 m., FRI 13730 (e) (A, kK, SAN, SING); Sik catchment area, FRI 16289 (e) (a, KEP, SAR); Langkawi, G. Raya, Corner s.n., 15 Nov. 1941 (sinc), Bukit Sawak F.R., KEP 66442 (kep), Sungei Batu Asap, 150 m., Ridley 15540 (BM, kK, SING); Perangin F.R., KEP 79283 (sinc); Kuala Muda, Gurun, Jerai F.R., KEP 17925 (sING); Bukit Malut, KEP 7683 (sinc). Pinanc: Muku head, 150-240 m., Curtis 418 (2e) (K, sinc); Pulau Penang, above waterfall garden, SF.N 3368 (Bo, sING), Penang Hill, 762 m., FRJ 20525 (ker), Pantai Acheh F.R., KEP 72565 (keEpP). Perak: Larut, 150-240 m., King’s collector 3378 (x, Pp); Lumut F.R., 120 m., FRI 977 (?e) (A, K, KEP, SAN, SING); Taiping Hill, J. W. Anderson 114 (sinc); Batu Undan F.R., 305 m., KEP 54223 (keEp). SELANGor: Ulu Gombak F.R., FRI 2213 (a, K, SAN, SING); 22 mile Ginting Simpah, KEP 12854 (sinc); Bukit Lagong F.R., KEP 80646 (ker). NEGRI SEMBILAN: Kuala Pilah, Serting F.R., KEP 62955 (a, K, KEP, SING); Senaling Inas F.R., KEP 62882 (KEP); Pedas, G. Angsi F.R., 457 m., FRI 14620 (KEP, SAR, SING). MALAccA: Pulau Besar, Stone 9034 (BISH, G, KLU, MO); Pulo Pusar, Griffith, Kew dist. 877 (Bo, E, G, K, P); Bukit Bruang, 0 m., Derry 393 (BM, kK, sinc); Bukit Sandanan, 0 m., m., FRI 20705 (kEP, SAN); Kemansul F.R., KEP 99583 (e) (KEP): Ulu Sat .R., Machang, KEP 100143 (a, kK, KEP, SING); Pasir Mas, Gual Periok, KEP 66804 (KEP); Ulu Temiang F.R., Gua Musang, KEP 104284 (a, K, KEP, SAN, sinc); G. Rabong, 610 m., FRI 20617 (kep); Kuala Krai, Sungei Durian F.R., KEP 104728 (k, KEP, SING); KemahangF. R.,18m., KEP 93588 (KEP). TRENGGANU: along Sungei Pelong, FRI 14841 (k, KEP, SAR, SING): Jerangau F.R., m., KEP 78562 (ker); Dungun, Ulu Chukai F.R., KEP 100059 (kEp); 40th mile Jerteh Road, Sungei Tong, FRI 2521 (k, KEP, SAR, SING); path to G. Tebu, Jabi, 610 m., Shah et al. 3306 (sinc). PAHANG: Menchali F.R., KEP 94898 (A, KEP, SAN, SAR, sinc); Aur F.R., FRI 3646 (a, KEP, sinc); Ulu Sungei Kelui, 610 m., FRI 10956 (e) (4, kK, KEP, — Ulu Sungei Anak Endau, 396 m. FRI 8174 (a, K, KEP, SAR, SING); Kerdau, KEP 29997 (ker); Rotan Tunggul F.R., KEP 23389 (k, KEP, SING); Lepar Hill F.R., 150 m., FRI 9186 (e) (KEP); Baloh F.R., KEP 66646 (ker); Ulu Perah, KEP 1121/2 (ker); Kuantan, Sungei Sawah Chini, KEP 2712 (?e) (K, KEP, ay Ginting Highlands, 915 m., Stone 7443 (BIsH, KLU); Bukit Simpul, KEP 15638 (sinc); Kemasul F.R., 45 m., KEP 78741 (kep). JoHore: Mawai, SF'N 34748 (a, BO, K, KEP, SING); Labis Puls, G. Besar Massif, FRI 14014 (2e) (A, K, KEP, SAN, SING); G. Panti Timor, 478 m., FRI 7770 (kep, sinc); Johore Coast, Tg. Penawar, 15 m., FRI 7629 (a, K, KEP, SING); Kangka Sedili Kechil, SFN 28599 (k, KEP, siNc); Jemaluang F.R., KEP 73463 (ker); Lenggor F.R., KEP 72652 (xep); G. Arong F.R., 30 m., KEP 71312 (kep). Singapore: Bukit Timah F.R., SFN 38848 (a, BO, LAE, SING); Botanic Gardens jungle, SFN 36255 (a, BO, K, KEP, LAE, P, SING); Changi, Goodenough 1958 (sinc); Jurong, Corner s.n., 14 Jan. 1932 (sinc); SW. side of Seletar Reservoir, upper Mandai, SFN 40014 (sinc); N. side of MacRitchie Reservoir, Sinclair 5372 (sinc). Sumatra and adjacent islands. Riau: Lingga Arch., Pulau Singkep, bb 27/4 (Bo); Koeantan, Tjerenti, 50 m., bb 25239 (2?) (Bo, L). Dsampr: Moera Tembesie, Soengei Roean, 15 m. _ bb 9250 (e) 510 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 (Bo). SELATAN: Koeboestreken, 10-20 m., Endert 256 (po, L). Barat: Loeobek Sikaping, Tandjong Boengo, 370 m., bb 6512 (e) (Bo); Sidjoengdjoeng, Moeara, 596 m., bb 6057 (Bo). UTARA: Laboehan Batoe, Kota Pinang, Langga Pajoeng, ny Toroes 3354 (A, NY, US). eens A: G. Permisan, Soengei Selan, Bunnemeyer 2046 (Bo, L); G. Maras, Soengei Liat, 500 m., bb 1954 (Bo, L): Gadoeng, 20 m., bb 10579 (2) ce Bevitunc: Tandjong Pandan, Bantan, 20 m., 6b 7376 (Bo). Borneo. Sarawak. Ist Division: Semengoh Arboretum, 15 m., S 5361 (Bo, K, KEP, L, P, SAN, SAR, SING); Lundu, G. Pueh, 160 m., S 13724 (a, BO, K, L, SAN, SAR, SING); G. Gading, 760 m., S 13325 (?) (a, Bo, SAN, SAR, SING); G. Santubong E., 762 m., S 13695 (k, L, SAN, SAR, SING). 4th Division: Bukit Mentagei, Bok-Tisam Marudi, 610 m., S 23286 (a, kK, KEP, SAN, SAR, SING); Bukit Mersing, Anap, 400 m., S 21926 (a, BO, K, SAN, SAR SING). BRUNEI: Berakas F.R., 60 m., S 7820(a, Bo, K, KEP, NY, SAR, SING); Andalau F.R., 15 m., SAN 17529 (a, Bo, K, KEP, SING); Kuala Belait, S 1945 (sar). SABAH. Kudat: Lokapas, Bengkoka, 72 m., SAN 1827 (a, Bo, K, Us). Keningau: Sook plain, mile 11/2 Tulid road, SAN 55615 (san). Labuk & Sugut: base of Bukit Mesasau, 180 m., SAN 25429 (Bo, K, KEP, L, SAN, SAR); Telupid, mile 87, 120 m., SAN 71474 (san). Sandakan: Leila F.R., A 27/2 (a, K, KEP, KLU, SING, US); Sepilok F.R., 120 m., A 3880 (a, kK, KEP, SAN, SING); Sungei Kapur, SAN 22940 (SAN, SAR); Batu Sapi, Mt. Walker, SAN 61592 (SAN). KALIMANTAN. Timur: Central Kutei, Belajan R., Kostermans 10254 (a, BO, K, LAE, P, SING); E. Kutei, Kostermans 7248 (a, B, BO, CANB, G, K, LAE, P, SING). Selatan: Bukit Besae, 305 m., Motley 304 (kx). Tengah: Beneden Dajak, 40 m., bb 8159 (po). Java a nd adjacent islands. Bawean Is.: Mt. Besar, 400 m., Buwalda 3224 (Bo, L). KARIMOENDJAWA: Japara, 150 m., Ja 1705 (Bo). Eco.ocy. Usually in well-drained mixed dipterocarp forest; often common on sandy, acid soil (eastern Sabah); sometimes on highly leached, giant podzols (elsewhere in Borneo); in swamp forest (Malaya) (Corner, loc. cit.. as C. floribundum; FRI 2521, Sinclair 7763); on ultramafic rock (Borneo): in evergreen forests (Thailand); to 975 m. alt. On mainland SE. Asia flowering mostly December to February, rarely in March, April, July, August, and November; data from the rest of Malesia scanty, but flowering perhaps more scattered. Flower scented, sometimes fetid (‘‘very foetid, [smelling] of garlic when decaying’—SFN 33143). Fruiting February to May, rarely at other times; fruit yellowish or bluish green, eaten by iban patong (SFN 28599, Johore). One atypical specimen has bilabiate galls (FRI 14014, Johore). GERMINATION AND YOUNG PLANT. The radicle breaks through the stone to one side of the base. The seedling has two pairs of leaves separated by well- developed internodes over | cm. long. Subsequently produced internodes are longer, the terminal bud is functional, and growth is erect. (Stevens et al. 121; Stone 9034.) 100b. Calophyllum tetrapterum Miq. var. obovale (Miq.) P. F. Stevens, comb. et stat. nov C. obovale Miq. Fl. Indiae Batavae, Suppl. 1(3): 498. 1861; F. Mueller in Walp. Ann. Syst. Bot. 7: 357. 1868; Vesque inC. DC. Monogr. Phanerog. 1980] STEVENS, CALOPHYLLUM 511 8: wa 1893. Type: Sumatra, ad on Siboga, Teysmann, HB 644 (holotype, u; isotypes, Bo, K, L, C. griffithii T. Anderson in ae f FL. Brit. India 1: 273. 1874; King, Jour. Asiatic Soc. Bengal, II. 59: 179. 1890; Vesque in C. DC. Monogr. Phanerog. 8: 602. 1893; Ridley, Fl. Malay Penin. I: 187. 1922, pro parte; M T. C. Whitmore, Tree Fl. Malaya 2: 168. 1973. Type: Malaya, Malacca, Griffith, Kew dist. 882 (holotype, kK; isotypes (sometimes s.n.), BO, E, H, L, M, P). C. globuliferum Ridley, Kew Bull. 1938: 121. 1938, paratypo excepto: Masamune, Enum. Phanerog. Born. 475. 1942. Type: Dutch SE. Borneo [Kalimantan], Bangarmassing, Motley 618 (holotype, k). C. elegans auct., non Ridley; Keith, N. Borneo Forest Rec. ed. 2. 2: 313. 1952, Tree 3-20(-27) meters tall, d.b.h. to 20 cm.; outer bark smooth or rather distantly fissured; latex clear yellow, fluid, becoming opaque when rubbed. Twigs flattened, usually not strongly angled; axillary innovations sometimes with basal scars. Lamina obovate (rarely elliptic to oblong), (2.2—)3.7-9.5(-14) by (1—)2-5(-6.5) cm., rounded to shortly and bluntly acuminate at apex, usually notably pale at margin, marginal thickening (0.2-)0.4-1 mm. wide, rarely with clear submarginal vein ca. 0.7 mm. from margin, thinly coriaceous to coriaceous, the venation + clear on both surfaces, (4 or) 5 to 8 (to 10) veins /5 mm. Inflorescences sometimes terminal; bracts sometimes foliaceous, up to 2 cm. long, subpersistent. Distripution. The Malay Peninsula, Sumatra, and Borneo; scattered (Map SELECTED SPECIMENS SEEN. Malaya. Perak: G. Besut, Soepadmo s.n., 28 Nov. 1976 (BISH, KLU). Sumatra and adjacent islands. Riau: Riouw Arch., P. Bintan, Pinang, Kaloebi, Goenoeng Si an si shee 3471 (A, US), Naa Pajoeng, si Toroes 3409 (intermediate) (A, k, y, US), s1 Mandi Angin, si Toroes 3983 (intermediate) (A, us). ae eens Ist Division: Bukit Jebong, Bau, S 25605 (A, K, KEP, L, SAN, SAR, SING); G. Pueh, Sematan, 628 m., S 34475 (A, SAN, SAR). Brunet: Anduki F.R., KEP 37144 (kx, kep); Bukit Puau, 12 m., Ashton s.n., July 1959 (sar); Temburong, Bukit Patoi, 281 m., SAN 17132 (A, BO, K, KEP, L, SAN, SING); Berakas F.R., 30 m., S 4926 (intermediate) (SAR). SABAH. Sipitang: Mile 5, Menggalong F.R., SAN 21806 (A, K, SAN). Beaufort: Weston, Siangan F.R., 69 m., SAN 78166 (SAN). Papar: Kimanis, 24 m., SAWN 41411 (k, L, SAN). Keningau: Sook, mile 21/2 Tulid road, 917 m., SAN 49521 (SAN). KALIMANTAN. Timur: W. Kutei, Mt. Palimasin near Tabang on Belajan R., 700 m., Kostermans 12899 (Bo, k, L); Boelongen, Sungei Binai, Rutten 20 (Bo, vu). Selatan: G. Pamatton, Korthals s.n. (L). Barat: between Mt. Klam and S. Djemela, Hallier 2526 (a, Bo, K, L, SING). EcoLocy. Primary or sometimes secondary mixed dipterocarp forest, in Sarawak sometimes on limestone; 12-917 m. alt. Flowering March, April, and June (flower scented); fruiting April, June, and October (fruit reddish yellow, pedicel red (Kestermans 12899)). 512 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 GERMINATION AND YOUNG PLANT. The seedling has two or three pairs of scale leaves and two pairs of expanded leaves, all separated by well-developed internodes. The young plant is erect, the terminal bud is functional, and the internodes are well developed. (Two plants only: Stevens et al. 219A.) The epithet obovale refers to the leaf shape of the type specimen. 100c. Calophyllum tetrapterum Miq. var. blumutense (M. R. Henderson & Wyatt-Smith) P. F. Stevens, comb. et stat. nov. C. blumutense M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 343. pl. 22. 1956; T. C. Whitmore, Tree Fl. Malaya 2: 171. 1973. Type: Malaya, Johore, Gunong Blumut, 3300 feet [1006 m.], 23 March 1923, KEP 5880 coll. Yeob (holotype, sinc; isotypes, K, SING). Shrub 1.5 meters to tree 12 meters tall; bark unknown. s flattened, not strongly angled; axillary innovations lacking basal scars. Lamina elliptic (rarely suboblong), 5.5-11.3 by 2-4.3 cm., acute to subacumi- nate at apex, pale at margin, marginal thickening 0.2-0.35 mm. wide, coriaceous, the venation clear above, obscure below, the latex canals impressed (G Blumut), 5 to 7 veins/5 mm. Inflorescences very rarely terminal; bracts not seen, (?)small, fugaceous. DistripuTion. Malay Peninsula, Johore (Map 34). SELECTED SPECIMENS SEEN. Malaya. Jounore: G. Blumut, 762 m., KEP 98052 (K, KEP); G. Panti, 488 m., SFN 18098 (Bo, K, SING). Ecotocy. Shrub or small tree of colline or lower montane rain forest, 488-1006 m. alt. Flowering February and March; fruiting in May. The name of the mountain on which the type specimen was collected suggested the epithet blumutense. Calophyllum tetrapterum is usually an easily recognizable species. It is characterized by its short terminal bud, its often strongly four-angled and soon glabrescent twigs, and its rather small, more or less chartaceous lamina with clear venation, a sharply raised midrib narrowing gradually from the base, and a pale-drying margin. The base of the dried lamina is minutely truncate where it joins the deeply concave petiole. The inflorescences and pedicels are usually slender, although the latter may become thickened in fruit; the more or less spherical fruits often dry grayish or pale brownish and sharply wrinkled. The epithet tetrapterum (‘‘four winged’’) is appropriate, since many specimens have strongly four-angled twigs. Infraspecific Variation Henderson and Wyatt-Smith adopted a wide view of this species (see loc. u sort out the variation within the species as here delimited, as well as to clarify the relationships between C. tetrapterum, C. rupicola, and C. pisiferum. 1980] STEVENS, CALOPHYLLUM 513 The type specimen of Calophyllum tetrapterum, Junghuhn s.n., differs from many others assigned to var. tetrapterum. The indumentum on the inflorescence axis and tepals in particular is relatively well developed (in some other (such as Forbes 3220a), as well as some from Malaya, agree with the type. Although there are a few specimens intermediate between Calophyllum tetrapterum vars. obovale and tetrapterum (cited under the former), the two are usually readily distinguishable by the characters given in the key above. There may also be differences in the field characters, although this needs to be confirmed. A tree of C. tetrapterum var. obovale on Bukit Jebong, near Kuching, Sarawak, had fluid latex that became opaque when rubbed, and venation that was visible on the upper surface of the lamina but invisible on the lower, the reverse of the usual case. There are also possible differences in the seedlings (see above). 4475 is a very robust specimen of Calophyllum tetrapterum var. obovale with twigs up to 3.5 mm. across and notably short terminal internodes. Kostermans 12899, from Kalimantan, is a very small-leaved specimen referred to var. obovale with some hesitation: it has a terminal bud 1.5-2 mm. long, coriaceous, obovate leaf blades 2.2-5.7 by 1-2.4 cm., and fruits only 7 by 6 mm. borne on red pedicels. KEP 48612 is an unlocalized specimen probably from Malaya that may belong here; C. fetrapterum var. obovale is less distinct in Malaya than elsewhere in its range. The taxon Calophyllum tetrapterum var. blumutense is a very local variant, and specimens from the two mountains from which it is known differ somewhat. The latex canals on the lower lamina surface of the Gunong Panti specimen are not impressed, while those of the Gunong Blumut specimens are. en Henderson and Wyatt-Smith described C. blumutense, they noted that it was similar to C. floribundum (= C. tetrapterum var. tetrapterum) but thought that it could be distinguished by the more distant venation of the lamina. However, specimens from the Malay Peninsula (e.g., KEP 100143, 104284) otherwise clearly assignable to C. tetrapterum var. tetrapterum have compara- bly distant venation. Henderson and Wyatt-Smith also described the fruits of C. blumutense as smooth, while those of C. floribundum were described as wrinkled. However, the fruits of the former are ripe, and there is a tendency in C. tetrapterum for ripe fruits to be smooth, with large air spaces disorganizing the outer layer and occupying most of the space between the stone an the skin (see also under C. foetidum, below). There are a number of specimens of Calophyllum tetrapterum var. tetrap- terum from the Malay Peninsula and Sumatra (cited as ‘‘e’’ in the list above) that are somewhat different from the others. They have a shorter, more congested inflorescence usually less than 2 cm. long, and the flowers are small. The lamina often dries darker brown on the upper surface; the margin does not dry notably paler; the venation is rather dense, with (8 to) 11 to 13 (to 15) veins/5 mm.; and the midrib on the upper surface of the leaf, although narrowing gradually from the base, is relatively little raised. The 514 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 fruit 1s spherical and is smooth when mature; the young fruit is at least sometimes prominently apiculate. The type specimen of C. rupicola var. elatum is a specimen of this type. In some characters (leaf color, venation density, flower, and inflorescence) these atypical specimens of C. tetrapterum var. tetrapterum approach C. calaba var. bracteatum. Yet other collections, notably some from Kelantan (e.g., KEP 104284, FRI 20705), have fruits like those mentioned above. More studies are needed to clarify the significance of this variation, and it would be premature to give these specimens formal recognition. Synonyms and Nomenclature As mentioned above, the type specimen of Calophyllum tetrapterum differs somewhat from many of the specimens assigned to var. tetrapterum. Miquel (1854, loc. cit.) described the pedicels as sometimes being four together, although they are in pairs, separated by an internode. Miquel’s description may have led Planchon and Triana (/oc. cit.) to compare C. fetrapterum with C. spectabile (= C. soulattri), a very different species that often has four flowers at a node. The type specimens of most of the names reduced to synonymy under Calophyllum tetrapterum var. tetrapterum are nearly all close matches with the common form of the variety. The specimens cited under C. floribundum in the original description, Maingay, Kew dist. 170, 172, and ?171, belong to three species. Maingay, Kew dist. 170 agrees with the original description in these last two species, which are very distinct, see the discussion after C. pulcherrimum. The type specimen of Calophyllum venustum, King’s collector 7763, has the broader leaf marginal thickening that is common in C. fefrapterum vat. tetrapterum, and the lamina tends to be rounded at the apex. In both of b ut 1 taxon based on this character (see also above under C. tetrapterum var. blumutense). 1980] STEVENS, CALOPHYLLUM 515 The type specimen of Calophyllum rupicola var. elatum is probably to be referred to C. tetrapterum var. tetrapterum, although it is a somewhat atypical specimen (see above). The type specimens of Calophyllum obovale and C. globuliferum are very similar, and both also have terminal inflorescences. However, C. griffithii is reduced to 1y under C. tetrapterum var. obovale with some hesitation. e key craracies used to separate this species was the presence of a submarginal vein (although this is obscured by the submarginal thickening in most specimens of C. tetrapterum var. obovale and is thus not obvious, it is sometimes visible). Ridley (1922, Joc. cit.) noted that C. griffithii was common on the riverbanks in Pinang, Malacca, Johore, and Singapore; I am not sure to what species he was referring, although it may have been C. tetrapterum sensu lato. 101. Calophyllum rupicola Ridley, Trans. Linn. Soc. Bot. II. 3: 278. 1893 (‘SC. rupicolum’’), Fl. Malay Penin. 1: 182. 1922; M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 346. 1956; T. C. Whitmore, Tree Fl. Malaya 2: 168. 1973. Type: Malaya, Pahang, Tahan River, anno 1891, Ridley 2636 (holotype, sinc; isotypes, BM, K). C. rupicola Ridley var. elatum T. C. Whitmore, Gard. Bull. Singapore 26: 270. 1973, pro parte, typo haud incluso, Tree Fl. Malaya 2: 168. 1973, pro parte. Shrub or tree 0.6-6 meters tall; bark unknown; latex yellow to cream. Twigs slightly flattened, 0.5-2(-3.5) mm. across, strongly 4-angled, drying brown, + puberulent when young; axillary innovations usually lacking basal scars; internodes 0.5-3.5 cm. long; uppermost pair of axillary buds rounded, to | mm. long, spreading, not very conspicuous; terminal bud plump, 1.2—4.5 mm. long, with short, grayish to brownish, adpressed indumentum (hairs, Ficure 31, 0, p), underdeveloped internode to 2.5 mm. long. Petiole 2—4(—8) mm. long, concave above, convex below, glabrescent; lamina elliptic to elliptico-oblong (rarely oblong), (1.9-)3-9(-16) by (0.35-)0.75-3(-4.7) cm., acute to acuminate at apex, narrowly cuneate at base, rather broadly and distantly undulate but slightly recurved at margin, coriaceous to thinly coriaceous, drying dark brick color to umber above and umber to fulvous below, + persistently puberulent on midrib above and below (on entire lower surface), the midrib above narrowing gradually from base, raised, 0.15-0.25 (-0.3) mm. wide at midpoint, below raised, striate, the venation apparent to subobscure above, usually subobscure below, raised, latex canals often impressed, (6 to) 12 to 15 veins/5 mm., angle of divergence 50-60(—80)°. Inflorescences from foliate axils, with (1 to) 3 to 9 flowers, unbranched, the axis 0.25-1(-2.2) cm. long, strongly angled, puberulent, lowest internode 2-5(-7) mm. long, upper internodes if present often very short; bracts ovate, ca. 1.8 mm. long, densely puberulent below; pedicels 1.5-7 mm. long, puberulent to glabrous, strongly angled even in fruit. Flower (?)hermaphroditic; tepals 4 or 6, the outer pair ovate, 2.7-3 by ca. 2.2 mm., with strip of puberulence down back, the inner ones elliptico-oblong, 4—5 by, 1.5-2.6 mm.; 516 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 stamens ca. 50, the filaments to 3 mm. long, the anthers subelliptic to oblong, 0.4-1 mm. long, + rounded at apex; ovary 1.2-1.6 mm. long, style 2-2.7 mm. long, stigma unknown. Fruit ellipsoid to subspherical, 0.8-1.2 cm. by 0.6-0.95 mm., usually strongly apiculate, drying brown, wrinkled when immature, later smooth; outer layer not detaching cleanly from stone, very thin, obscured by large air spaces developed; stone spherical to ellipsoid, 7-10.5 by 6-8.5 mm., rounded at apex, the walls ca. 0.15 mm. thick, smooth, unmarked; (?)spongy layer. DistripuTion. Northeastern Malaya and Peninsular Thailand; a form in eastern Malaya and scattered in Sumatra (Map SELECTED SPECIMENS SEEN (all specimens of the variant seen cited). Thailand: Peninsula, Narathiwat, Waeng, Sangkhachand et al. 1067 (Br, L). Malaya. KeLANTAN: Kuala Rek, SFN 10175 (a, sinc); Chan[n]ing Woods, Ridley s.n., 30 Jan. 1917 (var.) (Kk); Bukit Baka, Machang, 457 m., Shah & Shukor 3163 (sinc); Ulu Sat F.R., 210 m., FRI 2951 (kK, KEP, sING); G. Stong, 762 m., FRI 12416 (k, KEP); Mak Meh Nak, KEP 37975 (sinc). TRENGGANU: Mandi Angin, S. Loh near Kuala Datok, 30 m., FRI 8952 (k, Kep, L); Ulu Telamong, Bukit Rambai F.R., FRI 1140] (var.) (KEP, L, SAN, SING); Bukit Bauk F.R KEP 76091 (2) (a, ‘SING); Sungei Trengganu, near Jeram oe FRI 8350 (A, K, KEP, L, SAR, SING); Compartment 90, Gunong Tebu F.R., 150 m., FRI 2513 (KEP, L, SAR, SING); Ulu Brang, near Kuala Lallang, 90 m., FRI 12526 (A, K, KEP, L, SING); PAHANG: between Sungei Puteh and Teku Taman Negara, 1920 (k); Taman Negara, Sungei Tahan, Shah & Shukor 2670 (c, KEP, . SING): G. Tahan, Wong & Wyatt-Smith 22 (kep); Sungei Tembeling, 90 m., Stone 10889 (KLu); between Kuala Tahan and Kuala pga a m., Chin 1318 (k, KLU, L); Tahan woods, Kuala Teku, 120 m., FRI 4762 (k, KEP, L, sinc); Rompin, KEP 15461 (var.) (KEP); eee F.R., “Ahmad & Shukor 448 (var.) (Cc, KEP, SING); Bukit Cheras, ca. 180 m , Henderson g.n., 22 Oct. 193] (var.) (sING); Bukit Goh F.R., KEP 3132 (var.) (SING). Jouore: G. Blumut, Upper Camp, 549 m., FRI 8824 (var.) (A, KEP, L, SAR, SING); Sungei Kayu Ara, Mawai-Jemaluang Road, Corner s.n., 9 Feb. 1935 (var.) (sinc); Panti F.R., 6 m., Stevens et al. 104 (var.) (a), 104A (var.) (A). Sumatra. SELATAN: Toelang Bawang nabij Talung Batoe, 30 m., /denburg 44 (var.) (Bo); Moeko2, Lalang Loeas, 50 m., bb 23555 (var.) (Bo, L, Mo). Utara: Padang Sidempuan, near village Grunggan, 150 m., Kostermans 22040 (var.) (a, L). Eco.tocy. Usually rheophyte growing on banks of rivers and streams, hence liable to be inundated. Normally shrub less than 2.5 m. tall, but FRI 8350 apparently taken from tree 8 m. tall with trunk initially creeping along ground. Sometimes in forest away from streams—then small tree (e.g., FR/J 2516). Variant typically tree (see below). 30-762 m. alt., the variant 6-550 m. alt. Flowering January, April, May, July, August, and November; fruiting Janu- ary, February, April, and June to October (ripe fruit yellowish or orange). The typical form of Calophyllum rupicola can be characterized by its often rather small, (narrowly) elliptic, subcoriaceous lamina that dries grayish brown above and golden brown below, with the latex canals on both surfaces often 1980] STEVENS, CALOPHYLLUM aly impressed. The buds along the twigs are often markedly supra-axillary. Its inflorescence is short and congested, with the upper internodes often shorter than the lowest one; both the inflorescence axis and the short pedicels are strongly angled. The small fruits are apiculate, sometimes markedly so, and are reported to be yellow to orange when ripe. The epithet rupicola, ‘‘rock- dweller,’’ emphasizes the fact that this species is usually found growing among rocks by rivers and streams When Calophyllum rupicola grows away from streams, it looks rather different from the typical rheophytic form of the species; it is larger in all its parts and has a broader leaf blade. However, the two extremes are connected by intermediates. Scars at the bases of the innovations are found in the smaller, profusely branched rheophytic forms. Some of the specimens cited as C. rupicola var. elatum by Whitmore (loc. cit.; FRI 2513, 2516) belong to the forest form (see also the discussion after C. tetrapterum). Sangkhachand et al. 1067, from Peninsular Thailand, has stout twigs and large leaves (the upper limits of the measurements given) and was taken om a tree 10 meters tall. In such characters as inflorescence and color and venation of the dried leaf, it agrees with the typical form of Calophyllum rupicola. KEP 76091 was included in C. rupicola variety by Henderson and Wyatt-Smith (/oc. cit.). Itis retained in C. rupicola with hesitation: the specimen is superficially like C. dasypodum but differs in hair type and leaf. The thick outer layer of the immature fruit also invites comparison with C. dispar. The specimens cited above as ‘‘var.’’ are perhaps merely an extreme form of the forest-dwelling Calophyllum rupicola. They have not been included in the description above and can be characterized as follows. Shrub 0.5 meters (Kostermans 22040) to tree 6(-18) meters tall; inner surface of outer bark brick orange; latex opaque, pale yellow, only tardily becoming sticky. Petiole tending to dry dark brown, contrasting with paler brown twig; lamina elliptic-oblong, (3.5-)7-16.5 by (1.1-)3.2-5 cm., sharply acuminate, acumen (0.3-)0.5-1 cm. long, thinly coriaceous, margin closely and strongly undulate, angle of divergence of venation 65-80° In other characters these specimens seem close to Calophyllum rupicola: the latex canals of the lamina tend to dry impressed, the inflorescence axis and the short pedicels are strongly angled, and—according to field notes on Kostermans 22040—the fruit is yellow. The variant of C. rupicola grows with C. retrapterum in the Panti Forest Reserve, Johore (C. rupicola—Stevens et al. 105, 105A; C. tetrapterum—Stevens et al. 104), and the two are clearly different taxa. The inner surface of the outer bark of Calophyllum rupicola variant was brick orange in color and the latex opaque, pale yellow, and fluid; the margin of the lamina was undulate; and the very young plant was arched, with the stem initially plagiotropic. These bark characters are at most unusual in C. tefrapterum (the inner surface of the outer bark of C. tetrapterum in the Panti F.R. was brownish, and the latex was clear yellow and sticky). I have never seen C. tetrapterum with undulate margins to the leaf blades, nor have I seen young plants growing other than erect. Clearly, it will be of considerable interest to study germination and the young plant of C. rupicola sensu stricto > 518 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Calophyllum rupicola is apparently related to C. pisiferum, which in Thailand (although apparently not in Malaya) is also sometimes a rheophyte, and to the variable C. tetrapterum. Calophyllum pisiferum has stouter twigs, better- developed indumentum, and leaf blades that are less sharply pointed at the apex and that dry browner. Calophyllum tetrapterum is usually quite different from C. rupicola, having broader leaf blades with clearer, more distant venation and a margin that dries paler than the rest of the blade; the blade itself rarely dries golden brown below, and the latex canals are usually + invisible. The inflorescence and pedicels are longer and are not so clearly angled. However, a few specimens referable to C. tetrapterum at least superficially approach C. rupicola. FRI 10956 (610 m., SE. Pahang) is like C. rupicola in inflorescence type, flower size, and—to a certain extent—color of the dried leaf; it was taken from a tree 15 meters tall. FRI 13630 (425 m., Kedah) is perhaps like C. rupicola in fruit type, although in other respects it is not similar. Careful field observations of the Calophyllum rupicola—C. pisiferum—C. tetrapterum group as a whole in Malaya and Thailand are needed to assess the significance of all the variation discussed. 102. Calophyllum pisiferum Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 294. 1862; Ridley, Fl. Malay Penin. 1: 184. 1922; M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 345. 1956; T. C. Whitmore, Tree Fl. Malaya 2: 172. 1973. Type: Malaya, Malacca, Nov. 1837, Gaudichaud S&6 (lectotype, G; isolectotype, P). Ficure 32, k, 1. C. retusum Wall. ex Choisy var. cambodgense Pitard in Lecomte, Fl. Gén. Indo-Chine 1(4): 321. 1910; Gagnep. in Humbert, Fl, Gén. Indo-Chine Suppl. 1: 275. 1943 (incl. C. sangkae); Pham, Cay-C6 Mién Nam Viét-Nam. ed. 2. 2: 303. 1970. Type: Cambodge, Dom-Phaong, Hahn 86 (holotype, P). C. retusum Wall. ex Choisy var. cochinchinense Pitard in Lecomte, FI. Gén. Indo-Chine 1(4): 321. 1910; Gagnep. in Humbert, Fl. Gén. Indo-Chine Suppl. 1: 275. 1943; Pham & Nguyen, Cay- Co Mién Nam Viet-Nam, 179. bang 62bis D. 1960; Pham, Cay-Cé6 Mién Nam Viét-Nam. ed. 2. 2: 303. fig. 1970. Type: Cochinchine [Vietnam], Ti Tinh, Thorel 1395 (lectotype, p; isolectotypes, B, k). C. sangkae Craib, Kew Bull. 1925: 18. 1925, Fl. Siam. Enum. 1: 122. 1931. Type: Siam [Thailand], ae Sangka, ca. 300 m., 4 Jan. 1924, Kerr 8283 (holotype, k; isotype, C. motleyi Ridley, Kew Bull. 1938: 122, 1938; Masamune, Enum. Phanerog. Born. 476. 1942. Type: Dutch SE. Borneo [Kalimantan], Bangarmassing, Motley 865 (holotype, k). C. retusum auct., non Wall. ex Choisy; T. Anderson in Hooker f. FI. Brit. India 1: 272. 1874, pro parte; Pierre, Fl. Forest. Cochinch. 1: pl. 102, 1885; Vesque, Epharmosis 2: r. 25. 1889; King, Jour. Asiatic Soc. Bengal, II. 50: 176. 1890, pre majore parte. C. amoenum auct., non Wall. ex Choisy; Vesque in C. DC. Monogr. Phanerog. 8: 576. 1893, pro parte. C. rupicola Ridley variety; M. R. Henderson & Wyatt-Smith, Gard. Bull. 1980] STEVENS, CALOPHYLLUM 519 Singapore 15: 347. pl. 27. 1956, pro parte. C. rupicola Ridley var. elatum T. C. Whitmore; Corner, Gard. Bull. Singapore Suppl. 1: 104. 1978, pro parte. Shrub 0.1 meter to tree 30 meters tall, d.b.h. to 60 cm.; trunk without buttresses, knee roots sometimes present; outer bark yellowish and brownish mottled, or pale yellow, hoop marked or not, somewhat cracked longitudinally or lenticellate, the inner surface dirty greenish and dull straw mottled; under bark dark red; inner bark dark red; latex opaque yellow, sticky (pale yellow—SAWN 51649; bright yellow—Motley 865); sapwood dirty white or pale ochre; heartwood red and beautifully marked, or light brown. Twigs slightly to strongly flattened, 1.5-2.5 mm. across, sharply 4-angled, drying grayish brown to blackish, shortly and usually persistently tomentose; axillary innovations lacking basal scars or not; internodes 0.3-3 cm. long; uppermost pair of axillary buds rounded, less than 0.7 mm. long, + spreading, inconspicuous; terminal bud plump, 2-4.5 mm. long, with adpressed to spreading, brown indumentum (hairs, Ficure 31, f, h-k), underdeveloped internode absent. Petiole 2.5-5 mm. long, deeply concave above, convex t oblong or (narrowly) elliptic, (1.9-)3-7.5(-9.5) by (0.9-)1.6-3.2(-4) cm., round- ed to acute at apex, broadly rounded (shallowly cordate) to narrowly cuneate at base, almost flat to distantly undulate and at most slightly recurved at margin, thinly coriaceous to coriaceous, drying sepia to olivaceous above and umber to sabelline below, subpersistently puberulent on midrib above and especially below (soon glabrescent), the midrib above rather narrow at base, further narrowing gradually, sharply raised (+ depressed at first), 0.15-0.35 mm. wide at midpoint, below raised, striate or rounded, the venation on both surfaces + apparent, raised, latex canals sometimes impressed, (3 to) 5 to 8 (to 11) veins/5 mm., angle of divergence 55-75°. Inflorescences from (de)foliate axils along stem, with 5 to 15 flowers, unbranched, the axis (0.2-)1-3.5 cm. long, short-tomentose at base, otherwise + puberulent, lowest internode to 5(-10) mm. long, next internode often shorter than those at end of inflorescence; bracts ovate to subelliptic, 2.2-3 by 0.8-1.5 mm., deciduous; pedicels 0.3-1.3(-1.8) cm. long, glabrous or almost so, slender and 4-alate. Flower (?)hermaphroditic; tepals 4, the outer pair broadly elliptic to ovate, 3.5-5 by 1.5-3 mm., sometimes with short hairs on back near apex, the inner ones elliptic to obovate, 5-6 by 2-4(-5) mm.; stamens 30 to 70, the filaments to 3.5 mm. long, the anthers elliptic, 0.4-0.6 mm. long, (sub)retuse at apex; ovary 1.2-1.3 mm. long, the style 2.5-3.5 mm. long, the stigma expanded, 0.3-0.4 mm. across. Fruit spherical to ellipsoid, 6-9 by 5.5-7 mm., sharply apiculate, drying sharply wrinkled, smooth when quite 0.5 mm rounded at apex, the walls less than 0.1 mm. thick, smooth, unmarked; spongy layer thin DistriBuTION. Southern Vietnam to Borneo, scattered, excluding Java (Map 520 JOURNAL OF THE ARNOLD ARBORETUM [voL. 6 _— AP 35. Distribution of Calophyllum pisiferum (circles), C. dispar (squares), C. lineare (half-circle), and Calophyllum sp. 98 (stars in solid circles) in Southeast Asia—Malesia. SELECTED SPECIMENS SEEN. Burma: Martaban, Griffith s.n. (BM, ©, FI, M, P). Vietnam: flumen Saigon prope Beu Chuc, Pierre 3648 (p); Phu Thoe. Sinus Siamica, Pierre 3648 (pe); Cai Cong, Thorel 1395 (p). Cambodia: Koh Kong, presqu’ile Smach, Vidal 5000 (L, P); pres de Thmar Banf, 400 m., Martin 1495 (p); route de Sré Ambel, Martin 1017 (rp); Kampot, cascade Tuk Sap, Martin 1133 (rp); Pnomh Penh, Béjaud 458 (x, pe); Kg. Thom, Kg. Loa Bejaud 215 (rv); Siem Reap, Phnom Kulen, 250 m., Martin 1298 (P). Thailand: Chantaburi, Makam, Ban-ang, Chit 161 (BKkF); E. of Makam, 100 m., Sorensen et al. 464 (c); Krat, Kao Saming, Kerr 9435 (Bm, c, K, L, Pp); Kao Kuap, 700 m., Kerr 17774 (a, BM, c, L, P)} SE. Ko Kut, Charoenphol et al. 5097 (aau); Trat, Kaw Kut, 20 m., Smitinand 5723 (pkF). Malaya. Macacca: sine loco, Griffith s.n. (coll. Lemann) (BM, BO, G, GH, K, L, MO, P). JoHoRE: Mawai, ‘“‘low”’ alt., SFN 34706 (?) (Bo, LAE, SING), SFN 34747 (?) (a, L, sinc); Kangka Sedili Ketchil, Corner s.n., 17 June 1934 (?) (sinc). Sumatra and adjacent islands. 1980] STEVENS, CALOPHYLLUM 52] SELATAN: Meranjat, Sebetan, Teysmann, HB 3451 (po). BaANGKa: Lobok Besar, 5 m., bb 34028 (L, sinc); Rias, 10 m., bb 1540] (Bo, L). BeLituNG: Tandjong Pandan, Teysmann s.n. (Bo). Borneo. Sapan. Labuk & Sugut: Sungei Mangkan- yoh, N. of Klagan R., SAN 51649 (san); Lower Sugut, SAN 27622 (san). Sandakan: Ulu Dusun F. R.,2m., Stevens et al. 354 (a). Kinabatangan: opposite Gabang Camp, SA WN 23096 GG KEP, SAN, SAR, SING). KALIMANTAN. Timur: Central (Bo); Kaiteng, Sungei Sekunyir, Kumai, Anderson (1975) I (Bo). Tengah: Djoeking Koempai (Koelo Kapaeos), bb 2180 (Bo); Beneden Dajak, Mangkoe- top, bb 2957 (Bo). Barat: Mandor, 50 m., Dunselman 101 (so); Pontianak, sine loco, bb 16373 (Bo, kK, L, SING, u). Ecotocy. On SE. Asian mainland: shrub or small tree frequently on stream banks, usually rocky places (narrower-leaved specimens typically rheophytes), also on sandy soil; 20-700 m. alt. Flowering October and January (flower scented); fruiting November to February (ripe fruit apparently orange to light chestnut, although reported by Pierre (/oc. cit.) to be white). In Malesia: in peat swamps, periodically inundated flat land, or riverine forest; below 50 m. alt. Flowering November, January, and March (flower scented; corolla reported to be pink (Anderson (1975) 1)); fruiting March and April (fruit yellow and sour, or orange) GERMINATION AND YOUNG PLANT. The radicle probably breaks through the stone to one side of the base. The seedling is small (less than 7 cm. tall), and has two or three pairs of leaves separated by internodes ca. | cm. long. During subsequent growth, the internodes produced are successively longer, the plant grows erect, and the terminal bud is functional. (Stevens 353, from Sabah.) Loca uses. A decoction of the bark is used to cure diarrhea (Cambodia); the branches are used for house and boat poles (Thailand). Calophyllum pisiferum can be recognized when sterile by its sharply four-angled twigs that are often persistently short-tomentose. Its rather small leaf blades vary considerably in size and sometimes also in shape on one specimen and are sepia above in older specimens; their venation is relatively distant and prominent. Internode length increases toward the top of the inflorescence axis, and the axis is usually covered with indumentum for its entire length. The pedicels are slender, and the flowers always have four tepals. The fruits are small and wrinkled and have a thin outer layer; they look rather like dried, wrinkled peas (pisiferum means ‘‘pea-bearing’’). Calophyllum pisiferum is quite a variable species and BAS a disjunct ne tribution'® (Map 35), especially in mainland Southeast Asia. Its been troublesome. With only one exception, the specimens from this area were collected from shrubs or small trees less than 9 meters tall (Smitinand 5723 is reported to have been taken from a tree up to 30 meters tall—probably '°The single specimen from Burma, Griffith s.n., may be incorrectly labeled. 522 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 an error) often growing alongside rivers. The leaf blades vary from narrowly elliptic to ovate. The Malesian specimens were taken from medium-sized to fairly large (to 30 m.) trees generally growing in wet places; variation in lamina shape is less, and the blades are usually more or less ovate. In 36 in specimens from the Southeast Asian mainland (although they occur on some innovations in Charoenphol 5106). Nevertheless, C. pisiferum is broadly circumscribed because in other vegetative details, hair type, flowers, and fruit, all specimens are similar. Calophyllum lineare is very close to C. pisiferum and may be merely an extremely narrow-leaved form of it. There are other, perhaps minor, dif- ferences between the taxa: C. lineare has slender twigs that dry grayish brown, and the stone is separated from the base of the fruit by a fibrous zone 1-2 mm. long. Such a fibrous zone is much less well developed in but so are those of such specimens of C. pisiferum as Martin 1133 and Kerr 17774. Further collections of C. pisiferum and C. lineare from Cambodia and Thailand may lead to a reduction in rank of the latter, but such a reduction is at present premature. The type (and only) specimen of C. lineare cannot be satisfactorily accommodated in C. pisiferum. Calophyllum pisiferum is also related to C. dispar; for the differences between the two, see C. dispar. Calophyllum rupicola and C. tetrapterum are also somewhat similar to C. pisiferum; both these species are variable, and C. rupicola is also a rheophyte. The indumentum of both species is less prominent than that of C. pisiferum. Calophyllum rupicola often has denser venation, the latex canals are frequently (rather than rarely) impressed, the buds along the stem are often supra-axillary, the fruit is ovoid-elliptic, and the apex of the leaf blade is acute to acuminate. When dry the leaf is characteristically greenish-grayish brown above and golden brown below, rather than the sepia color of C. rupicola. The flowers and the fruits of C. tetrapterum are larger than those of C. pisiferum, and the upper internodes of the inflorescence axis are usually not markedly longer than the lower ones. The leaf blade generally dries a different color, and the margin is more or less recurved. Nitrasirirak 206 (probably C. tetrapterum, from S. Thailand) has the inflorescences of C. pisiferum. The specimens from Johore cited above are included with some hesitation; they were included in an unnamed variety of Calophyllum rupicola 4 KEP in other characters they agree with C. pisiferum, and the field notes attached to Corner s.n. suggest that the bark characters are similar to the trees of C. pisiferum seen in Sabah, although the plants are somewhat smaller (less than 10 m. tall). The specimen of Gaudichaud 86 at Geneva is designated as the lectotype 1980] STEVENS, CALOPHYLLUM 323 of Calophyllum pisiferum. Planchon and Triana cited another specimen, “Griffith [Kew dist. 876] in herb. Planchon, ex herb. Hook.,’’ also collected in Malacca. Of the two syntypes of C. retusum var. eee (Thorel 1395 and 1407), Thorel 1395 at Paris is designated as the synt The type specimens of Calophyllum motleyi and C. retusum var. ene ense are good matches with the type of C. pisiferum. The type specimens of C. retusum var. cambodgense and C. sangkae represent a rather long-leaved variant common in the Cambodia-Thailand area. 103. Calophyllum lineare Kostermans, Adansonia, II. 13: 333. pl. 1, 1, 2. 1973. Type: Cambodia, Kah Kong, Khbal Kah, 12 Feb. 1970, Martin 1783 (holotype, P; isotypes, L, P). Shrub 1.5 meters tall; bark not known. Twigs slightly flattened, 0.7-1 mm. across, 4-angled, drying grayish brown, subpruinose, puberulent when young; axillary innovations lacking basal scars; internodes 0.3-1.5 cm. long; uppermost pair of axillary buds rounded, ca. 7 mm. long, suberect; terminal bud conical, 1.8-2.4 mm. long, with short, tomentose, brown indumentum (hairs, Ficure 31, s), underdeveloped internode absent (to 1 mm. long). Petiole 2-2.5 mm. long, concave above and convex below, glabrous when mature; lamina narrowly elliptic, 1.8-6 by 0.18-0.6 cm., Cuneate to rounded at apex, narrowly cuneate at base, not undulate but slightly recurved at margin, coriaceous, drying umber to sepia above and below, glabrous when mature, the midrib above narrowing gradually from base, flat to slightly raised, 0.08-0.14 mm. wide at midpoint, below raised, + rounded, the venation above and below subapparent, slightly raised, 7 to 11 veins/5 mm., angle of divergence 45—50°. Infructescences from foliate axils, with scars of 3 to 7 flowers, unbranched, the axis 0.8-2 mm. long, sparsely tomentose toward base, lowest internode 3-7 mm. long; bracts unknown, their scars sometimes 2 mm. below pedicel they subtend; pedicels 1-1.5 cm. long, glabrous. Flower unknown; tepals probably 4 (from scars). Fruit broadly ovoid, 8-10 by ca. 7 mm., apiculate, drying russet, smooth, when submature obovoid and sharply wrinkled; outer layer not detaching cleanly from stone, ca. 0.3 mm. thick, disorganized by air spaces: stone ellipsoid, ca. 8 by 6.3 mm., rounded at apex, with fibrous zone ca. 2 mm long at base, the walls ca. 0.1 mm. thick, smooth, unmarked; spongy layer thin. DistriBuTION. Cambodia (Map 35); known only from the type locality. EcoLocy. Rheophyte with narrow leaves, among rocks at edge of river. Fruiting in February; fruit orange. Locat use. The fruit is used as bait for fish. Calophyllum lineare can be recognized by its narrowly elliptic, coriaceous leaf blades, short terminal bud, and small fruits borne on slender pedicels. The specific epithet /ineare alludes to the shape of the leaf blades. Calophyllum lineare is closely related to C. pisiferum; for the differences 524 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 between the two species, see C. pisiferum. It also shows a superficial similarity to the more or less linear-leaved specimens of C. nodosum from Brunei However, C. nodosum has a shorter terminal bud with much less conspicuous indumentum, axillary innovations with basal scars, and a midrib impressed on the lower surface of the lamina but raised and continuous with the leaf surface on the upper. Kostermans (/oc. cit.) noted that the lamina of Calophyllum lineare was densely and finely marked by pores on both surfaces. I have seen pores (stomata) only on the lower surface. 104. Calophyllum dispar P. F. Stevens, sp. nov. Figure 32, m-o. C. molle auct., non King; H. Keng, Gard. Bull. oe 28: 244. 1976; Corner, Gard. ae Singapore Suppl. 1: 104. 1978 A Calophyllo pisifero, quo aliter simile est, in gemma terminali (3.5—)4.5-7 mm. longa (versus 2-4.5 mm. longa), lamina 6-15 cm. longa et 1.8-5 cm. lata (versus (1.9-)3—7.5(-9.5) cm. longa et (0.9-)1.6—3.2(-4) cm. lata), et fructu strato exteriore (0.5—)I—-1.5 mm. crasso (versus 0.2-0.5 mm. crasso), differt. Tree 15-18 meters tall, d.b.h. to 30 cm.; trunk without buttresses; outer bark yellowish to grayish, smooth or with diamond-shaped fissures; inner bark pinkish to brown; latex yellow or yellowis Twigs slightly flattened, |.5-3 mm. across, 4- angled (with 4 additional raised lines), drying blackish, subpersistently brown-tomentose or gray-brown-— puberulent; axillary innovations sometimes with basal or subbasal scars; internodes 0.5-3.5 cm. long; uppermost pair of axillary buds rounded, ca. | mm. long, suberect, inconspicuous; terminal bud plump, (3.5-)4.5—-7 cm. long, with brown, tomentose to subappressed indumentum (hairs, Ficure 31, l, r), underdeveloped internode absent. Petiole 4-6 mm. long, broadly and deeply concave above and convex below, transiently to subpersistently puberulent to tomentose; lamina elliptic to ovate, 6-15 by 1.8-5 cm., obtuse to short-acuminate at apex, cuneate to broadly rounded at base, strongly and closely undulate but at most slightly recurved at margin, coriaceous, drying bay to umber above, often with grayish covering, and brick to umber below, transiently puberulent to persistently tomentose on midrib below (rarely angled), the venation apparent on both surfaces, raised, 5 to 9 (to 11) veins /5 mm., angle of divergence (65—)75-80°. Inflorescences from foliate axils along twigs, with 7 to 21 flowers, unbranched (rarely with 3-flowered branches ca. 3 mm. long), the axis 0.7-5.2 cm. long, puberulent to subtomentose apie toward apex), lowest internode 2-4 mm. long; bracts subovate, m. in fruit to 10 mm. long, usually obviously incrassate, to 2 mm. across. Flower (?)hermaphroditic; tepals 4, sometimes glabrous, narrowly ovate, 3-4 by |.5—2 mm., outer pair sometimes tomentose, inner pair sometimes with tuft of hairs at apex; ovary ca. | mm. long, the style 2.2-2.5 mm. long, the stigma 1980] STEVENS, CALOPHYLLUM 325 Ficure 32. a, b, Calophyllum incumbens: a, Kostermans 12813, terminal bud, x 6; b, BRUWN 327, fruit, longitudinal section, x 1.5. c,d, C. clemensorum (Clemens 35038), fruit, x 1.5: c, from outside; d, longitudinal section. e-g, C. banyengii (Stevens et al. 296). e, terminal bud, pte dart : f, longitudinal section; g, from outside. h-j, C. dioscurii: h, FRI 19292, habit, x 0.5; 1, FRI 19292, terminal bud, x 3; j, FRI 6773, fruit, longitudinal section, x 1.k, 1, C. pisiferum (Griffith s.n.): k, terminal bud, x 6; 1, fruit, longitudinal section, x 3. m-o, C. dispar (SFN 37715). m, n, fruit, x 3: m, from outside; n, longitudinal section. 0, terminal bud, x 3. 526 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 peltate, ca. 0.6 mm. across, obscurely radiate. Fruit spherical to broadly ellipsoid, 7-10 by 7-9 mm., apiculate, drying pruinose-brown, strongly wrin- kled; outer layer not detaching cleanly from stone, (0.5-)1-1.5 mm. thick, at first compact, air spaces developing under skin; stone spherical, 4.5—7.5 by 4.5-7 mm., rounded at apex, the walls 0.1—-0.2 mm. thick, smooth, unmarked; spongy layer thin. Type: Singapore, Mandai Road, 29 July 1940, SFN 37715 coll. Kiah (holotype, A; isotypes, BM, BO, KEP, SING). Disrripution. Malaya to Borneo, scattered, excluding Java (Map 35). ADDITIONAL SPECIMENS SEEN. Malaya. TRENGGANU: 55 m. [88 km.] S. Kuantan toward K. Trengganu, Phytochem. Survey Malaya S05 (1, sinc). PAHANG: Rompin, Bukit Serdang State Land, KEP 29864 (ker). Singapore. W. end of Seletar Reservoir, upper Mandai, SFN 39698 (po, F, kK, sING), Sinclair 7727 (©), 15 m., Stevens et al. 716 (a). Sumatra and adjacent islands. SELATAN: Lematang Ilr, 75 m., T 3 P 894 (T[horenaar| 894) (po, L, LAE, P), T 3 P 859 (T [horenaar] 859) (Bo). Barat: Solok, Loeboek Soelasik, 1150 m., bb 6544 (Bo). Atsen: Is. Simaloer, Landschap Tapah (Defajan), Achmad 1453 (Bo, L), 1664 (Bo, L), 1751 (Bo, L). Borneo. Saban. Keningau: Mile 231/2, Tulid Road, Sook Plain, SA N 55545 (san). KALIMANTAN. Timur: peak of Balikpapan, G. Beratus, 760 m., Kostermans 7592 (a, BO, K, L, LAE, P, SING). Selatan: Bandjermasin, Dachlan 20b (Bo). Sine loco (?Sumatra), Buwalda 6751 (Bo, Kc) Ecotocy. In swamp forest (SFN 37715); on sandy, blackish soil (SAN 5545); on sandstone (Kostermans 7592), 15-1150 m. alt. Flowering July, October, and November; fruiting February, July, August, and November. Calophyllum dispar can be recognized by its plump, usually tomentose terminal bud; its slightly flattened, strongly angled, and often tomentose twigs; and its medium-sized lamina with a short petiole and with the venation on both surfaces clear and relatively distant. The inflorescence axis 1s strongly four-angled, and the flowers have four tepals. The ripe fruits are small (1 cm. or less long) with the outer layer relatively very thick ((0.5—)1-1.5 mm.). The epithet dispar was chosen to emphasize the disparity in size between the medium-sized leaf blades and the small fruits. The closest relative of Calophyllum dispar is C. pisiferum, the two species being separable by the characters given in the diagnosis. Both C. pisiferum (at least in Malesia) and C. dispar (at least in Singapore) may grow in swamp forest, and apart from the characters given above, the two are very similar. More collections of both species in Malesia are needed to understand the relationship between them—it is possible that the two taxa are only varietally distinct. Of the specimens of Calophyllum dispar cited above, those from Malaya (Trengganu), Singapore, and Borneo have similar tomentose indumentum on twigs, midrib of the lamina (often a few hairs persist on the lower surface of the lamina), and at least the lower part of the inflorescence axis. Specimens from Sumatra and Malaya (Pahang) have less well-developed, more or less 1980] STEVENS, CALOPHYLLUM 527 puberulent indumentum, and there is also a tendency for the midrib to narrow more gradually from the base of the blade. Variation in hair type is considerable (Ficure 31, 1, r); some hairs in the tomentose form have only a few cells, yet are similar in size to the more common hairs with many more cells. SFN 37715 was included in Calophyllum molle by Henderson and Wyatt- Smith (1956) and subsequent authors, albeit with considerable hesitation. Although the distribution of indumentum on the vegetative parts of C. molle and C. dispar is similar, they differ in many other characters (the ovary of C. molle is densely pubescent, the leaf blade has much closer venation, and the fruits are very much larger). Calophyllum molle and C. dispar are not at all closely related taxa. 105. Calophyllum incumbens P. F. Stevens, sp. nov. Ficure 32, a, b. A speciebus aliis Calophylli in lamina (per)coriacea elliptica-obovata margine plerumque valde undulata in siccitate saepe supra olivacea infra mellea-cin- namomea, inflorescentiis axillaribus indumento subpuberulo praeditis, floribus cum 8 tepalis paribus exterioribus dorsalibus puberulis, et fructu strato exteriore plerumque I-1.8 mm. crasso, differt. Tree 12-30 meters tall, sometimes leaning, d.b.h. to 23 cm.; trunk ridged or with spur roots; outer bark yellowish, gray to brown, or orange, yellow, gray, and fawn mottled, (slightly) fissured; inner bark orange- to red-brown; latex yellow, clear, sticky. Twigs slightly flattened, 1.5-3.2 mm. across, 4-angled, drying dark brown, transiently farinose-puberulent; axillary innovations lacking basal scars or with single pair near base; internodes (0.5-)1.5-5 cm. long; uppermost pair of axillary buds rounded, 1-1.8 mm. long, spreading; terminal bud plump, 3.6-6.5 mm. long, with brown, puberulent to subfurfuraceous indumentum (hairs, Ficure 34, a, b; cf. 22, k, 1), underdeveloped internode (absent to) 2-3.5 mm. long. Petiole 0.5-1.5 cm. long, broadly and deeply concave above and convex below, puberulent when young; lamina elliptic to obovate, (4.4—)5.5-16.5 by (1.6—)2.8-6 cm., shortly acuminate to acute at apex, cuneate to attenuate at base, strongly undulate and slightly recurved at margin, coriaceous to very coriaceous, drying olivaceous to sepia above and cinnamon to near honey below, farinose-puberulent on midrib below when young, the midrib above gradually narrowed from base, raised to almost flat, 0.15-0.4 mm. wide at midpoint, below raised, slightly striate, the venation apparent above and obscure to subapparent below, raised, latex canals above sometimes slightly impressed, 7 to 17 veins/5 mm., angle of divergence 65—80°. Inflores- cences from foliate axils, with 5 to 9 flowers, unbranched, the axis 0.8-4.2 cm. long, ee lowest internode 2-8 mm. long; bracts not known; pedicels (0.3-)0.6-2.2 cm. long, puberulent, usually thickened in fruit, to mm. thick. Flower ferent tepals (4 to) 8, the outer pair elliptic to ovate, 4-5 by 3-4 mm., farinose-puberulent on back, especially at base, the next pair broadly ovate to elliptic, 6-7 by 3.5-5.5 mm., the inner ones elliptic to lingulate, (4-)5.5-8 by (0.7-)1.5-3.5 mm.; stamens I15 to (?)250, the filaments to 4.5 mm. long, connate for up to 1.3 mm., the anthers suboblong, 528 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 0.6-1.1 mm. long, retuse at apex; ovary |.5-2 mm. long, the style ca. 3.5 mm. long, the stigma peltate, ca. 0.8 mm. across, (?)2-radiate. Fruit ellipsoid to ovoid, 1.7-2 by 1.4-1.6 cm., apiculate, drying grayish brown, smooth, vinaceous-brown and smooth when young, strongly wrinkled when submature; outer layer detaching + cleanly from stone, (?0.5-)I-1.8 mm. thick, compact except for air spaces developing under skin; stone ellipsoid to subspherical, 0.9-1.35 by 0.7-1.3 cm., rounded at apex, the walls 0.15-0.6 mm. thick, smooth, unmarked; spongy layer (?)thin. Tyre: Borneo [Kalimantan], West Kutei, Mt. Palimasan near Tabang on Belajan River, 70 m., 9 Nov. 1956, Kostermans 12813 (holotype, L; isotypes, BO, CANB, K, KEP, L, NY, P, SING). DistRIBUTION. Sumatra and Borneo (Map 36). ADDITIONAL SPECIMENS SEEN. Sumatra. Barat: Bangkinang, Pedadit, 250 m., bb 23017 (go, L). AtseH: G. Leuser Nature Reserve, G. Bandahara, ca. 25 km. NNW. of Kutatjane, 1100 m., Wilde & Wilde-Duyfjes 12974 (KEP, L). Borneo. Bruner: Bukit Suang, ulu Batu Apoi, 610 m., BRUN 327 (xo, BRI, KEP, L, SAR, SING); R. aoe -ulu R. Belait watershed, 57 m., BRUN 158 (Bo, RI, KEP, L, SAR, SING). SABAH. Penampang: Sun Suran Trail, 320 m 37777 (san). Keningau: Mile 5 Rashna Road, Nabawan, SA N 83875 (a). Tawau: Vee ae abe £120 100 110 Map 36. Distribution of Calophyllum dioscurit (squares, localized specimens; large triangles, incompletely localized specimens), C. costulatum (small triangles), C. aff. senna (half-circle), C. banyengii (stars), and C. incumbens (circles) in Malesi 1980] STEVENS, CALOPHYLLUM 529 Brassey Range, 770 m., Stevens et al. 480 (A). KALIMANTAN. Timur: W. Kutei, Mt. Palimasan near Tabang on Belajan R., 600 m., Kostermans 13037 (Bo, CANB, K, KEP, L, NY, P, SING). EcoLocy. Usually in colline forest, (57-)320-1100 m. alt. In Kalimantan, locally abundant in Agathis forest on sandy, acid, waterlogged soil; in Penampang, Sabah, in primary (sic) forest on steep hillside with much lapotang (Gleichenia linearis C. B. Clarke); in Tawau, small tree on ridge with several other species of Calophyllum. Flowering June and December; submature fruit August and September (fruit white (Kostermans 13037) or greenish (SAN 83875)). Calophyllum incumbens is a rather variable species that nevertheless can readily be recognized by its somewhat short, plump terminal bud and its elliptic-obovate, often very coriaceous and rigid leaf blades that are strongly undulate at the margins and that often dry bicolored (olive above and honey to cinnamon below). The inflorescence axis, pedicels, and at least the bases of the backs of the outer pair of tepals are covered by farinose-puberulent leaning tree—hence the specific epithet (from incumbo, Calophyllum incumbens is superficially similar to C. depressinervosum: the leaves of the two species dry a similar color, but in C. incumbens the venation on the upper surface of the lamina is raised rather than more or less depressed, the axillary innovations lack basal scars, the inflorescence axis is puberulent rather than glabrous, and the outer layer of the fruit is much thicker. Calophyllum incumbens 1s similar to C. rupicola in color of the dried specimens and in indumentum distribution, but it is not a rheophyte, the leaves are larger and often thicker, the flowers have eight tepals, and the fruits are larger and have a much thicker outer layer. The two specimens collected by Kostermans from Kalimantan are the most robust and have thicker leaf blades than the others. The specimens from Sabah are somewhat less robust and are intermediate between the specimens from Kalimantan and those from Brunei and Sumatra. Only two specimens with flowers are known, Wilde & Wilde-Duyfjes 12974 and SAN 37777; the latter specimen has more stamens and smaller anthers. Wilde & Wilde-Duyfjes 12974 has vinaceous-brown immature fruits with an outer layer perhaps only 0.5 mm. thick; ripe fruits from Sumatra are needed to confirm the occurrence of the species there. FB 10650, a sterile specimen from the Philippines, dries a similar color to specimens of Calophyllum incumbens, but it has branched hairs and probably belongs to the C. blancoi complex, which characteristically has hairs of this type. Immature fruits of Calophyllum incumbens are smooth and dry vinaceous- brown. When submature they are wrinkled, but when mature they become smooth again, with air spaces just under the brittle skin. 530 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 106. Calophyllum dioscurii P. F. Stevens, sp. nov. Ficure 32, h-j. C. aaa Hooker f. Fl. Brit. India 1: 272. 1874, quoad Maingay, Kew dis ee sp. I. H. Burkill & Haniff, Gard. Bull. Straits Settl. 6: 174. 1930; Calophyllum sp. 45, M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 351. p/. 31. 1956; T. C. Whitmore, Tree Fl. Malaya 2: ere Eevee A speciebus aliis Calophylli in ramulo in siccitate albescenti et petiolo nigrescenti, lamina 3—9(-10.1) cm. longa apice valde acuminata nervis laterali- bus (sub)densis elevatis, et fructu mediocri strato exteriore |—1.8 mm. crasso, differt Tree 15-33 meters tall, d.b.h. to 55 cm.; trunk probably unbuttressed; outer bark grayish yellow, gray, or brown, smooth, with few lenticels, or fissured, sometimes with large, thin scales; inner bark dark brown; latex yellow or dark red (FR/ 20070), clear. Twigs slightly flattened, 1.2-1.6 mm. across, obscurely 4-angled, drying whitish to pale yellowish, transiently puberulent; axillary innovations lacking basal scars; internodes 0.5-3(-6.5) cm. long; uppermost pair of axillary buds + acute, to 1.5 mm. long, erect; terminal bud plump to conical, 2.5-5.5 mm. long, with reddish brown, puberulent indumentum (hairs, Ficure 34, c-g; cf. 34, a), underdeveloped internode absent. Petiole 3.5-6 mm. long, slender, broadly concave above and convex below, glabrescent, drying black; lamina elliptic to ovate (rarely suboblong), 3—9(-10.1) by |.2-3.4cm., acuminate (caudate) at apex, cuneate (rarely acute) at base, strongly undulate but not recurved at margin, coriaceous, drying olivaceous to bay above [sometimes shiny] and olivaceous to sepia below, subpersistently puberulent to farinose on midrib below (also above), the midrib above gradually narrowed from base, raised (depressed at first), 0.15-0.25(-0.4) mm. wide at midpoint, below raised, + rounded, the venation apparent on both surfaces, raised, [7 to] 12 to 20 (to 28) veins /5 mm., angle of divergence (55—)70-80°. Inflorescences from foliate axils, with 5 to 7 flowers, unbranched, the axis 4-9 mm. long, puberulent, lowest internode 0.5-3 mm. long; bracts not known; pedicels 4.5-6 mm. long, subpersistently puberulent, in fruit to 2 mm. thick. Flowers known only when old, (?)hermaphroditic; tepals (?)4; stamens (?)numerous, the filaments to 3 mm. long, the anthers oblong-elliptic, ca. 0.5 mm. long, retuse at apex; ovary 1-1.3 mm. long, the style ca. 2 mm. long, the stigma subpeltate, ca. 0.4mm. across. Fruit spherical, 1.2-1.6 by 1.2—-1.5 cm., rounded to apiculate at apex, drying pale pruinose-brown, deeply wrinkled when young, + smooth when older; outer layer not detaching cleanly from stone, 1-1.8 mm. thick, compact when young, large air spaces developing when older; stone spherical to ellipsoid, 0.9-1.1 by ca. 0.9 cm., rounded at apex, the walls 0.3-0.5 mm. thick, smooth, unmarked; spongy layer thin. Type: Malaya, Kedah, Ulu Muda F.R., 1000 feet [305 m.], 23 June 1969, FRI 11719 coll. Bray (holotype, a; isotypes, K, KEP, L, SAR, SING). 1980] STEVENS, CALOPHYLLUM eyes! DistripuTIoN. Malay Peninsula, Sumatra, perhaps also in Java and eastern Borneo (Map 36). ADDITIONAL SPECIMENS SEEN. Malaya. Kepan: Ulu Mudah F.R., 457 m., FRI OF 3 (Keke ee L, SAR), 120 m., FRI 11763 (kK, KEP, L); Bigia Enggang F. Ru 15 ae! OEP 73770: ay Sungkap F.R., KEP 59624 (kep). PERAK: Bintang Hijan F.R., KEP 39154 (x). Near SEMBILAN: Port Dickson, Sungei Menyala ER: a 103 (KEP), 45 m., KEP 64079 (ker), 30 m., KEP 66585 (KEP, sinc); Pasoh F ., anon, S.Nn., 27 Aug. 1974 (KEP). MALAcca: sine loco, Maingay 1644 (Ken ae 172) (kK). Pananc: Ulu Bertram at Kuala Mensum, 549 m., FRI 20070 (k, KEP, SAR); Lesong F.R., near Sungei Jekatih, ‘‘low hill,”’ FRI 15995 (a, K, KEP); King George Vth Natl. Park, 180 m., KEP 71956 (KEP); Kuala Lompat, 45 m., Chivers DCL 45 (kep), 406 (KEP); Gali, Raub, SFN 16220 (SING). JOHORE: Kluang, SFN 9424 (sinc). Sumatra and adjacent islands. Barat: Ond. afd. Oud Agam, Sipisang, 660 m., bb 2872 (Bo, L). Nan Angkola en Sipirok Panobasan, Dk. Poehoetan Lajan, 600 m., bb 31537 (a, Bo, Ny, sinc); Sibolga, Laboean Talang, P. Poenei (P. Morsala), ee is PD: 19292 (A, BO, L); Is. Morsala, Bukit Muara Talang, 40 m., Kostermans’ collector 24 (A, BO, K, L). ATJEH: Simaloer, Achmad 391 (?) (Bo), 517 (2) (Bo, L, U); 601 (?) (Bo, L, u); Asahan, Simpang Toba, 10 m., bb 8442 (no). CS prope Djeboes, Teysmann, “HB 3222 (?) (Bo). Borneo. KALIMANTAN. Tim Boeloengan, Kabiran, S. Simendoeroet, 200 m., bb 11760 (Bo). Java. see Passier Toegoe, Kikoehkoeran, Hasskarl 392 (L ). Ecotocy. Usually in colline mixed dipterocarp forest, 10-660 m. alt. Flowering in January; fruiting November (?ripe), January, February, April, and June (fruit green, finely mottled with yellow, or ‘“‘bronzed’’ (FRI 20070): reported to be eaten by siamang (Chivers DCL 406; Chivers, 1974)). Loca uses. In Pahang, Malaya, it is used as a medicine: ‘‘Sidauwai puteh [used] after child[birth] with a decoction of the root’? (SFN 16220). In Bangka the bast is used in atap thatching (as a tie?). Calophyllum dioscurii is a distinctive species that is easily recognized, even when sterile, by its small, prominently acuminate leaf blades that dry pale brownish green and have very dense, fine venation, and by its pale-drying twigs that contrast strongly with the dark-drying petioles. In addition, the internodes tend to be rather short (usually less than 3 cm.); the pedicels are short; and air spaces develop in the outer layer of the fruit. The epithet is taken from Dioscuri, the name given to Castor and Pollux, inseparable twins of classical mythology, in commemoration of the work of M. R Henderson and J. Wyatt-Smith on the genus. Henderson and Wyatt-Smith (/oc. cit.) compared this species with Calo- Phyllum gracillimum. However, in that species the venation is obscure, the indumentum is much better developed, and there are hairs on the an- thers; in fruit the two are not similar. They also thought that C. floribundum (= C. tetrapterum var. tetrapterum) was close to C. dioscurii. However, there are numerous differences in vegetative and fruit characters (venation density, twig color and type, thickness of the outer and woody layers of the fruit), and the two species are not closely related. ao JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 61 The specimens cited above that were collected from Simaloer, Sumatra, differ from the others in having a thicker lamina that is acute to acuminate at the apex, dries more or less brown, and has less dense venation; they also have stouter petioles. The characters in which they differ from the other specimens are enclosed in brackets in the above description. Hasskarl 302, from Java, is a sterile specimen, perhaps collected from a young tree; it has a lamina up to 12 by 3.1 cm 107. Calophyllum banyengii P. F. Stevens, sp. nov. A speciebus aliis Calophylli in ramulo tenui 0.8-1.2 mm. in transverso in siccitate flavido, petiolo tenui in siccitate nigrescenti, lamina chartacea mediocra valde acuminata (lamina 4.8-10.5 cm. longa acumine 1-1.5 cm. longo) margine valde undulata, fructu late ovoideo putamine tenuissimo (minus uam 0.1 mm. crasso), differt. Tree 8-12 meters tall, d.b.h. to 10 cm.; trunk without buttresses; outer bark yellowish, smooth, the inner surface straw brown; under bark reddish; inner bark pale red; latex yellow, (?)clear, sparse, sticky. Twigs flattened, 0.8-1.2 mm. across, rounded or with 4 raised lines, drying yellowish to pale brown, subpersistently brown-farinose; axillary innovations lacking basal scars (with pair of scars ca. 5 from base); internodes 1-7 cm. long; uppermost pair of axillary ae eabiene to 3 mm. long, erect; terminal bud narrowly conical, 3.5—4(-5.5) mm. long, with + appressed, brown indumentum (hairs, Ficure 34, h-j; cf. 34, c), underdeveloped internode absent (to 4 mm. long). Petiole 0.7-1.8 cm. long, + deeply concave above and convex below, subglabrous, drying blackish; lamina elliptic-oblong, 4.8-10.5 by 2.2-3.9 cm., acuminate at apex, acumen slender and to 1.5 cm. long, acute at base, strongly and closely undulate and slightly recurved at margin, thinly coriaceous, drying dark olivaceous-buff above and olivaceous below, subpersistently farinose on midrib above, the midrib above gradually narrowed from already narrow base, raised, 0.1-0.15 mm. wide at midpoint, below raised, striate to obscurely angled, the venation subobscure on both surfaces, slightly raised, latex canals impressed below, 8 to 14 veins/5 mm., angle of divergence 70-75°. Inflorescences from foliate axils along twigs, with 3 to 7 flowers, unbranched, the axis (0.1-)0.6-2 cm. long, puberulent, lowest internode (1-)4-7 mm. long; bracts subovate, to 3 mm. long, deciduous (subpersistent); pedicels (2-)5-12 mm. long, sparsely puberulent. Flower known only in bud, Carga cia tepals (?6 or) 8, the outer pair ovate, 1.6-3 by 1.4-2.2 mm., + puberulent on back, the inner ones persisting at the base of fruit, obovate, ca. 5 by 2.3 mm.; stamens 140 to 175, the filaments to 4.5 mm. long, the anthers oblong, ca. 0.6 mm. long, subretuse at apex. Submature fruit broadly ellipsoid to ovoid, ca. 1.2 by 1 cm., drying pruinose- brown, obscurely striate; outer layer not detaching cleanly from stone, ca. 0.8 mm. thick, compact; stone subspherical, ca. 1 by 0.85 cm., rounded at apex, the walls less than 0.1 mm. thick, smooth, (?)unmarked; spongy layer at first thick. 1980] STEVENS, CALOPHYLLUM 533 Type: Sarawak, Ist Division, Gunong Santubong, 54 m., 20 Dec. 1975, Stevens et al. 296 (holotype, a; isotype, SAR). DistripuTIoN. Northern Borneo, scattered (Map 36). ADDITIONAL SPECIMENS SEEN: Borneo. Sarawak. 4th Division: Miri, proposed Lambir Natl. Park, S 25074 (a, L, SAN, SAR, SING). SABAH. Kinabatangan: Brassey Range, 660 m., Stevens et al. 578 (?) (a). Ecotocy. On Gunong Santubong, in lowland dipterocarp forest, ca. 54 m. alt. Well-developed flower buds and submature fruits in December. Calophyllum banyengii can be recognized by its slender, yellowish-drying twigs; its slender, blackish-drying petioles; and its medium-sized lamina with a strongly acuminate apex and closely undulate margins. Its flowers have about eight tepals, and its ovoid fruits have a rather thin, but compact, outer layer and a very thin-walled stone. The epithet commemorates Banyeng ap Nudong, who found the tree from which the type collection was made beside the much-traveled track up Gunong Santubong. Calophyllum banyengii is at least superficially similar to C. dioscurii: the terminal buds of the species are similar, as are the yellowish-drying twigs and the acuminate leaves. However, C. dioscurii has smaller, more coriaceous leaf blades that are more shortly and less abruptly acuminate, stouter twigs, and shorter petioles. The fruits are smaller, large air spaces develop in the outer layer, and the stone has a perfectly distinct wall ca. 0.2 mm. thick. In some flowers of Calophyllum banyengii the outermost pair of tepals appears to be markedly smaller than the next pair. These outermost tepals are separated from the others by a short length of pedicel and appear to be bracts that lack axillary flowers. Stevens et al. 478, from the Brassey Range in Sabah, is similar to the specimens from Sarawak except that the lamina is narrower (up to 11.8 by 3.1 cm.) and is gradually acuminate at the apex, and the petiole is at least 6 mm. long. There is a tendency for the veins on the upper surface of the lamina to be impressed. However, the fresh leaf had a notably undulate margin, as did the individual from which the type specimen was taken. 108. Calophyllum novoguineense Kanehira & Hatusima, Bot. Mag. Tokyo 56: 562. fig. 3. 1942; P. F. Stevens, Austral. Jour. Bot. 22: 409. 1974. Type: Dutch New Guinea [Irian Jaya] , Boemi, 40 km. inward of Nabire, 300 m., 10 March 1940, Kanehira & Hatusima 12701 (isotypes, A, BO). C. warenense Kanehira & Hatusima, Bot. Mag. Tokyo 56: 564. fig. 5 1942; P. F. Stevens, Austral. Jour. Bot. 22: 409. 1974. Type: Dutch New Guinea [Irian Jaya], Waren, 96 km. south of Manokwari, 2 m., 23 March 1940, Kanehira & Hatusima 13708 (isotypes, A, BO). Tree 15-20 meters tall, d.b.h. to 20 cm.; outer bark yellow-brown, smooth except for fine vertical cracks, the inner surface dull straw-brown,; under bark red to deep red; inner bark red; latex cloudy, not sticky, white or 534 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 becoming white when rubbed (details from Papuan specimens). Twigs slightly flattened, 0.4-1.2 mm. across, 4-angled, drying brown (yellowish when old), with sparse, + adpressed hairs when young; axillary innovations lacking basal scars; internodes (0.2-)0.5-2 cm. long; uppermost pair of axillary buds rounded, 0.7-1.2 mm. long, suberect: terminal bud subconical, 1-2 mm. long, with brown, adpressed to spreading, puberulent indumentum (hairs, Figure 34, t, u; ef. 34, 1), underdeveloped internode to I mm. long. Petiole 1-4 mm. long, broadly concave above, convex below, + glabrescent; lamina elliptic to subcuneiform or subobovate, 0.6—5 by 0.35-3 cm., subacute to rounded at apex, cuneate at base, somewhat undulate and recurved at margin, entire lamina often + boat shaped, coriaceous, drying brown-vinaceous to umber above and umber to sabelline-sepia below, with subadpressed to erect hairs on midrib below (on entire lower surface), the midrib above narrowing gradually from base, + raised, (0.07-)0.15-0.3 mm. wide at midpoint, sometimes disappearing up to 5 mm. below apex, below only slightly raised, striate, the venation above subobscure, below apparent, raised, 5 to 9 (to 11) veins/5 mm., angle of divergence 50-70°. Inflorescences from foliate axils, with 3 to 7 flowers, unbranched, the axis 0.2-2.5 cm. long, pubescent, especially toward base, sometimes glabrous above, lowest internode (2—)4—7 mm. long; bracts elliptic, ca. 1.5 mm. long, caducous; pedicels 3-15 mm. long, glabrous or sparsely pubescent. Flower (?)hermaphroditic; tepals 4, glabrous or almost so, the outer pair suborbicular to ovate-elliptic, 2.5-3.2 by 2-2.5 mm., the inner pair subelliptic, 2-3.5 by 1.5-2.5 mm.; stamens 20 to 35, the filaments to 2 mm. long, the anthers elliptic to oblong, 0.4—1.5 mm. long, retuse at apex; ovary 0.4-1 mm. long, the style ca. | mm. long, the stigma peltate, 0.3-0.5 mm. across, + 3-radiate. Fruit spherical, 5-7 by 4.5-6 mm., apiculate, drying brown, + wrinkled: outer layer not detaching cleanly from stone, 0.3-0.5 mm. thick, becoming disorganized by air spaces; stone spherical to ellipsoid, 3.5-6.5 by 3-5.5 mm., rounded at apex, the walls less than 0.1 mm. thick, smooth, (?)unmarked: spongy layer thin. DistripuTioN. Western New Guinea (Map 45). ADDITIONAL SPECIMENS SEEN. Papuasia. IRiAN Jaya. Fakfak: Genofa, 750 m., bb 22569 (a, B, BO, L, SING). Djajapura: Cycloop Mts., Ifar, 400 m., van Royen & Sleumer 6592 (k, L). Papua New Guinea. Western: Kiunga Airstrip, 25 m., NGF 18302 (a, CANB, E, K, LAE, M, MO, SING), 20 km. from Kiunga on Rumginae road, 30 m., Stevens et al. 919 (a). Ecotocy. Varied habitats: NGF 18302 at edge of lowland freshwater swamp; Stevens et al. 919 in well-drained lowland forest; Kanehira & Hatusima 13708 in dry thicket by seashore; Kanehira & Hatusima 12701 in colline, A gathis- dominated forest; bb 22569 on chalk; 2-750 m. alt. Flowering March, April, and August; fruiting April, August (submature), and September. YOuNG PLANT. Young plants ca. 30 cm. tall have short (less than | cm.) internodes and narrowly elliptic leaf blades; the terminal bud is functional. and growth is erect. Innovations from the terminal bud lacked basal scars, while innovations from axillary buds had them. (Observations made on numerous plants beneath the tree from which Stevens et al. 919 was collected.) Taste 14. Differences between western Papuasian species of the Calophyllum novoguineense complex. C. novoguineense C. bicolor C. caudatum C.undulatum C. bifurcatum C. parvifolium TWIG THICKNESS (mm.) 0.4-1.2 1.5-2.5 0.7-1 Ca. | 1-1.4 0.7-1.2 TERMINAL BUD LENGTH (mm.) ]-2 4-7 2-3 1.5-3 3-5.5 1.8-3 FUNCTIONAL + + + + (?)- + LAMINA LENGTH (cm.) 0.6—-5 (2-)4-8(-13) PP 5.3-10.5 (4—)8-15 2.5-4 APEX SHAPE Subacute-— Acute-— Acuminate— Long- Long- Acute rounded acuminate caudate acuminate acuminate (rounded) BASE SHAPE Acute Rounded- Cuneate Acute Rounded Cordate acute MarGIN CLOSELY DULATE i _— = + . 150 | 160 ze ae | = { z Sey, 3 | J et | OE S| Pome , ; So % S50, ®. 2, =e | : e ° _ | - | Q | @ | | , | “5 * | | mse “. 4 = cn aye ° 1 ey | | { = | ere? % | | | °| ial. me 4 | a “e! | eo. Ba, i | e p38. Distribution of Calophyllum neo-ebudicum (circles; not including doubtful records from the Celebes) and C. caledonicum (squares) in the western Pacific. Inset 1: C. vitiense var. vitiense. Inset 2: C. cerasiferum (circles) and C. leucocarpum (star in solid circle). 550 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 of the inflorescence and flowers: in C. gracillumum there are hairs on the pedicels, the backs of the tepals, and the anthers. 116. Calophyllum articulatum P. F. Stevens, sp. nov. Ficure 36, f-t. A speciebus aliis Calophylli quibus laminis cordatis basibus habent in innovationibus basibus tumidis cum ramulis articulatis, fructu strato exteriore tenui (circa 0.6 mm. crasso), et putamine haud signato, differt. Tree 8-25 meters tall, d.b.h. to 40 cm.; outer bark yellowish brown (grayish), cracked at first, becoming fissured; inner bark pale reddish to brown; latex clea r. Twigs flattened, 2-3.5 mm. across, obscurely 4- or 6-angled, drying pale to dark brown, subtomentose when young; axillary innovations lacking basal scars, but swollen at base; internodes (0.5—)l-3.5 cm. long; uppermost pair of axillary buds rounded, less than | mm. long, inconspicuous; terminal bud plump, 2.5-3 mm. long, with subtomentose, brown indumentum (hairs, Ficure 35, b-d), underdeveloped internode absent. Petiole 2-3.5 mm. long, + flat above and convex below, with few subpersistent hairs; lamina elliptic or obovate to subtrapeziform, 2.3-7(-11) by 0.9-3.8(-6.5) cm., + rounded at apex, cordate to narrowly rounded at base, broadly and distantly undulate but not recurved at margin, very coriaceous, drying chestnut above (midrib and margin notably paler) and umber below, glabrous at maturity, the midrib above gradually narrowed from base, almost flat to raised, surrounding lamina + raised, 0.3-0.6 mm. wide at midpoint, below slightly raised, ea the venation subobscure above and subapparent below, raised, 4 to 7 (to 9) veins /5 mm., angle of divergence 40-55°. Infructescences from foliate axils, with scars of (?5 to) 7 to 9 flowers, unbranched, the axis 2.4-6.2 cm. long, glabrous, lowest internode |.2—3.2 cm. long; bracts not known; pedicels 0.6-1.2 cm. long, glabrous. Flower not known. Fruit ellipsoid, ca. 1.15 by 0.9 cm., apiculate when young, becoming rounded, drying pruinose-brown, smooth; outer layer not detaching cleanly from stone, ca. 0.6 mm. thick, air spaces developing beneath skin; stone ellipsoid, ca. 9 by 7.5 mm., rounded at apex, the walls ca. 0.15 mm. thick, smooth, unmarked; spongy layer thin. Type: Seram (West), Waiselang, 100 m., 23 June 1959, Kuswata & Soepad- mo 219 (holotype, Bo; isotypes, A, CANB, K, KEP, L, LAE, NY, P, SING). DistriBuTION. The Moluccas (Map 39), ADDITIONAL SPECIMENS SEEN. Moluccas. AMBon: Waai, slope of Mt. Salahatu, 300-600 m., Kuswata & Soepadmo 301 (Bo, 1); Mt. Salhoetoe, Teysmann S.n. (BO). Ecotocy. Colline forest, 100-600 m. alt. Ripe fruit reported to be blue. alophyllum articulatum can be readily recognized by its very coriaceous leaf blades that are rounded to cordate at the base, its partly enclosed axillary buds, and its articulations where the axillary branches join the stem. The epithet articulatum (‘‘jointed’’) was chosen because of this last feature. 1980] STEVENS, CALOPHYLLUM 551 129 (bb pee ee bud, » longitudinal section; Ficure 36. a, Calophyllum sp. bu d: h, axillary bud, x 6; 1, leaf base, & Hatusima 13980). j, k, fruit, 13 l, terminal bud, x 6. m-o, Calo phylum sp. m, terminal bud, x 6. n, 0, fruit, X 1: n, transverse section; , longitudinal Seclio n; k, from outside. 121 Merrill’s collector 142). o, from outside. doz JOURNAL OF THE ARNOLD ARBORETUM [voc. 61 Sho ea : y S Map 39. Distribution of Calophyllum archipelagi (stars), Calophyllum sp. 129 (erect triangles), C. articulatum (erect half-circles), C. savannarum (inverted half-circles), C. laticostatum (inverted triangles), C. carrii var. carrii (solid circles), and C. carrii var. longigemmatum (open circles) in Malesia. Inset: Calophyllum sp. 121. Ae ie ot hs Calophyllum articulatum differs from both C. venulosum and C. calcicola, which may have at least superficially similar leaves, in having an inflorescence with more flowers (in both of the latter species, there are rarely more than five). The lowest internode of the inflorescence is about half the length of the entire axis and is much longer than the upper internodes (in the other two species the relationship is reversed). The swollen bases of the innovations, which then become noticeably constricted where they join higher-order branches, and the partly enclosed axillary buds, distinguish sterile material of C. articulatum from that of the other two species. Unlike C. venulosum, but like C. calcicola, the outer layer of the fruit is thin; however, C. calcicola has an inconspicuous submarginal vein that is lacking in C. articulatum. The color of the dried leaf blade (especially in the type specimen), the articulations at the bases of the innovations, the partly enclosed axillary buds, and the midrib type suggest that Calophyllum articulatum is closely related to C. savannarum. However, the functional terminal bud, the cordate base of the leaf blade, and the inflorescence type of C. articulatum all readily distinguish the two species Teysmann s.n. has leaf blades that are up to II by 6.5 cm. and are broadly cordate at the base. 117. Calophyllum savannarum A. C. Smith, Jour. Arnold Arb. 22: 352. 1941; Stevens, Austral. Jour. Bot. 22: 394. 1974. Type: Netherlands New Guinea [Irian Jaya], Hollandia [Djajapura] and vicinity, 20-100 m., 29 June 1938, Brass 8888 (holotype, 4; isotypes, BO, BRI, K, L, LAE). Ficure 4, b-d. 1980] STEVENS, CALOPHYLLUM 550 Tree 10-12 meters tall, d.b.h. to 19 cm.; trunk and bark not known. Crown with hanging branches. Twigs strongly flattened, 1-2.5 mm. across, 2-angled or rounded, drying (pale) brown, glabrous when mature; axillary innovations lacking basal scars, 5 ie of axillary buds rounded, less than | mm. long, spreading, inconspicuous (hairs, Figure 35, e); terminal bud not seen, apparently not functional and soon falling off, underdeveloped internode 3-7(-13) mm. long. Petiole 0.7- 1.5(-2) cm. long, slightly concave above, convex below, glabrous; lamina ovate to elliptic, (5.5—)7-12.5 by (2—)2.3-4.5 cm., cuneate to bluntly acuminate at apex, acute to decurrent at base, moderately and distantly undulate and slightly recurved at margin, coriaceous, drying chestnut to sepia above, shiny, midrib and margins (veins) notably paler, and umber to sepia below, glabrous, the midrib above narrowed gradually from base, raised, continuous with surrounding blade, 0.3-0.6 mm. wide at midpoint, below slightly raised, angled, venation apparent on both surfaces, raised, 5 to 7 veins/5 mm., angle of divergence 45-60°. Inflorescences from foliate axils, with 5 to 9 flowers, unbranched, the axis 3-4 cm. long, glabrous, sola internode (0.3—)0.5-1.5 cm. long; bracts unknown; pedicels 1.1-1.7 cm. long, glabrous, somewhat incrassate in fruit. Flower (?)hermaphroditic; tepals 4, glabrous, the outer pair broadly ovate, ca. 5.5 by 5 mm., the inner pair + obovate, ca. 7 b 4 mm.; stamens 50 to 70, the filaments to 3 mm. long, + free, the anthers oblong, 2-2.7 mm. long, retuse at apex; ovary 1.2 mm. long, the style ca. 3 mm. long, the stigma peltate, ca. 1.5 mm. across, 3- to 5-radiate. Fruit ovoid, ca. 1.4 by 1.1 cm., rounded at apex, drying grayish olive, smooth; outer layer detaching cleanly from stone, 0.8 mm. thick, compact except for air spaces developing under skin; stone ovoid-ellipsoid, 1-1.1 by 0.6-0.7 cm., apiculate or rounded at apex, the walls ca. 0.15 mm. thick, smooth, unmarked; spongy layer thin. DistriBuTION. Northern Irian Jaya (Map 39). SELECTED SPECIMENS SEEN. Papuasia. IRIAN Jaya. Geelvink Bay: Seroei, Eil. Japen, Mariatoe, 500 m., bb 30477 (a, ae ay Hollandia [Djaja- pura], ‘“‘Dok 2,”’ BW 3915 (A, BO, K, L, LAE, NG). Eco.ocy. Secondary forests on steep slopes; principal component of forest clumps in secondary savanna; 20-500 m. alt. Flowering in August; fruiting June and August (fruit blue). Calophyllum savannarum is an easily recognized species, even when sterile, because of its nonfunctional terminal bud and consequent profuse branching, and its axillary shoots that are noticeably swollen at the base but lack the scars of perulae. The axillary buds, which are almost glabrous, are incon- spicuous due to the way they are protected (Ficure 4, b, c). Its leaf blades are coriaceous, dry dark brown (but with lighter-colored midrib, margins, and veins), and have rather distant, steeply ascending veins. The epithet savannarum (‘‘of the savannas’’) reflects the habitat of this species around Djajapura. 554 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Calophyllum savannarum is most closely related to C. articulatum. For the differences between the two species, see C. articulatum. 118. Calophyllum vitiense Turrill, Jour. Linn. Soc. Bot. 43: 17. 1915. Type: Fiji [Viti Levu, Mba], Nandarivatu, some three miles [4.8 km.] along the road to Suva, 2 Dec. 1906, im Thurn 297 (lectotype, kK; isolectotype, K). Tree 5-30 meters tall, d.b.h. to 50 cm.; trunk without buttresses; outer bark brown, slightly fissured or flaking, exposing greenish yellow patches; latex brownish, smelling strongly of green ginger; sapwood grayish brown; heartwood deep reddish brown with dark brown lines. Twigs slightly flattened, 1.7-4 mm. across, persistently 4-angled to 4-alate (with additional, less prominent raised lines, rarely coarsely striate), drying dark brown, farinose to subtomentose when young; axillary innovations lacking basal scars; internodes 0.5—3 cm. long; uppermost pair of axillary buds rounded, to 1.5 mm. long, + spreading; terminal bud plump (conical), (4.5—)7-12 mm. long, with brown to grayish, subcrustose to short-tomentose indumentum (hairs, Figure 35, f, g, k, 1; cf. 35, n), underdeveloped internode absent (to 3 mm. long). Petiole 0.5-3 cm. long, broadly and deeply concave above, convex below, glabrous when mature; lamina ovate to elliptic or oblong, (4-)11-21 by (1.2-)3-5.8 cm., shallowly retuse to acuminate at apex, acute to attenuate (rarely broadly rounded and then attenuate) at base, undulate and at most slightly recurved at margin, thinly coriaceous, drying sepia to grayish sepia above, margin and midrib paler, and umber below, sparsely farinose on midrib below when young, the midrib above gradually narrowed from base, depressed, clearly demarcated from lamina by often raised margins, 0.25-0.7 mm. wide at midpoint, center clearly sulcate, becoming raised toward apex, below raised, angled toward apex, striate toward base, the venation subobscure to apparent above and below, raised, (3 to) 5 to 9 (to 11) veins /5 mm., angle of divergence 65—80°. Inflorescences from foliate axils, with 5 to 13 (to ca. 21) flowers, unbranched, the axis (1.3-)4-12 cm. long, sparsely brown-farinose to tomentose toward base, or subglabrous, lowest internode (0.3—)0.7-2.2(-4.5) cm. long; bracts elliptic to ovate, 2.5-5 by 1.7-2 mm., caducous; pedicels (0.4-)1-3.2 cm. long, glabrous or sparsely farinose. Flower (?)hermaphroditic; tepals (7 or) 8 (to 12), the outer pair broadly ovate to suborbicular, 4.5-6 by 2.8-3.5 mm., the inner ones elliptic to oblong, 7.5-12.5 by 3.5-10 mm.; stamens 275 to 340, the filaments to 5.5 mm. long, the anthers oblong to elliptic, 0.5-1 mm. long, retuse at apex; ovary 2.5-2.8 mm. long, the style 2.7-S mm. long, the stigma peltate, 0.9-1.3 mm. across, 3- or 4-lobed. Fruit + ellipsoid, 2.2-3.3 by 1.8-2.2 cm., rounded at apex, drying dark Indian red, pruinose or not, + broadly wrinkled, the wrinkles sharp and deep in submature fruit, rounded and shallower in mature fruit; outer layer not detaching cleanly from stone, ca. 0.5(-1.5) mm. thick, + fibrous, with numerous air spaces; stone ellipsoid, 2—2.7 by 1.5—2 cm., rounded at apex, the walls 1-3 mm. thick, smooth, unmarked, with basal plug ca. 10 mm. across; spongy layer thin. 1980] STEVENS, CALOPHYLLUM 555 Key to the Varieties of Calophyllum vitiense 1. Terminal bud (6-)8-12 mm. long, usually with subcrustaceous indumentum; lamina acute to acuminate at apex........... 118a. var. vitiense. 1. Terminal bud 4.5-7 mm. long, usually short-tomentose; lamina shallowly retuse to rounded (acute) atapex........ 118b. var. amblyphyllum. 118a. Calophyllum vitiense Turrill var. vitiense C. vitiense Turrill; Watkins, Dept. Agr. Jour. Agr. Fiji 31: 15. fig. 1961; Parham, FI. Fiji Is. 134. 1964, ibid. ed. 2. 192. 1972; A. C. Smith & Darwin, Jour. Arnold Arb. 55: 227. figs. 9, 11. 1974, pro majore parte. C. tenuicrustosum A. C. Smith & Darwin, Jour. Arnold Arb. 55: 236. fig. 22. 1974, pro parte. Tyre: Fiji, Viti Levu, Mba, slopes of Mt. Nairosa, eastern flank of Mt. Evans range, 700-1050 m., 28 April 1948, A. C. Smith 4058 (holotype, a; isotypes, K, NY, P, US). C. spectabile auct., non Willd.; A. Gray, U. S. Expl. Exped. 15(1): 218. 1854, pro parte, Proc. Am. Acad. 5: 315. 1862, Bonplandia 10: 34. 1862; Seem. Fl. Vit. 11. 1865; Horne, Year Fiji, 258. C. (?)polyanthum auct., non Wall. ex Choisy vel C. (?lanceolatum auct., non BI.; Seem. Viti, 433. 1862. Tree 5-30 meters tall, d.b.h. to 50 cm. Twigs with brown, farinose to tomentellous indumentum when young; terminal bud plump, (6—)8-12 mm. long, with brown to grayish, crustaceous (rarely puberulo-tomentose) fadumeaninn: Lamina ovate to elliptic or subob- long, (4-)11-21 by (1.2-)3-5.8 cm., acute to acuminate at apex, acute to attenuate (very rarely rounded) at base. Inflorescence axis (2.5—)5-12 cm. long, glabrous or farinose at base; pedicels (0.4-)1.7-3.2 cm. long. DistripuTIoN. The Fiji Islands (Map 38). ELECTED SPECIMENS SEEN. Pacific Islands. Fut. bage Levu. Mba: vicinity of Nandarivatu, 900 m., Gillespie 4229 (a, BO, GH, K, NY, P, US); Nausori Highlands, Mt. Evans Range, 579 m., Damanu E118 os “Nesana Soa 457 m. C.S.I.R.O. S 1404/6 (k). Serua: Nabukavesi, . Mls CasthehkOy 15 1404/4 (k); Naboutini, 150 m., C.S.J.R.O. S 1404/15 (k ‘nee sine loco, FDA 14236 (a, CHR, K, NY). Tailevu: E. of Wa aa ka R., near Ndakuivuna, A.C. Smith 7201 (Gu, k, P, us); Colo [Tholo] -i-Suva, Watkins 713 (kK); Tamavua, Gillespie 2407 (Bo, GH, K, NY, P, US); Kalambo, FDA 16410 (mass); Naisinu, 150 m., Gillespie 3647 (BisH). Ovalau: oe Korotolutolu, W. of Thawathi, C. Sm ith 8021 (GH, Kk, By p, us); W. of Lovoni valley, ridge S. of Mt. Korolevu, A. C. Smith 2 (GH, K, NY, Pp, US); E. of Lovoni valley, A. C. Smith 7269 (GH, K, Ny, P, US). Wanua Levu. Mathuata: near Lambasa, K, us), Motuyagaikaveta, 90 m., CS.LR. oO. § 1404/2 (x); Ne Fi. Fiji. [FDA] 12910 (kx); Seaqaqa, coffee plantation, Fl. Fiji [FDA] 13935 (cnr, K, LAE). Sine loco: Howard 187 (us). Eco.tocy. Dense or open forest, rarely in grassland; 90-1050 m. alt. Flower- ing August, and October to January; fruiting mostly May to August (fruit purple). 556 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Locat use. The wood is used for general construction (A. C. Smith & Darwin, Joc. cit.). 118b. oe vitiense Turrill var. amblyphyllum (A. C. Smith & Darwin) . F. Stevens, comb. et stat. nov C. amblyphyllum A. C. Smith & Darwin, Jour. Arnold Arb. 55: 232. figs. 17, 18. 1974, pro minore parte. Type: Fyi, Viti Levu, Serua, hills west of Waivunu creek, between Ngaloa and oo 50-150 m., 23 Sept. 1953, A. C. Smith 9243 (holotype, US} isotypes K, NY, P). C. tenuicrustosum A. C. Smith & Darwin, i pine Arb. 55: 236. 1974, pro minore parte. Tree 9-15 meters tall. Twigs short-tomentose when young; terminal bud plump to conical, 4.5-7 mm. long, with short, tomentose (rarely subcrustaceous), brown indumentum, hairs to ca. 0.25 mm. long. Lamina oblong to subelliptic, (4.3-)6.5-13.6 by 2.7-4.6(-5.2) cm., minutely retuse to rounded (acute) at apex, acute at base, or toward base broadly rounded and at base short-attenuate. Inflorescence axis (1.5-)2.3-6.5 cm. long, short-tomentose toward base; pedicels 0.9-2 cm. long. DistRiBuTION. Fiji, Viti Levu. ADDITIONAL nee SEEN. Fiji. Vit1 Levu. Serua: near Taunovo Creek, E. of Wainiyambia, -50 m., A. C. Smith 9576 (GH, kK, NY, P, US); Queen’s road, littoral forest, ce 7036 (BISH). Rava: Qoyo F.R., 45 m., FDA 13764 (BISH); Goya, 30 m., Damanu E121 (k). Eco.ocy. Forests near coast, sometimes by sea; below 50 m. alt. Flowering in November. The epithet amblyphyllum means ‘blunt leaf,’’ so is appropriate for this taxon. Calophyllum vitiense is a fairly easily recognizable species, having charac- teristic leaf blades that dry sepia on the upper surface, with the margin and the depressed midrib brighter brown; the venation is rather distant and clear. The fruit has a thin (usually ca. 0.5 mm.) outer layer. The upper surface of the leaf blade is minutely bullate when viewed at a magnification of x 30 or greater; this is most clearly visible in the younger leaves. The epithet vitiense is derived from a Latinized form of ‘‘Fiji.”’ The description of the bark is taken from Watkins (/oc. cit.) and is probably that of Calophyllum vitiense var. vitiense. The specimens of Calophyllum amblyphyllum included in C. vitiense have the characteristic leaf type described above; however, the majority of the specimens included in this species by Smith and Darwin (loc. cit.) are here placed in C. cerasiferum and C. neo-ebudicum. Calophyllum vitiense vat. amblyphyllum differs from var. vitiense in having leaf blades that are more or less rounded at the apex, rather than acute to acuminate, and a short terminal bud that often, but not always(e.g., FDA 13764), has short, tomentose 1980] STEVENS, CALOPHYLLUM By, indumentum. The specimens cited under C. vitiense var. amblyphyllum were all collected at low altitudes, while var. vitiense is usually a plant of colline orest. There is substantial similarity between the large-leaved species of Calophyl- lum from Fiji and Samoa (C. vitiense, C. neo-ebudicum, C. cerasiferum, and perhaps also C. inophyllum). All have similar hairs, fruits with stones that have basal plugs, flowers with usually eight tepals, and internodes that are rather short for such large-leaved plants. Calophyllum vitiense has sepia-drying leaves with rather distant venation and a minutely bullate upper epidermis; C. neo-ebudicum has rather shiny leaves drying other than sepia and with dense venation drying a different color; C. cerasiferum has leaves drying dull sepia and with dense, concolorous venation. Calophyllum inophyl- lum is the only species of the group with leaves that are basically rounded at the apex, and it has rather distant venation. Calophyllum tenuicrustosum was characterized by its flowers, which have more than eight tepals, and by its fruits, which have a thin outer layer. It was compared with C. cerasiferum, in part probably because Howard 195, which was included in C. tenuicrustosum, is C. cerasiferum. Although the type specimen of C. tenuicrustosum has a rather narrowly elliptic leaf blade, it has all the other leaf characteristics of C. vitiense, including the minutely bullate upper surface. The thin outer layer of the fruit is also characteristic of C. vitiense. The fruits of C. tenuicrustosum are rather small for those of C. vitiense (in the lowest part of the size range for the species), but as the illustration in Smith and Darwin (Joc. cit.) shows, they are immature: they lack an embryo, and the woody layer is poorly organized. The fruits of FDA 7036 are somewhat more mature and have a basal plu It is not certain that Calophyllum vitiense grows on Vanua Levu; I have seen only the single unlocalized specimen cited above from there. The three complementary sheets of im Thurn 297 at Kew were designated the lectotype of Calophyllum vitiense by Smith & Darwin (loc. cit.); Turrill also cited a sterile specimen (Seemann 47) in the protolog. 119. Calophyllum neo-ebudicum Guillaumin, Jour. Arnold Arb. 12: 227. 1931; St. John & . Smith, Pacific Sci. 25: 326. 1971; A. C. Smith & Darwin, Jour. Arnold Arb. 55: 230. figs. 14-16. 1974. Type: Hebrides, Aneityum, Anelgauhat Bay, 25 m., 4 Feb. 1929, eee 705 (lectotype, p; isolectotypes, A, BO, BRI, NY, US). C. samoense Christophersen, Bishop Mus. Bull. 128: 147. fig. 20. 1935; Yuncker, Bishop Mus. Bull. 184: 52. 1943; Parham, New Zealand Dept. Arnold Arb. 55: 229. figs. 12, 13. 1974. Type: Samoa, Savaii, Vaipouli- Manase, cia 21 Sept. 1929, Christophersen 720 (holotype, BIsH (n.v.); isotypes, A, US). G. eRe P. F. Stevens, Austral. Jour. Bot. 22: 389. fig. 12. 1974. Tyre: Solomon Islands, Guadalcanal, Gold Ridge, 690 m., 16 Oct. 1962, BSIP 648 coll. Whitmore (holotype, LAE; isotype, SING C. spectabile auct., non Willd.; Planchon & Triana, Ann. Sci. Nat. Bot. 558 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 IV. 15: 266. 1862, guoad A badie 29; Drake, Illus. Fl. Insul. Maris Pacifici, fasc. 6. 116. 1890, pro parte; Drake, Fl. Polynesie Frang. 10. 1893; Hemsley, Jour. Linn. Soc. Bot. 30: 169. 1894; Reinecke, Bot. Jahrb. 25: 656. 1898; Denkschr. Akad. Wiss. Wien. Math.-Naturw. 81: 313. 1910; Lloyd & Aiken, Bull. Lloyd Libr. Bot. 4: 72. 1934; Burrows, Bishop Mus. Bull. 138: 136, 154. 1936; Yuncker, Bishop Mus. Bull. 184: 52. 1945 C. vitiense auct., non Turrill; Setchell, Carnegie Inst. Wash. Publ. 431: 69. 1924; A. C. Smith, Jour. Arnold Arb. 22: 347. 1941: F. S. Walker, Forests Brit. Solomon Is. Protect. 124. 1948; T. C. Whitmore, Guide Forests Brit. Solomon Is. 78. 1966, Gard. Bull. Singapore 22: 12. 1967: Sykes, New Zealand Dept. Sci. Industr. Res. Bull. 200: 100. fig. 9. 1970; Foreman, Check List Vasc. PI. ae oe 42, 86, 87. fig. 1972; . Smith & Darwin, Jour. Arnold Arb. 55: 227. fig. 10. 1974, pro Cc. "inophy tm auct., non L.; Guillaumin, Jour. Arnold Arb. 12: 227. C opin A. C. Smith & Darwin, Jour. Arnold Arb. 55: 232. fig. 19. 1974, pro minore parte C. leptocladum . c. Smith & Darwin, Jour. Arnold Arb. 55: 221. 1974, pro minore part Calophyllum sp. Burkill, Jour. Linn. Soc. Bot. 35: 27. 1901. Tree 6-58 meters tall, d.b.h. to 184 cm.; trunk rarely with buttresses to 1.3 meters tall; outer bark gray to yellowish mottled at first, with brown and pinkish patches intermingled, later brown, deeply fissured, not hoop marked, the inner surface blackish; under bark reddish; inner bark reddish, fibrous; latex yellow (rarely cream), clear, but tending to become turbid, sticky. Crown pyramidal when young, becoming cauliflower shaped with age. Twigs slightly flattened, 1.5-4 mm. across, obscurely 4-angled, drying brown to blackish, puberulent to short-tomentose when young; axillary innovations lacking basal scars; internodes 0.7—4 cm. long; uppermost pair of axillary buds rounded, to 2 mm. long, + erect; terminal bud plump (rarely conical), mm. long. Petiole 0.6-3.7 cm. long, broadly concave above, convex below, often drying blackish, glabrous when mature; lamina rather narrowly ovate to elliptic (rarely suboblong), (4-)5—19.5 by 1.5-7.3 cm., acuminate (obtuse) at apex, cuneate to narrowly acute or attenuate at base, undulate but slightly or not recurved at margin, coriaceous, drying olivaceous to umber above, often nitid, margin and midrib often paler, and fulvous to near dark olive-buff below, glabrous to subpersistently puberulent on midrib below, the midrib above gradually narrowed from base, depressed, margins usually raised, becoming raised toward or above midpoint, (0.1—)0.2—0.5(—0.8) mm. across at midpoint, below raised, angled toward apex, striate toward base, the venation + apparent above and below, raised, latex canals + raised or not, 7 to 14 veins /5 mm., angle of divergence 60-80°. Inflorescences from upper foliate axils (rarely terminal), with 7 to 17 flowers, unbranched (very rarely with 1980] STEVENS, CALOPHYLLUM Bok 3-flowered branches to | cm. long, or flabellate), the axis 1.5-9.2(-13) cm. long, puberulent or short-tomentose, especially toward base (glabrous above), lowest internode 0.4—3.3(-4.5) cm. long; bracts ovate, to 5 mm. long, not persistent; pedicels 0.7-2 cm. long, glabrous to puberulent. Flower (?)her- maphroditic; tepals 8 (to 12), the outer pair ovate to suborbicular, 2.5-5.5 by 2.7-4 mm., back sometimes puberulent toward base, the inner ones elliptic to obovate, 7-10 by 3.5-6 mm.; stamens 150 to 275, the filaments to 5.5 mm. long, the anthers elliptic or suboblong, 0.4-1.2 mm. long, retuse at apex; ovary 1.5-2.5(-3) mm. long, the style 2.5-3 mm. long, the stigma peltate or infundibular, 0.5-1 mm. across, 3- or 4-radiate or not. Fruit ovoid or ellipsoid to subspherical, 2.1-3.7(-4.3) by 1.5-2.8 cm., + rounded at apex, drying vinaceous-brown to blackish and pruinose, broadly and + deeply wrinkled to smooth; outer layer usually not detaching cleanly from stone, (0.5—)l-3 mm. thick, compact, usually with air spaces developing adjacent to skin and stone (throughout); stone ovoid or ellipsoid to subspherical, (1.5-)1.8-3.2(-4) by (1-)1.3-2.5 cm., rounded to obtusely pointed at apex, the walls (0.7—)1.3-2.1(-3.3) mm. thick, smooth, unmarked, with basal plug 6-10 mm. across; spongy layer thin. DistriBuTION. (?)Celebes, New Britain, and Bougainville, to the New He- brides, Fiji, Samoa, and Tonga (Map 38). SELECTED SPECIMENS SEEN. Celebes. SALasar: Lembang, Djampea, 245 m., bb 2947 (?) (Bo). Papuasia and adjacent islands: see C. rainy Uae Stevens, loc. cit. Pacific Islands. Fist. Moala: near Maloku, 200 m . C. Smith 1334 (BO, GH, K, NY, P, US). Naitamba: sine loco, FDF L 12384 oo Mango: sine loco, 50-100 m., Bryan 569 (a). Viti Levu. Mba: mountains near Lautoka, 360 m., Gieeniood 907 (a); Mt. Evans Range, 610 m., Greenwood 1232 (us). Sine loco, Graffe s.n. (Ny, w). Ovalau: sine loco, Horne 43 (x). Koro: main ee 300-500 m., A. C. Smith 1047 (Bo, GH, K, NY, US). Wanua Levu. Mathuata: . Numbiloa, E. of Lambasa, 500-590 m., A. C. Smith 6449 (a, k, Ny, P, an Thakaundrove: near Valeni, FDA 15722 (?) (BISH); sane of Navonu Creek, 305 m., Howard 104 (BisH, CHR, US). ToNGA. Vava’u, sine loco, Crosby 204 (kK); Kao, Yuncker 15883 (Bm, vu, us, w); ’Eua, near ee us peas 240 m., Yuncker 15326 (BISH, BM, US). HorNE ISLANDS AND SAM eCas nse, Smith & Darwin, loc. cit. Also, Samoa: Upolu, Ve ores “1399 (w); bei Tiaril, Rechinger 1317 (w), bei Utumapu, Rechinger 941 (w). Nive: Ana, near Hakapu-Liku road, Sykes 821A (sisH, CHR); Huvalu Forest, aa 399 (cHR). New Hesribes: see C. neo-ebudicum, Smith & Darwin, loc. Eco.Locy. Usually primary lowland to colline rainforest, to 825 m. alt. Often abundant, especially in Solomon Islands and New Hebrides, favoring ridges and other well-drained habitats. In New Hebrides common in Kauri (A gathis) forests; Agathis regenerates in gaps resulting from death of mature C. neo-ebudicum (Beveridge, 1975). Sometimes on coralline limestone in Fiji and the Horne Islands, rarely so elsewhere. Flowering in Solomon Islands February, April, May, July to December; elsewhere September to February (flower scented). Fruiting more or less year around in both areas (fruit pale green at first, turning bluish to purple-black when ripe). 560 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Details of the establishment of Calophyllum neo-ebudicum on Kolombangara (the Solomon Islands) are given by Whitmore (1974, as C. vitiense). The fruits are well dispersed, and the seedling grows to adult size in closed forest as well as in gaps. GERMINATION AND YOUNG PLANT. The radicle pushes out a basal plug during germination. The seedling has two (Sykes 823), three, or four pairs of leaves; when there are four pairs, some leaves are always very much reduced (1-4 mm. long) and soon fall off; the internodes are 1-2.2 cm. long. Subsequent growth is erect, the terminal bud is functional, and the internodes gradually become longer. (Sykes 823; LAE 50491.) LocaL NAME AND USES. ‘‘Gwarangwaro’’ (Kwara’ae). Through much of the range of the species, the timber is used for building canoes; more locally it is used for building houses or in making spears or bowls. Hair oil is made from the flowers (Fauro Islands). In Malaita small saplings, with the outer layer of the bark removed to give bright, saffron-colored sticks, are reported to have been used in bride purchase. Calophyllum neo-ebudicum can be recognized by its frequently dark-drying twigs and leaf stalks and by its ovate-elliptic lamina, the upper surface of which often dries somewhat nitid greenish and has a depressed midrib. The predominantly eight-tepaled flowers have small anthers and a short (less than 3 mm.) style. The spherical to ovoid-ellipsoid fruits dry more or less smooth and have a rather thick outer layer and a moderately thick stone; the stone has a basal plug. Indumentum on parts of the plants other than the terminal bud is usually inconspicuous. The specific epithet is derived froma Latinization of ‘‘New Hebrides.”’ The single specimen cited from the Celebes, bb 22947, may not belong here. The specimen is sterile, and fertile material, preferably in fruit, is needed to confirm the record. A number of specimens from throughout the range of the species have terminal inflorescences, but are not otherwise unusual. FDA 15722, cited by Smith and Darwin as C. leptocladum, has very small, coriaceous leaf a like C. neo-ebudicum. In vegetative characters BSP 8876 is like C. neo-ebudi- cum, but the thin, sharply wrinkled outer layer of the fruit is like that of C. inophyllum; fruit shape is intermediate between those of the two species. Greenwood 1232 has exceptionally thick-walled stones (measurements in parentheses); the specimen consists only of fallen leaves and fruit. Calophyllum pseudovitiense and C. neo-ebudicum cannot be separated since they have similar leaves, inflorescences, and flowers, and the anthers of both are small. Smith and Darwin (Joc. cit.) separated Calophyllum samoense from C. neo-ebudicum predominantly on fruit characters. Calophyllum neo-ebudicum had fruits that were definitely longer than broad and a firmly fibrous mesocarp 1-4 mm. thick that did not fall away from the endocarp (their terminology) 1980] STEVENS, CALOPHYLLUM 561 and leave air cavities, while C. samoense had ovoid or subglobose fruits and a fibrous-spongy mesocarp 3.5-6 mm. thick with prominent cavities. The fruits of C. neo-ebudicum figured by Smith and Darwin (from Kajewski 399) are immature; those of Kajewski 953, which are submature, have cavities immediately under the skin. The fruits of Bristol 2248 (originally determined as C. samoense) have a subcompact outer layer with cavities under the skin and adjacent to the stone, although in most other fruiting specimens of C. samoense the cavities have invaded the entire outer layer. All these specimens have ripe or nearly ripe fruits, and the development of cavities seems to be a part of the ripening process. However, both in C. neo-ebudicum from the Solomon Islands (e.g., BSIP 79; seed possibly not developing) and in specimens placed by Smith and Darwin (Joc. cit.) in C. amblyphyllum, but properly identified as C. neo-ebudicum (e.g., A. C. Smith 6449; nearly mature), the outer layer may be almost compact. Although the fruits of C. samoense tend to be broader than the others, this is not an absolute difference, specimens with similar fruits having been collected from the Solomon Islands (e.g., BSIP 6255, Ulawa Island). Of the Zea originally included in C. amblyphyllum but correctly placed here, A. C. Smith 6449 (Fiji) has fruits about 1!/4 times longer than broad, ie those of Yuncker 15583 (Tonga) are almost spherical. Fruits from Niue seem to be longer than broad (Sykes 821A). Specimens of all taxa discussed have stones with thick walls and a plug at the base that presumably becomes detached during germination, and the ripe fruits are reported to be (pruinose) dark purple, black, or sometimes brown. There are no vegetative differences between the specimens of the various taxa placed in synonymy, and in most details of flower and inflores- cence they are also similar. However, in specimens of C. neo-ebudicum the anthers are 0.4-0.8 mm. long, while in those referable to C. samoense they are (0.5-)0.7-1.2 mm. (Anther length is known from only a single Fijian collection (Graffe s.n., from Viti Levu); the anthers are ca. | mm. long.) Specimens in flower have not yet been collected in Niue. Smith and Darwin (loc. cit.) note that C. neo-ebudicum and C. samoense were reported to have differences (unspecified) in general aspect, bark, and wood, and that samoense grew on coralline limestone, while C. neo-ebudicum never did. However, on Samoa itself C. samoense does not seem to have been collected on limestone, and it certainly grows on other rocks Although specimens assignable to Calophyllum samoense show certain differences when compared to other specimens placed in C. neo-ebudicum, such as somewhat longer anthers, relatively broader fruits, and twigs that more frequently dry brown, there is overlap in these characters between specimens from Samoa, Tonga, and the Horne Islands and those from the rest of the range of the species. Furthermore, the length of the anthers is not known in some parts of the range of the species (see above). It thus seems unwise to recognize any infraspecific taxa within C. neo-ebudicum at present. Variation in hair type in Calophyllum neo-ebudicum is considerable. The variation in BSIP 429 alone bridges the gap between the large, erect hairs of Kajewski 2643 and the more normal, small, papillate, adaxially curved 562 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61] hairs of most other specimens, including those that have been called C. samoense (Ficure 35, h, n). I have not seen A badie 29, on which Planchon and Triana (loc. cit.) based their record of Calophyllum spectabile in the Society Islands, but it probably belongs here. It should be noted that Calophyllum neo-ebudicum must be lectotypified on the sheet of Kajewski 705 in the Museum d’Histoire Naturelle at Paris (cf. Smith & Darwin, Joc. cit.). Guillaumin worked on the set of Kajewski’s specimens deposited in the herbarium there. 120. Calophyllum cerasiferum Vesque, Epharmosis 2: ¢. 32. 1889, in C. DC. Monogr. Phanerog. 8: 585. 1893; Parham, Pl. Fiji Is. ed. 2. 192. 1972; A. C. Smith & Darwin, Jour. Arnold Arb. 55: 235. figs. 20, 21. 1974. Type: Fiji [Viti Levu, Namosi Province], Woma Peak, 24 August 1860, Seemann 49 (lectotype, G; isolectotypes, BM, G, GH, K, P). C. burmannii auct., non Wight; Seem. Bonplandia 9: 254. 1861, Viti, 433. 1862; Horne, Year Fiji, 258. 1881; Drake, Illus. Fl. Insul. Maris Pacifici, fasc. 6. 116. C. burmannii Wight var. parvifolium auct., non Wight; Seem. Fl. Vit. 11. 1865. Calophyllum sp. nov. Horne, Year Fiji, 258. 1881. C. amblyphyllum A. C. Smith & Darwin, Jour. Arnold Arb. 55: 232. 1974, pro parte. Tree 12.5-25 meters tall, d.b.h. to 60 m.; trunk (?)without buttresses; outer bark yellowish, rough to smooth, with wade vertical erodes (?fissures); inner bark red; latex yellow (A. C. Smith 4944), white (A. C. Smith 8378), or opaque (Howard 336). Twigs slightly flattened, (1.7-)2.5-4.5(-7) mm. across, + 4-angled at first, becoming coarsely striate, with ca. 8 raised lines, drying dark brown when young, later brown and yellowish white, sparsely farinose when young; axillary innovations lacking basal scars; internodes 0.5—2.5(—3.5) cm. long; uppermost pair of axillary buds rounded, to 2 mm. long, + spreading; terminal bud plump, 0.5—1.6 cm. long, with subcrustose, grayish brown indumentum (hairs, Ficure 35, q-s, z), underdeveloped internode absent. Petiole (0.5—)0.8—2(-2.8) cm. long, flat or broadly to rather narrowly concave above, convex or + angled below, often drying blackish, glabrous when mature; lamina elliptic to oblong, (3.2-)4.5-16 by (1.5-)2-6.8 cm. (ca. 15 by 4 cm.), shortly and rather abruptly acuminate to rounded at apex, attenuate at base, or broadly rounded toward base and attenuate at very base, undulate and slightly to strongly recurved at margin, coriaceous, drying + sepia above and honey to umber below, farinose on midrib below when young, the midrib above + abruptly narrowed near or gradually narrowing from base, subdepressed, raised, (6 to) Il to 18 veins/5 mm., angle of divergence 65—80(-85)°. 1980] STEVENS, CALOPHYLLUM 563 Inflorescences from foliate axils, with (3 to) 5 to 9 (to 13) flowers, unbranched (rarely with 3-flowered branches up to 3.5 cm. long), the axis (1.1-)2-5.8(-10) cm. long, farinose (rarely subglabrous) when young, lowest internode 1-3 (—4.5) cm. long; bracts ovate, ca. 5 mm. long, soon deciduous; pedicels 0.4-2.3 cm. long, sparsely farinose when young. Flower (?)hermaphroditic; tepals 8 to 12, the outer pair broadly ovate to suborbicular, 4.75-7.5 by 4.5-6.5 mm., the next pair broadly ovate to oblong or elliptic, 4.5-9 by 4-7 mm., the inner ones ovate to elliptic or oblong, 6.5-11 by 2.5-7 mm.; stamens (80 to) 120 to 250, the filaments to 4.5 mm. long, connate for up to 1 mm., the anthers elliptic to suboblong, 0.3-1.4 mm. long, retuse at apex; ovary I-2 mm. long, the style 1-2.5 mm. long, t the stigma peltate to infundibular, ca. 1 mm. across, not lobed. Fruit spherical to ellipsoid, 1.8-3.2 by 1.7-2.4 cm., rounded to subacute at apex, drying vinaceous-brown to orange-cinnamon, smooth to sharply wrinkled; outer layer detaching + cleanly from stone, (0.8-)l-2.5 mm. thick, compact but skin often breaking off easily in ripe fruits; stone ellipsoid to subspherical, (1.6—)2.1-2.5 by 1.5-2.1 cm., + rounded at apex, the walls 1-1.6 mm. thick, smooth, unmarked, with basal plug to | cm. across; spongy layer thin. DistRiBUTION. Fiji Islands (Map 38). ADDITIONAL SPECIMENS SEEN. Pacific Islands. Fut. Sine loco: Horne s.n. (Gu). Viti Levu. Mba: vicinity of Nandarivatu, S. slopes of Mt. Ndelainathovu, 870-890 m., A. C. Smith 4944 (a, k, Ny, Pp, US), Sovutuwambu, 750-800 m Degener 14664 (A, K, NY), Waimogi, 823 m., FDF 1075 (kx), Vunidawa Path, 23 m., Vaughan 3431 (BM, kK), Waimongge Creek, 610 m., Berry 82 (mass, us), 85 (mass), 88 (Mass), Damanu K&8 (cur, US). Nandronga & Navosa: S. of Mt. Victory [Tomanivi], 914 m., FDA 14292 (sisH, kK); track to Vanua Levu village, 610 m., Berry 79 (cur, Mass); Rairamatuku Plateau, between Nandrau and Nanga, 725-825 m., A. C. Smith 5461 (a, k, Ny, P, US). Serua: Nabukelevu, FDA 15654 (cur, Mass); Naboutinini, 150 m., Damanu R22 (xk), FDF 573 (BIsH); Ngaloa, 210 m., Damanu G22 (x, ny), FDF 574 (nisn). wie hills bordering Wainavindrau Creek, near Wainimakatu, 150-250 m . C. Smith ee (GH, K, NY, P, US), 8553 (GH, kK, NY, Us); hills near Navua io 200-300 , Greenwood 1036 ( (A, K); Mt. Vakarongasiu, Gillespie 3267 (BIsH); Mt. Voma Track. 610 m., FDA 604 (suva). Naitasiri: Nanubu R., Rewasau, 914 m. Howard 308 (uve us); Mendrausuthu Range, 751 m., FDA 15463 (cHR, MASS, SUVA), 1547] (CHR, LAE); Mt. Naitarandamu, Gillespie ao 32 (GH). Sine loco Koroturanga, 600-700 m., A. C. Smith 8286 (GH, kK, Ny, P, US). Sine loco: Milne 244 (x). Eco.ocy. Colline and lower montane forest, on ridges and summits at higher altitudes (stunted tree in exposed conditions); 150-1250 m. alt. Flowering March, June, and November; fruiting June, and August to November (fruit reddish tinted—A. C. Smith 8525). 564 JOURNAL OF THE ARNOLD ARBORETUM [voc. 61 Locat use. The wood is good timber and is used in construction. Although Calophyllum cerasiferum is a variable species, it can be charac- terized by its rather plump terminal bud with more or less crustaceous, grayish brown indumentum; its twigs, which dry yellowish brown and coarsely striate; its coriaceous leaf blades, which have dense venation and dry sepia on the upper surface; and its fruits, which have a more or less compact outer layer and a thick-walled stone with a basal plug. The epithet cerasiferum (‘‘bearing cherries’) alludes to the dried fruits, which are about the size of large cherries. The circumscription of Calophyllum cerasiferum has been somewhat trou- blesome. It was originally decided to recognize two taxa: one including the large-leaved and -fruited specimens, with leaf blades sometimes strongly recurved at the margins and acuminate at the apex; the other, C. cerasiferum sensu stricto, including the smaller-leaved and -fruited specimens with the leaf blades only slightly recurved at the margin and more or less rounded at the apex. The fruits of the former group of specimens are wrinkled, while those of the latter are smooth. However, there are all intermediates in leaf size and type: Damanu G22 (large, strongly revolute blades); A. C. Smith 8525 (large, not very revolute blades); A. C. Smith 5461 (smaller blades); A. C. Smith 4944, FDF 1075, FDA 14292, and Berry 79 (small leaves acute at the apex; fruit wrinkled); Seemann 49 (leaves obtuse, fruit + smooth). There is considerable variation in the width of the leaf blade in both small- and large-leaved specimens. In details of terminal bud, hair type, leaf (apart from its size and apex), and fruit (except for the surface), the specimens are all basically similar. A. C. Smith 4944 is the only large-leaved specimen that has flowers; the flowers have eight tepals, and the filaments appear to have a yellow pigment. Of the smaller-leaved specimens, Howard 195, Berry 82, Berry 88, and FDA 15463 have flowers; all have ten to twelve tepals, and there is no pigment in the filaments. Within these latter specimens there is considerable variation in anther size: the anthers of Berry 88 are up to 1.5 mm. long, while those of FDA 15463 are only 0.3-0.5 mm Thus, despite the considerable variation in the group, all specimens have a basic similarity. In some characters the variation is continuous and is independent of variation in other characters; in others (notably those of the flower), there is too little material to evaluate the variation. Hence, it seems sensible to adopt broad limits for Calophyllum cerasiferum, at least for the time being. Greenwood 1036, which was apparently collected from a young tree of ew cerasiferum, has leaves with a lamina up to ca. 22.5 by 5.5 _ A. C. Smith 1789 is included in C. cerasiferum only with hesitation. With its broad midrib (figure in parentheses in the description above) and its rather distant venation (6 to 8 veins/5 mm.) it approaches C. vitiense; however, it has a coriaceous lamina with a recurved margin, and its twigs, although much stouter than those in other specimens of either species, are more similar to those of C. cerasiferum than to those of C. vitiense. Its 1980] STEVENS, CALOPHYLLUM 565 inflorescences are extremely robust (measurements in parentheses in the description above). Calophyllum cerasiferum has been lectotypified on the duplicate of Seemann 49 once in the Boissier Herbarium in Geneva and now in the general herbarium there. It seems that Vesque based his original illustration of C. cerasiferum on this specimen, rather than on the duplicate at Kew Herbarium. Thus, the typification of C. cerasiferum by Smith and Darwin (Joc. cit.) has to be slightly emended. 121. Calophyllum sp. Ficure 36, m-o. Tree ca. 12 meters tall; trunk and bark unknown. Twigs slightly flattened, 1.2-2.5 mm. across, obscurely 4-angled, drying blackish, grayish- to brown-puberulent; axillary innovations lacking basal scars; internodes 0.5-2.5 cm. long; uppermost pair of axillary buds ca. 1.5 mm. long, rounded, subspreading; terminal bud plump, 5-9 mm. long, with grayish to brown, crustaceous to puberulent indumentum (hairs, Ficure 35, p; ef. 35, 0), underdeveloped internode to 2 mm. long. Petiole 0.3-1.2 cm. long, shallowly concave above, convex below, glabrescent; lamina ovate to rhombi- form, 2.8-8.6 by 1.3-2.5 cm., acuminate at apex, acute to narrowly acute at base, somewhat undulate and slightly recurved at margin, coriaceous, drying + sepia above and sepia to sabelline below, glabrous when mature, the midrib above narrowing gradually from base, + level or slightly depressed, 0.3-0.4 mm. wide at midpoint, below rather inconspicuous, raised, striate, the venation above and below subobscure to apparent, raised, 10 to 1S veins/5 mm., angle of divergence 55-70°. Inflorescences from foliate axils, with 7 to 15 flowers, unbranched, the axis 1.4—3 cm. long, puberulent, lowest internode 3-7 mm. long; bracts unknown; pedicels 0.5-1 cm. long, puberulent. Flower (?)hermaphroditic; tepals 8, the outer pair broadly ovate, 4.5-5 by 3.5-4.5 mm., the next pair elliptic, 7 by 5 mm., to suborbicular, 5.3 by 6 mm., the inner ones elliptic, 6-7.5 by 3-4 mm.; stamens 120 to 145, the filaments to 4.5 mm. long, the anthers oblong, ca. 1 mm. long, retuse at apex; ovary ca. 1.5 mm. long, the style 2.5-3 mm. long, the stigma peltate, 0.6-0.8 mm. across, obscurely radiate. Fruit ellipsoid, 1.9-2.3 by 1.4-1.8 cm., persistently apiculate or not, drying vinaceous-brown, + wrinkled; outer layer detaching + cleanly from stone, ca. | mm. thick; stone ellipsoid, 1.3-1.8 by 1.3-1.5 cm., rounded at apex, the walls 0.4-0.9 mm. thick, ca. 2 mm. at base and ca. 0.25 mm. to one side of base, smooth, unmarked; spongy layer thin. DistriBuTION. Philippine Islands, Luzon (Map 39). SELECTED SPECIMENS SEEN. Philippines. Luzon. Bataan: Lamao R., Mt. Marive- les, Whitford 257 (F, G, K, NY, P, US), 1017 m., Williams 424 (GH, K, NY, us); Lamao F.R., BS 1577 (us). EcoLtocy. Lower montane forest, ca. 1000 m. alt. Flowering in January; fruiting March, May, July, October, and November. Calophyllum sp. 121 can be characterized by its plump terminal bud and 566 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 its medium-sized lamina that is acute to acuminate at the apex, with the midrib on the upper surface drying reddish brown. The inflorescence is rather short, and the flowers have eight tepals. The fruit is relatively large (ca. 2 cm. long) and has a rather thin outer layer and a stone that lacks a basal plug. Although in general appearance Calophyllum sp. 121 is similar to C. neo-ebudicum, the stone lacks the basal plug characteristic of that species: the hairs suggest a comparison with C. whitfordii (cf. Ficures 35, p, and 43, q, s, y). Until Calophyllum sp. 121 is better known, it is premature to describe it formally. In leaf and twig there is considerable variation among the specimens cited. BS 1577 and Whitford 1190 have short, rather thickly coriaceous leaf blades with rather dense venation, and stout twigs with short internodes, while Williams 424 has longer, thinner leaf blades with more distant venation, and thinner twigs with longer internodes. There is no comparable variation in the fruit, and in all specimens the midrib on the upper surface of the lamina narrows gradually from the base and is reddish. 122. Calophyllum caledonicum Vieill. ex Planchon & Triana, Ann. Sci. Nat. t. IV. 15: 291. 1862; Vesque, Epharmosis 2: ¢. 3. 1889, in C. DC. Monogr. Phanerog. 8: 546. 1893; Guillaumin, Ann. Mus. Colon. Mar- seille, I]. 9: 102. 1911; Baker f. Jour. Linn. Soc. Bot. 45: 273. 1921: C. T. White, Jour. Arnold Arb. 7: 95. 1926; Daniker, Mitt. Bot. Mus. Zurich 112: 271. 1933; Guillaumin, Fl. Nouv. Caléd. 217. 1948: Sarlin, Bois Foréts Nouv. Caléd. 206. pl. 91. 1954. Type: New Caledonia, bords des ruisseaux a Gatope, anno 1886, Vieillard 175 (lectotype, P; (?)isolectotypes, a, K, NSW). C. montanum Vieill. ex Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 292. 1862; Vieill. Ann. Sci. Nat. Bot. IV. 16: 74. 1862; Schlechter, Bot. Jahrb. 39: 193. 1906; Guillaumin, Ann. Mus. Colon. Marseille, II. 9: 102. 1911; Sarlin, Bois Foréts Nouv. Caléd. 308. pl. 93. 1954. Type: New Caledonia, montagnes a Balade, Vieillard 174 (holotype, p; iso- types, A, FI, G, L, P). Tree 3-10(—?20) meters tall, d.b.h. to 30 cm.; trunk without buttresses: outer bark brown to gray, fissured, thick; wood reddish. Twigs slightly flattened, 3-4.5 mm. across, 4- (very rarely 2-)angled when young, soon becoming coarsely striate, drying brown when young, later often with yellowish patches, sparsely farinose-puberulent when young; axillary innovations lacking basal scars; internodes 0.7-3.5 cm. long; uppermost pair of axillary buds rounded, to 1.5 mm. long, suberect, inconspicuous; terminal bud plump, 0.75-1.2(-1.8) cm. long, with short-tomentose to subcrustaceous, grayish to brown indumentum (hairs, Ficure 35, t-w), underdeveloped internode absent (to 3.5 mm. long). Petiole (0.2-)0.5—1.5(-2.5) cm. long, flat to broadly concave above, convex below, glabrous when mature; lamina oblong to elliptic, 5-14.3 by 1.8-3.6(-4.5) cm., rounded (rarely acute to subacuminate) at apex, acute to broadly rounded at base, slightly undulate, 1980] STEVENS, CALOPHYLLUM 567 slightly or not recurved at margin, coriaceous, drying umber to olivaceous above and below, glabrescent or sparsely and subpersistently farinose on midrib below, the midrib above narrowing rather quickly near base, broadly depressed at first, 0.2-0.6 mm. wide at midpoint, becoming raised in top '/g of lamina, below raised, striate (rarely angled), the venation above and below apparent, raised, (6 to) 8 to 13 veins/5 mm., angle of divergence 70-85(-90)°. Inflorescences from foliate axils (very ee terminal), with (5 to) 11 to 25 flowers, unbranched (rarely with 5-flowered branches up t 0 4 cm. long), the axis (3-)5-10(-21) cm. long, transiently puberulent or subglabrous, lowest internode |.5-4(-8) cm. long; bracts narrowly ovate to elliptic, to 11 by 3 mm., not persistent; pedicels 0.7—2(-3.5) cm. long, glabrous to puberulent. Flower (?)hermaphroditic; tepals (9 to) 12 (to 16), the outer pair + orbicular, 4-8 by 2.7-7 mm., the inner ones elliptic to obovate, (7-)10-12.5 by (2-)3.5-7 mm., outer tepals larger and inner ones glabrous; stamens 330 to 430, the filaments to 4.5 mm. long, the anthers elliptic to suboblong, 0.7-1.1 mm. long, rounded to retuse at apex; ovary ca. 1.8 mm. long, the style 2-3 mm. long, the stigma subpeltate, ca. 0.7 mm. across. Fruit spherical to ellipsoid, 1.7-2.7 by 1.2-2.2 cm., rounded to obtuse at apex, drying + vinaceous-brown, smooth or broadly wrinkled; outer layer detaching cleanly from stone, 1-4.5(-10) mm. thick, compact, air spaces developing under skin or not; stone spherical to ellipsoid, 1.2-1.8 by 0.9-1.6 cm., rounded to acute at apex, the walls 0.2-1 mm. thick, smooth, unmarked, at base poorly defined semifibrous (?)plug ca. . mm. across; spongy layer thin. DistriBUTION. New Caledonia (Map 38). SELECTED SPECIMENS SEEN. New Caledonia: Balade, Vieillard 175 (Gc, Pp); som- met de Puébo, Vieillard 174 (c, Gu, k, L); Wagap, Vieillard 173 (rp); So. base of Mt. Kaala, McKee 7985 (cANnB, K, L, P, US); vallée de Pouai (entre Ta et la Ouaime), 20-100 m. — 15618 (re); Chagrin Mine, Koumac Distr., McKee 7996 (a, CANB, K, L, P, US); Hienghene, McKee 26918 (rp); Kone, Pinjen, 10 m., McKee 29360 (P); er de la Oumbea, au dessus de Koe, Balansa 1335 (a, K, P, US); Bourail, col des Rouseilles, Sarlin 151 (pr); Haute Boulari, flancs du Pic Buse, 500 m., Virot 522 (a, Pp); banks of Toutouta R. near junction of the Kalouéhola, 50 m., McKee 7732 (a, CANB, K, L, NSW); Kanala, Vieillard Me (Nsw, NY, P); Bergen am Ngoye, 100 m., Schlechter 15241 (8M, ES EAT Gaks M, NSW, P, W); am Sudwestfusse des Mt. Humboldt, Daniker 573 (P); Paita, Le Rat 82 (pr); foréts de la Caricouye, 200-400 m., Franc 39 (a, G, NY); Noumea, Gandoger s.n., June 1906 (m); montagnes de St (a); route de la montagne des Sources, 500 m., McKee 24840 (p); Plaine Lacs, Mois de Mai forest, 250-350 m., Buchholz 1383 (a, BISH, K, P, us); Route de Yaté, Sarlin 165 (rp); Prony, 2 m., Balansa 584 (BM, Kk, P); Baie de Sud, Raoul s.n. (P) Eco.tocy. On serpentine rocks by rivers, on schists away from rivers; to 500 m. alt. Flowering November to February, and August (flower sweetly scented (like those of oranges—Virot 1340)); fruiting September to January (fruit blue to black, glaucous). 568 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 GERMINATION AND YOUNG PLANT. The seedling has three pairs of leaves separated by well-developed internodes at least | cm. long. (McKee 24841.) Calophyllum caledonicum can be recognized by its plump terminal bud; stout twigs, which dry yellowish and striate when old; more or less oblong leaves that have clear venation and a depressed midrib; and flowers, which have at least nine tepals. The specific epithet comes from the Roman name for Scotland. Calophyllum caledonicum is perhaps related to the C. vitiense—C. cerasiferum group of species; particularly in its terminal bud, twig, and flower, it is most similar to C. cerasiferum. However, it is not clear that C. caledonicum has a plug at the base of the stone like the other members of the C. vitiense group. Vesque (1893, loc. cit.) thought that Calophyllum inophyllum and C. engl were close, since both have flowers with a large number of tepals. However, C. inophyllum ae has only eight nes and it further differs from C. caledonicum in having a shorter terminal bud, a more or less obovate, thinner lamina, longer pedicels, and a larger fruit that differs in structure and is pale green when ripe. The two species are probably not particularly closely related (but see below). Variation within Calophyllum caledonicum Planchon and Triana noted that Calophyllum montanum had leaves in whorls see below), the leaves are alternate. Few subsequent collections have whorled leaves. Those of Sarlin 278 are in whorls of four, while those of McKee 20797 and McKee 15618 are in whorls of three. Five shoots of McKee 20797 were seen at Paris: two of the five had opposite leaves, and three had leaves in threes; all shoots came from the same tree (McKee, pers. comm.). The arrangement of the leaves in whorls on one of the shoots had broken down in an older part of the shoot, with scars of alternate, opposite, and ‘‘adjacent”’ leaves (1.e., leaves borne two at a node, but adjacent, rather than opposite, in insertion). Although specimens with whorled leaves tend to have long petioles and blades that are more or less acuminate at the apex, there are intermediates in these characters as well. The fruits of McKee 20797 are broadly ellipsoid, ca. 2.4 cm. long, and with a stone ca. 1.7 cm. long. The as yet immature fruits of McKee 15618 are ovoid and ca. 2.7 cm. long, but the stone is only ca. 1.4 cm. long, the outer layer being up to 1 cm. thick—over twice as thick as is usual in the species. Although the whorled arrangement of the leaves of the specimens discussed is a remarkable character, it is unstable and does not correlate with other characters. Calophyllum montanum is left in synonymy under C. caledonicum, as was first suggested by Vesque (1893, loc. cit.). Virot 511 and 522, both sterile specimens, have the midrib on the lower surface of the lamina angled rather than striate; the young stem is strongly two- (vs. four-)angled. 1980] STEVENS, CALOPHYLLUM 569 Nomenclature The sheet of Vieillard 175 at Paris with a locality label ‘‘bords des ruis- seaux a Gatope’’ is designated the lectotype of Calophyllum caledonicum. Other sheets, possibly isolectotypes, bear a label ‘‘bords d’un torrent a Gatope.”’ Possible Hybridization with Calophyllum inophyllum Two specimens collected from the southern end of New Caledonia (Goro, Sarlin 182 (pe); Baie du Prony, Balansa 585 (p)), are intermediate between Calophyllum caledonicum and C. inophyllum. The lamina is up to 20 by 5-8 cm. and has 5 to 8 veins/5 mm. It is more coriaceous and narrower than is usual for C. inophyllum, but in both venation density and midrib type it approaches that species. Both specimens are in fruit (those of Balansa 585 are mature, the stone being filled by the embryo); the fruit is of a lighter color than is usual in C. caledonicum. The fruits approach those of C. inophyllum, although the spongy layer does not seem to be particularly thick. Sarlin 182 has on its field label ‘‘tamanu de bord de mer,”’ the French name for C. inophyllum! 123. Calophyllum carrii P. F. Stevens, Austral. Jour. Bot. 22: 360. fig. 2. 1974. Type: Papua [Northern Province], Isuarava, 3500 feet [1530 m.], 15 Feb. 1936, Carr 15599 (holotype, LAE; isotypes, A, CANB, K, L, NY, SING). Tree; for details of trunk and bark, see varieties. Twigs slightly flattened, 3.5-7 mm. across, + 4-angled, or with ca. 8 raised lines, becoming striate when older, drying grayish brown, puberulo-farinose when young; axillary innovations lacking basal scars; internodes 1-3 cm. long; uppermost pair of axillary buds rounded, ca. 1.5 mm. long, suberect, inconspicuous; terminal bud plump, I-3 cm. long, with grayish brown, crusta- ceous indumentum (hairs, Ficure 35, aa, bb, ff; also submoruloid), underde- veloped internode to 3 mm. long. Petiole 1-3 cm. long, broadly concave above, convex to angled below, glabrescent; lamina obovate to suboblong, (5.5—)7-12 by 2.9-6 cm., obtuse to shallowly retuse at apex, acute at base, undulate and recurved at margin, marginal thickening sometimes up to | m wi oriaceous, drying sabelline above and fulvous-umber below, pinche eat puberulent to furfuraceous on midrib below, the midrib above narrowing gradually from base, depressed at first, 0.2-1.5 mm. wide at midpoint, becoming raised in top half of leaf, below raised, rounded to striate, the venation rather obscure above, + apparent below, raised, 7 to 13 (to 15) veins /5 mm., angle of divergence 65-75°. Inflorescences from lower foliate (rarely defolits) axils, with 5 to 18 flowers (with 3-flowered branches to 1.2. cm. long), the axis 3.3-4.5 cm. long, farinose-puberulent, lowest internode 2-4 mm. long; bracts unknown; pedicels 0.5—1.9 cm. long, farinose-puberulent. Flower known only in bud; tepals 8 (rarely 6), the two outer broadly ovate, ca. 6.5 by 5.5 mm., sparsely farinose on back at very base; stamens 90 to 120, the anthers oblong, 0.5-1 mm. long, + retuse at apex; ovary ca. 570 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 1.8 mm. long, the style unknown, the stigma excentrically peltate, ca. 1.5 mm. across. Immature fruit (from var. longigemmatum) subspherical, c 1.8 cm. long, drying smooth. Key to the Varieties of Calophyllum carrii _ . Twigs 3.5—4.5(-5.5) mm. across; terminal bud 1—2.2 cm. long; lamina lacking an obviously hhickened PAE Mace: zg os, a gs Rc a eB a. var. carril. 1. Tw wigs 4 4.5-7 mm. across; terminal bud 2-3 cm. long; lamina with thickened MAPeinsea. AN, WIM og: ee Ge ee a 123b. var. longigemmatum. 123a. Calophyllum carrii P. F. Stevens var. carrii Tree ca. 35 meters tall, d.b.h. to 100 cm.; trunk with buttresses to 0.9 . tall; outer bark dark brown, fissured: under bark light brown; latex golden, clear. Twigs 3.5—4.5(-5.5) mm. across, + 4-angled or with ca. 8 raised lines; terminal bud 1.3-2.2 cm. long; lamina with slightly recurved margin, marginal thickening not obvious. Distripution. Central and Northern provinces of Papua New Guinea (Map SELECTED SPECIMENS SEEN. Papuasia: see Stevens, loc. cit. Ecotocy. Lower montane rain forest, 1050-1525 m. alt. Flowering in February (late bud). 123b. Calophyllum carrii P. F. Stevens var. longigemmatum P. F. Stevens, Austral. Jour. Bot. 22: 361. 1974. Type: Nederlands Nieuw-Guinea [Irian Jaya], Hollandia, mouth of Tami River, IS m., 8 July 1956, BW 2813 coll. Schram (holotype, LAE; isotypes, BO, CANB, K, KEP, L, SING). Tree ca. 30 meters tall; trunk without obvious buttresses; outer bark dark gray, flaking off in irregular, rectangular scales; under bark reddish and cream mottled; inner bark creamy red; latex yellow. Twigs 4.5-7 mm. across, usually with 8 raised lines; terminal bud 2-3 cm. long; lamina with strongly recurved margin, marginal thickening ca. | mm. wide. DistriBuTION. Mainland New Guinea; known only from a small area along the northern coast (Map 39). SELECTED SPECIMENS SEEN. Papuasia: see Stevens, loc. cit. Ecotocy. Lowland rainforest, 15-300 m. alt. Conoidal galls ca. | mm. tall sometimes occur on the lower surface of the lamina (BW 2813). The epithet /Jongigemmatum refers to the distinctive terminal buds of this variety. 1980] STEVENS, CALOPHYLLUM 571 OD Ten carrii can be recognized by its long, plump terminal bud, obovate to leaf blades with the midrib on the upper surface depressed for at least half its ne and flower with (usually) eight tepals. Its indumentum is not well developed, and the older twigs at least are striate. The specific epithet commemorates C. E. Carr, who collected the type specimen during his fatal trip to New Guinea. Although the two varieties are readily distinguished by the characters given above, neither is well known. 124. Calophyllum collinum P. F. Stevens, sp. nov. piers sp. D, aff. C. sil Lauterb.; P. F. Stevens, Austral. Jour. _ 22: 397. 1974. A speciebus aliis Calophylli in internodiis brevibus 0.3—2 cm. longis, lamina parva costa supra valde sulcata venulis 11 usque ad 20 per 5 mm., fructu subsphaerico strato exteriore 1.5-3 mm. crasso e putamine nad munde secedenti, differt Tree 22-36 meters tall, d.b.h. to 85 cm.; trunk without buttresses: outer bark brown (gray), fissures at first short and distinct, becoming confluent, inner surface brown (orange-brown); under bark red; inner bark red or reddish; latex yellow, clear, viscous (becoming green). Twigs slightly flattened, 1.2-3.5 mm. across, strongly 4-angled, often with 4 additional raised lines, drying dark brown, transiently subfarinose to subpersistently tomentose; axillary innovations lacking basal scars; internodes 0.4-2 cm. long; uppermost pair of axillary buds rounded, ca. | mm. lon subspreading; terminal bud plump, 6-9 mm. long, with brown, Seromena. to subcrustaceous indumentum (hairs, Ficure 35, x, y, cc—-ee), underdeveloped internode to | mm. long. Petiole (0.45-)0.6-1.2 cm. long, broadly concave above and convex below, glabrescent to subpersistently tomentose; lamina obovate to elliptic (rarely suboblong), (1.8—-)2.3-8 by (1—-)1.3-4.5 cm., rounded to subretuse or subcuneate at apex, acute to cuneate at base, slightly undulate and not recurved or slightly so at margin, coriaceous, drying umber to olivaceous above and fulvous to olivaceous below, transiently farinose to subpersistently puberulo-tomentose on midrib below, the midrib above grad- ually narrowed from base, strongly sulcate but at least margins raised, 0.2-0.4 mm. wide at midpoint, becoming raised toward apex, below raised, striate, the venation obscure to subapparent on both surfaces, raised to + flat, [1 to 20 veins /5 mm., angle of divergence 65—80°. Infructescences from foliate axils along twigs, with scars of 7 to 21 flowers, unbranched, farinose to subtomentose, lowest internode 1-10 mm. long; bracts not known; pedicels 4-9 mm. long, glabrous. Flower not known. Fruit subspherical (ovoid), 1.2-1.8 by 1.1-1.7 cm., rounded (rarely acute—Stevens et al. 921) at apex, drying brown, smooth; outer layer not detaching cleanly from stone, 1.5-3 mm thick, compact apart from air spaces developing under skin; stone subspherical to ellipsoid, 0.9-1.4 by 0.8-1 cm., rounded (rarely obtuse) at apex, the walls 0.3-0.5 mm. thick, to 0.7 mm. thick at base, smooth, unmarked; spongy layer thin. oe) JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Type: Papua, Western District, Ningerum, 110 m., 29 August 1970, NGF 42992 coll. Henty & Barlow (holotype, a; isotypes, K, L, LAE, SING). DistripuTiON. Eastern New Guinea (Map 41). ADDITIONAL SPECIMENS SEEN. Papuasia. IRIAN JAYA. Snow Mountains: Bernhard Camp, Idenburg R., 550 m., Brass & Versteegh 13576 (a, 2 Me Guinea. Morobe: near Garaina 520 m., NGF 45057 (a, E, K, L, NG). Western: 8 km. from Kiunga, 25 m., Stevens et al. 888 (a), $90 “a oe ns from Kiunga, 30 m., cen et al. 921 (A EcoLocy. Colline forest or lowland forest with colline aspect, 25-520 m. alt. Fruiting March, July, and August (fruit blue to blackish—field notes of NGF 45057 mention dark green fruit, but it is likely that these would finally turn bluish). Calophyllum collinum can be recognized by its short internodes, its strongly four-angled twigs that usually have an additional four raised lines, and its small, oblong-obovate leaf blades with a strongly sulcate midrib and rather dense venation (11 to 20 veins/5 mm.). Its fruits have a thick outer layer that is compact except for air spaces that sometimes develop under the skin. The species looks like a plant from forests at higher altitudes, as Y. Lelean remarked when we first found the plant; this suggested the specific epithet. Calophyllum collinum is a fairly common tree around Kiunga, especially noteworthy because of its clean, brown-colored bole that becomes deeply fissured only in very large specimens. Its relationships are not clear (see also Stevens, Joc. cit.), but it can be readily distinguished from C. sil by the characters given in the diagnosis. Although the high-altitude variant of C. sil may have a midrib similar to that of C. collinum, it has larger leaves with less dense venation and twigs with longer internodes. 125. Calophyllum costatum J. F. Bailey, Queensland Agr. Jour. 5: 392. pi. 136. 1899 (1 Oct.); F. M. Bailey, Queensland FI. 1: 104. 1899 (Dec.), Comprehens. Catal. Queensland Pl. 54. fig. 40. 1913. Type: Australia, Queensland, Evelyn [North Kennedy], 6 July 1894, J. F. Bailey s.n. (holotype, BRI; isotype, LAE (frag.)). Tree to 25-33 meters tall, d.b.h. to 60 cm.; trunk without buttresses; outer bark dark brown or dark grayish, flaky or peeling off and cracked only in places, inner bark pink, layered, fibrous; latex pink-cream to whitish (turning brown on exposure (Hyland 8912). Twigs flattened, 2-4 mm. across, at most obscurely 4-angled, drying brown when young, whitish gray to yellowish white when older, subpersistently puberulent; axillary innovations lacking basal scars; internodes 0.5-2(-3) cm long; uppermost pair of axillary buds pointed, to 3.5(-6.5) mm. long, erect, usually inconspicuous; terminal bud plump, 6.5—11 mm. long, with brownish, puberulo-tomentose indumentum (hairs, Ficure 35, gg, hh), underdeveloped internode 1-3(-6) mm. long. Petiole 0.7-1.9 cm. long, broadly and shallowly concave above, convex below, glabrous; lamina elliptic to obovate, 4.7-8.8 by 2.2-3.9 cm., subacute to rounded at apex, acute to cuneate at base, distantly 1980] STEVENS, CALOPHYLLUM Be and shallowly undulate but not recurved at margin, thinly coriaceous, drying umber above and slightly paler below, subpersistently farinose on midrib below, the midrib above narrowing gradually from base, strongly sulcate in bottom 1/3 of lamina, becoming raised, 0.2-0.3 mm. wide at midpoint, below raised, obscurely angled to striate, the venation above and below + clear, raised (rarely subobscure), 6 to 11 (to 14) veins/5 mm., angle of divergence 55-65°. Inflorescences from foliate axils, with 5 to 7 flowers, unbranched, the axis to 1.7 cm. long, puberulent, lowest internode ca. 8 mm. long; bracts ovate, to 8 mm. long, deciduous; pedicels 5-7 mm. long, puberulent. Flower (?)hermaphroditic; tepals 8, the outer pair ovate, ca. 7.5 by 6.5 mm., puberulent on back, the inner ones suborbicular, ca. 7 by 7 mm., outer pair puberulent dorsally down middle; stamens 95 to 140, the filaments to 2.5 mm. long, the anthers oblong, ca. 2 mm. long, acute at apex; ovary ca. 1.3 mm. long, the style unknown, the cient peltate, ca. 1 mm. across. Fruit ellipsoid to ovoid, 2.8-3.5 by 2.3-3.8 cm., + acute apex, drying + vinaceous-brown, shallowly to rather sharply and deeply longitudinally wrinkled; outer layer detaching cleanly from stone, 1-2.5 mm. thick, compact; stone ellipsoid to ovoid, 2.3-3 by 1.9-2.2 cm., subacute to rounded at apex, the walls 1.2-1.5 mm. thick, ic 3 mm. thick near base, smooth, unmarked, apparently with basal plug 3 mm. across; spongy layer (?)thin. DistripuTion. Northeastern Queensland, Australia (Map 40). SELECTED SPECIMENS SEEN. Australia. QUEENSLAND: Bellenden Kerr, 1075 m., L. S. Smith 14698 (pri); Ravenshoe, Manuell 41 (pri); Middle Creek, Herb. Flecker 6605 coll. Flecker; Atherton, Rhys s.n., Feb. 1916 (sri); Evelyn, Mar 40. Distribution of Calophyllum sil (circles), C. sil variant (squares), C. leleanii (st ars), C. waliense (star in solid circle), and C. streimannii (half-circles) in Malesia. Inset: C. costatum. 574 JOURNAL OF THE ARNOLD ARBORETUM [voL. 6] _ F. Bailey s.n., 15 April 1908 (x); S.F.R. [State Forest Reserve] 265 (near Crater), 1000 m., Stocker 670 (a, CANB, K, KEP, LAE); S.F.R. [State Forest Reserve] 1203, Dirran, 1140 m., Irvine 1203 (a, Kk, LAE); Boonjie L.A. [Logging ea], 680 m., Hyland 8912 (a); Cook Distr., Hugh Nelson Range, L. S. Smith 10507 (a). EcoLocy. Canopy tree of colline forest, 680-1140 m. alt. Flowering in November; fruiting January and March. Calophyllum costatum can be recognized by its plump, moderate-sized terminal bud; its fairly small lamina that is more or less cuneate at the apex and has relatively clear and distant venation; and its rather large fruits (ca. 3 cm. long) that are broadly wrinkled. The outer layer of the fruit is comparatively thick (1-2.5 mm. across); the stone is thick walled and may have a basal plug. The midrib of the dried leaf is very prominent, and this fact suggested the specific epithet. Calophyllum costatum is perhaps related to C. obscurum (q.v.). The original description of Calophyllum costatum must be attributed to Bailey filius, whose publication antedates that of his father by two months. 126. Calophyllum obscurum P. F. Stevens, Austral. Jour. Bot. 22: 380. fig. 1974. Type: Solomon Islands, Malaita, Are Are District, west coast, Kiu, 13 Feb. 1963, BSIP 3875 (holotype, LAE; isotypes, K, L, SING). Tree to 30 meters tall, d.b.h. to 90 cm.; outer bark yellow to gray-brown, scaly, fissured; latex yellow, opaque. Twigs slightly flattened, 2-2.5 mm. across, 4-angled, drying grayish brown, subpersistently grayish farinose-puberulent; axillary innovations lacking basal scars; internodes 0.5—1.75(-3) cm. long; uppermost pair of axillary buds rounded, to 1.5 mm. long, + spreading, inconspicuous; terminal bud plump, 0.7—-1(-1.7) cm. long, with subcrustaceous, grayish indumentum (hairs, Ficure 37, a, b), underdeveloped internode 1-4 mm. long. Petiole 0.9-2.5 cm. long, flat to shallowly concave above, convex below, persistently farinose; lamina elliptic, ovate, or obovate, 7-11.5 by 3.3-6 cm., obtuse to subacuminate at apex, acute at base, undulate but slightly recurved at margin, coriaceous, drying near umber above and sepia below, subpersistently farinose on midrib below, the midrib above narrowing gradually from base, rather indistinct, margins slightly raised near base, 0.25—0.35 mm. wide at midpoint, becoming Ficure 37. Hairs “(from Berea bud, unless otherwise noted). a, b, Calo- phyllum obscurum (BSIP 7741): b, from above. c-f, C. sil: c, d, NGF 33731; e, NGF 36754; f, bb 24368. g, h, C. laticostatum: g, NGF 3284, from stem; h, LAE 58797. 1, C. archipelagi (Backer 27827). j, k, Calophyllum sp. 129 (bb 19445). 1, m, C. peekelii: 1, NGF 18330, from above; m, BSIP 8804, from above. n, 0, C. leleanii: n, Brass 3447; 0, Musser et al. S-20a. L ene bape 59253). q-t, C. vexans: q, Brass a tr, LAE 51230, from below; s, bb 30923, from below; t, NGF 18307. C. papuanum (NGF 93956). w-y, C. euryphyllum: w, NGF 27335; x, ene 4662; y, LAE 59189. Scale = 120 um. (in w, scale = 240 pm.). 576 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 raised, below raised, subangled, the venation obscure above, obscure to subapparent below, 9 to 16 veins/5 mm., angle of divergence 60-75°. Inflorescences from foliate axils, with 5 to 11 flowers; unbranched (rarely with 3-flowered branches to 0.7 cm. long), the axis 0.7-2.8 cm. long, puberulent, lowest internode (0.3-)I-1.7 cm. long; bracts unknown; pedicels 0.8—1.2 cm. long, puberulent. Flower known only in late bud, (?)hermaphroditic; tepals 8, the outer 4 suborbicular, ca. 5.5 by 5.5 mm., farinose-puberulent on back; stamens 200 to 250, the anthers oblong, |.5-2.3 mm. long, acute at apex; ovary ca. | mm. long, the style ca. 1.5 mm. long, the stigma peltate, 1.6-2.4 mm. across, irregularly fringed. Fruit + spherical, 2.4—3.1 by 2.2-2.7 cm., sharply apiculate, drying grayish brown, wrinkled; outer layer detaching + cleanly from stone, 0.5-1.3 mm. thick, compact; stone subspherical, 2.2-2.9 by 2-2.6 cm., rounded at apex, the walls 1.5-2 mm. thick (3-4 mm. thick at base), smooth or slightly rugose, unmarked; spongy layer thin. DistripuTION. The Solomon Islands (Map 43). SPECIMENS SEEN. Papuasia. SoLoMoN IsLANps: see Stevens, Joc. cit. Eco.ocy. Ridges in primary rainforest, or raised, sometimes flooded, coral platforms; 60 m. alt. Flowering February and March (flower scented); fruiting August and December (fruit green to grayish). Calophyllum obscurum can be recognized by its plump terminal bud and its medium-sized leaf blades that dry dull umber-sepia and have obscure, fairly close venation and a rather indistinct midrib on the upper surface. The inflorescence is covered in grayish, farinose-puberulent indumentum, and the wrinkled fruits have a thin outer layer and a thick-walled stone that appears to lack a basal plug. In two collections the flower is reported to be yellow; this color may be caused by the presumably brownish indumentum covering the white tepals. The specific epithet alludes both to the obscure venation and to the generally undistinguished appearance of the specimens. The thick-walled stone that lacks a basal plug and the indumentum covering the entire inflorescence immediately distinguish Calophyllum obscurum from the other species of Calophyllum in the Solomon Islands. It is perhaps related to C. carrii, from mainland New Guinea, although that species has more robust twigs, leaf blades with a depressed midrib on the upper surface, clearer venation, and longer pedicels. Ripe fruits of C. carrii are unknown. Calophyl- lum obscurum is also similar in many points to C. costatum, from Australia; that species, however, has yellowish twigs, smaller leaf blades with less dense venation and a more prominent midrib, and a fruit that, although of similar size, has a thicker outer layer and a stone apparently with a basal plug. The hairs of all these species are rather similar (Ficures 35, aa, bb, ff—hh; 37, a, b). 127. Calophyllum sil Lauterb. Bot. Jahrb. 58: 14. 1922; O. Schwarz, Repert. Sp. Nov. 24: 89. 1927; T. C. Whitmore, Gard. Bull. Singapore 22: 12. 1967; P. F. Stevens, Austral. Jour. Bot. 22: 395. 1974. Type: Deutsch 1980] STEVENS, CALOPHYLLUM a1 Neu-Guinea [Irian Jaya], Siidkuste bei Gelieb, 31 Oct. 1907, Brander- horst 179 (holotype, 8, destroyed; isotypes, Bo, K, L, U). C. ramiflorum O. Schwarz, Repert. Sp. Nov. 24: 88. 1927. Synrtypes: Australia [Northern Territory], Port Darwin, cliffs near Hospital, Bleeser 502 (Nsw, MEL); Finniso River District, Bleeser A27 (n.yv.). C. procerum A. C. Smith, Jour. Arnold Arb. 22: 344. 1941. Type: Papua, are Fly puee Lake Daviumbu, Aug. 1936, Brass 7589 (holotype, sotypes, L). Casi is seat. Nova Guinea Bot. 8: 309. 1910. C. warburgii auct., non Engler; A. C. Smith, Jour. Arnold Arb. 22: 345. 1941, quoad Brass 7724. Tree 4-30 meters tall, d.b.h. to 139 cm.; trunk without buttresses (with spurs); outer bark yellowish at first, becoming rusty brown to gray, + deeply fissured, with transverse cracks between fissures, scaling or not; under bark bright red; latex yellow or greenish yellow, clear, sticky, sometimes aromatic. Twigs slightly flattened, 2.6—-4 mm. across, strongly 4-angled (rounded or with 6 raised lines), drying brown, sparsely brown-farinose; axillary innovations lacking basal scars; internodes 0.3-3.5 cm. long; uppermost pair of axillary buds rounded, ca. | mm. long, spreading, inconspicuous; terminal bud plump, 5-10(-13) mm. long, with brown, puberulent indumentum (hairs, Ficure 37, c-f), underdeveloped internode to 2.5(-3.5) mm. long. Petiole 0.6-1.5(-2) cm. long, broadly concave above, convex below, glabrous when mature; lamina obovate to elliptic (suboblong), (2.8—-)5-12.5 by (1-)2.5-5.5 cm., acute to cuneate or rounded at apex, cuneate, acute, or broadly rounded and ultimately shortly acute at base, broadly undulate but slightly (rarely strongly and broadly) recurved at margin, coriaceous, drying + cinnamon-sepia on both surfaces or greenish olivaceous to fuscous-black above and olivaceous below, glabrous or sparsely farinose-puberulent on midrib below (also above when young), the midrib above gradually narrowed from base, depressed, mm. long), the axis 0.3-4.5 cm. long, farinose-puberulent, lowest internode (0.1-)0.2-0.7(-1.2) cm. long; bracts ovate, to 3.5 mm. long, densely puberulent below, deciduous; pedicels 3-7 mm. long, farinose-puberulent. Flower (?)her- stamens 45 to 80, the filaments to 2.5 mm. long, the anthers oblong, 0.5-1.3 mm. long, + retuse at apex; ovary |.3-1.5 mm. long, the style 1.5-1.7 mm. long, the stigma peltate, 0.5-0.7 mm. across, slightly radiate. Fruit spherical to ovoid, 1.3-1.9 by 1-1.6 cm., + acute at apex, drying vinaceous-brown, thick, to 2 mm. thick toward apex, compact; stone + spherical, 0.8-1.3 578 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 by 0.8-1.2 cm., rounded at apex, the walls 0.15-0.4 mm. across, smooth, (?)unmarked; spongy layer thin. DistRiBUTION. Southern mainland New Guinea and northern Australia; forms of uncertain status from the Moluccas, northern mainland New Guinea, and New Britain (Map 40). SELECTED SPECIMENS SEEN. Moluccas. HALMAHERA: Tobelo, N. Totodokoe, 30 m., bb 33807 (?) (Bo, L, sinc); Weda, Tilope, 25 m., bb 24843 (?) (Bo, L, SING). SerAM: Kiandarat, 60 m., bb 25830(?) (Bo, L, SING). TANIMBAR: Otimmer, 100 m., bb 24368 (?) (Bo, L, sinc). Papuasia: see Stevens, Joc. cit. Also, IRIAN Jaya. Vogelkop: Beriat (+ 12 km. S. of Tenimaboean), 10 m., BW 6259 (2) (L). Papua New Guinea. Madang: Imbron R., 30 m., NGF 49237 (2?) (A, L, LAE). Western: 2 km. W. of lokwa, 10 m., LAE 60448 (0). Australia. WesTeRN Australia: Mitchell Plateau, N. Kim mberley, Beard 7013 (DNA, NSW, PERTH); Prince Regent R. Reserve, Gariyeli Creek, Fern Gulley, George 12822 (PERTH); Bonaparte Archipelago, Osborn Is., P. G. Wilson 11061 (PERTH). NortHeRN TERRITORY: Cannon Hill, Must 791 (cANB, DNA, K, LAE); W. face of Mt. Brockman Range, 30 m., Schodde AE2 (cans, DNA, kK); Bay no. 3 [Arnhem Bay] is., R. Brown ‘‘6302’’ (Bm, Kk); Sampit Creek Gorge, Arnhem Land, McKean B796 (cans); 65 km. NE. Pine Creek, W. Arnhem Land, Balgooy & Byrnes 1348 (pNA, K, L, LAE); 3 km. S. of E. Alligator R. crossing, Byrnes 2187 (CANB, DNA, LAE); Jim Jim Falls area, R. E. Fox 521 (pna); Se Bay, Byrnes 973 (pna); Lee Point, Byrnes 1734 (pNa); Gulmarri, Elcho Dunlop 3949 (pna); Giddy R. crossing, Byrnes 2592 (DNA, CANB); Port oe Holtze 38 (MEL). QUEENSLAND: Jardine R., Gittins 1844 (Nsw); Iron Range, 20 m., Brass 19294 (a, CANB, K, SING). W. of Bamaga, ca. 2.7 cm. SW. of Cape York, L. S. Smith 12485 (a, CANB, K, L, LAE); Scrubby Creek, 91/2 km. N. of Rocky R., 50 ia , Hyland 5433 oF Claudie R., 80 m., Dockrill 564 (k, L); Arbor. Res. between MclIvor R. and Cape Flattery, 20 m. Hyland 6552 (k, 1); Cook Distr., Annam = junction with Parrot Creek, 215 m., L. S. Smith 14321 (4, CANB, L, LAE, MEL); Weipa Concession, Marmoss reek, 10 m., Dockrill 859 (LAr); Granite Creek, Lower Bloomfield R., L. S. Smith 11086 (v); Etty Bay, Innisfail, L. S. Smith & Webb 3265 (a); Trinity Harbour, W. Hill 54 (MEL). EcoLtocy. Savanna woodland or riverine gallery forest (see also Beard, 1976, as C. australianum), rarely secondary forest; 7-215 m. alt. Flowering April, and June to August (flower scented); fruiting May, and September to November (fruit bluish to purple-black). Paijmans 376 and L. S. Smith 11086 have galls on the leaf; these are slightly transversely elongated pustules at right angles to, and often near, the midrib. The variant grows in rainforest, 10-630 m. alt. LocaL use. The wood is used for larger dugout canoes on the Fly River (Western Province, Papua New Guinea). Calophyllum sil can be recognized by its plump terminal bud; its moderate- sized and more or less elliptic lamina that has a depressed midrib on the upper surface and that dries more or less concolorous; its four-tepaled flowers; and its fruits, which dry sharply wrinkled and vinaceous-brown, with the 1980] STEVENS, CALOPHYLLUM 579 outer layer moderately developed, compact, and not detaching from the thin-walled stone. The specific epithet, si/, is a name given to this species in the southern part of Irian Jaya. Variation within Calophyllum sil is discussed in Stevens (loc. cit.). The specimens from New Guinea cited above with a question mark belong to the variant that usually grows at a moderate elevation (but BW 6259 was collected at 10 m. alt.) in rainforest; the status of this variant is unclear, since all except one of the specimens (NGF 36754) are sterile. The specimens cited above from the Moluccas region are also all sterile. Some (e.g., bb 33807) have rather thickly coriaceous leaf blades with broadly recurved margins, and the older leaves tend to dry fuscous-black on the upper surface. Calophyllum sil is probably related to C. laticostatum and perhaps also to C. collinum. All have basically similar hairs (Ficures 35, x, y, cc-ee; 37, c-h), blue fruits, and flowers often with four tepals (flowers of C. collinum are not known). In addition, both C. si/ and C. laticostatum sometimes have latex canals on the lower surface of the leaf obliquely ascending over the normal latex canal/vein system. Calophyllum silis not close to C. australianum (= C. calaba var. australianum), as was suggested earlier (Stevens, loc. cit.). Although young plants of C. sil have leaves similar in shape to those of C. calaba var. australianum, the twigs of the former are almost square in transverse section, while those of the latter are flattened; the hairs of the two are not notably similar (cf. Ficures 37, c-f, and 12, q). 128. Calophyllum laticostatum P. F. Stevens, Austral. Jour. Bot. 22: 375. fig. 8. 1974. Type: Papua, Milne Bay, about 6 miles [8 km.] up the Dawa Dawa River, 250 feet [75 m.], March 1945, NGF 1327 coll. L. S. Smith (holotype, Lar; isotypes, A, BRI, CANB, L, NSW). Tree 15-43 meters tall, d.b.h. to 91 cm.; trunk without buttresses or spurs (?rarely with aerial roots); outer bark yellowish in young tree, becoming gray to brown, shallowly to rather deeply fissured, scaling or not, the inner surface dark red, or orange-brown and orange or red and brown mottled: under bark dark red; inner bark dark red; latex yellow (greenish yellow), clear, sticky. Twigs slightly flattened, 2-5.5 mm. across, 4-angled, drying deep brown, subpersistently brown farinose-puberulent; axillary innovations lacking basal scars; internodes (0.4—)1-3 cm. long; uppermost pair of axillary buds rounded, ong, spreading, inconspicuous; terminal bud plump, (0.8—)1.3-2.7 cm. long, with + crustaceous, gray-brown indumentum (hairs, Ficure 37, h), underdeveloped internode to 4 mm. long. Petiole (0.4-)0.7-1.4 cm. iene broadly concave above, glabrescent below; lamina elliptic to narrowly obovate, (3-)6.5-16 by (0.8-)1.4-4.5 cm., acute (rarely acuminate (NGF 18373)) at apex, narrowly cuneate at om very base often shortly rounded, not undulate to rather distantly so and narrowly to broadly recurved at mar- gin, coriaceous, drying + umber above and sabelline-sepia below, transiently farinose on midrib (rarely near margin) below, or glabrous, the midrib above narrowed gradually from base, broadly depressed, 0.2-1.6 mm. wide at 580 JOURNAL OF THE ARNOLD ARBORETUM [vo. 61 midpoint, below raised, striate, the venation above obscure to subapparent, below + apparent, raised, sometimes with latex canals ascending over veins on lower surface, (9 to) 11 to 20 veins /5 mm., angle of divergence (40-)60-70 (-80)°. Inflorescences from foliate axils along twigs, with 5 to 21 flowers, unbranched (rarely with 3-flowered branches to 3 mm. long), the axis 1.2-5.5 cm. long, + puberulent, especially toward base, lowest internode 3-11 mm. long; bracts ovate, ca. 4mm. long, deciduous; pedicels 2-11 mm. long, sparsely puberulent to glabrous. Flower known only from buds, (?)hermaphroditic; tepals 4 or 6, the outer pair ovate, ca. 5 by 4 mm., stamens ca. 168, the anthers oblong, ca. | mm. long, + retuse at apex; Ovary Ca. 1 mm. long, the style ca. 1.2 mm. long, the stigma excentrically peltate, ca. 0.8 mm. across. Fruit ovoid to subspherical, 1.3-1.9 by 1.2-1.6 cm., rounded at apex, drying mid-brown, broadly and obscurely wrinkled; outer layer not detaching cleanly from stone, (0.5-)1-3 mm. thick, compact or with air spaces developing under skin; stone subspherical, 0.65-1.1 by 0.6-1 cm., rounded at apex, the walls 0.2-0.8 mm. thick, smooth, unmarked; spongy layer thin. DistripuTION, Western New Guinea (Map 39), possibly the Philippine Islands (Luzon). SELECTED SPECIMENS SEEN. Philippine Islands. Luzon. Mt. Calvario, Dolores, 600 m., FB 30019 (uc). Papuasia: see Stevens, loc. cit. Eco.ocy. Well-drained lowland or colline rainforest, 20-1370 m. alt. Flower- ing in January; fruiting October, January, and March, submature fruit in May (fruit bluish). YounG PLANT. The young plant is erect, and the terminal bud is functional. Calophyllum laticostatum can be recognized by its fairly long (usually more than | cm.), plump terminal bud and its rather narrowly elliptic, coriaceous leaf blades that have a broadly sunken midrib narrowing gradually from the base. The fruits are small (less than 2 cm. long) and have a well-developed outer layer. The epithet /aticostatum (‘‘broad midrib’’) emphasizes one of the most prominent features of the species. Calophyllum laticostatum is apparently related to C. archipelagi and Calo- phyllum sp. 129. The three taxa can be separated as shown in TaBe 15. Stevens (loc. cit.) compared C. laticostatum with the New Caledonian species C. caledonicum and C. montanum (= C. caledonicum). Calophyllum cale- donicum is perhaps most closely related to the C. vitiense-C. neo-ebudicum complex and has larger fruits and flowers with eight or more tepals. Relatively little material of Calophyllum laticostatum in flower and fruit is known, and there is a considerable amount of vegetative variation (see also Stevens, loc. cit.). The single specimen cited from the Philippine Islands (FB 30019) has a small terminal bud (figures in parentheses in the description above) and rather small leaves, and its pedicels are ca. 1 cm. long (usually less than 7 mm. in other specimens). However, when compared with the Papuasian specimens, it agrees well in other vegetative characters, and its submature fruits seem to be of the same general type—albeit with a slightly thinner outer layer. Tas_e 15. Comparison of differences between C. laticostatum, Calophyllum sp. 129, and C. archipelagi. C. laticostatum Calophyllum sp. 129 C. archipelagi TERMINAL BUD LENGTH (mm.) (8-)13-27 9-13 6-7.5 TWIG THICKNESS (mm.) 2-5.5 1.5-3 (1.3-)2-3(-3.5) EAF PETIOLE WIDTH (mm.) (1.2-)2-3.5 1-2(-2.5) 1-2 LAMINA TEXTURE Very coriaceous Coriaceous Coriaceous VENATION DENSITY (veins /5 mm.) (9 to) 11 to 20 6 to Il (4 or) 5 to9 LOowEST INTERNODE OF INFLORESCENCE, LENGTH (mm.) 3-11 12-18 8-15 TEPAL NUMBER 4to6 Unknown 8 PEDICEL LENGTH (mm.) 2-8(-11) 7-13 5-16 UIT LENGTH (cm. 1.3-1.9 Ca. 1.4 Caz3 OUTER LAYER THICKNESS (mm. (0.5-)1-3 | aus AIR SPACES DEVELOPING IN OUTER LAYER Under skin Throughout + Throughout WONTIAHdOTVO ‘SNHARLS loge! 18s 582 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 NGF 18373, from Kiunga, Western Province, Papua New Guinea, has leaf blades that are acuminate at the apex. Field notes of this specimen mention ‘‘much branched aerial roots.’’ I have not seen such roots on plants of Calophyllum laticostatum either at Kiunga or on New Britain. 129. Calophyllum sp. FiGure 36, a. C. celebicum Koord. (quoad Koorders 17305) et C. wawaroenti Koord. in Koord.-Schum. Syst. Verzeich. (Fl. N.O. Celebes) 3: 87. 1914. Nomina. Tree to 31 meters tall, d.b.h. to $5 cm.; trunk and bark unknown: latex yellow, pale yellow, or greenish. Twigs slightly flattened, 1.5-3 mm. across, 4-angled, drying mid- to darkish- brown, brown farinose-puberulent when young, axillary innovations lacking basal scars; internodes 0.7-3 cm. long; uppermost pair of axillary buds rounded, ca. 0.5 mm. long, spreading, very inconspicuous; terminal bud plump, 0.9-1.3 cm. long, with subcrustose, grayish brown indumentum (hairs, Figure 37, J, k), underdeveloped internode to I(-5) mm. long. Petiole |.3-1.8 cm. long, broadly concave above, convex below, glabrous: lamina elliptic to subobovate, 9-20 by 2-4.8 cm., acute to subacuminate at apex, narrowly cuneate to subacute at base, rather strongly and closely undulate but not recurved at margin, thinly coriaceous, drying umber to sabelline-olivaceous above and umber to sabelline below, glabrous when mature, the midrib above narrowed gradually from base, broadly depressed, 0.35-0.6(-1) mm. wide at midpoint, below raised, subangled to striate, the venation usually apparent on both surfaces, raised, 6 to I! veins/5 mm., angle of divergence 65-—75(—80)°. Infructescences from foliate axils, with scars of 11 to 15 flowers, unbranched, the axis to 5.5 cm. long, sparsely puberulent toward base, lowest internode |.2-1.8 cm. long; bracts unknown; pedicels 0.7-1.3 cm. long, glabrous. Flower unknown. Fruit subspherical, ca. 1.4 by 1.4 cm., rounded at apex, drying vinaceous- brown, smooth; outer layer not detaching cleanly from stone, (?)thin, com- pletely disorganized by large air spaces; stone ellipsoid, ca. 1.2 by | cm., minutely apiculate, the walls ca. 0.5 mm. thick, smooth, unmarked: spongy layer thin. DistripuTion. Celebes, Moluccas (Soela Island), Poeloe Laut off southeastern Borneo (Map 39). SPECIMENS SEEN. Borneo. KALIMANTAN. Selatan: Poeloe Laut, Sei Paring, 100 m., 6b 13248 (Bo). Celebes. SuLawesi: Minahassa, Menado, by Bivak Totok, Ratatotok, 200 m., Koorders 17305 (Bo, 1), bij Kajoewatoe, 200 m., Koorders 17293 (go, L), Gorontalo, Poso, 100 m., bb 19445 (A, BO, L, SING). Sacasar: Ond. Saleier, Lembang?, eil. Djampia, 245 m., bb 22947 (1). Moluccas. Mancote: Kimakol, Lampaoe, 30 m., bb 29806 (a, Bo, L, SING). Ecotocy. (?)Forest, 30-200 m. alt. Fruiting January (almost ripe); fruit e Calophyllum sp. 129 can be recognized by its only slightly flattened twigs; its large, plump terminal buds; and its rather thinly coriaceous, usually elliptic 1980] STEVENS, CALOPHYLLUM 583 lamina. The midrib on the upper surface of the lamina is broadly depressed and narrows gradually from the base. The pedicels are relatively long and slender, and the fruits are subspherical, with the outer layer probably thin and completely disorganized by air spaces. Calophyllum sp. 129 is closely related to C. laticostatum and C. archipelagi, but can be separated from them by the differences given in TasLe 15 130. Calophyllum archipelagi P. F. Stevens, sp. nov. Ficure 33, d. A speciebus aliis Calophylli in costa in pagina superiore laminae depressa, lamina tenuiter coriacea, axe inflorescentiae 2.5—6.5 cm. longo, tenui, et fructu submagno circa 3 mm. longa, differt. Tree 10-30 meters tall, d.b.h. to 55 cm.; outer bark dark brown, rough; inner bark red-brown. Twigs flattened, (1.3—-)2-3(-3.5) mm. across, + strongly 4-angled, drying blackish to brown, brown-farinaceous when young; axillary innovations lacking basal scars; internodes (0.5-)1-5 cm. long; uppermost pair of axillary buds rounded, less than | mm. long, suberect, inconspicuous; terminal bud plump to conical, 6-7.5 mm. long, with grayish to ferrugineous, subcrustaceous indumentum (hairs, Ficure 37, i), underdeveloped internode to I(-6) mm. long. Petioles 0.7-1.6 cm. long, broadly concave above, convex below, farinose when young; lamina elliptic, 4.6-15 by 2.3-5.1 cm., + acute at apex, acute to narrowly cuneate at base, strongly but distantly undulate and slightly recurved at margin, coriaceous, drying sepia to olivaceous above and cinnamon to sepia below, very transiently sparsely farinose on midrib below, the midrib above gradually narrowed from base, depressed, 0.35-0.7 mm. wide at midpoint, below raised, angled toward apex and striate toward base, the venation apparent on both surfaces, raised, (4 or) 5 to 9 veins/5 mm., angle of divergence 65-75(-80)°. Inflorescences from foliate axils along twigs, with 9 to 17 flowers, usually not branched (rarely with 3-flowered branches to 1 cm. long), lowest flowers sometimes flabellately arranged, the axis 2.5—-6.5 cm. long, sparsely farinose, lowest internode 0.8-1.5 cm. long; bracts not known; pedicels 0.5-1.6 cm. long, sparsely farinose. Flower known only in bud, (?)hermaphroditic; tepals 8, the outer pair broadly ovate, ca. 5 by 4.5 mm., the inner ones + glabrous; stamens 180 to 190, the anthers oblong, 1.3-2.2 cm. long, retuse at apex; ovary ca. 1.5 mm. long, the style ca. 2.7 mm. long, the stigma peltate, ca. | mm. across. Fruit ellipsoid to ovoid, ca. 3 by 2 cm., rounded at apex, drying brown to vinaceous, smooth to broadly and irregularly wrinkled; outer layer not detaching cleanly from stone, ca. 0.6 mm. thick, with large air spaces developing; stone ellipsoid, 2.1-2.4 by 1.2-1.8 mm., rounded at apex, the walls 0.6-0.9 mm. thick, smooth, unmarked; spongy layer (?)thin. Tyre: Java, Kangean, G. Eteng bij Tambajangan, 30 March 1919, Backer 27827 (holotype, Bo; isotypes, A, Bo (several), L). DistripuTion. Kangean Island off northeastern Java, the Lesser Sunda Islands (Map 39) 584 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 ADDITIONAL SPECIMENS SEEN. Java and adjacent islands. Timur: Kangean Is., G. Eteng, Tambajangan, Backer 27596 (Bo). Lesser Sunda Islands. SumBawa: Mangkar Moenteh, 70 m., bb 14016 (Bo), 350 m., bb 11477 (Bo). Komopo: sine loco, Saakov 39 (po). Flores: Maomere, Managadjoeng, 500 m., bb 9654 (Bo); W. part, near Reo, Kostermans & Wirawan 921 (aau, G, L); Nunang, See, 650 m., Schmutz 775 (tL). Ecotocy. Hilly ground (sometimes limestone) or ground flooded during rainy season, 70-650 m. alt. Flowering (almost open buds) April and May; submature fruit August and October. Calophyllum archipelagi can be recognized by its elliptic leaves with a broadly depressed miirib on the upper surface, its relatively short (less than 8 mm.), plump terminal buds, and its rather large (ca. 3 cm.) fruits with thin outer and woody layers. The specific epithet alludes to the fact that all the specimens of this taxon have been collected from the Sunda Archipelago. Calophyllum archipelagiis perhaps related to C. laticostatum (from Papuasia) and Calophyllum sp. 129 (mostly from the Celebes and Moluccas). Both of the latter taxa have less flattened young twigs and older twigs that dry striate (vs. more or less terete as in C. archipelagi). Other differences between these taxa are given in Tas_e 15 Two specimens of Calophyllum archipelagi have fruits. The fruits of Saakov 39 dried vinaceous, and the lamina margin dried much paler than the rest of the leaf; bb 9654 has fruits that dried brown, although they are identical in structure to those of Saakov 39, and the lamina margin did not dry paler than the rest of the leaf. In this latter character bb 9654 is like the other specimens cited above. Schmutz 775 is a sterile specimen, perhaps collected from a sapling. The midrib is narrowly depressed only at the base of the lamina. 134; aoe an Lauterb. Bot. Jahrb. 58: 11. 1922; P. F. Stevens, ral. Jour. Bot. 22: 386. 1974. Synrypes: Neu-Mecklenburg [New eree eer Jan. 1909, Peekel 781, Namarodu, Jan. 1909, Peekel 132; Key Inseln, Pulu Ubur, Warburg 20041 (all p, destroyed). Ficure 6, h, 1. C. kajewskii A. C. Smith, Jour. Arnold Arb. 22: 353. 1941; F. S. Walker, Forests Brit. Solomon Is. Prot. 123. 1948; T. C. Whitmore, Guide Forests Brit. Solomon Is. 77. 1966, Gard. Bull. Singapore 22: 9. 1967; Foreman, Check List Vasc. Pl. Bougainville, 42, 84. fig. 1974. Type: Bougainville, Buin, Koniguru, 800 m., 6 Aug. 1930, Kajewski 2024 (holotype, a; isotypes, BO, BRI, C, G, L, NSW, P, SING). Tree 20—40(—?63) meters tall, d.b.h. to 180 cm.; trunk sometimes with small, thick buttresses or spurs to 1.6 meters tall; outer bark dark brown, deeply fissured, scaling, thick, the inner surface blackish, or brownish black and yellow mottled; under bark pink-brown to deep red; inner bark pink-brown to deep red; latex clear (opaque) yellow, very sticky. Crown broad, branches spreading, leaves suberect. 1980] STEVENS, CALOPHYLLUM 585 Twigs not to somewhat flattened, 3.8-6.5 mm. across, strongly 4-angled to alate (rarely obscurely angled), drying dark brown, glabrous or sparsely transiently farinose when young; axillary innovations lacking basal scars; internodes (0.5-)1-7 cm. long; uppermost pair of axillary buds rounded, less than 1 mm. long, spreading, inconspicuous; terminal bud plump, 0.9-1.5 cm. long, with crustaceous, grayish brown indumentum (hairs, Ficure 37, |, m), underdeveloped internode 2-5 mm. long. Petiole 1.1-2 cm. long, broadly concave above, convex to subangled below, glabrous when mature; lamina obovate to oblong or subelliptic, 8.5-17.5(-21.5) by 3.5-10 cm., rounded to shallowly retuse at apex, acute at base, distantly undulate and not recurved to slightly so at margin, rather thickly coriaceous, drying sepia to vinaceous- buff above and sepia to sabelline below, sparsely farinose on midrib when young, soon glabrescent, the midrib above narrowing rather gradually from base, slightly raised, center + sulcate, below raised, angled to striate, the vena- tion apparent above and below, raised, 6 to 9 (to 12) veins/5 mm., angle of divergence 65-75°. Inflorescences from foliate axils, with 7 to 21 (to 31) flowers (with 5-flowered branches to 1.7 cm. long), the axis 1.5-8 cm. long, glabrous (rarely farinose toward base), basal internode (0.1—-)0.4—1.1(-2) cm. long; bracts ovate, to 4 mm. long, soon deciduous; pedicels 0.7—1.5(-2) cm. long, glabrous. Plants (?)dioecious; tepals 8, sometimes glabrous, the outer pair broadly ovate, 4-5 by 3.5-5.5 mm., the inner ones elliptic to obovate, 4.5-7.3 by 6-6.3 cm.; stamens 210 to 290, in staminate flower the filaments to 6 mm. iong, the anthers oblong, 1.7-2.5 mm. long, apiculate, in pistillate flower filaments to 2 mm. long, anthers I-1.5 mm. long; ovary ca. 2.5 mm. long, the style 2.7-3.5 mm. long, the stigma peltate, in pistillate flower ca. 4 mm., or in staminate flower ca. 2 mm. across, irregularly dentate at edge. Fruit spherical to ovoid, 4.5-7 by 4.2-6 cm., rounded to obtusely pointed at apex, drying brownish, with broad, shallow, longitudinal wrinkles; outer layer detaching cleanly from stone, 2-4 mm. across, compact; stone spherical to + ovoid-ellipsoid, 3.8-6 by 3.7-5.2 cm., rounded to obtusely pointed at apex, the walls 3-5 mm. thick, smooth, unmarked, basal plug to 7 mm. across; spongy layer thin. DistriBuTION. Japen Island; scattered on mainland New Guinea; New Britain and New Ireland to the Solomon Islands (Map 41). SELECTED SPECIMENS SEEN. Papuasia: see Stevens, Joc. cit. Also, Papua NEw Guinea. Gulf: Aird [Kikori] R., [7. F.] Bevan s.n., MEL 75535 (fruit only) (MEL). Eco.ocy. Usually in well-drained primary lowland rainforest, also in season- ally inundated rainforest, swamps, forest growing over limestone; to 311 m. alt. Flowering (or late bud) February, April, August, September, and December (flower scented); fruiting January, February, June, September and November (fruit dull green or blue-green (NGF 18330), perhaps eaten by flying foxes (LAE 66044)). Details of regeneration of Calophyllum peekelii on Kolombangara (the Solomon Islands) are given by Whitmore (1974, as C. kajewskii): dispersal 586 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Map4l. Distribution of Calophyllum peekelii (squares) and C. collinum (triangles) in Papuasia of the seed is poor (the fruits are reported to be blue-black—see above), and development of trees from seedlings takes place in high forest or in gaps. The growth rate of the tree seems to increase with size (Whitmore, 1974, fig. 7.2). There are small, crateriform galls on the upper surface of the leaf, mostly near the midrib, in NGF 39344 GERMINATION AND YOUNG PLANT, The radicle pushes out the basal plug from the stone during germination. The seedling has three pairs of leaves separated by well-developed internodes; the lowest pair is only ca. | cm. long and soon drops off. Subsequently produced internodes are also well developed, growth is erect, and the terminal bud is functional. (NGF 12352; Stevens, LAE sheet no. 125333.) LocaL NAMES AND USES. ‘‘Baula,’’ ‘‘bau’ula’’ (Kwara’ae, Solomon Islands). The wood is strong and is excellent for building canoes or houses. The bark can be removed from the log, dried, and used as a fuel; the flammable latex in the bark ensures a good blaze. Calophyllum peekelii is a very distinctive species with its stout, strongly four-angled twigs, its obovate to oblong, coriaceous leaf blades that are rounded to retuse at the apex, and its large fruits the size of a small orange. The fruits have a thick-walled stone with a basal plug that is pushed out during germination. The specific epithet commemorates the missionary G. Peekel, who lived in New Guinea for over forty years. There is little variation within Calophyllum peekelii, but as with the other possibly dioecious taxa, more material and field observations are needed to seme a her or not the species is dioecious or has some other breeding sy Saplings - Calepiiliin peekelii have leaves with blades up to 42 by 14 cm. that are rather abruptly acuminate at the apex. 1980] STEVENS, CALOPHYLLUM 587 The only record of Calophyllum peekelii from the Gulf Province of Papua New Guinea is based on detached fruits. However, as noted above, these are very distinctive. A neotype of Calophyllum peekelii is not selected since it is not clear that all of the syntypes have been destroyed. The correct application of the name C. peekelii is discussed in Stevens ae Cit.) 132. Calophyllum leleanii P. F. Stevens, sp. nov. Ficure 40, e-g. C. solomonense A. C. Smith, Jour. Arnold Arb. 22: 346. 1941, typo excepto; T. C. Whitmore, Guide Forests Brit. Solomon Is. 78. 1966, pro parte, Gard. Bull. Singapore 22: 9. 1967, pro parte; P. F. Stevens, Austral. Jour. Bot. 22: 398. 1974, pro parte. C. grandifolium Koord. ex Koord.-Schum. Syst. Verzeich. (Fl. N. O. Celebes) 3: 87. 1914. Nomen. A speciebus alis Calophylli in gemma terminali aliquanto longo, lamina plerumque 10-28 cm. longo et 4.5-10 cm. lato, nervis lateralibus manifestis, et fructu strato exteriore 0.6—2 mm. crasso, putamine parietibus |.3-2 mm. crassis et obturamento circa 10 mm. in transverso proviso, differt. Tree 8-25 meters tall, d.b.h. to 54 cm.; trunk without buttresses; outer bark orange to yellowish when young, becoming brown to gray, scaly, the inner surface yellow to orange; under bark red; inner bark red; co yellow, clear or opaque Twigs flattened: (3-)4-6 mm. across, slightly 2-, 4-, or 6-angled, with obscure horizontal lines at nodes, drying brown, farinose-puberulent when young; axillary innovations lacking basal scars; internodes (0.7—)1.5-l10 cm. long; uppermost pair of axillary buds subacute, 1-6 mm. long, spreading to suberect; terminal bud often narrowly conical, 0.8-2 cm. long, with brown, crustaceous to adpressed indumentum (hairs, Ficure 37, n, 0; cf. 37, d), underdeveloped internode to 4.5 mm. long. Petiole |.1-2.5 cm. long, concave above and convex to angled below, puberulent when young; lamina elliptic or ovate to suboblong, (5.5-)10-28 by (2.5—)4.5-10 cm., obtuse to short-acuminate at apex, acute to cuneate at base, narrowly recurved and undulate at margin, coriaceous, drying umber to grayish sepia above and fulvous to olivaceous below, sparsely puberulent on midrib below when young, the midrib above abruptly narrowed at to gradually narrowed from base, + raised, center sulcate, 0.2-0.9 mm. wide at midpoint, below strongly raised, angled, the venation apparent on both surfaces, especially so above, raised, latex canals sometimes raised, 5 to 10 veins/5 mm., angle of divergence 50-80°. Inflo- rescences from foliate axils near ends of twigs, with 5 to 11 flowers (with 3-flowered branches to 3 mm. long), the axis 0.3-2.5 cm. long, subglabrous, lowest internode 1-10 mm. long; bracts not known; pedicels 0.8-1.7 cm long, glabrous. Flower staminate or hermaphroditic; tepals 4, 7, or 8, glabrous or almost so, the outer pair suborbicular, 6-7.5 by 5.5-8 mm., fleshy, strongly concave, the next pair suporpicular, ca. 7 by 6 mm., the inner ones spathulate- lingulate, 8.5-11 by 3-4.2 mm.; stamens 330 to 420, the filaments to 7 mm. 588 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 long, the anthers elliptic-oblong, 0.8-1.5 mm. long, retuse at apex; ovary in hermaphroditic flower ca. 2 mm. long, in staminate flower absent, the style ca. 2.7 mm. long, the stigma peltate, ca. 0.9 mm. across. Fruit subspherical, 1.8-3.5 by 1.6-3.1 cm., + apiculate, drying grayish brown, wrinkled; outer layer not detaching cleanly from stone, 0.6-2 mm. thick, to 4 mm. thick when fresh, compact; stone spherical to ellipsoid, 1.6—2.5 by 1.4-1.9 cm., rounded to obscurely apiculate at apex, the walls 1.3-2 mm. thick, to 3 mm. thick at base, smooth, unmarked, basal plug ca. 10 mm. across; spongy layer thin. Type: Papua New Guinea, West New Britain District, Hoskins Sub-District, Mt. Lollo, 1260 feet (386 m.), 13 Feb. 1971, LAE 51145 coll. Lelean & Stevens (holotype, a; isotypes, K, L, LAE). DistRiBuTION. Scattered: Sulawesi (probable), New Britain, and the Solomon Islands (Map 40). ADDITIONAL SPECIMENS SEEN. Celebes. SuLAwesi: Minahassa, Koorders 1729] (BO, LL, P), 17298 (Bo, K, L, P); Sungei Sadaunta, Musser et al. S-20a (a). Papuasia. Papua New Guinea. New Britain: Kandrian Subdistr., Fullerborn village, 60 m., NGF 21788 (a, canB, L, LAE); road W. of Fullerborn Harbour, about | mile, 100 m., NGF 12957 (L); near Kandrian, Stevens s.n., LAE sheet no. 140450 (Lae); Mt. Lollo, NGF 6607 (a, Bo, CANB, K, L, LAE), 30510 (LAE). SOLOMON IsLANpDs. Guadalcanal: Makina area, Marau, 270 m., BSIP 9458 (k, L); Rere R., BSIP 2794 (x, L), 3317 (L); Sandfly Is., Florida, 45 m., BSJP 18090 (sinc). Santa Isabel: Tataba, 50 m., Brass 3447 (a, BRI, L); '/2 mile due W. of Tatamba, BS/P 2580 (xk, L); Nahao Bay, 51 m., BSIP 7308 (K, L). Ecotocy. Locally common in primary slope and ridge forest; on Santa Isabel on soil derived from ultramafic rock; 45-914 m. alt. Flowering in December; fruiting December and April (fruit greenish). Loca uses. The wood is used for making spears, and a dye is made from the plant (West Nakanai, Hoskins); it is a good timber tree. Calophyllum leleanii can be recognized by its fairly long terminal bud and suberect to spreading uppermost axillary buds; its rather large lamina with the midrib on the upper surface often relatively inconspicuous; its large flowers, which have numerous stamens (more material is needed to confirm this); and its sharply and shallowly wrinkled fruits. The fruits have a fairly thick outer layer and stone wall; the stone has a basal plug. It is a pleasure to name this taxon after Mr. Y. Lelean, who has accompanied me on many field trips in Papua New Guinea. Calophyllum leleanii seems most closely related to C. waliense; this is discussed further under C. waliense. The specimens of Calophyllum leleanii from Sulawesi are included with some hesitation. They are all sterile, but in all details including hair structure (Ficure 37, n, 0) they agree with specimens from New Britain and the Solomons. However, more collections of C. /eleanii are needed, both to confirm its range and to establish the distribution of staminate and hermaphroditic flowers 1980] STEVENS, CALOPHYLLUM 589 in the population (see also Stevens, Joc. cit., as C. solomonense). The type specimen of Calophyllum solomonense has long seemed to me rather different from the other specimens that were included in the species: the terminal bud has tomentose indumentum, with the hairs long and often branched at the base (cf. Ficures 37, n, o, and 14, a—m), and the pedicels of the submature fruits are rather long and slender. In both these and other characters the type agrees with C. soulattri; therefore, C. solomonense is reduced to synonymy under C. soulattri. 133. Calophyllum waliense P. F. Stevens, sp. nov. Ficure 38, e-g. A speciebus aliis Calophylli quibus laminis cordatis habent a costa in pagina superiore laminae depressa et putamine signato parietibus circa 2 mm. crassis obturamento basali proviso, differt. Tree 20-25 meters tall; trunk without buttresses; outer bark cream-brown to dark brown, slightly fissured; latex yellow, clear, stic Twigs flattened, 2.5-5.5 mm. across, 6-angled, soon becoming rounded, with obscure horizontal lines at nodes, drying brown, sparsely brown-farinose when young; axillary innovations lacking basal scars; internodes 1.5-3.5(-7) cm. long; uppermost pair of axillary buds subacute, to 2.5 mm. long, erect to spreading; terminal bud plump to narrowly conical, 0.8-1.3 cm. long, with gray-brown, crustaceous indumentum (hairs, Ficure 37, p), underdeveloped internode absent (-2 mm. long). Petiole 3-5 mm. long, concave above and convex below, glabrous; lamina elliptic to suboblong, (9—)14.5-27 by (3.8-)6- 10.3 cm., rounded to acute at apex, cordate at base, undulate and narrowly recurved at margin, coriaceous, drying dark brick to olivaceous above and sabelline below, farinose on midrib below when young, the midrib above gradually narrowed from base, depressed, 0.5-1.3 mm. wide at midpoint, below raised, becoming raised toward apex, angled (becoming striate toward base), the venation manifest on both surfaces, raised, latex canals sometimes raised, 4 to 7 veins/5 mm., at base to 14 veins/5 mm., angle of ee 75-80°. a eee from foliate axils near ends of twigs, with 5 flowers, unbranched, the axis |.9-3.2 cm. long, flattened and strongly pee glabrous, lowest internode 3-9 mm. long; bracts not known; pedicels 1.3-1.5 cm. long, glabrous. Old flower only known, (?)hermaphroditic; outer tepals broadly ovate, ca. 6 by 5 mm., strongly concave, inner ones obovate, to 12.5 by 4.5 mm.; stamens (?)numerous, the anthers oblong, ca. 1. m. long, subrounded at apex; ovary ca. 2.5 mm. long, the style ca. 2.5 mm. corrugate-wrinkled; outer layer not detaching cleanly from stone, ca. 1.5 mm. thick, compact; stone spherical, ca. 3 by 2.8 cm., subobtuse at apex, the walls ca. 2 mm. thick, smooth, with 4 or 5 longitudinal markings, outer layer persisting over them, basal plug ca. 10 mm. across; spongy layer absent. PE: Papua New Guinea, Bismarck Archipelago, Manus Island, Wali River near Derimbat, 100 m., 3 July 1973, LAE 5241] coll. Foreman (holotype, A; isotypes, CANB, L, LAE). 590 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 DistriBuTION. Papua New Guinea, known only from Manus Island (Map ) ADDITIONAL SPECIMENS SEEN. Papuasia. Papua New Guinea. Manus: ridge peee Liap and Derimbat, 100 m., LAE 59253 (a, 1, Laz), LAE 59255 (A, L, LAE). Ecotocy. Lowland forest on ridges ca. 100(—?550) m. alt. Young fruit July and October; fruit greenish. GERMINATION AND YOUNG PLANT. The radicle pushes out the basal plug. The seedling has two pairs of leaves separated by an internode ca. 1.3 cm. long. Subsequent internodes are well developed, the plant is erect, and the terminal bud is functional. (Seedlings from LAE 52411.) Calophyllum waliense is a distinctive species. Its robust twigs are six-angled, and its terminal buds have furfuraceous indumentum. The lamina is large, cordate at the base, and with a depressed midrib on the upper surface. The fruits are large, and the thick-walled stone has four or five longitudinal stripes and a basal plug. The specific epithet is derived from the Wali River, where the type specimen was found. Calophyllum waliense is superficially similar to C. grandiflorum, from Java, but in the latter species the twigs dry yellowish, rather than brown, the thinner leaf blade lacks a depressed midrib and has less dense venation, and the inflorescence and flowers are much larger. The structure of the fruit is different, and the seed of C. grandiflorum germinates by breaking the stone to one side of the base, the seedling having only a single pair of leaves. Calophyllum waliense is related neither to C. grandiflorum nor to members of the C. venulosum group. The closest relative of Calophyllum waliense is probably C. solomonense: their twigs and terminal buds are similar, and their leaf blades dry a similar color with the very margin often brighter than the rest of the lamina. The stones of both species have thick walls and a basal plug. Calophyllum waliense differs from C. solomonense most obviously in its cordate lamina with a depressed midrib on the upper surface and in its longitudinally marked stone. Calophyllum waliense may grow at 550 meters on Mt. Dremsel, Manus. The sheet of LAE 53644 at Leiden has a shoot of C. waliense (with a number tag) as well as one of C. euryphyllum; all other duplicates of this number that I have seen are of the latter species. 134. Calophyllum euryphyllum Lauterb. Bot. Jahrb. 58: 14. 1922; T. C. ard. Bull. Singapore 22: 12. 1967, pro parte; P. F. Stevens, ral. i Bot. 22: 365. fig. 4. 1974. Type: Nordost-Neu-Guinea sre New Guinea], Lagerberg, Augusta Fluss Station, Sept. 1887, Hollrung 761 (holotype, B (destroyed); isotype, wrsL). C. peekelii auct., non Lauterb.; T. C. Whitmore, Gard. Bull. Singapore 22: 11. 1967, pro parte. 1980] STEVENS, CALOPHYLLUM 59] Ficure 38. a, b, Calophyllum goniocarpum: a, bb 17942, habit, x 0.5; b, bb 33771, terminal bud, x 3. c, d, Calophyllum sp. 143 (NGF 24444): c, terminal bud, x 6; d, habit, x 0.5. e-g, C. waliense (LAE 52411). e, g, stone, < 0.75: e, from outside; g, longitudinal section, plug displaced. f, terminal bud, x 3. 592 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Tree 12-26 meters tall, d.b.h. to 100 cm.; trunk without or with only very short buttresses; outer bark brown, becoming fissured, + scaling, the inner surface dark red; under bark buff, reddish buff, or red-brown; inner bark red; latex clear yellow (rarely yellowish white), sticky Twigs strongly flattened, 2.5-6.5 mm. across, fancied and with tw additional raised lines, drying dark brown, rather persistently farinose to short-tomentose; axillary innovations lacking basal scars; internodes (1-)3- 10(-16) cm. long; uppermost pair of axillary buds pointed, (1—)3—10(-14) mm. long, spreading, supernumerary axillary buds often present; terminal bud subconical, 0.7-1.6 cm. long, with brown, puberulo-tomentose indumentum (hairs, Figure 37, w-y), underdeveloped internode (2-)4-11 mm. long. Petiole 1.6-3.5(-4.2) cm. long, broadly and usually deeply concave above, angled below, usually persistently puberulent; lamina ovate to subelliptic, (6.5—)8.5-19 by (1.5-)5.4-12 cm., subretuse to subacuminate at apex, broadly rounded to cuneate or acute at base, at most slightly and distantly undulate and slightly recurved at margin, coriaceous, drying sabelline to near olivaceous (with grayish covering) above and fulvous to sabelline-sepia below, glabrescent or subpersistently farinose-puberulent on midrib on both surfaces, the midrib above quickly narrowed near base, becoming + raised, the center strongly sulcate at first, 0.3-0.6 mm. wide at midpoint, below strongly raised, angled, the venation + obscure on both surfaces, or subapparent below, slightly raised, latex canals sometimes also slightly raised below, 9 to 13 veins/5 mm., angle of divergence 60-80°. Inflorescences from upper foliate axils, sometimes 2 per axil, with 5 to 15 flowers, usually unbranched (flabellate and/or with 3-flowered branches to 1.2 cm. long), the axis 1.2-5 cm. long, densely puberulent, lowest internode 0.5-1.3 cm. long; bracts ovate, 3-5.5 mm. long, deciduous; pedicels 0.7-1.2 cm. long, densely puberulent. Flower (?)hermaphroditic; tepals 4, the outer pair ovate, 8-9.5 by 6-7.5 mm., densely puberulent on back, the inner pair elliptic-ovate, 8-10 by 7-8 mm., sometimes puberulent in band down back; stamens 70 to 180, the filaments to 6 mm. long, the anthers oblong, 2-3 mm. long, apiculate; ovary to 3 mm. long, densely puberulent, the style ca. 3.5 mm. long, glabrous, the stigma neltate, ca. 3 mm. across, irregularly lobed. Fruit subspherical, 2.8-6 by 2.8-6 cm. rounded at apex, drying snuff-brown, rather closely and shallowly wrinkled; outer layer not detaching cleanly from stone, 2-5 mm. thick, compact; stone spherical, 2.5-5.5 by 2.5-5.5 cm., rounded at apex, the walls 0.5-1. thick, smooth, unmarked; spongy layer thin. DistripuTion. Northern New Guinea, the Bismarck Archipelago, and the Aru Islands (Map 43). SELECTED SPECIMENS SEEN. Papuasia. IRIAN Jaya. Aru: Wakatoebi, P. Oedjir, 20 m., bb 25490 (L, Mo). Vogelkop: Dore, van Hasselt 6 (po). Geelvink Bay: Meos ‘Num, 10 m., BW 15513 (L). Papua New Guinea. Morobe: Umboi Is., Liplip-Mambi R. area, 200 m., LA E 66098 (L, Lar). For additional specimens, see Stevens, Joc. cit. 1980] STEVENS, CALOPHYLLUM 593 Ecotocy. Usually primary rainforest, sometimes over coral; to 610 m. alt. Flowering August and September; fruiting May, June, and November (fruit greenish). GERMINATION AND YOUNG PLANT. The radicle breaks the stone wall to one side of the base. The seedling has two pairs of leaves separated by an internode 0.5-2 cm. long. Subsequently produced internodes are much longer, the terminal bud is functional, and the plant is erect. (LAE 53644, 58539, 59189, Noona Dan Exp. 2014; see also Stevens, loc. cit.) Calophyllum euryphyllum is a very distinctive species with its strongly flattened, usually puberulent twigs; well-developed and spreading uppermost pair of axillary buds; rather large, ovate leaf blades with inconspicuous venation; flowers with four tepals and a puberulent ovary; and spherical fruits that have a compact outer layer and a thick-walled stone lacking a basal plug. The leaf blades are quite broad; hence Lauterbach coined the epithet euryphyllum (Greek—eurys, ‘‘broad,’’ and phyllon, ‘‘leaf’’). UPNG 2019, from the Kiriwina Islands, has the facies of Calophyllum euryphyllum, but the midrib on the upper surface of the lamina narrows gradually from the base, and the flowers have nine tepals. The specimen is rather poor, and further collections are needed to establish the identity of this plant. The relationships of Calophyllum euryphyllum are unclear. It is possibly close to C. papuanum and its relatives, which have very different terminal buds, fruits, and germination; to C. obscurum, which has much smaller leaves and fruits; and to C. costatum, which also has small leaves and fruits that probably have a basal plug. Calophyllum carrii may also be a member of this group; its fruits are unknown, but its leaf has a depressed midrib. Hairs of all of these taxa are much branched and are generally similar. 135. Calophyllum papuanum Lauterb. Bot. Jahrb. 58: 9. fig. 2. 1922; Engler in Engler & Prantl, Nat. Pflanzenfam. ed. 2. 21: fig. 82. 1925; A. C. Smith, Jour. Arnold Arb. 22: 348. 1941; Hartley et al. Lloydia 36: 276. 1973; P. F. Stevens, Austral. Jour. Bot. 22: 382. 1974. Type: Sud-Neu-Guinea [Irian Jaya], sudliches Hellwig-Gebirge, 1450 m., 23 Dec. 1912, Pulle 799 (holotype, B; isotypes, Bo, K, L). Ficures 4, a; 6, a, b. Tree (5—-)16-40 meters tall, d.b.h. to 92 cm.; trunk sometimes with spurs or buttresses to 3 meters; outer bark dark gray to brown, becoming strongly fissured and flaking when mature, the inner surface purplish to red-black; under bark dark red to pink; inner bark dark to pale red; latex clear yellow, very sticky (light yellow, milky). Crown conical at first, becoming irregular, spreading. Twigs strongly flattened, 2-5(-8) mm. across, (rarely 2-) 4- or 6-angled, often with inconspicuous transverse raised line at nodes, drying brown, subpersistently tomentose (rarely farinose-puberulent); axillary innovations 594 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 —— basal scars; internodes |-6 cm. long; uppermost pair of axillary buds + pointed, 2.5-4.5 mm. long, erect; terminal bud strongly flattened, 7-15 mm. long, with brown, tomentose indumentum (hairs, Figure 37, u, v), underdeveloped internode to 3 mm. long. Petiole 0.6-2.5 cm. long, narrowly concave above, angled below, usually subpersistently tomentose; lamina ovate to subobovate, subcuneiform or suboblong, (2.4—)6.5—17(-22) by (1.4—)3.5-8 (-11) cm., subacute to rounded at apex, cuneate to rounded at base, slightly undulate and not recurved to slightly so at margin, coriaceous, drying umber to olivaceous (shiny) above and umber to olivaceous below, when young with indumentum on both surfaces, often persistently subtomentose on lower surface, the midrib above usually narrowing gradually from base, + depressed at first, soon becoming + raised, (0.2-)0.4-0.8 mm. wide at midpoint, below raised, angled, the venation above subobscure, below + apparent, raised, 5 to 12 veins/5 mm., angle of divergence 60-80°. Inflorescences from foliate axils near ends of twigs, with (1 to) 3 to 7 (to 11) flowers, usually unbranched, the axis (0.3-)0.7-1.5 cm. long, short-tomentose to sparsely puberulent, lowest internode 0.3—1.2 cm. long; bracts ovate, to 5.5 mm. long, soon deciduous; pedicels 0.5-1.3 cm. long, puberulent to subtomentose. Plant (?)dioecious: tepals usually 8, the outer pair broadly ovate, 4.5-9.2 by 4.5-9 mm., short- tomentose on back, the inner ones oblong to obovate, 6-14 by 4-12 mm., at least outer pair puberulo-tomentose on back; stamens (70 to) 150 to 300, in staminate flower the filaments 3.5-10 mm. long, the anthers oblong, 2.3—5 mm. long, apiculate, sometimes papillate on apiculus, in pistillate flower filaments ca. 3 mm. long, anthers 0.5-1 mm. long; ovary 2.3-5.5 mm. long in pistillate flower, smaller in staminate flower, tomentose, the style 1.5—2.5 mm. long, the stigma peltate, 3-5.5 mm. across (absent in staminate flower). Fruit + spherical, 2-4 by 2-4 cm., rounded at apex, drying brown, smooth; outer layer not detaching cleanly from stone, 1.5-5 mm. thick, + compact but air spaces developing near stone; stone subspherical, 1.4—3 1.4-3 cm., rounded (obscurely 2- to 4-angled) at apex, the walls |-2.5 mm. thick, barely developed just to one side of base, irregularly pock marked; spongy layer thin. DisrripuTion. Moluccas, mainland New Guinea, and Fergusson and Gooden- ough islands (Map 42). SELECTED SPECIMENS SEEN. Moluccas. Morotai: G. Sangowo, 800 m., Koster- mans 1073 (a, Bo, L). Papuasia: see Stevens, Joc. cit. Also, PAPUA NEW GUINEA. Papuan islands: E. Slopes Mt. Kilkerran (Woiatabu), NE. Fergusson Is., 1600 m., LAE 68957 (a). Eco.ocy. Usually canopy tree of colline or montane forest often dominated by Fagaceae, rarely in more or less swampy forest or depleted Agathis forest over limestone with thick clay cover; (2—)120—1830 m. alt. Flowering January to March, May, July, September, November, and December (flower scented); fruiting January, April, and September (fruit greenish). GERMINATION AND YOUNG PLANT. The radicle pushes through the area of very thin stone just to one side of the base (cf. Stevens, Joc. cit.). The seedling 1980] STEVENS, CALOPHYLLUM 595 Map 42. Distribution of Calophyllum papuanum aati C. vexans (open squares), and C. vexans variant (solid squares) in Papuasia has three (rarely two) pairs of leaves separated by well-developed internodes; the lowest pair of leaves may drop off well before the others. Subsequent internodes are also well developed, the terminal bud is functional, and growth is erect. Germination is similar in fruits with angled and with rounded stones (NGF 10331, LAE 54757, Pullen 1125—rounded;, LAE 58071—angled). Locat use. The wood is used in building. Calophyllum papuanum is a very distinctive species with a flattened terminal bud and medium-sized leaf blades that usually have subpersistent, puberulo- tomentose indumentum over the lower surface. The flower has a tomentose ovary, and the rather large, spherical fruits are 2-4 cm. long and have a pock-marked stone. The specific epithet is taken from ‘‘Papua,’’ which once referred to the entire island of New Guinea. Calophyllum papuanum is related to C. pauciflorum and C. vexans: for the differences separating these species, see TaBLe 16. Plants of Calophyllum papuanum growing at higher altitudes have angled stones; those growing at lower altitudes have rounded ones (Stevens, Joc. cit.). Other interesting variation is shown by specimens from the hills south of the Sepik River. These specimens have rather small, narrow, subovate leaf blades that are minutely rounded at the base; fruits are not known from plants with leaves of this type. 136. Calophyllum pauciflorum A. C. Smith, Jour. Arnold Arb. 22: 348. 1941; Hartley et al. Lloydia 36: 276. 1973; P. F. Stevens, Austral. Jour. Bot. 22: 384. 1974. Type: New Guinea, Mt. Aloki, Yunzaing, 4500 feet [1372 m.], 21 April 1936, J. & M. S. Clemens 2378 (holotype, A) 596 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 C. congestiflorum A. C. Smith, Jour. Arnold Arb. 22: 349. 1941. Type: Netherlands New Guinea [Irian Jaya], 15 km. south-west of Bernhard Camp, Idenburg River, 1800 m., Jan. 1939, Brass & Versteegh 11902 (holotype, a; isotypes, BO, BRI, L, LAE). Calophyllum sp. Lane-Poole, Forest Res. Terr. Papua New Guinea, 117. 1925. Tree 8-36 meters tall, d.b.h. to 48 cm.; trunk not buttressed (fluted at base); outer bark brown to dark brown (light gray), fissured and scaling, the inner surface brown to dark brown; under bark dark red; inner bark reddish brown; latex clear yellow, sticky (gray, cloudy). Twigs strongly flattened, 1.25-3 mm. across, 4- or 6-angled (2-angled when older), with inconspicuous transverse raised line at nodes, drying brown, glabrous or sparsely brown-farinose; axillary innovations lacking basal scars; internodes 0.3-5 cm. long, to 10 cm. long on leader shoots; uppermost pair of axillary buds + pointed, 1-4.5 mm. long, at first inconspicuous; terminal bud strongly flattened, 5-11 mm. long, with brown, farinose-puberulent indumentum (hairs, Ficure 39, a), underdeveloped internode absent (-2 mm. long). Petiole 2-8 mm. long, deeply channeled above, angled below, + glabrous when mature; lamina rhombiform, cuneiform, elliptic, or obovate, 2.5—-5.5 (-9.5) by 0.9-2.5(-4) cm., obtusely pointed to rounded at apex, cuneate at base, slightly undulate or recurved at margin, coriaceous, drying umber to cinnamon-sepia above and fulvous-umber to sabelline below, when young sparsely puberulent on both surfaces, glabrescent or persistently sparsely farinose near midrib below, the midrib above narrowed gradually from base, + depressed at first with margins raised, becoming + raised with center sulcate, (0.3-)0.4-0.55 mm. wide at midpoint, below raised, angled, the venation subobscure on both surfaces, slightly raised (rarely impressed), 7 to 12 veins /5 mm., angle of divergence 50-70°. Inflorescences from upper foliate axils (very rarely terminal), with 3 to 5 flowers, unbranched, the axis 0.3-1.2(-1.5) cm. long, farinose-puberulent at least toward base, lowest internode 2-4 mm. long; bracts ovate to elliptic, 2.5-4.5 mm. long, deciduous; pedicels 2-6 mm. long, glabrous or sparsely farinose-puberulent. Plant (?)dioecious; tepals 8 (rarely 7), the outer pair ovate, ca. 3.75 by 3.5 mm., sometimes farinose on back toward base, the inner ones oblong to obavates to 5.5 by 6.5 mm.; stamens 70 to 125, the filaments to 4.5 mm. long, the anthers suboblong, 0.7-1.7 mm. long, apiculate; ovary 1-2 mm. long, the style ca. | mm. long, the stigma peltate, 1.5-2.5 mm. across, radiate. Fruit spherical, 1.5-1.9 by 1.7 cm., rounded at apex or apiculate, drying brown-vinaceous, smooth or finely striate; outer layer not detaching cleanly from stone, 3-4.8 mm. thick, + compact; stone spherical, |-1.3 by 1-1.3 cm., usually rather obscurely 2- or 3-angled, rounded at apex, the walls 0.7-1.5 mm. thick, thinner just to one side of base, pitted; spongy layer thin. DistripuTIoN. New Guinea, excluding the Vogelkop Peninsula (Map 43). SELECTED SPECIMENS SEEN. Papuasia: see Stevens, loc. cit. EcoLocy. Lower or sometimes upper montane forest, often dominated by TaBLe 16. Some differences between Calophyllum papuanum, C. pauciflorum, and C. vexans. C. papuanum C. pauciflorum C. vexans TERMINAL BUD STRONGLY FLATTENED + + are TWIG THICKNESS (mm.) 2-5(-8) 1.25-3 b3-25 LAMINA LENGTH (cm.) (2.4-)6.5-17(-22) 2.5-5.5(-9.5) 3.4-12.5 INDUMENTUM ON LOWER SURFACE + PERSISTENT, SUBTOMENTOSE +(-) = = VENATION ON UPPER SURFACE OBSCURE + + —(+) FLOWER ANTHER LENGTH, STAMINATE FLOWER (mm.) Zou 0.7-1.7 8-2. Hairs ON PISTIL Dense on ovary, None Sometimes few some on style FRUIT LENGTH (cm.) 2-4 1.5-1.9 1.8-2.7 WOATIAHdOTVO ‘SNHARLS [0861 L6S 598 JOURNAL OF THE ARNOLD ARBORETUM [voc. 61 pia 43. Distribution of Calophyllum euryphyllum (solid circles), C. hirasi- m (open circles), . ee ncraaels (squares), and C. obscurum (stars in solid circles) in Papuas Castanopsis or Nothofagus; 1550-2900 m. alt. Flowering January and April: fruiting January, February, and September (fruit greenish). GERMINATION AND YOUNG PLANT. The radicle probably breaks through the stone wall just to one side of the base. The seedling has three pairs of leaves separated by well-developed internodes. Subsequently produced inter- nodes are longer, the young plant is erect, and the terminal bud is functional. (Pers. obs.) Loca use. The wood is good as a green fuel. Calophyllum pauciflorum is a distinctive species recognizable by its flattened terminal buds and its small, often cuneiform-rhombiform leaf blades that appear glabrous when mature. The midrib on the upper surface of the blade FIGURE 39. Hairs (from terminal bud, unless otherwise noted). a, Caine lum er nei AS 13122), from above. b, c, C. hirasimum, hairs to ca. 420 ym. long: b, BW 14295; c¢, Kanehira & Hatusima 13980. d-f, . pe aa bases birefringent: d, Brass 7385; e, Kalkman 4554: f, Brass 7099. g-j, C. goniocarpum, base or entire hair birefringent: g, LAE 56132; h, NGF 1387; i, Brass 27669; j, LAE 68580. k, s, C. morobense: k, NGF 46766; s, NGF 24490. 1, m, C. trachycaule (Brass 5654), slightly birefringent, walls sometimes rough. n, Calophyllum sp. 143 (NGF 24444), basal projection sometimes lacking. 0, C. suberosum (NGF D719? oo py. persimile (NGF 18303), from stem. q, r, t, C. streimannii: qd; LAL. 52775; t, t, NGF 24285. u, v, C. rufinerve (Kanehira & Hatusima 12218), pep birefringent, apex and base of hair ca. 900 um. long. Scale = 120 w 1, scale = 240 pm.). 600 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61] narrows gradually from the base, and the venation is more or less obscure. The inflorescences have only three to five flowers, and the fruits are less than 2 cm. long. The few-flowered inflorescences asian the specific epithet. Calophyllum pauciflorum is related to C. vexans and especially to C. papuanum; for the differences separating nee taxa, see TaB_e 16. The differences are mostly in the size of parts and the prominence of the indumentum, but there is never any trouble in identifying specimens. Although specimens of C. papuanum from higher altitudes have smaller leaves and fruits with angled stones (and in these characters approach C. pauciflorum), in indumentum development, venation prominence, and midrib type, they are like specimens of C. papuanum from lower altitudes. The anthers of Carr 14143 (C. papuanum, collected at 1585 m. alt.) are 2.3-2.5 mm. long—considerably longer than those of C. pauciflorum. In most characters of wood anatomy studied by van der Graaff and Baas (1974), the two were indistinguishable, except that C. papuanum had wider vessels. The similarities between the three taxa (flattened terminal bud, twig type, branched to almost stellate hairs, more or less well-developed dioecy, mushroomlike stigma, and pocked, often angled, stone) are considerable, and it is of interest to see that the three taxa replace one another geographically and/or ecologically. Although C. papuanum and an imperfectly known form of C. vexans grow within a few hundred yards of each other in apparently similar habitats near Kiunga in the Western Province of Papua, the identity of the form of C. vexans is in doubt (see the discussion after that species). Specimens of Calophyllum hirasimum have been confused with C. pauci- florum; this is discussed further under C. hirasimum. More collections of Calophyllum pauciflorum are needed to evaluate the variation in stamen and pistil size and to ascertain whether or not the species is dioecious (see also Stevens, Joc. cit.). There are a few stamens persisting at the base of an immature fruit on Kalkman 4318; these have anthers ca. 1.7 mm. long—the size of anthers in staminate flowers. However, the anthers have apparently not dehisced, and so the fruits may have been produced by functionally pistillate flowers. 137. Calophyllum vexans P. F. Stevens, Austral. Jour. Bot. 22: 407. fig. 16. 1974. Type: Solomon Islands, New Georgia group, North-west Shortland Island, Kupala Point, 10 feet [3 m.], 7 March 1965, BS/P 13206 coll. Runikera et al. (holotype, LAE; isotype, L). C. gaimanum P. F. Stevens, Austral. Jour. Bot. 22: 367. fig. 5. Type: Papua, east bank of Lower Fly River, Nov. 1936, Brass ae (holotype, LAE; isotypes, A, BO, BRI, G, K, L). Calophyllum sp. Lane-Poole, Forest Res. Terr. Papua New Guinea, 117. 1925; F. S. Walker, Forests Brit. Solomon Is. Prot. 123. 1945. C. cerasiferum auct., non Vesque; T. C. Whitmore, Guide Forests Brit. Solomon Is. 77. 1966, Gard. Bull. Singapore 22: 13. 1967. Tree 12-35 meters tall, d.b.h. to 60 cm.; trunk sometimes with buttresses or stilt roots to 1.7 meters tall; outer bark brown, smooth at first, becoming 1980] STEVENS, CALOPHYLLUM 601 fissured and + scaly, the inner surface blackish; under bark red to red-brown; inner bark reddish, fibrous; latex yellow (white or greenish), clear (turning cloudy), sticky. Twigs flattened, 1.5-2.5 mm. across, 4-angled to subalate, with inconspicu- ous transverse raised line at nodes, drying brown, farinose-puberulent at least when young, or subglabrous; internodes 0.5-8 cm. long; axillary innova- tions lacking basal scars; uppermost pair of axillary buds + pointed, |-4 mm. long, erect; terminal bud strongly flattened to conical, (3.5-)6-12 mm. long, with subcrustaceous to puberulent, brown indumentum (hairs, Figure 37, q-t), underdeveloped internode 0.5-3.5 mm. long. Petiole 0.4-1.4 cm. long, + deeply concave above, convex to angled below, glabrous when mature; lamina elliptic to trapeziform, subovate, subobovate, or oblong, 3.4—-12.5 by 1.2-4.7 cm., acute to subacuminate at apex, acute to cuneate at base, or rounded toward and attenuate at very base, not undulate to moderately so and recurved at margin, coriaceous, drying sepia to near sabelline-olivaceous above and sepia to honey below, glabrescent or subpersistently farinose on scure), raised, (6 to) 8 to Il (to 16) veins/5 mm., angle of divergence (40—-)55-75°. Inflorescences from foliate axils (rarely terminal on short, leafy axillary shoots), sometimes 2 per axil, with 3 to 5 (rarely to (?)7) flowers, unbranched, the axis 0.1-1.3 cm. long, subglabrous or farinose-puberulent, lowest internode 1-6 mm. long; bracts ovate, ca. 3.5 mm. long, deciduous; pedicels 2-8.5 mm. long, glabrous or sparsely farinose. Plant (?)dioecious; tepals 4 to 8, the outer pair ovate, 2.7-5 by 2.8-4 mm., sparsely subpuberulent on back toward base, the inner ones elliptic to obovate, 5-6 by 2.5—4.5 mm.; stamens 30 to 105, the filaments 3-4 mm. long, connate at very base, the anthers oblong, 1.8-2.6 mm. long, apiculate, sometimes papillose on apiculus in pistillate flowers; ovary 1-2 mm. long, the style 1.2-1.7 mm long, sometimes with sparse hairs, at least in staminate flowers, the stigma peltate, 1.8-2.5 mm. across, radiate, sometimes absent in staminate flowers. Fruit spherical to ovoid-ellipsoid, 1.8-2.7 by 1.4-2.2 cm., rounded at apex, drying brown-vinaceous, smooth to sharply and shallowly wrinkled; outer layer not detaching cleanly from stone, 1.5-2.75 mm. across, compact; stone ellipsoid to subovoid, (1-)1.4-1.7 by (0.75-)0.9-1.4 cm., rounded at apex, sometimes subangled, the walls 0.7-2.2 mm. across, pocked; spongy layer thin. DistrisuTION. Throughout Papuasia at suitable elevations (Map 42). SELECTED SPECIMENS SEEN. Moluccas. SerAM: Kiandarat, 60 m., bb 25839 i j . ve : Jaya. Geelvink Bay: Eil. Biak, Seroei, 50 m., bb 30923 (L, sinc). Papua New Guinea. Morobe: Umboi Is., Lip Lip- ~Mambi R. area, 200 m., LAE 66100 (K, L, LAE). 602 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Ecotocy. Usually well-drained lowland or colline rain forest, but swampy forest in Western Province; to 900(-1450) m. alt. Flowering November to July (flower scented); fruiting January, April, June, and October (fruit greenish). GERMINATION AND YOUNG PLANT. The radicle emerges through the area of thin wall just to one side of the base of the stone (NGF 12351). The young plant is erect, and the terminal bud is functional. LocaL NAME. “‘Kaumanu’”’ (Kwara’ae, Solomon Islands). Calophyllum vexans can be recognized by its often flattened terminal bud, its angled twigs, and its leaf blades, which are widest near the middle and which have rather distinct venation and a midrib that is prominent on the upper surface and that narrows gradually from the base. The inflorescence is small, and the plants appear to be dioecious; the fruits have a pock-marked stone. Circumscribing the limits of this taxon has been difficult; hence the epithet vexans (‘‘vexing, The differences separating Calepiplian vexans from its relatives, C. Papuanum and C. pauciflorum, are listed in TaBLe 16 Although Calophyllum gaimanum is reduced to synonymy under C. vexans, it could represent a taxon that may have to be recognized formally when the variation within this rather variable species is better understood (see also Stevens, loc. cit.). e type of C. gaimanum and NGF 17736 (from the Western Province), and specimens from Umboi Island, New Britain, and New Ireland (but not from Mussau Island) all have a strongly flattened terminal bud and rather narrowly elliptic leaves. In what are presumably staminate flowers, the pistil is relatively well developed; there may be hairs on the style (e.g., Brass 8337, LAE 51230) or not (e.g., NGF 46049). I have not seen hairs on the styles of pistillate plants (e.g., LAE 52191). NGF 27406, from New Britain, lacks a strongly flattened terminal bud but in other respects agrees with the rest of this group. On Brass 8337 and NGF 17736 the midrib usually disappears up to 5 mm. below the apex of the lamina, but this seems to be the only difference between the Papua specimens and those from the Bismarck Archipelago. Some axillary is cences of Brass 83371 short, leafy lateral shoots; similar inflorescences were noticed on Noona Dan Expedition 1127, collected on Mussau Island. Specimens from the Solomon Islands have a broader lamina and lack an obviously flattened terminal bud; specimens from Irian Jaya have a similar terminal bud, but can be distinguished by other characters such as color of the dried plant and venation density of the lamina. All specimens of Calophyllum vexans collected in Papua (except Brass 8337 and NGF 17736) form another distinct group that has subalate twigs of the form on New Britain is only slightly raised and is yellowish white; the blade is more coriaceous, and the margin is not undulate. bb 25839 (Seram) 1980] STEVENS, CALOPHYLLUM 603 probably belongs to the same group as the Papuan specimens; flowers of the Papuan specimens are not known. There is discussion of further minor variation within Calophyllum vexans in Stevens (/oc. cit.), as well as details of size variation of stamens and pistils in staminate and pistillate flowers. 138. Calophyllum hirasimum P. F. Stevens, sp. nov. Ficure 36, j-l. C. congestiflorum auct., non A. C. Smith; Kanehira & Hatusima, Bot. Mag. Tokyo 56: 560. fig. I. 1940. A speciebus aliis Calophylli in lamina suboblonga, inflorescentiis pauciflori- bus bracteis subpersistentibus foliaceis, floribus cum 4, raro 6, tepalis, et fructu ellipsoideo circa 1.4 cm. longo putamine parietibus circa 2 mm. crassis, differt. Tree 4-18 meters tall, d.b.h. to 30 cm.; bark not known. Twigs flattened, 1-2 mm. across, 4-angled when young, becoming subterete, drying dark brown, subpersistently farinose; axillary innovations lacking basal scars; internodes |-6 cm. long; uppermost pair of axillary buds + rounded, (1-)1.5-2 mm. long, erect; terminal bud conical, 2—6(—9) mm. long, with brown, tomentose to subcrustaceous indumentum (hairs, Figure 39, b, c), underdevel- oped internode to | mm. long. Petiole 5-10 mm. long, usually broadly concave above, convex below, farinose when young; lamina elliptic to oblong-elliptic or obovate, 2.8-6.7 by 1.4-3.2 cm., rounded at apex, acute at base, slightly undulate and recurved at margin, coriaceous, drying dark brick to umber (shiny) above and sepia to sabelline below, the midrib subpuberulent on both surfaces, + persistently so below, the midrib above narrowed near base, raised, 0.07-0.15 mm. wide at midpoint, below raised, angled to striate, the venation subapparent above and apparent below, raised, 9 to 12 (to 15) veins/5 mm., angle of divergence 65—70°. Inflorescences from foliate axils near ends of twigs, with 3 to 7 flowers, not branched, the axis 0.8-2.8 cm. long; + puberulent toward base, lowest internode 0.4—-1.4 cm. long; bracts often foliaceous, to 2.8 by 1.5 cm., subpersistent; pedicels 0.5—1 cm. long, glabrous. Flower (?)hermaphroditic; tepals 4 (or 6), sometimes glabrous, . outer pair suborbicular, 3.5-4 by 3.5-4 mm. (rarely 2.5 by 1.5 mm.), inner ones elliptic to obovate, to 6.5 by 4mm.; stamens 50 to 85, the ae 2.5 mm. long, almost free, the anthers elliptic. ca. | mm. long, retuse t apex; ovary ca. 1.7 mm. long, the style ca. 1.5 mm. long, the stigma 1.4 by 1 - ca. 1.15 by 0.95 cm., rounded at apex, the walls to 2 mm. thick, somewhat thinner at base, smooth, unmarked; spongy layer thin. Tyre: Netherlands New Guinea [Irian Jaya], Vogelkop Peninsula, S. slope Mt. Nettoti, path Anjai-Wekari River, 1750 m., 30 Nov. 1961, van Royen & Sleumer 7896 (holotype, a; isotype, L). 604 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 DistriBuTION. Western New Guinea (Map 43). ADDITIONAL SPECIMENS SEEN. Papuasia. IRIAN Jaya. Vogelkop: Arfak Mts., Angi, by Iray, Lake Giji, 2100 m., Kanehira & Hatusima 13980 (a, Bo): Res. Manokwari, Iraay (Anggi Gigi Meer), 150 m., BW 265 (CANB, L, LAE); Anggi Gita Lake, Mt. Kongremottie, 2040 m., BW 14100 (a, L), Summit of Mt. Misjnuk (= Mesenuk), 2150 m., BW 14068 (a, L), Mt. Tidjei, 2100 m., BW 14295 (a, L), mountain summit, 2100 m., Kostermans 2490 (xo); Ransiki, Moendi-Berg, 1900 m., BW 2249 (cans, KEP, L, LAE). Snow Mountains: Wissel Lake region, Is. Maiarie en Tage riviertje, 1750 m., Eyma 4953 (a, BO, K, L, SING). Ecotocy. Locally abundant small tree of primary Nothofagus-conifer forest, remnant Ericaceae-Myrtaceae-Nothofagus scrub, or secondary forest; 1750- 2150 m. alt. Flowering January, July, and November (floral axis reportedly red); fruiting January and April (fruit purple). Calophyllum hirasimum can be recognized by its small, oblong-elliptic- obovate leaf blades with rather evident venation; its inflorescences, whic have subpersistent, foliaceous branches; and its ellipsoid fruits, which have a thin outer layer that is disorganized by air spaces, and a smooth, thick- walled stone. The specific epithet combines the names of R. Kanehira and S. Hatusima, who made very important collections, including the only fruiting collection of this species, in western Irian Jaya. Although confused with Calophyllum pauciflorum (BW 265 and 2249 were cited under that species with hesitation by Stevens, 1974a), C. hirasimum is not closely related to it. In C. hirasimum the terminal bud is not flattened (vs. flattened in C. pauciflorum) in the plane of the expanded lamina; the young twigs are basically four-angled (vs. two-angled); the lamina is oblong to obovate (vs. rhombiform); the midrib above narrows quickly (vs. gradually) near the base; the inflorescence axis is extended (vs. compact); the bracts are foliaceous and subpersistent (vs. small and deciduous); the fruit is purple (vs. greenish); and the stone is smooth (vs. pock marked). The hairs of the two are also quite different (cf. Figures 39, b, c; and 39, a); C. pauciflorum uppermost axillary buds; in other respects these specimens are not unusual. BW 2249 and Kanehira & Hatusima 13980 have rather dense venation (11 to 15 veins/5 mm.), and these specimens have somewhat shorter leaf blades than the others; both specimens have the inflorescence typical of the species. The description of the fruit above is taken from Kanehira & Hatusima 13980. 139. Calophyllum trachycaule Lauterb. Bot. Jahrb. 58: 13. fig. 3. 1922; A. . Smith, Jour. Arnold Arb. 22: 346. 1941; P. F. Stevens, Austral. Jour. Bot. 22: 405. 1974. Type: Nordost-Neu-Guinea [Papua New Guinea], Sepikgebiet, Augusta-Flussgebiet, Etappenberg, 850 m., 30 Oct. 1912, Ledermann 9572 (isotypes, kK, L). 1980] STEVENS, CALOPHYLLUM 605 Tree 10-20 meters tall, d.b.h. to 52 cm.; spurs and buttresses absent; outer bark light or straw brown, with many shallow fissures, not peeling, the inner surface orange and greenish straw mottled; under bark deep red; inner bark red, with darker spots and lines in outer part; latex white, not sticky (Stevens et al. 910), or copious, yellowish, milky (BW 12877). Twigs somewhat flattened, 1.8-3.5 mm. across, 4-angled or rounded, drying brown when young, often yellowish when older, (sub)persistently brown- tomentose; axillary innovations lacking basal scars; internodes (0.5—)1-6 cm. long; uppermost pair of axillary buds subrounded, 1.5-3(-5) mm. long, erect; terminal bud plump, 6-11 mm. long, with brown, tomentose indumentum (hairs, Ficure 39, 1, m), underdeveloped internode to 1.5 mm. long. Petiole 0.6-1.5 cm. long, narrowly to broadly concave above, convex below, persis- tently tomentose; lamina elliptic or oblong to subovate, (5-)7-14 by 1.5-4.5 cm., subacuminate to rounded at apex, rounded to cuneate or acute at base, obscurely undulate and slightly recurved at margin, coriaceous, drying umber to near sabelline above and near sabelline to sepia below, + persistently tomentose on midrib below (also above), the midrib above gradually narrowed from base, raised, 0.2-0.35 mm. wide at midpoint, below raised, slightly striate, angled toward base, the venation obscure on both surfaces (subapparent below), slightly raised, latex canals often depressed, 12 to 20 veins/5 mm., angle of divergence 70-80°. Inflorescences from foliate or defoliate axils, with 7 to 19 flowers (with 3-flowered branches to 6 mm. long), the axis 0.8-3.7 cm. long, tomentose, lowest internode 3-6 mm. long; bracts unknown; pedicels 2.5-6 mm. long, tomentose. Flower (?)hermaphroditic; tepals 4, the outer pair ovate, 5-6 by 4-5 mm., the inner pair + elliptic, 6-8 by 4.5-6 mm.; stamens ca. 180, the filaments to 3.75 mm. long, the anthers elliptic, 0.5-0.7 mm. long, retuse at apex; ovary 1.5-2 mm. long, the style ca. 2.5 mm. long, the stigma peltate, ca. | mm. across, 4-radiate. Fruit ellipsoid, ca. 2.4 by 2 cm., rounded at apex, drying vinaceous, smooth; outer layer not detaching cleanly from stone, 2.2-3 mm. thick, compact apart from air spaces developing under skin; stone ellipsoid, ca. 1.8 by 1.2 cm., acute at apex, 3- or 4-angled, the walls 1.3-1.7 mm. thick, 2.3-2.8 mm. thick at angles and ca. 0.15 mm. thick just to one side of base, rather rough; spongy layer thin. DistRIBUTION. Occasional in New Guinea (Map 44). ADDITIONAL SPECIMENS SEEN. Papuasia. IRIAN Jaya. Vogelkop: N. slopes of Upper Aifat Valley, between Senopi and Aifatfekaan, W. of Kebar Valley, 1050 m., BW 12877 (?) (a (frag.), L, LAE). PapuA New Guinea. Western: Kiunga, 18 m., Stevens et al. 910 (a). Central: Kubuna, 100 m., Brass 5654 (A, BO, K, LAE, NY, US); Koitaki, 457 m., Carr 12887 (k, L, LAE, NY, SING). EcoLocy. Lowland or colline forest, 18-850 m. alt. Flowering in October; fruiting in November (fruit blue). Calophyllum trachycaule can be recognized by the well-developed, tomen- tose indumentum that covers the inflorescence axis and pedicels (but not the tepals), stem, leaf midrib, and terminal bud; by its leaves, which have 606 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 Map 44. Distribution of Calophyllum goniocarpum (squares), C. trachycaule (solid Stars), C. rufinerve (triangle), C. piluliferum (circles), C. se a (stars in solid circles), and Calophyllum sp. 143 (open star) in Papuasia dense, obscure venation and impressed latex canals; by its short pedicels; and by its medium-sized fruits with angled stones. The epithet trachycaule means ‘‘rough stemmed,”’ an allusion to the indumentum on the twigs. Calophyllum trachycaule is closely related to C. goniocarpum; some dif- ferences between these and other related taxa are summarized in TaB_e 17. All these taxa have leaf blades of a similar texture and color when dry, the margins are barely undulate and slightly revolute, and the venation is dense and somewhat obscure. Flowers are known from three taxa (C trachycaule, C. goniocarpum, and C. rufinerve); all have four tepals. The outer layer of the fruit has a distinctive texture when dry: in young fruit it is compact, with a few prominent, but not dense, fibers; in fruit near maturity it is more or less disorganized by air spaces. In all taxa (except for C. rufinerve, fruit of which I have not seen) the stones are moderately to very strongly angled, the walls are often rough or pock marked, an there is a thin area just to one side of the base (Ficure 6, c-e). Hairs in all taxa are similar, with thick-walled, often verruculose, basal cells and more or less thin-walled apical cells; there is usually a single basal branch, and the hairs are frequently completely birefringent. Plants of both C. heterophyllum and C. goniocarpum vary considerably in size. The limits and number of the taxa in the Calophyllum trachycaule complex are difficult to judge. There is great variation in leaf shape and size and in general robustness of the plant (cf. Ficure 38, a, and 38, c; see especially C. goniocarpum) and considerable variation in fruit size, superimposed on a general similarity in leaf type (which extends to anatomical details of the Tas_e 17. Variation of some characters in Calophyllum trachycaule and its relatives. Calophyllum C. trachycaule C. heterophyllum C. rufinerve C. goniocarpum sp. 143 C. piluliferum TERMINAL BUD LENGTH (mm.) 6-11 6-10(-12) 5-11 -14 2-4 -6 INDUMENTUM Tomentose Crustaceous— Tomentose Furfuraceous-— Subcrustaceous + Subcrustaceous subtomentose short-tomentose AMINA LENGTH (cm.) (5-)7-14 4.5-9.5 6.8-12.5 (4.2-)6-23 2.2-5.2 4.3-8.8 APEX SHAPE Subacuminate- Truncate- Acute Subacuminate-— Acute- Acute- subacute subcuneate bluntly acuminate Veins /5 mm. 12 to 20 6 to 11 (to 14) 12 to 16 9 to 17 10 to 14 12 to 22 INFLORESCENCE TERMINAL INTERNODE SHORTER THAN OTHERS No Yes —_ No No No PEDICEL LENGTH (mm.) 2.5-6 4-16 Ca. 10 0.35-)5-9 5-10 6-12 (-18 in fruit) INDUMENTUM Tomentose None to sparse- “*Pilose”’ None None None puberulent FRUIT LENGTH (cm.) Ca. 2.4 Ca. 1.2 Ca. 0.5 1.7-2.8 Ca. 0.8 Ca. 1.1 WOTIAHdOTVO ‘SNAAALS [0861 LOO 608 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 blade) and fruit structure. More intensive field work, together with knowledge of germination and seedling, may help to clarify the limits of the taxa involved; few collections with flowers are known. Around Kiunga (Western Province), C. trachycaule sensu stricto, the local form of C. goniocarpum, and what may be a form of C. piluliferum grow in close proximity in well-drained forest on ridges, while near Buso (Morobe Province), the local form of C. goniocarpum and Calophyllum sp. 143 apparently grow in the same general area Lamina shape variation in Calophyllum trachycaule has been discussed earlier (Stevens, Joc. cit.). BW 12877 and Stevens et al. 910 have blades that are more or less acute at the apex and base and so approach the type specimen; the specimens from the Central Province have suboblong blades that are more rounded at the apex and the base. BW 12877 is also somewhat similar to the type (and only) specimen of C. rufinerve and is only provisionally included in C. trachycaule. 140. a a pea eae F. Stevens, Austral. Jour. Bot. 22: 371. ma pE: Nederlands Nieuw-Guinea [Irian Jaya], Star Mts., Sibil- -valley, 1200- 1300 m., 26 May 1959, Kalkman 4151 (holotype, LAE; 1sotypes, A, BM, BO, CANB, L). More or less prostrate shrub | meter tall to canopy tree 29 meters tall, d.b.h. to 35 cm.; trunk not buttressed; outer bark brown, smooth: latex yellowish, copious. Twigs flattened, 2-3.5 mm. across, usually obscurely 4-angled, drying yellowish brown, brown-puberulent when young; axillary innovations lacking basal scars; internodes 1-3(-5) cm. long; uppermost pair of axillary buds rounded, to 1.5 mm. long, suberect; terminal bud plump, 0.6—1(-1.2) cm long, with brown, crustaceous to puberulo-tomentose indumentum (hairs, Ficure 39, d-f), underdeveloped internode to 1(-3.5) mm. long. Petiole 0.35-1.1 cm. long, concave above and convex below, subpersistently puberulent; lamina oblong, elliptic, ovate, or obovate, 4.5-9.5 by 2-4 cm., broadly truncate to subacute at apex, rounded to cuneate at base, obscurely undulate and flat to recurved at margin, coriaceous, drying dull brick to sepia above and umber to honey-sabelline below, subpersistently puberulent on midrib above, margin, and especially midrib below, the midrib above narrowing gradually to rather quickly near base, flat or subdepressed at first with margins raised, becoming raised, 0.25-0.3 mm. wide at midpoint, below raised, striate, the venation subobscure above and subobscure to apparent below, raised to depressed, 6 to II (to 14) veins/5 mm., angle of divergence 75-85°. Inflores- cences from foliate axils along twigs, with (3 to) 5 to Il flowers (sometimes flabellately arranged), unbranched (rarely with 3-flowered branches to 0.6 cm. long), the axis 1-3.2 cm. long, puberulent (?glabrous toward apex), lowest internode 0.2-1.3(—1.7) cm. long, uppermost internode often very much shorter than others, terminal five flowers appearing umbellate; bracts not known; pedicels 0.4—1.6 cm. long, glabrous or sparsely puberulent. Flower (?)her- 1980] STEVENS, CALOPHYLLUM 609 maphroditic; tepals 4 (rarely 5), the other pair broadly ovate, ca. 5.5 mm by 3.5-4 mm., the inner pair elliptic-obovate, 6-7.5 by ca. 3.5 mm.; stamens 100 to 110, the filaments to 3 mm. long, the anthers oblong, ca. | mm. long, truncate to slightly retuse at apex; ovary ca. 1.1 mm. long, the style m. long, the stigma excentrically peltate, 0.7-1 mm. across, 4-radiate. Immature fruit subspherical, ca. 1.2 by 1.2 cm., drying maroon, smooth; outer layer soon disorganized by air spaces; stone (?)angled. DisTRIBUTION. Western New Guinea (Map 44); little collected. ADDITIONAL SPECIMENS SEEN. Papuasia. IRIAN JAYA. Vogelkop: N. slopes of upper Aifat Valley, between Senopi and Aifatfekaan, W. of Kebar Valley, 810 m., BW 12831 (?) Ae Snow Mountains: Star Mts., Sibil Valley, 1200-1300 m., Konan 4554 L, LAE, SING). PapuA New GuINneEA. Western: Palmer R., 3 km. below Paclioh Black R., 100 m., Brass 7385 (a, Bo, L). EcoLocy. Variable: shrubs or treelets, in heathy vegetation developed over poor, white, badly drained loam, 1200-1300 m. alt. (Star Mts.); large tree in primary forest, 810 m. (Kebar Valley); canopy tree common on lower ridges, 100 m. alt. (Palmer R.). pened as aad can be recognized by its plump terminal bud and its medium-sized lamina with obscure venation, at least on the upper surface, and cent only 7 to 11 veins/5 mm. The terminal internode of the inflorescence is often notably shorter than the others, or even undeveloped, so there often appears to be a terminal umbel of five flowers. The flowers have four tepals, with the outer pair rather thick. The stone is possibly angled. The epithet heterophyllum was chosen because of the considerable variation in leaf size and shape occurring on a single specimen. The delimitation and relationships of Calophyllum heterophyllum are unclear. Brass 7385 was collected at much lower altitudes and from a much larger tree than the other specimens; it has immature fruits. The margin of the lamina is considerably more recurved than that of the other specimens, but the venation density and inflorescence are similar, and the specimen has a similar facies. The fruits are still very immature, but the center part (stone plus embryo) appears to be angled. This suggests a relationship of C. heterophyllum to the C. trachycaule complex, from which the venation density and inflorescence type most readily distinguish it (TaBLe 17). BW 12831 has crustaceous indumentum on the terminal bud, and there are up to 13 veins/5 mm. Brass 7099 (a, Bo, K, L) was earlier cited as Calophyllum sp. B, aff. C. heterophyllum (Stevens, loc. cit.); it was collected in the same area as Brass 7385. It has elliptic-oblong leaf blades only 2.2—4.5 cm. long, and its immature fruits, which have dried spherical, smooth, and the same color as those of Brass 7385, are less than 8 mm. across. However, it is possibly to be included in C. heterophyllum. The field notes of Brass 7099 mention a spurred trunk and thick, flaky, suberose bark. As with other members of the C. trachycaule complex, C. heterophyllum needs careful study in the field. 610 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 141. Calophyllum rufinerve Kanehira & Hatusima, Bot. Mag. Tokyo 56: 563. fig. 4. 1942; P. F. Stevens, Austral. Jour. Bot. 22: 409. 1974. Type: Dutch New Guinea [Irian Jaya], Dalman, 45 km. inward of Nabire, 400 m., 2 May 1940, Kanehira & Hatusima 12218 (isotype, A). Tree 20 meters tall; bark unknown. Twigs somewhat flattened, |.5-2 mm. across, strongly 4-angled, drying brown when young, yellowish when older, subpersistently brown-tomentose; axillary innovations lacking basal scars; internodes 0.5-3 cm. long; uppermost pair of axillary buds rounded, to 2 mm. long, erect; terminal bud plump, 5-Il1 mm. long, with brown, tomentose indumentum (hairs, Ficure 39, u, v), underdeveloped internode absent. Petiole 0.8-1.3 cm. long, deeply concave above, convex below, tomentose; lamina subovate to elliptic, 6.8-12.5 by 1.9-3.4cm., acute at apex, cuneate to subrounded at base, obscurely undulate and slightly recurved at margin, coriaceous, drying fulvous-umber on both surfaces, subpersistently tomentose on midrib below, the midrib above gradually narrowed from base, raised, 0.2-0.35 mm. wide at midpoint, below raised, striate, the venation obscure above, subapparent below, slightly raised, latex canals + impressed, 12 to 16 veins/5 mm., angle of divergence 75-80°. Inflorescences from (?)defoliate axils, + fasciculate, with | to 3 flowers, axis absent; bracts unknown; pedicels ca. | cm. long, pilose. Tepals 4, glabrous, rounded-elliptic, ca. 2.5 mm. long; stamens ca. 25, ca. 1.2 mm. long; ovary ca. | mm. long, stigma dilated. Fruit spherical, ca. 5 mm. across. DistripuTion. Irian Jaya (Map 44); known only from the type location. EcoLocy. Agathis forest, 400 m. alt. Flowering and fruiting in May; fruit black. Calophyllum rufinerve is a very poorly known taxon that can be characterized by the tomentose indumentum on the terminal bud, twig, and midrib, the pilose pedicels, the fasciculate inflorescence, and the small, spherical fruits only about 5 mm. across. The epithet rufinerve means ‘‘reddish nerved,”’ an appropriate name because of the reddish indumentum that persists on the midrib. The fruits of Calophyllum rufinerve may have angled stones, but only Calophyllum sp. 143 of the C. trachycaule complex has fruits comparable in size to those of C. rufinerve, and that taxon has very much smaller leaves and a different inflorescence (see TaBLe 17). Stevens (/oc. cit.) suggested d and Hatusima’s original description. They cite the date and altitude of collection of the type specimen as March 2 and 500 meters; on the isotype at the Arnold Arboretum, the date is given as May 2, and the altitude as 400 meters. 142. Calophyllum goniocarpum P. F. Stevens, Austral. Jour. Bot. 22: 369. fig. 6. 1974. Type: Papua, Misima Island, Quartz Mountain, 150 m. 1980] STEVENS, CALOPHYLLUM 611 8 Aug. 1956, Brass 27669 (holotype, LAE; isotypes, A, BO, K, L) Ficures 6, c-e; 38, a, b. (Shrub or) tree, to 36 meters tall, d.b.h. to 60 cm.; buttresses and spurs absent, outer bark brown, gray-brown, pale yellowish brown, olive, or yellow-green, with long, shallow fissures or small lenticels, the inner surface yellowish straw; under bark brownish, yellow, or red with yellow on back; inner bark reddish; latex opaque yellow, not very sticky, to pale honey, sticky, or watery, resinous. Twigs slightly flattened, 2-4.5 mm. across, usually rather strongly 4-angled, drying (dark) brown (yellowish when older), with brown, + transient to subpersistent, puberulent to short-tomentose indumentum; axillary innovations lacking basal scars; internodes 0.5—5(-7.5) cm. long; uppermost pair of axillary buds rounded, 1-4 mm. long, suberect to erect; terminal bud plump, 0.5-1.4 cm. long, with gray to brown, furfuraceous to short-tomentose indumentum (hairs, Ficure 39, g-j; cf. 39, d), underdeveloped internode absent (-2.5 mm. long). Petiole 0.4-3 cm. long, + concave above, convex below, glabrescent; lamina obovate to oblong or elliptic, (4.2-)6-23 by (2-)2.5—9.5 cm., obscurely acuminate to retuse at apex, acute to cordate at base, distantly undulate and slightly recurved at margin, coriaceous, drying near bay or umber to near sabelline above to umber, honey, or sabelline below, soon glabrescent or subpersistently puberulent to subtomentose on midrib below, the midrib above + abruptly narrowing at base, becoming + raised, 0.2-0.5 mm. wide at midpoint (disappearing up to 5 mm. below apex), below raised, striate (weakly angled), the venation usually obscure on both surfaces, slightly raised, latex canals sometimes also raised, 9 to 17 veins /5 mm., angle of divergence 60—-80°. Inflorescences from foliate axils, with 5 to 17 flowers (with 3-flowered branches to | cm. long), the axis 1-6 cm. long, puberulent to subtomentose at base (also elsewhere), lowest internode 0.3-3.6 cm. long; bracts ovate, to 5 mm. long, soon deciduous; pedicels (0.35-)0.5-0.9 cm. long, to 1.8 cm. long in fruit, glabrous. Flower ager as ube tepals 4 (rarely 5), the outer pair ovate, 4.5-5 by 3.5-4 ., the inner pair elliptic-obovate, to 6 by 4 mm.; stamens 50 to 180, shes ree to 3 mm. long, the anthers oblong to elliptic, 0.5-2 mm. long, truncate to retuse at apex; ovary 1.2-1.5 mm. long, the style ca. 2.5 mm. long, the stigma peltate, ca. 0.7 mm. across, 3-radiate. Fruit ellipsoid to spherical, 1.7-2.8 by 1.2-2.6 cm., rounded at apex, drying pruinose- to vinaceous-brown, smooth; outer layer not detaching cleanly from stone, 1.5-5 mm. thick, air spaces often developing especially under skin; stone ellipsoid to subspherical, |.4-2 by 0.8-2.6 cm., acute to obtuse at apex, (2- or) 3- or 4- (or 5-)angled, the walls 0.5-1.4 mm. thick, to 2 mm. thick in angles, ca. 0.3 mm. thick just to one side of base, rough or pock marked; spongy layer thin. DistrIBUTION. New Guinea and the Moluccas (Map 44). SELECTED SPECIMENS SEEN. Moluccas. Mororai: subdistr. Tobelo, N. Totodo- koe, 40 m., bb 33914 (a, Bo, BRI, K, L, LAE, NY, P, SING), 30 m., bb 3377] (A, BRI, K, LAE, SING). Papuasia. [R1iAN Jaya. Vogelkop: Warnapi, N. ‘of Ransiki 612 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 (80 km. S. of Manokwari), 10m., Kostermans 4757 (Bo, L). Papua New Guinea. West Sepik: August R., NGF 3837 (2) (BO, CANB, K, L, LAE, ia Morobe: Buso R., 5 m., NGF F 244 74 (A, CANB, LAE, M, SING), Conn 318 (a); Natter Bay logging area, 93 km. SE. of Lae, 100 m., LAE 68580 (a). Southern Highlands: Mt. Bosavi, No. side, near mission station, 700-800 m., Jacobs 8961 (L, LAE, a Western: SW. corner of airstrip, Kiunga, NGF 18312 (a, CANB, L, LAE, M, MO). Milne Bay: Mayu Is., junction of Mayu and Ugat rivers, 330 m., "LAR 56132 (A, CANB, E, K, L, LAE, M, MO, Us); about half way along the N. shore of Milne Bay, NGF 1387 (?) (£, L, LAE, SING). Papuan Islands: Tagula Is., Mt. Riu, 700 m., Brass 27842 (A, k, L, LAE, US), Rambuso, Om., Brass 28042 (a, K, L, LAE, US). Eco.tocy. Well-drained, often colline forest, rarely swamp forest (Koster- mans 4757); 10-800 m. alt. Flowering in August; fruiting March, April, July, August, and October (fruit bluish to blackish, sometimes apparently dull green (LAE 68580)). Calophyllum goniocarpum can be recognized by its plump terminal buds, its usually only moderately well-developed indumentum, its medium-sized to large leaf blades with fine, often rather dense venation, its inflorescence axis at least | cm. long, and its fruits with angled stones. The epithet goniocarpum was chosen because of this last characteristic. Some of the characters by which Calophyllum goniocarpum can be distin- guished from other members of the C. trachycaule complex are given in TasLe 17. Calophyllum goniocarpum is a large-fruited member of the complex and lacks very well developed indumentum. There is considerable variation in the specimens assigned to Calophyllum goniocarpum. Variation in floral characters is poorly known; however, it may be noted that specimens from the Moluccas and the Vogelkop Peninsula (clearly more similar to one another than to any other specimens with angled stones) have flowers with 140 to 180 stamens, while those from the Papuan Islands have 50 to 80. Specimens from around Kiunga have stones that are subspherical in transverse section, the angles being poorly marked; elsewhere in the range of the species, the stones are almost triangular or square in cross section. NGF 1387, from the Milne Bay area, has rounded twigs ca. 1.3 mm. across that dry yellowish, the terminal bud is as short as 3.5 mm., and the lamina is as little as 4.3 cm. long. Although probably belonging to a taxon with angled stones, NGF 1387 is included here mainly for convenience; it has not been included in the description. Possible confusion between Calophyllum brassii and C. goniocarpum 1s discussed under C. brassii. 143. Calophyllum sp. Ficure 38, c, d. Tree 6-10 meters tall, d.b.h. to 10 cm.; trunk without buttresses; outer bark grayish brown, fissured, the inner surface orange-yellow; under bark orange-reddish; inner bark reddish; latex yellow. Twigs flattened, 0.7—1.3(-2) mm. across, + 4-angled, drying dark brown when young, yellowish brown when old, transiently subfarinose; axillary 1980] STEVENS, CALOPHYLLUM 613 innovations lacking basal scars; internodes 0.5-1.5(-3) mm. long; uppermost pair of axillary buds rounded, to | mm. long, erect; terminal bud plump, 2-4 mm. long, with brown, adpressed, subcrustaceous indumentum (hairs, Ficure 39, n), underdeveloped internode absent. Petiole 4-9 mm. long, strongly concave above, convex below, glabrous; lamina elliptic, 2.2-5.2 by 0.5-1.5 cm., acute to subcuneate at apex, cuneate at base, not undulate but slightly recurved at margin, coriaceous, drying umber above and sabelline below, with transient, adpressed indumentum on midrib below (also above), the midrib above gradually narrowed from base, raised, 0.1-0.15 mm. wide at midpoint, below raised, striate or subangled, the venation subobscure above, subapparent below, raised, 10 to 14 veins/5 mm., angle of divergence ca. 65°. Old inflorescences from foliate (rarely defoliate) axils, with 3 to 7 flowers, unbranched, the axis 0.5-1.8 cm. long, puberulent toward base, lowest internode 2-4 mm. long; bracts not known; pedicels 5-10 mm. long, glabrous. Flower not known. Fruit spherical, about 8 by 8 mm., drying purplish brown, smooth; outer layer not separating cleanly from stone, thin, air spaces developing; stone subspherical, ca. 6 by 6 mm., + rounded at apex, 3- (rarely 2-)angled, the walls 0.2-0.25 mm. thick, to 0.5 mm. thick in angles, + smooth, spongy layer thin. DistriBUTION. Known only from Papua New Guinea (Map 44). SPECIMENS SEEN. Papuasia. PapuA NEw Guinea. Morobe: Mt. Kawea, Buso, 800 m., NGF 24444 (a, CANB, E, K, L, LAE, M, SING), 600 m., LAE 52306 (L, LAE). Ecotocy. Rather stunted forest on soil derived from ultramafic rock, 600-800 m. alt. Fruiting in April; fruit blackish. Calophyllum sp. 143 can be distinguished from the other taxa known to have angled stones by its short terminal buds, small, elliptic leaf blades, and very small fruits (see Taste 17). The fruits seen (from LAE 52306) were probably ripe, to judge by both their color and the fact that the embryo filled the stone. Unfortunately, they did not germinate. 144. Calophyllum piluliferum P. F. Stevens, Austral. Jour. Bot. 22: 387. fig. 11. 1974. Type: Papua New Guinea, Western District, Kiunga, 45 m., 8 Aug. 1971, LAE 51772 coll. Streimann & Katik (holotype, LAE; isotypes, A, CANB, L, SING). Tree ca. 18 meters tall, d.b.h. ca. 23 cm.; trunk without buttresses, but pneumatophores perhaps present (Pullen 7531); outer bark light gray, slightly vertically cracked, with few pustules; under bark reddish; inner bark paler red; latex yellow. Twigs flattened, 1-1.5 mm. across, obscurely 4-angled, drying brown when young, yellowish or whitish brown when older, transiently farinose-puberulent; axillary innovations lacking basal scars; internodes 0.5-3 cm. long; uppermost pair of axillary buds rounded, to 1.5 mm. long, + erect; terminal bud plump, 4-6 mm. long, with brown, + subcrustaceous indumentum (hairs, Ficure 614 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 43, a, b), underdeveloped internode to | mm. long. Petiole 0.6-1.2 cm. long, shallowly concave above, convex below, glabrescent, drying blackish; lamina elliptic to suboblong, 4.3-8.8 by 1.4-3.2 cm., acute to bluntly acuminate at apex, attenuate at base, barely undulate but slightly recurved at margin, coriaceous, drying sabelline on both surfaces, + transiently farinose-puberulent on midrib below, the midrib above gradually narrowed from base, raised, 0.1-0.2 mm. wide at midpoint (disappearing just below apex), below raised, striate, venation obscure to apparent, the latter especially below, raised, 12 to 22 veins/5 mm., angle of divergence 65—75°. Infructescences from foliate axils, with scars of 3 to 11 flowers, rarely branched, the axis 0.15-1.6 cm long, glabrous, lowest internode 2-4 mm. long; bracts unknown; pedicels 0.6-1.2 cm. long, glabrous. Flower unknown; few stamens persisting at base of fruit, the filaments to 2 mm. long, the anthers oblong, ca. 0.5 mm., retuse at apex. Fruit spherical, ca. 1.1 by 1.1 cm., rounded at apex, drying pruinose-brown, smooth; outer layer not detaching cleanly from stone, ca. 0.5 mm. thick, with large air spaces developing; stone ellipsoid, ca. 9.5 by 7.5 mm., obtuse at apex, 3- or 4-angled, the walls ca. 0.4 mm. thick, to 1 mm. thick in angles, + smooth; spongy layer thin. DistriBUTION. Southern New Guinea (Map 44). ADDITIONAL SPECIMENS SEEN. Papuasia. IR1AN Jaya. Digul: Asmat region, Erma, O m., BW 6577 (L). Papua New Guinea. Western: Upper Fly R. along the Binge R., 10 m. Pullen 7531 (cans). Ecotocy. Forest in or near swamps, to 40 m. alt. Fruiting in August; fruit blackish red. Calophyllum piluliferum can be recognized by its rather small leaves that are acute to bluntly acuminate at the apex and attenuate at the base, and by its fairly small, spherical fruits with angled stones. Older twigs dry pale brown. The epithet piluliferum was chosen because the round fruits are like small balls (pilus, Latin ‘‘ball’’). Calophyllum piluliferum is related to C. trachycaule; some characters by which it can be differentiated from this and related taxa are given in TABLE lee The leaf blades of BW 6577 are somewhat thin, and their venation is notably prominent; however, it seems that the specimen was collected soon after a flush of growth. There are three collections from Papuasia in which the leaf base is so narrowly attenuate that the petiole appears to be much longer than it actually is (cf. also Calophyllum venulosum var. tenuivenium); they are perhaps close to C. piluliferum. Stevens et al. 913 was also collected at Kiunga, but its pale yellow—drying twigs are about 2 mm. across, and the lamina is apparently 11.5 by 3.4 cm., with the ‘‘petiole’’ ca. 2 cm. long. Two collections from Japen Island, Irian Jaya (bb 30480 (a, L, sinc) and bb 30481 (a, L)), have a similarly long ‘‘petiole’’? but differ in facies; the lamina of bb 30481 is up to 22 by 4.3 cm. More collections are needed to assess the status of these collections. 1980] STEVENS, CALOPHYLLUM 615 145. Calophyllum streimannii P. F. Stevens, Austral. Jour. Bot. 22: 401. fig. 14. 1974. Tyre: Papua New Guinea, Morobe District, 24 km. SW. of Morobe on the Mo River, 300 m., 28 Jan. 1972, NGF 24285 coll. Streimann (holotype, LAE; isotypes, A, CANB, K, L, SING). Tree 15-35 meters tall, d.b.h. ca. 30 cm.; trunk without buttresses; outer bark yellowish to dark gray, with longitudinal fissures; under bark reddish, inner bark cream; latex yellow. Twigs slightly flattened, 2-4 mm. across, + strongly 4-angled, drying dark brown, transiently farinose to subpersistently puberulo-tomentose; axillary innovations lacking basal scars; internodes 0.5-5 cm. long; uppermost pair of axillary buds rounded, to 2 mm. long, suberect, not very conspicuous; terminal bud plump, 5.5-10 mm. long, with crustaceous to puberulo-tomentose indumentum (hairs, Ficure 39, q, r, t), underdeveloped internode to | mm. long. Petiole (0.4-)1.5-2.5 cm. long, concave above, convex below, glabrous or subpersistently puberulent; lamina suboblong to elliptic or subobovate, 5.2-14 by 2.2-6 cm., acuminate to subacute at apex, acute to attenuate at base, rather distantly undulate and sharply recurved at margin, coriaceous, drying sabelline- to hazel-olivaceous above and fulvous-umber to sepia below, glabrescent or with subpersistent indumentum on midrib below, the midrib above narrowing gradually from base, becoming + raised, surrounding blade raised or not, 0.3-0.5 mm. wide at midpoint, below raised, + angled, the venation above subobscure to apparent, below apparent, raised, 6 to 10 veins /5 mm., angle of divergence 65-80°. Infructescences from foliate axils, with the scars of 7 flowers, unbranched, the axis ca. 2 cm. long, persistently puberulent to subtomentose, at least toward base, lowest internode 0.7-1 cm. long; bracts unknown; pedicels ca. 6 mm. long, (?)glabrous. Flower known only in bud, (?)hermaphroditic; tepals 8; stamens ca. 80, the anthers oblong, ca. 1.3 mm. long, acuminate at apex. Fruit ovoid to ellipsoid, 3.6-5 by 2.8-3.8 cm., pointed at apex, drying dark brown, smooth, outer layer detaching cleanly from stone or not, 4-5 mm. thick, compact; stone ovoid, ca. 2.5 1.9 cm., obtuse at apex, the walls 1-1.75 mm. thick, m. thick at base, smooth, unmarked, with basal plug ca. 8 mm. across; spongy layer thin. DistripuTion. Morobe Province, Papua New Guinea (Map 40). SELECTED SPECIMEN SEEN. Papuasia. Papua New GUuINEA. Morobe: Buso, 30 m., LAE 52755 (a, CANB, K, L, LAE). Eco.ocy. Ridges or hillsides, often with dipterocarps, 30-300 m. alt. Fruiting in January; fruit blue. GERMINATION AND YOUNG PLANT. The radicle pushes out a plug from the base of the stone. The seedling has two pairs of leaves separated by an internode of ca. 1 cm. Subsequently produced internodes are longer, the terminal bud is functional, and the plant is erect. (NGF 24285, 24491.) Calophyllum streimannii can be recognized by its suboblong to elliptic, medium-sized to rather large leaf blades that are acuminate at the apex, 616 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 somewhat sharply recurved at the margin, and with only 6 to 10 veins/5 mm. The fruit is large (3.6-5 cm. long) and is pointed at the apex; the outer layer is thick (4-5 mm.), and the stone has a basal plug. The epithet commemorates the collector, H. Streimann. Calophyllum streimannii is a rather heterogeneous taxon. NGF 28084 is included with hesitation; it differs in a number of details from the other specimens (see Stevens, /oc. cit.); the details of the flower bud in the description above are taken from this specimen. Calophyllum streimannii may also occur in the Western Province (Foreman n., LAE sheet 129036, 8 km. N. of Kiunga on road to Rumingae, 30 m (LAE)). Calophyllum streimannii is perhaps close to C. morobense, but it has larger leaves with more distant venation and fruits about twice as large that dry smooth rather than sharply wrinkled. The hairs of the two species are fairly similar (cf. Ficure 39, q, r, t, and 39, k, s) 146. Calophyllum morobense P. F. Stevens, Austral. Jour. Bot. 22: 378. fig. 4 (°C. morobensis’’). Type: Papua New Guinea, Morobe District, Sunkwep road, 220 feet [65 m.], 19 April 1971, NGF 46766 coll. Katik (holotype, LAE; isotypes, a, CANB, FE, G, K, M, SING). Tree to 21 meters tall, d.b.h. to 30 cm.; trunk without buttresses or spurs; outer bark dull greenish with slightly raised, elongated orange patches, or light gray, with shallow, boat-shaped fissures, the inner surface dull orange; under bark dull dark red; inner bark cream to reddish; latex whitish near outer bark, brownish near wood, or yellow (NGF 24490). Branching horizontal, crown o Twigs | ee: ca. 1.5 mm. across, 4-angled, drying dark brown, sparsely puberulent when young; axillary innovations lacking basal scars; internodes |-3.5(-7.5) cm. long; uppermost pair of axillary buds rounded, ca. 0.5 mm. long, spreading, inconspicuous; terminal bud plump to narrowly conical, 2.5-5 mm. long, with brown indumentum (hairs, Ficure 39, k, s), underdeveloped internode absent. Petiole 0.6-1 cm. long, concave above, convex below, glabrous at maturity; lamina elliptic to ovate or suboblong, 4.5-9 by 2-4.5 cm., acuminate at apex, acute to cuneate at base, slightly undulate and not recurved or slightly so at margin, coriaceous, drying bay to cinnamon above and bay to umber below, glabrous at maturity, the midrib above gradually narrowed from base, + depressed at first, eventually becoming slightly raised, 0.1—-0.2(-0.3) mm. wide at midpoint, below slightly raised, striate, the venation subobscure to apparent on both surfaces, slightly raised or not, latex canals sometimes impressed, 13 to 16 veins/5 mm., angle of divergence 75-80(-85)°. Infructescences from foliate axils, with scars of 7 flowers, unbranched, the axis |-1.5 cm. long, glabrous, lowest internode ca. 3 mm. long; bracts unknown; pedicels 5—7 mm. long, glabrous. Flower unknown. Fruit ovoid to subspherical, ca. 2.8 by 2.2-2.8 cm., acute at apex, drying brown, sharply and closely wrinkled; outer layer detaching cleanly from stone or not, 2-3.5 mm. thick, compact; stone ovoid to ellipsoid, 2.2-2.7 by 1.8-1.9 cm., + obtuse at apex, 1980] STEVENS, CALOPHYLLUM 617 the walls 1.3-1.5 mm. thick, smooth apart from shallow, elongated pock marks, probably with basal plug ca. 6 mm. across; spongy layer thin. DistripuTion. Morobe Province, Papua New Guinea (Map 45). SELECTED SPECIMEN SEEN. Papuasia. PapuA New Guinea. Morobe: near Buso Village, 5 m., NGF 24490 (a, CANB, E, K, LAE, M, SING). Ecotocy. Alluvium in lowland rainforest, 5-35 m. alt. Fruiting in April; fruit bluish to almost black. Calophyllum morobense can be recognized by its medium-sized, strongly acuminate leaf blades with a relatively inconspicuous midrib on both surfaces and fairly dense venation (13 to 16 veins/5 mm.). The fruits are relatively large, more or less acute at the apex, and dry strongly wrinkled; the stone walls are 1.2-1.5 mm. thick. Plants referable to this taxon have been collected only from Morobe Province, Papua New Guinea—hence the epithet. It was earlier thought (Stevens, Joc. cit.) that Calophyllum morobense was similar in leaf to C. pulcherrimum and C. floribundum (= C. tetrapterum); however, the midrib on the upper surface of the lamina is not surrounded by raised blade as in C. pulcherrimum, nor is it sharply raised as is usual in C. tetrapterum. Calophyllum morobense does not seem to be particularly closely related to these or any other West Malesian species but is most similar to C. streimannii. For the differences separating these species, see C. streimannii. Map 45. Distribution of Calophyllum macrophyllum (triangle), C. persimile (squares), C. novoguineense (open circles), C. suberosum (solid circles), C. morobense (solid stars), C. insularum (open stars), and C. acutiputamen (star in solid circle) in Papuasia. 618 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 147. Calophyllum acutiputamen P. F. Stevens, Austral. Jour. Bot. 22: 359. fig. I. 1974. Type: Papua, Rossel Island, Abaleti, 300 m., 5 Oct. 1956, Brass 28325 (holotype, Lak; isotypes, A, BO, K, L, US). (?)Large tree; bark unknown; latex reported to be ‘‘not milk Twigs slightly flattened, 2-2.5 mm. across, 4-angled and with two additional raised lines, drying dark brown, with subadpressed indumentum when young; axillary innovations lacking basal scars; internodes 0.5-3(-5.5) cm. long; uppermost pair of axillary buds + pointed, ca. | mm. long, erect and adpressed to terminal bud; terminal bud conical, 7-9 mm. long, with subadpressed, brown indumentum (hairs, Figure 43, c, d), underdeveloped internode to 1 mm. long. Petiole 0.7-1.2 cm. long, narrowly channeled above, angled below, + glabrous when mature; lamina elliptic to subobovate, 5.5-8 by 1.5-2.4 cm., acute at apex, cuneate at base, not undulate and not recurved to slightly so at margin, very coriaceous, drying cinnamon-sepia above and cinnamon below, subpersistently puberulent on midrib below, the midrib above narrowed gradually from base, raised, 0.3-0.35 mm. wide at midpoint, below raised, angled, the venation on both surfaces obscure, slightly raised, 9 to 11 (to 13) veins/5 mm., angle of divergence 55-60°. Infructescences from foliate axils, with scars of 3 to 5 flowers, unbranched, the axis 0.5-I cm. long, with sparse, subadpressed hairs, lowest internode ca. 4 mm. long; bracts unknown; pedicels 5-6 by 3 mm., sparsely puberulent. Flower unknown. Fruit ellipsoid, 4-4.3 cm. by 2.6-2.8 cm., + apiculate, drying brown, with shallow, longitudinal corrugations; outer layer not detaching cleanly from stone, ca. 3.5 mm. thick, with large air spaces developing; stone oe to subobovoid, ca. 3.5 by 1.5 cm., acute at apex, the walls 0.8-1.2 m thick, thinner to one side of base, + smooth, unmarked; spongy layer ae, DistriBUTION. Papuasia; known only from the type collection from Rossel Island (Map 45) EcoLocy. Canopy tree on ridges in colline forest, ca. 300 m. alt. Submature fruit in October. Calophyllum acutiputamen can be recognized by its more or less conical terminal bud; its unbranched, simple hairs; its elliptic, flat-drying leaves with obscure venation; and its relatively large (ca. 4 cm. long), ellipsoid fruits. The outer layer of the fruit is almost disorganized by air spaces although the skin is practically smooth; the stone is very sharply pointed at the apex. This latter feature suggested the specific epithet (acutiputamen means ‘‘pointed stone’’). 148. Calophyllum suberosum P. F. Stevens, Austral. Jour. Bot. 22: 403. fig. 74, pro parte. Type: Papua New Guinea, Western District, Oriomo River, 70 feet [21 m.], 19 Jan. 1959, NGF 10410 coll. White & Gray (holotype, LAE; isotypes, A, BO, BRI, CANB, K, L, SING). Se maga auct., non Lauterb.; T. C. Whitmore, Gard. Bull. Singapore 22: 11. 1967, pro parte. 1980] STEVENS, CALOPHYLLUM 619 Tree 15-35 meters tall, d.b.h. to 60 cm.; trunk with stilt roots to 2 meters tall; outer bark (yellowish) gray-brown, deeply fissured, with flakes or scales, the inner surface straw brown; under bark pale reddish brown; inner bark pale reddish brown; latex clear yellow, very sticky. Twigs flattened, 5-7(-9) mm. across, 4- or obscurely 6-angled, with obscure transverse lines at nodes, drying yellowish, sparsely brown-farinose when young; axillary innovations lacking basal scars; internodes 1-7.5 cm. long; uppermost pair of axillary buds acute (rounded), 2-5 mm. long, + erect; terminal bud narrowly conical, 0.8-1.3 cm. long, with brown, crustaceous to subfurfuraceous indumentum (hairs, Ficure 39, 0; also moruloid), under- developed internode to 2 mm. long. Petiole 2—-3.3 cm. long, broadly concave above, convex below, glabrous when mature; lamina ovate to elliptic, 16-35 by 7.5-14.2 cm., acute to rounded at apex, broadly rounded and finally acute at base, undulate and slightly recurved at margin, coriaceous, drying umber to sabelline-olivaceous above and near sabelline below, transiently brown- farinose on midrib below, the midrib above gradually narrowed from base, raised, surrounding blade raised, 0.4-0.7 mm. wide at midpoint, below strongly raised, + angled, the venation apparent above and especially below, raised, latex canals sometimes also raised, 5 to 8 (or 9) veins/5 mm., angle of divergence 70-80°. Infructescences from foliate axils, with scars of 3 to 5 flowers, unbranched, the axis 1.5—2.8 cm. long, farinose at least toward base, lowest internode 1-1.5 cm. long; bracts unknown; pedicels in fruit 1.53 cm. by 6 mm., glabrous. Flower unknown. Fruit in spirit spherical to ovoid, 8-9.5 by 7-8 cm., rounded to very obtusely pointed at apex, with broad, longitudinal ridges; outer layer not detaching cleanly from stone, 4-7 mm. thick, compact; stone spherical, 6-7 by 6-7 cm., rounded at apex, the walls ca. 0.7 mm. thick, smooth, (?)unmarked; spongy layer ca. 2 cm. thick. DistRIBUTION. Southern New Guinea (Map 45). ADDITIONAL SPECIMENS SEEN. Papuasia. [RIAN JAyA. Mimika: Asmat subdistr., Erma, BW 3231 (cans, L), 6501 hee 6504 (L), 6547 (L). Digul: along R. Digoel, near Koeweh, 5 m., BW 4839 (L). Papua New Guinea. Western: Kiunga, Tuidemasuk Road, NGF 18335 (A, CANB, L, LAE), near airstrip, Stevens et al. 780 (a); Fly R., d’Albertis s.n., anno 1877 (herb. Becc. 1144) (Fi); 32 km. from mouth of Oriomo R., 45 m , NGF 37159 (a, L, LAE, SING); near aly eee 20 m., LAE 60418 a. L, LAE). Gulf: Kikori R., NGF 4551 Ke. hy LAE, SING); Purari R. near Ravi Kavai, Pullen 6466 (CANB), See & Schodde 832 (A, CANB, L, LAE). Ecotocy. Swamp forest; river banks sometimes subject to tidal influence; below 45 m. alt. Fruiting January, March, and November (fruit green, embryo bright purple (NGF 10410; pers. obs.); fruit floats in water (NGF 37159) and may be dispersed by rivers). GERMINATION AND YOUNG PLANT. The radicle breaks the stone just to one side of the base. The seedling has three, or sometimes two, pairs of leaves separated by internodes 4-17 cm. long; the lowest pair may fall off before the others. Subsequently produced internodes are also long, the terminal bud is functional, and the plant is erect. (NGF 18335, Stevens et al. 780.) 620 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Calophyllum suberosum is a distinctive species that can be recognized by its large, ovate, coriaceous leaf blades in which the midrib on the upper surface is surrounded by raised blade. The very large fruits and the yellow-dry- ing twigs are also characteristic. The epithet suberosum (‘‘corky’’) was chosen eee the fruit has a well-developed spongy layer and is apparently able 0 float Calophyiiun: suberosum is superficially similar to C. persimile; specimens of the two species were included in the original description of C. suberosum. The differences between the two species are discussed under C. persimile and listed in TasLe 18. 149. Calophyllum persimile P. F. Stevens, sp. nov. Ficure 40, k. C. suberosum P. F. Stevens, Austral. Jour. Bot. 22: 403. 1974, pro parte. Calophyllum sp. C, P. F. Stevens, Austral. Jour. Bot. 22: 395. 1974. A Calophyllum suberoso, quo ut videtur simili est, in lamina angustiore elliptica-oblonga percrassa apice retusa vel obtusa basi plerumque acuta, trunco radicibus gralliformibus haud proviso, et latice opaco, differt. Tree 20-25 meters tall, d.b.h. to 40 cm.; trunk sometimes with small buttresses; outer bark yellow, or gray, red-brown, and yellowish mottled, or pale brown (rarely brownish black), fissured (scaly), the inner surface bright yellow (brownish under fissures); under bark red-brown or red and pale red mottled; inner bark red to cream; latex yellow, opaque, slightly viscous or not. Twigs slightly flattened, [1.5-]3-6.5 mm. across, strongly 4-angled to subalate, or 6-angled, with obscure transverse lines at nodes, drying brown and shiny when young, later yellowish, transiently and sparsely brown-farinose; axillary innovations lacking basal scars; internodes |-7.5(-9) cm. long; upper- most pair of axillary buds subacute, [l-]2.5-3.5 mm. long, erect; terminal bud plump to conical, [4-]7-9 mm. long, with brown, crustaceous to furfuraceous indumentum (hairs, Ficure 39, p; cf. 39, 0; also moruloid), underdeveloped internode to 2 mm. long. Petiole [l-]1.6-3.7 cm. long, shallowly concave to flat above and concave below, glabrous; lamina elliptic to oblong (rarely subovate), [5-] 10-30 by [2.5-]3.5-9(-11.3) cm., retuse to rounded (rarely obtuse) at apex, acute to decurrent (rarely broadly rounded and ultimately acute) at base, slightly and distantly undulate and not recurved midrib above gradually narrowed from base, raised, surrounding blade raised, 0.3-0.8 mm. across at midpoint, below raised, rounded to angled, the venation subapparent on both surfaces, raised, 4 to 10 veins /5 mm., angle of divergence [S50—] 65-80°. Inflorescences from foliate axils, with ca. 5 flowers, unbranched, the axis 1-2.1 cm. long, farinose toward base, lowest internode 0.4—1.3 cm. long; bracts ovate, 5-6.5 mm. long, farinose-puberulent beneath, subpersistent; pedicels 3-8 mm. long, glabrous. Flower (?)hermaphroditic; tepals 4, glabrous, the outer pair suborbicular, ca. 5 by 6 mm., the inner pair suboblong to 1980] STEVENS, CALOPHYLLUM 621 Ficure 40. a-c, Calophyllum rugosum. a, Poilane 3494, terminal bud, x 3. b, c, Poilane 30978, fruit, x 1: b, longitudinal section; c, from outside. d, C. insularum (bb 30941), habit, x 0.5. e-g, C. leleanii. e, NGF 6607, terminal bud, x 3. f, g, Brass 3477, fruit, x 1: f, longitudinal section; g, from outside. h, j, C. molle (KEP 77929), fruit, x 0.75: h, stone, from apex; j, entire fruit, longitudinal section. i, C. rubiginosum (FRI 8709), fruit, longitudinal section, x 1. k, C. persimile (NGF 18303), terminal bud, x 3. 1, Calophyllum sp. 151 (Stevens et al. 46), terminal bud, Calophyllum sp. 163 (S 19958). m, n, fruit, x 1.5:m, from outside; n, transverse section. 0, terminal bud, x 3. 622 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 suborbicular, ca. 9 by 8 mm.; stamens ca. 90, the filaments to 3 mm. long, the anthers oblong, 2-2.5 mm. long, acute at apex; ovary ca. 2.5 mm. long, the style ca. 2.5 mm. long, the stigma peltate, 3.5-5 mm. across, 3-radiate. Immature fruit ovoid, ca. 3 by 2.2 cm., smooth; outer layer ca. 3 mm. thick; stone not known. Type: Papua New Guinea, Western District, SW. corner of Airstrip, Kiunga, 25 m., IS Sept. 1972, NGF 18303 coll. Streimann & Lelean (holotype, a; isoines: E, CANB, K, L, LAE, M, MO, SING). DistripuTion. Western Papuasia (Map 45). ADDITIONAL SPECIMENS SEEN (* denotes specimen with leaf anatomy examined). Papuasia. IRIAN Jaya. Vogelkop: Warsamson Valley, E. of Sorong, 50 m. BW 12327 (1, Lae*). Geelvink Bay: Japen Is., Aisaoe, bivouac Sebosiar 210 m., BW 9229 (cans, L, ns 180 m., BW 10007 (L, LaE*), , BW 10539 (L, sinc), 200 m., BW 10547 jc, LAE*), Djajapura: ae R. 60 km. SE. of Sarmi, 70 m., BW 8092 (L*, LAE, CANB); Kwansem R., 200 km. W. of Hollandia, 138 m., BW 2378 (L, LAE*, sinc); Cycloop Mts., So. slope of Makanoi Range, W. of Kujabu R., 560 m., van Royen & Sleumer 6561 (A*, CANB, L, LAE); Hollandia, 50 m., bb 25081 (a*, Bo, L), Kostermans & Soegeng 287 (k*, L). Fak-Fak: Anakasi (Babu), 50 m., bb 32697 (L). Papua New Guinea. West Sepik: Vanimo, 30 m., LAE 52952 (Laz*), 30 m., LAE 53835 (L, LAE), VTA 2184 (Lae). Western: Rumginae Road, 8 km. from Kiunga, 45 m., Stevens et al. 808 (a*), 45 m., Stevens et al. 809 (a*), 12 km. from Kiunga, 30 m., Stevens et al. 843 (a*). EcoLocy. Well-drained rain forest (Papua New Guinea); moist valleys, or rain forest inundated in wet season (Irian Jaya); 20-560 m. alt. Flowering in September. Van Royen & Sleumer 6561 has rather irregular spherical galls ca. 2.5 mm. across (the center is raised) near the midrib on the lower surface of the lamina; in Kostermans & Soegeng 287 similar galls occur near the midrib and the margin. GERMINATION AND YOUNG PLANT. The seedling has (two or) three pairs of leaves separated by internodes 1-2.5 cm. long. Subsequently produced internodes are less than 4 cm. long, the young plant is arched, the leaves are held in one plane, and the terminal bud is functional; the stem straightens later. In older plants the internodes are considerably more than 4 cm. long. (Stevens et al. 809.) Calophyllum persimile can be recognized by its usually strongly four-angled twigs; its large, very coriaceous, usually elliptic to oblong leaf blades with the midrib on the upper surface surrounded by raised lamina; and its four-tepaled flowers. The young plant does not grow very fast, and the stem is initially arched. The epithet persimile (‘‘very similar’’) was chosen because of the considerable similarity of dried specimens of this species to those of C. suberosum. When I described Calophyllum suberosum, 1 included in it specimens of the species here described as C. persimile. However, I found plants of two TaBLe 18. Differences between Calophyllum suberosum and C. persimile. Calophyllum suberosum* Calophyllum persimilet STILT Roots £ COLOR OF OUTER BARK AND ITS INNER SURFACE LaTex ¢t LENGTH : BREADTH RATIO UPPER EPIDERMIS THICKNESS OF CUTICLE PLUS OUTER PERICLINAL CELL WALL TOTAL HEIGHT OF CELL (um.) Lower EPIDERMIS THICKNESS OF CUTICLE PLUS OUTER PERICLINAL CELL WALL TOTAL HEIGHT OF CELL (um.) INTERNODE LENGTH IN SEEDLING (cm.)£ YOUNG PLANT GrowTH¢ INTERNODE LENGTH (cm.)+ Large Grayish or brownish Clear yellow, sticky Ovate to elliptic Coriaceous 2.5 Lacking or very small Often yellowish or bright yellow Opaque yellow, + fluid Elliptic to oblong (rarely subovate) ery coriaceous (2-)2.5- 18-26.4 28.6-39.5 16.5—22 22-33 1-2.5 Arching To 4 Hasirat ¢ Swamp or riverside Ridge forest or periodically inundated forest *Leaf anatomy of specimens oes in Stevens (1974a) has been examined. + Leaf anatomy of specimens denoted by an asterisk in specimen citation has been examined. +Based mainly on personal mene at Kiunga. WOTIAHdOTVDO ‘SNAAALS [0861 €79 624 JOURNAL OF THE ARNOLD ARBORETUM [voL. 6] species with large, coriaceous leaves at Kiunga that differed most obviously in bark, seedling, and ecological preferences. Although it is not as easy to separate sterile material in the herbarium, this can be done using the characters listed in TaBLe 18. Calophyllum suberosum and C. persimile have very similar hairs (Figure 39, 0, p), twigs that dry yellowish, and large, coriaceous leaf blades with the midrib on the upper surface surrounded by raised blade. These characters alone distinguish the two species from all others in the genus. However, flowers of C. suberosum, ripe fruits of C. persimile, and observations on the ecology of both species from throughout their ranges are needed to clarify the relationship between them. The differences in the seedling and young plant listed in TasLe 18 are independent of the environment. The first young plant of Calophyllum persimile that I saw was growing in a swamp with numerous seedlings and young plants of C. suberosum. The young plant of C. persimile had the characteristic arching and slow growth of the numerous young plants later seen in better- drained ridge forest. There is some variation within Celophyllum persimile. The specimens cited from Japen Island (previously Calophyllum sp. C, aff. C. savannarum—see Stevens, /oc. cit.) are similar in anatomy to the other specimens from Irian Jaya but are smaller in most of their parts (measurements in brackets in the description above); they are tentatively included in C. persimile. Leaves of C. persimile from the Western Province of Papua New Guinea lack a hypodermis, which is present in leaves of specimens from elsewhere in the range of the species (with the exception of LAE 53835). LAE 53835, from a sapling, has exceptionally long leaf blades (ca. 56 by Il cm.) (Stevens, loc. cit.); it also has a rather thin cuticle plus outer periclinal epidermal wall complex on both the upper (ca. [5 wm. thick) and the lower surface (ca. 10.5 wm. thick) of the lamina. Van Royen & Sleumer 6561 has leaf blades that are bluntly pointed at the apex, and its bark is reported as being blackish brown, but it probably belongs to C. persimile. 150. Calophyllum gracillimum M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 348. pl. 28. 1956; T. C. Whitmore, Tree Fl. Malaya 2: 172. 1973; Corner, Gard. Bull. Singapore Suppl. 1: 104. 1978. Type: Malaya, Pahang, Bukit Balai, Rompin, 9 May 1919, KEP 2715 coll. Lambak (holotype, KEP; isotypes, kK, SING). Tree to 15 meters tall, d.b.h. to 12 cm.; trunk without buttresses; outer bark pale greenish yellow-gray to biscuit colored, or pale brown and orange- brown mottled, with small lenticels in vertical rows, the inner surface dirty brown; under bark reddish; inner bark pale red; latex white or not apparent. Twigs slightly flattened, 0.5-1 mm. across, + 4-angled when young, soon becoming rounded, drying yellowish (rarely brown), transiently farinose to subtomentose; axillary innovations lacking basal scars; internodes 0.5-3.5 cm. long; uppermost pair of axillary buds pointed, to 1.5 mm. long, erect; terminal bud (narrowly) conical, 1.5-4 mm. long, with brown, subtomentose indumentum (hairs, Ficure 43, k), underdeveloped internode absent. Petiole 1980] STEVENS, CALOPHYLLUM 625 2.5-4.5(-7) mm. long, deeply concave above, convex below, glabrescent, often drying blackish; lamina ovate to elliptic, (1.7-)3-7(-10.3) by (0.9-)1.4- 3(-3.8) cm., bluntly acuminate to acute at apex, cuneate at base, slightly undulate but not recurved at margin, thinly coriaceous, drying umber to sepia above and fulvous to sabelline below, glabrescent except for few hairs on midrib below, the midrib above gradually narrowed from base, raised, or level except for raised basal 2 mm., ca. 0.15 mm. wide at midpoint, below raised, subangled to + flat, sometimes with at least margins impressed, the venation obscure on both surfaces, sometimes latex canals impressed above and especially below, 6 to 9 veins/5 mm., angle of divergence 60-70°. Inflorescences from foliate axils near ends of twigs, with | to 3 (to 5) flowers, unbranched, the axis to 2(-14) mm. long, short-tomentose, lowest internode to 2(-10) mm. long; bracts ovate, ca. 3.5 mm. long, soon deciduous (rarely foliaceous, ca. 1.4 cm. long); pedicels 3.5-5(-9) mm. long, short-tomentose. Flower (?)hermaphroditic; tepals 6 (rarely 7), the outer pair broadly ovate to suborbicular, 4-5.5 by 4-5 mm., puberulo-tomentose on back, the next pair elliptic, ca. 6 by 4.5 mm., with + broad strip of indumentum down back, the inner pair suboblong to elliptic, ca. 5. by (?2—) ca. 3.5 mm.; stamens eae the filaments to 3 mm. long, the anthers subelliptic to oblong, . long, retuse at apex, with persistent hairs to 0.1 mm. long; ovary 1.3- es mm. long, stigma and style unknown. Fruit ea cal: ca. 1.1 by 1.1 cm., apiculate, drying vinaceous-brown, closely and sharply wrinkled; outer layer not detaching cleanly from stone, 0.5-0.9 mm. thick, compact; stone spherical, ca. 9 by 9 mm., rounded at apex, the walls ca. 0.2 mm thick, smooth, unmarked; spongy layer thin. DistriBuTION. Malay Peninsula (Map 46). ADDITIONAL SPECIMENS SEEN. Malaya. SELANGor: near the Gap, 800-900 m. Ando et al. 96 (probable) (KEP). gee Bukit Bauk F.R., FRI 2609 (?) (KEP, L, SING). PAHANG: Ulu Perah, KEP 11209 (KeEp); Fraser’s Hill, near Map 46. Distribution of Calophyllum rotundifolium (stars), C. aureo-brun- nescens (circles), Calophyllum sp. 151 (half-circles), C. gracilipes (squares), and C. gracilipes variant (stars in solid circles) in Malesia. Inset: C. gracillimum. 626 JOURNAL OF THE ARNOLD ARBORETUM [voL. 6] Jeriau waterfall, 975 m., Stevens et al. 20 (aA). Jonore: 1312 mile Mawai-— Jemaluang road, SFN 28998 (A, BO, K, KEP, LAE, NY, P, SING). Eco.ocy. Usually well-drained lowland or lower montane forest, also drier parts of swamp forest; to 975 m. alt. Flowering and fruiting February and May. YOuNG PLANT. The plant is erect, and the terminal bud is functional. Calophyllum gracillimum can be readily distinguished from other species of Calophyllum by its pale-drying twigs that often contrast with the blackish- drying petioles, and its small leaf blades that are bluntly acuminate at the apex, cuneate at the base, and with indistinct venation and often impressed latex canals. The inflorescences have one or a few flowers, and the anthers have hairs. The almost smooth, pale bark of the mature tree is also notable. The epithet gracillimum (‘‘very slender’’) is appropriate for this species. Calophyllum gracillimum is without obvious relatives. The small size of all its parts distinguishes it readily from the other taxa known to have hairs on their anthers (C. canum (sometimes); C. molle, and C. rubiginosum). It is superficially similar to the Fijian species C. leu he two can easily separated as discussed under the latter species l lum gracillimum may be close to Calophyllum sp. 151, although the latter lacks hairs on its anthers and differs ina number of vegetative details (most obviously in its rounded twigs, longer petioles, midrib on the upper surface of the lamina surrounded by raised blade, and denser venation). More collections of both taxa are needed, especially from low altitudes in Johore where both occur; fruits of Calophyllum sp. 151 are not known. Both FRI 2609 and Ando et al. 96 have twigs that dry dark brown, and although the latter specimen is probably Calophyllum gracillimum, the former may not be. It has rather prominent venation, an inflorescence with an axis to 1.4 cm. long bearing five flowers, and pedicels up to 9 mm. long. There is considerable variation in the development of the indumentum in C. gracillimum: specimens from Johore are short-tomentose on the twigs, while others are only farinose. The uppermost measurements of the petiole and lamina in the description are nearly always axillary (cf. Henderson & Wyatt-Smith, Joc. cit.), although when borne in the uppermost leaf axils they may obscure the terminal bud, thus appearing to be terminal. 51. Calophyllum sp. Ficure 40, |. Tree 15-21 meters tall, d.b.h. to 25 cm.; trunk without spurs or buttresses; outer bark mottled brown-yellow, to pale brown, almost smooth (hoop marked), lenticels in vertical rows, the inner surface + orange to pale brownish; under bark orange-reddish to orange-green; inner bark pale reddish; latex usually not obvious, slight moistening of cut surface, or colorless, not sticky (rarely opaque yellow, sticky). Twigs somewhat flattened, (1.2—)1.6-2 mm. across, rounded, drying whitish 1980] STEVENS, CALOPHYLLUM 627 to grayish brown, sparsely puberulent when young; axillary innovations lacking basal scars; internodes 1.5—4cm. long; uppermost pair of axillary buds rounded, 0.5-2 mm. long, erect; terminal bud plump to narrowly conical, 2.5-5 mm long, with brown, + adpressed indumentum (hairs, Ficure 43, 1, j), underdevel- oped internode absent. Petiole (0.4-)0.5-1.3 cm. long, broadly concave above, convex below, glabrescent; lamina ovate to subelliptic, (2.5-)4-6(-8.3) by (0.9-)1.7-3.2(-3.5) cm., rounded to subacute and + decurved at apex, acute to decurrent at base, slightly and distantly undulate but not recurved at margin, coriaceous, drying umber to near rosy buff above and sabelline to honey below, puberulent on midrib beneath when young, the midrib above narrowing gradually from base, initially strongly depressed, becoming raised, surrounding blade also raised, 0.2-0.3 mm. wide at midpoint, below raised, + angled, the venation above subobscure, apparent to subobscure below, flat or slightly raised, especially below, latex canals often impressed below, 8 to 13 veins /5 mm., angle of divergence 65-80°. Inflorescences from foliate axils near ends of twigs, with 5 (to 7) flowers, unbranched, the axis 0.6-1.4 cm. long, puberulent, lowest internode 3-7 mm. long; bracts elliptic-ovate, 1.8-2.8 mm. long, deciduous; pedicels 1-6.5 mm. long, puberulent. Flower (?)hermaphro- ditic; tepals 4, the outer pair suborbicular to elliptic, ca. 4.5 by 3.5-4 mm puberulent on back, the inner pair elliptic-obovate, 5-6.5 by 4-5 mm., with puberulent strip down back; stamens 105 to 160, the filaments to 6.5 mm long, perhaps with few minute hairs when young, the anthers elliptic, 0.35-0.6 mm. long, retuse at apex; ovary ca. 1.3 mm. long, the style unknown, the stigma peltate, 0.6-0.8 mm. across, 3-radiate. Fruit unknown DistripuTION. Southern Malay Peninsula, Singapore (Map 46). ee SPECIMENS SEEN. Malaya. JoHore: G. Pulai, 150 m., Stevens et 98 (a); Lenggor F.R., 30 m., Stevens et al. 60 (a); Renggam F.R., FRI Bie (KEP), 240 m., Stevens et al. 46 (a), Stevens et al. 53 (= FRI 23440) (A), KEP 71267 (kep); 42 miles, Mersing Road, Kluang, FR/J 21621 (keEp); Panti F.R., KEP 70320 (?) (ker), 5 m., Stevens et al. 112 (a). Singapore: Mandai forest (by zoo), 30 m., Stevens er al, 719 (a). Ecotocy. Weli-drained lowland, mixed dipterocarp forest; sometimes in seasonally inundated forest; 5-240 m. alt. Flowering in November (all Malayan specimens); flower scented. GERMINATION AND YOUNG PLANT. The seedling has one or two pairs of leaves separated by an internode 5-10 mm. long. Subsequently produced internodes are longer, the terminal bud is functional, and the plant is erect. The leaves of the seedling are 1.5-2.3 cm. wide; those produced in the first and immediately succeeding flushes are about half as wide. (Stevens et al. 46 Calophyllum sp. 151 can be characterized by its pale-drying, rounded twigs and its small leaves with relatively long petioles and blades that are rounded to subacute at the apices. The apex is rather strongly decurved, at least in the living leaf. The midrib on the upper surface of the lamina is initially depressed, but it soon becomes raised and is surrounded by raised blade. The inflorescence axis, pedicels, and backs of the outer and (partly) inner 628 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 tepals are all covered with indumentum; the flowers have four tepals and glabrous anthers. Calophyllum sp. 151 may be related to C. gracillimum, but until both taxa are better known, it is premature to describe the former. The differences between the two are discussed under C. gracillimum. 152. Calophyllum leptocladum A. C. Smith & Darwin, Jour. Arnold Arb. 55: 221. figs. 4, 5. 1974, pro majore parte. Tyre: Fiji, Ngau, slopes of Mt. Ndelaitha, on north spur, toward Navukailangi, 350-500 m., 22 June 1953, A. C. Smith 7874 (holotype, us; isotypes, A, K, NY). Tree 9-22.5 meters tall; outer bark yellow, with narrow, longitudinal cracks. Twigs flattened, 1.2-1.6 mm. across, + 2-angled or rounded, drying brown to yellowish, brown-farinose when young, or glabrous; axillary innovations lacking basal scars; internodes 0.5-2.5 cm. long; uppermost pair of axillary buds rounded (pointed), ca. 0.2(-0.5) mm. long, suberect to spreading; terminal bud narrowly conical, 2-3.7 mm. long, with brown to grayish, puberulent to subcrustose indumentum (hairs, Ficure 43, e-h; cf. 37, 0), underdeveloped internode absent (-1.5 mm. long). Petiole 4-10 mm. long, deeply concave above, convex below, glabrous when mature; lamina elliptic to suboblong, (2.8-)3.6-7(-8) by 1.4-2.4(-3.1) cm., acuminate at apex, acute to attenuate at base, slightly undulate or recurved at margin, thinly coriaceous, drying sepia to olivaceous above with midrib, margin, and venation often paler, umber to olivaceous below, glabrous or sparsely farinose on midrib below, the midrib above narrowed gradually from base, + raised, center strongly sulcate, surrounding lamina usually obscurely raised, 0.15-0.3 mm. wide at midpoint, below raised, subangled, the venation + apparent above and below, raised to subdepressed above, raised below, (10 to) 12 to 18 veins/5 mm., angle of divergence 60-75°. Inflorescences from foliate axils (very rarely terminal), with ca. 5 flowers, unbranched, the axis 0.6—1.5 cm. long, puberulent toward base, lowest internode 0.25—0.8(-1.2) cm. long; bracts narrowly ovate, to 2.5 mm. long, soon deciduous (rarely foliaceous, persistent); pedicels 0.2-1(-1.3) cm. long, glabrous. Flower (?)hermaphroditic; tepals 4, glabrous, the outer pair subelliptic, ca. 3 by 2.5 mm., the inner pair ne to 4 mm. mm.; stamens ca. 60, the filaments to 2 mm. long, the anthers oblong, 0.6-1 mm. long, slightly retuse at apex; ovary ca. 1.3 mm. long, style ca. 1.5 mm. long, stigma peltate. Fruit ellipsoid, ca. 1.6 by 1.35 cm., rounded at apex, drying + vinaceous-tawny, irregularly wrinkled; outer layer detaching + cleanly from stone, 0.5-0.8 mm. thick, compact; stone ellipsoid, 1-1.35 by 0.85-1.15 cm., minutely apiculate, the walls 0.4-0.5(?-0.8) mm thick, smooth, unmarked; spongy layer (?)thin. DistripuTion. Fiji Islands, Viti Levu, rare on Vanua Levu. SELECTED SPECIMENS SEEN. See Smith & Darwin, loc. cit., excl. Howard 104 and FDA 15722. Also, Viti Levu. Namosi: Mt. a ee 671 m., FDA 16127 (cur). TatLevu: Colo-i-Suva, 210 m., Bola I] (k). Navua Kadawa, side of Kadawa Trig. Hill, 150 m., C.S..R.0O. S Pi (K); Sociri 1980] STEVENS, CALOPHYLLUM 629 ee (Kadava), Nabukelova distr., [FDA] H 419 (k). Kanpavu: Kandavu , 150 m., C.S..R.O. S 1404/3 (kK). Ecotocy. Usually colline forest, (?18—)50-670 m. alt. Flowering December and March; fruiting in June (fruit nearly white). Galls—small, narrow projections ca. 1.5 mm. high on the upper surface of the lamina—are common and occur on over half the collections. Loca use. The wood makes good timber (Smith & Darwin, loc. cit.). Calophyllum leptocladum can be characterized by its small, narrow terminal bud and its rather small leaf blades with dense venation. It has small, four-tepaled flowers borne on few-flowered, axillary inflorescences. Its fruits are comparatively quite large (ca. 1.5 cm. long) and have a compact (0.5-0.8 mm.) outer layer that separates cleanly from the relatively thick (walls 0.40.5 mm. thick) stone. The epithet /eptocladum means ‘‘slender (delicate) twigs.” Calophyllum leptocladum is at least superficially similar to C. whitfordii (from the Philippine Islands), and the two species have similar galls on the leaves. Calophyllum whitfordii can be most easily distinguished by its predomi- nantly terminal inflorescences that nearly always have more than five flowers; the flowers have eight tepals. Calophyllum whitfordii also has leaves with less dense venation and a somewhat larger fruit with a thinner stone wall; the indumentum on the inflorescence axis extends to the pedicels and to at least the bases of the outer pair of tepals. Smith and Darwin (loc. cit.) cited four specimens of Calophyllum leptocla- dum from Vanua Levu. Howard 104 is a specimen of C. neo-ebudicum and so, probably, is FDA 15722. Berry 4and FDA 15733 seem to be C. leptocladum. 153. Calophyllum whitfordii Merr. Philip. Jour. Sci. 1(Suppl.): 96. 1906, Enum. Philip. Fl. Pl. 3: 81. 1923, pro majore parte. Tyre: Philippine Islands, Luzon, Bataan Province, Lamao River, Mt. Mariveles, 2300 feet [701 m.], Feb. 1905, FB 2613 coll. Meyer (isotypes, K, NY, SING, US). Tree to 28 meters tall, d.b.h. to 57 cm.; outer bark yellowish brown, fissured. Twigs flattened, 1-2 mm. across, rather bluntly 4-angled. drying brown when young, later yellowish, at first farinose (puberulent); axillary innovations lacking basal scars; internodes 0.4-3.5 cm. long; uppermost pair of axillary buds pointed, to 1.5 mm. long, erect; terminal bud + conical to plump, 2.5-4.5(-6) mm. long, with grayish brown to brown, puberulo-tomentose long. Petiole (2.5-)5-11 mm. long, broadly concave above, convex below, subpersistently puberulent; lamina elliptic, 3.6-8.5 by 1.9-4.3 cm., bluntly acuminate to obtuse at apex, cuneate to acute at base (minutely rounded at very base), not recurved but slightly undulate at margin, coriaceous, drying umber to sepia above and cinnamon-sepia to sabelline-olivaceous below, subpersistently puberulent on midrib above and especially below, the midrib above narrowing gradually from base, slightly to strongly raised, sulcate or not, 0.07-0.25 mm. wide at midpoint, below raised, + angled, the venation above and below + apparent, raised, (6 to) 9 to 13 (to 15) veins/5 mm., 630 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 61 angle of divergence 60-70(-75)°. Inflorescences terminal and/or axillary, with 7 to IS flowers, often flabellate, usually with branches up to 2 cm. long and with 5 flowers, the axis 2-6 cm. long, puberulent, lowest internode at least 1.2 cm. long; bracts unknown; pedicels 5-11 mm. long, puberulent. Flower (?)hermaphroditic; tepals 8, the outer pair broadly ovate, 4.5-5 by ca. 4 mm., puberulent on back, the inner ones oblong to obovate or elliptic, 8-9 by 2.5-5 mm., outer two sometimes puberulent in strip down back; stamens 125 to 230, the filaments to 4.5 mm. long, the anthers oblong, 0.7-1.5 mm. long, retuse at apex; ovary 1.7-2 mm. long, the style to 3 mm. long, the stigma peltate, ca. 0.8 mm. across, 3- or 4-radiate. Submature fruit spherical to ellipsoid, I-1.5 by 0.8-1.4 cm., apiculate, drying fuscous-black, finely striate to wrinkled; outer layer not detaching cleanly from stone, 0.5-0.8 mm. thick, compact; stone subspherical, 0.9-1.4 by 0.7-1.3 cm., rounded at apex, the walls 0.15-0.3 mm. thick, at base 0.5-1 mm. thick, smooth, (?)unmarked; spongy layer initially well developed. DistRIBUTION. The Philippine Islands (Map 47). SELECTED SPECIMENS SEEN. Philippine Islands. Luzon. Cagayan: sine loco, FB 6667 (us). Isabela: Mt. Moises, Clemens 16766 (?) (uc). Zambales: Subig, Merrill 1759 (xk, Nsw, Ny, US). Bataan: Mt. Mar iveles, Lamao R., FB 785 Map 47. Distribution of Calophyllum rugosum (triangles), C. aureum (open stars), C. rubiginosum (circles), C. recurvatum (stars in solid circles), Calo- phyllum sp. 164 (solid star), and C. whitfordii (squares) in Southeast Asia— Malesia 1980] STEVENS, CALOPHYLLUM 631 (BM, BO, F, K, LY, NSW, NY, P, SING, US). Laguna: Famy, Minayotan, 442 m., PNH 39394 (a, PNH, SING). Quezon: Malikboy, FB 31478 (Ny). MINDoRo: sine loco, FB 6855 (Bo, Ly, Ny, us). MinpANAo. Davao: Mt. Galintan, 150 m., BS 48913 (Ny, uc). EcoLocy. Colline forest, 150-700 m. alt. Flowering January to March; submature fruit May to July Conoid galls 1-1.5 mm. tall are frequent on the lower and sometimes on the upper surface of the lamina. Calophyllum whitfordii can be recognized by its small, conical, terminal buds usually less than 4.5 mm. long; its elliptic, rather small lamina that dries umber to sepia and often more or less nitid on the upper surface; its puberulent, ee inflorescences; and its spherical, shallowly wrinkled fruit with a compact outer layer. Small, conoidal galls on the lamina are common. The epithet commemorates H. N. Whitfor Although there is considerable variation in the prominence of the midrib on the upper surface of the lamina (in specimens such as BS 4891/3 it is almost flat), this variation is not correlated with that of other characters. Clemens 16766 has very short petioles (figures in parentheses in the description above). The circumscription of Calophyllum whitfordii has given me much trouble. I originally thought that it was part of the variable C. blancoi complex, and it may be related to that species. However, its leaves do not dry grayish or so strongly bicolored as in C. blancoi; they are often more or less nitid on the upper surface and are smaller than is usual for C. blancoi. The conoidal galls so common in C. whitfordii are uncommon in C. blancoi, and the hairs of C. whitfordii are less branched than is usual in C. blancoi. The terminal bud of C. blancoi is plump and nearly always considerably larger than that of C. whitfordii, and there are minor differences in the fruit. Calophyllum whitfordii is also superficially similar to C. leptocladum, from the Fiji Islands, but these two species can be more easily separated (see C. leptocladum). 154. Calophyllum rugosum P. F. Stevens, sp. nov. Ficure 40, a-c. A speciebus aliis Calophylli in gemma terminali 2.5-4.5 mm. longa, lamina mediocri venulis lateralibus 3 usque ad 7 per 5 mm. saepe prominentioribus in pagina supra quam in pagina infra, inflorescentia terminali, et fructu in siccitate valde rugoso parietibus putaminis tenuis, differt. Tree 5-12 meters tall, d.b.h. ca. 20 cm. ae unknow Twigs flattened, 1.2-2 mm. across, 2- or + rarer 4-angled, drying dark brown to blackish, puberulent when young; axillary innovations lacking basal scars; internodes 0.6—6 cm. long; uppermost pair of axillary buds rounded, ca. 1 mm. long, erect; terminal bud conical to plump, 2.5-4.5 mm. long with subadpressed, brown indumentum (hairs, Ficure 41, a, b), Ree internode absent (to | mm. long). Petiole [1.5—] 4-10 mm. long, strongly concave above, convex below, glabrous at maturity; lamina elliptic to obovate (rarely suborbicular) [lingulate], 1.6-10.5 by 1.4-5.4 cm., rounded to shallowly retuse [voL. 61 JOURNAL OF THE ARNOLD ARBORETUM 632 1980] STEVENS, CALOPHYLLUM 633 at apex, cuneate to broadly rounded [cordate] at base, not undulate to distantly so and slightly recurved at margin, coriaceous to very coriaceous, drying cinnamon-sepia above and sabelline to cinnamon below, puberulent on midrib below when young, the midrib above abruptly narrowed at base, at first slightly depressed, becoming flat or slightly raised, 0.15-0.25 mm. wide at midpoint, below slightly raised, margins depressed, becoming subdepressed toward apex, the ven ee apparent (obscure) above, often obscure below, raised (subdepressed), 2 to 7 veins/5 mm., angle of divergence 50-75°. Infructescences aaa and from adjacent axils, with scars of up to 19 flowers (with 5-flowered branches up to 5 mm. long), the axis up to 5 cm. long, subtomentose to puberulent especially near base, lowest internode 1-2 cm. long; bracts unknown; pedicels 1.3-2.1 cm. long, glabrous. Damaged flower only known; tepals 6, the outer pair ovate-elliptic, ca. 6 by 5 mm. Fruit spherical to ellipsoid, 2-2.2 by 1.7-2 cm. (but see discussion), rounded to apiculate, drying brown-vinaceous, deeply wrinkled; outer layer detaching more or less cleanly from stone, 0.5-1.5 mm. thick, compact; stone ellipsoid to spherical, ca. 1.9 by 1.6-1.75 cm., rounded at apex, the walls ca. 0.25 mm. thick, smooth, not marked; spongy layer at first well developed. Tyee: Indochine [Vietnam], Annam, Massif du Bi-Douppres du Haut Donai, 2000 m., 14 Oct. 1940, Poilane 30918 (holotype, A; isotypes, Pp (several)). DisTRIBUTION. Vietnam (Map 47). ADDITIONAL SPECIMENS SEEN. Vietnam: Nhatrang, 1800 m., Poilane 3494 (a, p), massif du Hon Ba, 1000-1500 m., Chevalier 38865 (Pr). coLocy. Open, shrubby montane vegetation, rocky soil or “‘bordure grande forét et celle de pin’’; 1000-2000 m. alt. Submature fruit in October; fruit and peduncle pale, watery green. Calophyllum rugosum can be recognized by its rather small terminal bud, its medium-sized lamina that is variously shaped at the base but always more or less rounded at the apex, its terminal inflorescence, and its deeply wrinkled fruits. The venation is often more prominent on the upper side Ficure 41. Hairs (from terminal bud, unless otherwise noted). a, b, Calo- Phyllum rugosum: a, Poilane 3494; b, Poilane 30918. c-f, C. rubiginosum: c, f, Soepadmo 86, from stem; d, Maingay, Kew dist. 168; e, Samat bin Abdullah 2174. g, C. rotundifolium (Robinson s.n.). h-j, C. sakarium (SAN 21612), axillary bud: i, from above. k-m, C. obliquinervium: k, SAN 33609; l, m, PNH 14132, apex and base of hair ca. 330 wm. eae n, 0, C. molle (FRI 8285), apex and base of hair ca. 510 pm. long. p-r, bb, cc, C. gracilipes. p, S 22401. q, Hallier 2992. 1, Williams 2193. bb, cc, Fe deemans 6013: bb, apex of hair (base similar to portion below arrow in cc); cc, basal part of hair ca. 1230 wm. long (apex similar to w). s, y, C. aurantiacum (Stevens etal, 195).t,u, C. echinatum: t, Cel. /II-407; u, Cel. /H-215.v,w, C. recurvatum o 7737), hairs to 720 wm. long, base and apex of hair ca. 500 pm. long. z, C. brachyphyllum (BS 34482). aa, o ardens (BRUN 297), ae Baal Scale = 120 um. (in c-e, scale = 240 p 634 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 of the lamina than on the lower, and the veins are relatively distant (3 to 7/5 mm.). The epithet rugosum (‘‘wrinkled’’) was considered appropriate because the dried fruits are so strongly wrinkled. The closest relative of Calophyllum rugosum is probably C. rotundifolium. The leaves of Chevalier 38865 are almost identical to those of Stone 7174 (C. rotundifolium), and the general color on drying and venation prominence of the leaves of Poilane 3494 are also similar. The two species have similar hairs, with rather thick-walled cells about as broad as long. Lamina anatomy is also similar, although the anticlinal walls of the upper epidermis of C. rotundifolium (Stone 7174) are at most inconspicuously sinuous near the top, while those of all collections of C. rugosum are characteristically sinuous for much of the height of the cell (this may be only a trivial difference). Both species have a short palisade mesophyll, lignified spongy mesophyll, and vascular bundles that are not transcurrent abaxially (although they are transcurrent in Poilane 30918). The inflorescences of the two are quite different: in C. rotundifolium the axis is absent, while in C. rugosum it is well devel- oped; the characteristic thick outer tepals of the flowers of C. rotundifolium do not occur in C. rugosum. The fruits of C. rotundifolium are unknown. The three specimens assigned to Calophyllum rugosum are at first sight very different. Poilane 30918 has long internodes and relatively large leaf blades that are cuneate or rounded at the base. Poilane 3494 has short internodes and smaller leaf blades that are broadly rounded at the base. Chevalier 38865 has long internodes and rather large, subsessile leaf blades that are cordate at the base; it is superficially very different from the other specimens (the characters in which it differs are enclosed in brackets in the description above). However, in anatomy, terminal bud, indumentum, general leaf type, and inflorescence position, all three are similar (although Poilane 30918 has veins more or less prominent on the lower surface of the lamina, and—as might be expected—vascular bundles transcurrent abaxially in transverse section). The description of the flower is taken from a very damaged bud found between the leaves (not attached to the shoot) of Poilane 3494. Chevalier 38865 has immature fruits that would possibly be about 1.5 cm. long when mature; in drying characteristics and general type they are similar to those of the type specimen. Poilane 5083 (Nhatrang, Mere et l’Enfant, 2000 m. (p)) has a facies similar to that of Calophyllum rugosum, but it has a terminal bud ca. 6 mm. long, axillary inflorescences, flowers with eight tepals, a better-developed palisade mesophyll, and a thinner-walled upper epidermis with the anticlinal walls only very slightly sinuous near the top. 155. Calophyllum rotundifolium Ridley, Jour. Fed. Malay States Mus. 5: 29. 1915, Fl. Malay Penin. 1: 188. 1922; M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 337. 1956; T. C. Whitmore, Tree Fl. Malaya 2: 168. 1973. Tyre: Malaya, Selangor, Gunong Mengkuang [Lebah], 5000 feet [1524 m.], 17 Jan. 1913, Robinson s.n. (holotype, sinc; isotypes, BM, K). 1980] STEVENS, CALOPHYLLUM 635 Tree ca. 9 meters tall, d.b.h. ca. 23 cm.; outer bark ochre, smooth, or surface cracked, with large lenticels; latex translucent. Twigs flattened, 1.5-2 mm. across, slightly angled, drying dark brown, puberulent when young; axillary innovations lacking basal scars; internodes 1-4 cm. long; uppermost pair of axillary buds + rounded, 1-2 mm. long, erect, conspicuous; terminal bud plump, 2-3 mm. long, with short, brown, tomentose indumentum (hairs, Figure 41, g), underdeveloped internode not apparent. Petiole ca. 0.5 mm. long, concave above, convex below, + glabrous; lamina + orbicular, (1-)2.5-6.5 by (0.7-)2.4-5.5 cm., + retuse at apex, cordate at base, neither undulate nor recurved at margin, coriaceous, drying umber above and cinnamon-sabelline to umber below, subpersistently puberulent on midrib below, the midrib above narrowing quickly at base, + depressed at first, becoming raised, 0.1-0.3 mm. wide at midpoint, below impressed or slightly raised, + angled, the venation above subapparent, raised, below obscure to invisible, 7 to 10 veins/5 mm., angle of divergence (50-)60-70°. Inflorescences terminal, with 1 (rarely 2) flower(s), axis absent; bracts unknown; pedicels 1.1-1.6 cm. long, + glabrous. Flower (?)hermaphroditic; tepals 8, the outer pair elliptic to ovate, 0.6-1.2 by 0.4-0.9 cm., flat to subconcave, coriaceous, glabrous, the next pair + orbicular, 1.3-1.4 by ca. 1 cm., concave, the inner ones obovate, |1.2-1.4 by 0.8-1 cm.; stamens ca. 200, the filaments 4-8 mm. long, at base connate for 1.5-2 mm., the anthers oblong, 1-1.8 mm. long, rounded to retuse at apex; ovary ca. 3 mm. long, the style ca. 3 mm. long, the stigma peltate, ca. 1.5 mm. across, + 3-radiate. Fruit unknown. DistripuTIon. Malaya, the southern part of the Main Range (Map 46). SELECTED SPECIMENS SEEN. Malaya. PAHANG/SELANGOoR: G. Ulu Kali, 1738 m., FRI 12585 (kep), 1800 m., Stone 8416 (kLU). SELANGoR: G. Mengkuang Lebah, 1646 m., Derry s.n., 4 Mar. 1907 (sING). EcoLocy. Stunted montane forest, 1525-1800 m. alt. Flowering in January. Calophyllum rotundifolium is a distinctive species characterized by its suborbicular leaf blades with the venation less prominent on the lower surface than on the upper, and by its usually single, terminal flowers with eight tepals, the outer pair of which are coriaceous and almost leaflike. The epithet rotundifolium (‘‘round leaves’’) is very appropriate for this species. Calophyllum clemensorum, from Borneo, has been confused with C. rotundifolium, but the two are probably not close since the former has leaf blades with clearer venation about equally prominent on both surfaces and a terminal inflorescence with an axis 2-4 cm. long. The closest relative of C. rotundifolium is probably C. rugosum, from Vietnam (for the differences separating them, see C. rugosum). The thick, rather leaflike outer tepals of Calophyllum rotundifolium are distinctive. Although exceeding some of the leaves in size, they have steeply ascending (rather than widely divergent) venation and lack a midrib. It is possible that these outer tepals are bracts, but no buds were seen in their axils. Ridley (1915, Joc. cit.) described the ‘‘inner sepals’? as being only 636 JOURNAL OF THE ARNOLD ARBORETUM [vo. 61 2 mm. long, and although later authors have followed him in this, they appear to have been mistaken. Henderson and Wyatt-Smith (/oc. cit.) described the venation on the upper side of the leaf as less prominent than that on the lower. This is normally the case in Calophyllum, but it is clearly otherwise in this species. Leaf blades of saplings are suboblong and are up to 13 by 6.6 cm. (Derry s.n., Stone 8416). In such specimens the terminal bud is up to 7 mm. long. 156. Calophyllum aureo-brunnescens M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 341. p/. = 1956; T. C. Whitmore, Tree Fl. Malaya 2: 172. 1973. Type: Malaya, Pahang, Gunong Tahan, 5000 feet [1524 m.], 11 June 1922, SFN 7855 coll. Haniff & Nur (holotype, sinc; isotypes, K, KEP). C. venustum auct., non Ridley; Ridley, Jour. Fed. Malay States Mus. 2: 110.1909. Tree 3—4.5 meters tall; trunk and bark unknown. Twigs flattened, 1.5-3 mm. across, rounded, drying dark brown, sparsely puberulent when young; axillary innovations lacking basal scars; internodes (2-)3-5.5 cm. long; uppermost pair of axillary buds subacute, to 2.5 mm. long, suberect to spreading; terminal bud narrowly conical, 3.5-8 mm. long, with pale brown, crustaceous indumentum (hairs, Ficure 43, |; also mionilgia). underdeveloped internode |—6(—10) mm. long. Petiole 0.8-1.3 cm. long, broadly and rather shallowly concave above, convex below, glabrous when mature; lamina elliptic (rarely subovate or subobovate), (2.7—)3.5—6.7 by (1.1-)1.8-3.3 cm., acute to obtuse (retuse) at apex, acute at base, neither undulate nor recurved at margin, coriaceous, drying mid to dark brown above, paler and + pruinose below, sparsely puberulent on midrib below when young, the midrib above narrowing gradually from base, margins not clear, + depressed at first, becoming level, 0.2-0.35 mm. wide at midpoint, below raised, + striate (angled toward apex), the venation subapparent to obscure above, subapparent below, raised, (9 to) 11 to 16 veins/5 mm., angle of divergence (45—)50-70°. Inflorescences terminal and from adjacent foliate axils, with 7 to 11 flowers, branched or not, often flabellate, the axis 2.5-5 cm. long, inconspicuously farinose, lowest internode |.3-3.5 cm. long; bracts + obovate, ca. 4mm. long, soon deciduous; pedicels 0.7-1.1 cm. long, puberulent. Flower (?)hermaphroditic; tepals 8 (rarely 9), the outer pair broadly ovate, 6.5-7 by ca. 5 mm., the inner ones elliptic to obovate, 7-9 by 2.2-4 mm.; stamens ca. 200, the filaments to 3.5 mm. long, the anthers elliptic-oblong, 0.6-1.1 mm. long, retuse to apiculate; ovary |.8-2 mm. long, style and stigma unknown. Fruit unknown. DistriBuTION. Northeastern Malaya (Map 46). SELECTED SPECIMENS SEEN. Malaya. PaHanc: G. Tahan, Ridley 16015 (BM, K, SING); Padang Luas, 1676 m., F.M.S. Museum 12240 (sinc); G. Ulu Kechau, 1829 m., KEP 42911 (k, KEP). EcoLocy. Montane forest, 1525-1830 m. alt. Flowering May to July. 1980] STEVENS, CALOPHYLLUM 637 Calophyllum aureo-brunnescens is an imperfectly known species that can be characterized by its rounded twigs, its narrowly conical terminal buds, its suberect to spreading uppermost pair of axillary buds, its rather small, basically elliptic lamina that is usually other than retuse at the apex, and its terminal inflorescence with eight-tepaled flowers. The epithet aureo- brunnescens means ‘‘gold-brownish’’; I am not sure why it was coined. It has been suggested that Calophyllum aureo-brunnescens may be close to C. aureum; for discussion see C. aureum. 157. Calophyllum aureum Symington ex M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 340. p/. 20. 1956; T. C. Whitmore, Tree Fl. Malaya 2: 171. 1973. Type: Malaya, Pahang, Cameron Highlands, Gunong Beramban, 5 June 1933, KEP 31007 coll. Symington (holotype, SING; 1SOtypes, K, KEP). Shrub or tree 2.5-15 meters tall, d.b.h. to 38 cm.; trunk (?)unbuttressed; outer bark yellowish to ochre, smooth at first, becoming fissured; inner bark red-brown; latex clear golden or opaque cream Twigs strongly flattened, 1.2-2.5 mm. across, 4-angled, drying dark brown, yellowish brown when much older, transiently subglabrous or farinose- puberulent; axillary innovations lacking basal scars; internodes 2-6.5 cm. long; uppermost pair of axillary buds pointed, 1.5-3.5 mm. long, erect; terminal bud narrowly conical to somewhat plump, (3.5-)4.5-6 mm. long, with subfur- furaceous grayish brown to subtomentose rufous indumentum (hairs, Ficure 43, p), underdeveloped internode absent. Petiole 4.5-9.5 mm. long, concave above, convex below, glabrescent; lamina obovate Gane subelliptic), 2-5 by 1.3-3.6 cm., retuse (very rarely rounded) at apex, cuneate to acute at base, broadly undulate or not and not recurved to slightly so at margin, entire lamina drying + concave, coriaceous, sabelline-olivaceous above and sepia below, when young farinose on midrib below, the midrib above abruptly narrowed at base, raised to subdepressed, with raised margins, 0.15—0.25 mm. wide at midpoint, disappearing just short of apex or not, below raised, striate (subdepressed toward apex), the venation above apparent, below subobscure to apparent, raised to level, 12 to 19 veins/5 mm., angle of divergence (55-)65-75°. Inflorescences terminal (also from uppermost pair of foliate axils), with 7 to 13 flowers, often with 3-flowered branches to 1.5cm. long, frequently flabellate, the axis to 4 cm. long, farinose to puberulent at least near base, lowest internode 1.5—3.2 cm. long; bracts foliaceous, to 2cm. long, subpersistent; pedicels 0.6-1.1 cm. long, glabrous (sparsely farinose toward base). Flower (?)hermaphroditic; tepals 8, the outer pair broadly ovate, 5.5-6.5 by 4.5-5.5 mm., the inner ones elliptic to obovate, 9-10 by 4-6 mm.; stamens ca. 150, the filaments to 4.5 mm. long, the anthers oblong, 1.2-1.5 mm. long, subacute to retuse at apex; ovary ca. 2 mm. long, the style ca. 3 mm. long, the stigma peltate, ca. 0.8 mm. across, 3- or 4-radiate. Immature fruit ovoid, ca. 1.8 by 1.4 cm., apiculate, drying brown, smooth; outer layer ca. 1.2 mm. thick; stone walls probably very thin. DistripuTiIon. Malaya, the Main Range (Map 47). 638 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 SELECTED SPECIMENS SEEN. Malaya. Perak: G. Korbu, 1524-2134 m., KEP 32118 (k, KEP). PaHaNc: Cameron Highlands, Bukit Mentigi, 1524 m., Chew 860 (A, AAU, G, K, KEP, SING, UC), path to G. Beramban, 1494 m., FRI 15675 (keP, L); G. Benom, 1829 m., FRI 3305 (kep), Kluang Terbang, ‘Barnes SN. (sinc). SELANGOR: Fraser’s Hill, KEP 11468 (ker, sinc); G. Bunga Bua, FRI 307 (KEP, SING). Eco.ocy. Lower montane forest (trees on ridges very stunted), 1490-2135 m. alt. Flowering June and October; flower cream, fragrant. Barnes s.n. and Chew 860 have leaves modified as bilabiate galls, perhaps caused by coccids. Calophyllum aureum can be characterized by its uppermost pair of axillary buds, which are long and erect; its long internodes; its small, usually obovate and retuse lamina that dries grayish green above with the midrib brownish, and brown to golden brown below; and its relatively few-flowered, farinose- puberulent terminal inflorescences with widely spreading branches and persis- tent bracts. The epithet aureum (‘‘gold’’) alludes to the yellowish bark. Barnes s.n. was cited under Calophyllum aureo-brunnescens by Henderson and Wyatt-Smith (op. cit.); they thought that it was intermediate between that species and C. aureum. However, Barnes s.n. is merely a poor, galled specimen of C. aureum; FRI 3305, also from G. Benom, is more obviously this species. The fruits of both C. aureum and C. aureo-brunnescens are still unknown; the two species may not be closely related. 158. Calophyllum rubiginosum M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 308. p/. 5. 1956; Smythies, Common Sarawak Trees, 61. 1965; Kochummen, Malayan Forest Rec. ed. 2. 17: 215. 1965: T. C. Whitmore, Tree Fl. Malaya 2: 189. 1973; H. Keng, Gard. Bull. Singapore 28: 245. 1976; Corner, Gard. Bull. Singapore Suppl. 1: 104. 1978. Type: Malaya, Negri Sembilan, Berembun-Angsi Reserve, Bukit Tangga, 25 Feb. 1918, KEP 1953 coll. Kinsey (lectotype, KEP; isolecto- type, K, SING). Ficure 40, 1. C. wallichianum auct., non Planchon & Triana; King, Jour. Asiatic Soc. Bengal, II. 59: 179. 1890, quoad descr.; Ridley, Jour. Straits Branch Roy. Asiatic Soc. 33: 48. 1900, Fl. Malay Penin. 1: 187. 1922, pro parte: . R. Henderson, Gard. rn — Settl. 4: 224. 1938; Foxworthy, Malayan Forest Rec. 3: fig. | C. muscigerum Boerl. & Lae ex Heyne, Nutt. Pl. Nederl.-Indié. ed. 2. 2: 1085. 1927, ibid. es bs 1085. 1950; Pukol & Ashton, Checklist Brunei Trees, 93. 1964. Calophyllum sp. Koord. ae at Verzeich. 2: 40. 1910. Tree 7.5-39 meters tall, d.b.h. to 77 cm.; trunk without buttresses (with small spurs); outer bark brownish at first, becoming grayish to yellowish or ochre, with long, shallow fissures, or rugulose with minute cracks, flaking or scaling, hoop marked, the inner surface yellow to orange; under bark orange-straw to deep red; inner bark red; latex white, sticky or not Twigs flattened, 2-3.5 mm. across, strongly 2- or 4-angled, drying whitish 1980] STEVENS, CALOPHYLLUM 639 or yellowish, subpersistently brown-tomentose (rarely puberulent); axillary innovations lacking basal scars; internodes (l-)2—7 cm. long; uppermost pair of axillary buds pointed, 1-2 mm. long, erect; terminal bud plump, 4-8 mm. long, with brown, short-tomentose to puberulent indumentum (hairs, Ficure 41, c-f), underdeveloped internode absent (-2 mm. long). Petiole 0.5-3 cm. long, slender, narrowly to broadly concave above, convex and + persistently short-tomentose below; lamina elliptic to suboblong or ovate, (4.5—)7-16 by (1.6-)2-5.7 cm., acute to acuminate at apex, acute to subdecurrent (rarely rounded) at base, distantly and + deeply undulate and plane to slightly recurved at margin, coriaceous, drying olivaceous to umber above and clay or cinna- mon-sabelline to sienna-umber below, color often obscured by ferrugineous indumentum, usually tomentose (rarely puberulent) on both surfaces initially, becoming glabrescent above, the midrib above narrowing gradually from base, sharply raised, 0.2-0.5 mm. wide at midpoint, below strongly raised, + striate, the venation subobscure on both surfaces, raised to depressed, latex canals as clear as veins, 8 to 14 veins/5 mm., angle of divergence 70-80°. Inflores- cences terminal and from adjacent foliate axils, with 7 to 15 flowers (very rarely single, terminal flower), unbranched, often flabellate, the axis 2.5-7 cm. long, short-tomentose, lowest internode 0.5-1.7 cm. long; bracts + linear, to 3.2 cm. by 2 mm., soon deciduous; pedicels 3.5—16 mm. long, short-tomen- tose. Flower (?)hermaphroditic; tepals (4 or) 8, the outer pair ovate, 5.5-6.5 by 3.5-5.5 cm., tomentose on outer and inner surfaces, especially near apex, the next pair (broadly) elliptic, 7.5-9 by 4-5.5 mm., tomentose in band down back, the inner ones obovate to subelliptic, to 8 by 4 mm.; stamens 130 to 235, the filaments to 5.5 mm. long, the anthers suboblong, 0.6-1 mm. long, rounded to slightly retuse at apex, with subpersistent hairs to 0.2 mm. long; ovary |.5-1.7 mm. long, the style to 2.5 mm. long, the stigma sub- peltate, to 0.9 mm. across, 3-radiate. Fruit ovoid-ellipsoid, 2-2.2 by 1.4—-1.8 cm., + acute at apex, drying brown, sharply and prominently wrinkled, fur- furaceous; outer layer detaching cleanly from stone or not, ca. 1.5 mm. thick, compact; stone ellipsoid, ca. 1.7 by 1.4 cm., rounded at apex, the walls less than 0.2 mm. thick, smooth, unmarked; spongy layer thin. DistriBUTION. Southern Malay Peninsula to Sumatra and Borneo (Map 47). SELECTED SPECIMENS SEEN. Malaya. SELANGor: Bukit Lagong F.R., 502 m., KEP 83675 (ker); Bangi F.R., KEP 10974 (kep, Pp); Rantau Panjang, KEP 14772 (kep, P); Kuala Lumpur, Jalan Rajah, KEP 10470 (e£, kK, KEP, NY, SING); Kepong, F.R.I. plantation, KEP 38507 (k, KEP, sING, Us); Sungei Buloh F.R., 240 m., KEP 99637 (a, KEP, SAR); Kuang F.R., KEP 21962 (KEP, Ny, P); Ayer Hitam F.R., KEP 10903 (ker); Kanching, KEP 14782 (ker); Weld Hills F.R., KEP 2615 (ker); Ulu Gombak F.R., KEP 11645 (kep, Ny, SING); Bukit Cheraka F.R., KEP 65567 (ker); Bukit Tunggul F.R., KEP 23644 (ker); Ulu Langat F.R., KEP 53604 (ker); Ampang F.R., FRI 16054 (kep, sinc); Kajang, KEP 2649 (kep); 12th mile Bentong road, Samat bin Abdullah 174 (a, G, KLU). Matacca: 15th mile Sungei Udang F.R., Sinclair 8937 (B, E, K, SING); Ayer Panas F.R., KEP 25253 (kx, Ker); Batang Malaka F.R., KEP 2063 (k, SING); Ramuan China F.R., KEP 25257 (po, KEP). NEGRI SEMBILAN: Sungei Menyala F.R., KEP 64654 (kep, sinc); Senawang F.R., KEP 18505 (a, kK, KEP, SING); 640 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Gunong Angsi F.R., 457 m., FRI 17332 (k, KEP, SAR, SING). PaHaNnc: ca. 10 km. S. of Kampong Aur, FRI 15895 (kep); Lesong F.R., 150 m., FRI 19843 (KEP, SAR). JoHORE: Kluang Forest, FRI 8709 (a, K, KEP, SAR, SING); Jemaluang F.R., KEP 69995 (ker); Sungei Kayu, Kiah s.n., 28 Oct. 1936 (sinc). Singapore: Bukit Timah, Ridley 6196 (a, E, K, NSW, P, SING); NE. end MacRitchie Reservoir, SFN 39634 (Bo, E, K, P, SING); Chua Chu Kang, Ridley s.n., anno 1894 (sina); Botanic Gardens, SGN 1668 (sinc). Sumatra. Riau: Indragirische Bovenlanden, Danau Mengkoeang, 60 m., bb 27550 (Bo, L); Upper eee Pakanbaru, Tenajan , Soepadmo 8&6 (A, B, BO, C, E, LAE, NY, SING). SELATAN: Rawas, | Grashoff 970 (Bo, L). ‘‘Central, Nogul Droog,” Roarders 15772 (Bo). Borneo. Sarawak. 3rd Division: Melinau, Ulu Selada, path to Bukit Mabong, 305 m., S 24266 (L, SAN, SAR, SING). 4th Division: Baram, Sungei Beling, FA 636 coll. Egon, 24 May 1948 (sar); Poak, Hose s.n., 8 Nov. 1911 (Bm). BRUNEI: Kuala Beblong, 60 m., Ashton s.n., Oct. 1959 (sar). Saban. Beaufort: Beaufort Hill, SA N 36900 (SAN). KALIMANTAN. Timur: C. Kutei, Belajan R., G. Kelepok near Tabang, 250 m., Kostermans 10584 (Bo, CANB, K, KEP, L, NY, P, SING); W. Koetei, Long Bleh, 30 m., bb 16059 (a, Bo). Ecotocy. Lowland or colline forest, 30-500 m. alt. Flowering January to May, and July and August (flower scented); fruiting April to July, and Oc- tober to December (fruit brownish green) YounG PLANT. The young plant is erect, and the terminal bud is functional. Loca uses. The wood, although susceptible to termite attack, is used in construction. The latex is very poisonous and, mixed with rice, is used to exterminate rats (Heyne, Joc. cit.). It is also reportedly used as a poison to stun fish (Kostermans 10584, Kalimantan); in the 3rd Division of Sarawak the latex of C. rubiginosum, or another species with white latex, is placed in small dams and left overnight for this purpose (Banyeng ak Nudong, pers. comm.). Calophyllum rubiginosum is a distinctive species characterized by its pale whitish yellow—drying twigs, its usually well-developed and persistent indu- mentum, its terminal inflorescences, and its ellipsoid, deeply wrinkled fruits with a furfuraceous surface and a very thin-walled stone. The epithet rubiginosum (“‘rusty’’) is very appropriate since the conspicuous, rusty brown indumentum persists so long. Although Calophyllum rubiginosum is superficially similar to C. molle, the two species keying out adjacent to one another in Henderson and Wyatt-Smith (op. cit.), there are numerous differences between them, and they are probably not closely related. Calophyllum molle has yellow latex, clearer venation, a petiole less than | cm. long, axillary inflorescences, flowers with four tepals and at least 500 stamens, a densely tomentose ovary, and a spherical fruit that is finely wrinkled but not furfuraceous and that has a stone with walls about | mm. thick. However, C. molle and C. rubiginosum are, so far as is known, the only two large-leaved species of Calophyllum to have hairs on the anthers. (Hairy anthers are also found in a few specimens of C. canum.) Calophyllum rubiginosum has also been confused with C. wal- lichianum (= C. rufigemmatum) because both have conspicuous, reddish 1980] STEVENS, CALOPHYLLUM 641 eats Nia however, there is otherwise little similarity between the t tiers is relatively little variation within Calophyllum rubiginosum. Some specimens from Sumatra have somewhat smaller leaves that are more or less rounded at the base of the lamina, and strongly flattened, two-angled twigs; C. rubiginosum usually has four-angled twigs, and the base of the lamina varies from more or less rounded to decurrent. Grashoff 970 (from Sumatra) has inflorescences that consist of a single, terminal flower, but in other characters it is unremarkable. Some specimens from Borneo, as well as Sinclair 8937, from Malaya, have indumentum that is shorter and less prominent and persistent than is typical for the species. The description of the rather remarkable linear bracts is taken from KEP 38507 (Malaya); whether or not the bracts are always linear is unclea The type collection of Calophyllum rubiginosum, "OEP 1953, consists of two sheets, one with flowers and the other with fruits; they were collected at different times but in the same area. Calophyllum rubiginosum has been lectotypified on the flowering collection; the fruiting collection was made n May 22, 159. Calophyllum molle King, Jour. Asiatic Bengal, IJ. 59: 177. 1890; Curtis, Jour. Straits Branch Roy. Asiatic Soc. 25: 78. 1894; Ridley, Fl. Malay Penin. 1: 185. 1922; I. H. nee ll & M. R. Henderson, Gard. Bull. Straits Settl. 3: 347. 1925; M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 307. 1956; T. C. Whitmore, Tree Fl. Malaya 2: 188. 1973. Type: Malaya, Penang Hill, March (fl.) or June (fr.), 1888, Curtis 1426 (syntypes, K, SING). Ficure 40, h, j. Tree 12-30 meters tall, d.b.h. to 50 cm.; trunk without buttresses or spurs; outer bark pale brown to gray-yellow, shallowly and distantly fissured or with vertical lines of lenticels, hoop marked, the inner surface orange; under bark dark brown; inner bark orange-red to dark reddish brown, fibrous; latex clear yellow, not sticky (dirty colored, nonresinous); wood white, radially flecked, soft. Twigs slightly flattened, 1.7-2.2 mm. across, obscurely 4-angled, drying yellowish or whitish, densely and persistently brown-tomentose, hairs to 0.5 mm. long; axillary innovations lacking basal scars; internodes (1-)2-6 cm. long; uppermost pair of axillary buds rounded, 1-2 mm. long, spreading; terminal bud plump to elliptic, 4-10 mm. long, with dense, brown, tomentose indumentum (hairs, Ficure 41, n, 0), underdeveloped internode 1-2. m long. Petiole 5-9 mm. long, deeply concave above, convex below, subpersis- tently tomentose; lamina ovate to suboblong or subelliptic, 6-18.5 by (1.5-)2.2-5 cm., shortly acute to acuminate at apex, rounded to cuneate and subcucullate at base, undulate and narrowly recurved at margin, thinly coriaceous, drying cinnamon-drab-grayish sepia to vinaceous-hazel above and sabelline to ful- vous-umber below, rather persistently tomentose on midrib above and on entire lower surface, the midrib above narrowing near base, or already narrow at base and further narrowing gradually, raised, 0.15-0.3(-0.6) mm. wide 642 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 at midpoint, below raised, rounded to striate, the venation apparent above, slightly less so below, raised, (8 to) 11 to 17 veins /5 mm., angle of divergence 70-75(-80)°. Inflorescences from foliate axils along stem, with 3 to 5 flowers, unbranched, the axis 0.5-2.9 cm. long, persistently tomentose, lowest internode 0.5-1.8 cm. long; bracts narrowly ovate, ca. 4 mm. long, soon deciduous; pedicels 4-12 mm. long, tomentose, in fruit to 1.4 cm. long and 3.5 mm thick. Flower (?)hermaphroditic; tepals 4 (rarely 6 in terminal flowers), the outer pair ovate-oblong, 7-8.5 by 4-6 mm., short-tomentose on back (also on inner surface, at least near apex), the inner ones + obovate-elliptic, 7.5—9.5 5-6.5 mm., nearly always tomentose in band down back; stamens 510 to 580, the filaments to 5 mm. long, connate for ca. 0.4 mm., the anthers suboblong, 0.4-0.7 mm. long, rounded to subretuse at apex, with rather dense, persistent hairs to 0.3 mm. long; ovary 2-3 mm. long, tomentose, the style 2.5-3 mm. long, the stigma peltate, ca. 0.9 mm. across, 3-radiate. Fruit subspherical to ellipsoid, 2.2-3.4 by 2-2.1 cm., apiculate to acute at apex, pubescent, drying brown to orange-brown, shallowly wrinkled; outer layer detaching cleanly from stone, 2.3-3.5 mm. thick, compact; stone spherical to ellipsoid, 1.4-2.9 by 1.3-1.5 cm., + obtuse at apex, the walls 1-1.3 mm thick, sometimes with 3 shallow, longitudinal depressions running from apex, otherwise smooth, discolored area at base 5.5—-7 mm. across; spongy layer thin. DistrisuTion. The Malay Peninsula, possibly also western Borneo (Map 48). SELECTED SPECIMENS SEEN. Malaya. Kepan: Gunong Inas F.R., 60 m., KEP 79265 (KEP). Pinang: oe Penang, back of W. Hill, 610 m., Curtis = 1426, March 1901 (k, sinc). Perak: Larut, 310-465 m., King’s collector 6724 (BM, FI, K, P, SING); Sg. Me au, Tapah Road, KEP 4563 (kK, SING); Tronoh, KEP 12022 (Kkep); Ulu Piah, KEP 11970 (ker); Sungei Pangkor F.R., 36 m., KEP 75968 (KEP). SELANGOR: Kuang, KEP 14776 (kep, p); Bukit Lagong F. R. , 457 m., KEP 52233 (KEP, SING); eee R., KEP 21975 (ker); Panjang Rawang, KEP 24931 (Kep). TRENGGANU: Ulu Besut, near Bukit Jebak Puyoh, 150 m. FRI 8285 (a, k, Kep); Ulu Sungei Loh below E. face G. Mandi Angin, 701 m., FRI 12135 (kK, KEP). PAHANG: G. Tapis, 610 m., FRI 10903 (A, KEP, SAR, SING): Raub, Bilut F.R., KEP 23353 (Kep); Rasatan: Baloh F.R., 15 m., KEP 77971, pro parte (KEP); Temerloh, KEP 5458 (sinc). JoHore: Renggam Pa 45 m., KEP 71274 (ker); Mersing F.R., 15 m., KEP 77929 (a, BO, K, KEP, sING); Labis F.R., Ulu Endau, FRI 7878 (a, k, KEP, SAR, SING); NW. G. Blumut, 427 m., FRI 8757 (a, K, KEP, SAR, SING); Kota Tinggi, Panti F.R., KEP 53935 (KEP, SING). Borneo. KALIMANTAN. Barat: Sanggau, G. Keramas, 75 m., bb 14405 (?) (Bo). EcoLocy. Mixed dipterocarp forests, often on ridges and hillsides; 15-701 m. alt. Flowering March (most) and August (once); fruiting June, July, and September to November (fruit brown). GERMINATION AND YOUNG PLANT. The young plant is erect, and the terminal bud is functional. Calophyllum molle can readily be recognized by its well-developed, tomen- tose indumentum; its leaf blades, which are often slightly cucullate at the 1980] STEVENS, CALOPHYLLUM 643 Map 48. Distribution of Calophyllum molle (squares), C. ardens (stars), C. aurantiacum (erect triangle), Calophyllum sp. 163 (inverted triangles), C obliquinervium (circles), and C. sakarium (star in solid circle) in Malesia. base; its axillary inflorescences; its flowers, which have four tepals and anthers that are more or less covered by short hairs; and its large, shallowly wrinkled fruits with a thick outer layer and a thick-walled, triradiately marked stone. The soft, tomentose indumentum of this species presumably suggested the epithet molle (“‘soft’’). Calophyllum molle is superficially similar to C. rubiginosum, and both species have hairs on their anthers. However, the two are not otherwise very similar; for the differences separating them, see C. rubiginosum The report of Calophyllum molle from Sumatra (Ridley, loc. cit.; Beccari, PS 953—see Calophyllum sp. 54) is incorrect. However, bb 14405, from western Kalimantan, is possibly to be referred to C. molle. It has a terminal bud up to 1.5 cm. long, its stems dry mid-brown, its petiole is broad, and its lamina is up to 14.5 by 6.5 cm. However, in general leaf and indumentum type it is similar to C. molle, and its flowers have four tepals, hairy anthers, and a densely tomentose ovary Henderson and Wyatt- Smith (loc. cit.) suggested that Calophyllum molle was typified by Curtis 1426; this number and several (unspecified) numbers of King’s collector (Kunstler) had been mentioned in the protolog. In the Kew and Singapore herbaria there are some specimens of Curtis 1426 with flowers that were collected in March (the year is not mentioned), and others 644 JOURNAL OF THE ARNOLD ARBORETUM [voL. 6] with fruits that were collected in June, 1888. Not having seen material from Calcutta, I have not designated a lectotype. 160. Calophyllum obliquinervium Merr. Philip. Jour. Sci. 20: 409. 1922, Enum. hilip Pl. 3: 30. 1923; Keith, N. Borneo Forest Rec. ed. 2. 2: 314. 1952; J. Anderson, Gard. Bull. Singapore 20: 154. 1963, pro minore parte; Pukol & Ashton, Checklist Brunei Trees, 93. 1964, pro parte; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 7: 15. 1967, Field Guide Trees W. Malesia, 161. 1974. Type: Philippine Islands, Palawan, Feb. 1920, FB 27929 coll. Cenabre, Paras, & Gellidon (isotypes, BM, UC). C. benjamina Ridley, Kew Bull. 1938: 119. 1938; Masamune, Enum. Phanerog. Born. 475. 1942; Keith, N. Borneo Forest Rec. ed. 2. 2: 313. 1952. Type: Brunei, Limbang, Haviland 505 (holotype, kK; isotypes, sar, NG). C. pseudotacamahaca vel. aff. auct., non Planchon & Triana; Keith, N. Borneo Forest Rec. ed. 2. 2: 313. 19 Tree 4-20(-30) meters tall, d.b.h. to 64.5 cm.; without buttresses; outer bark yellowish to grayish brown at first, with diamond-shaped fissures, becoming brown to blackish with reddish or brownish tinge, cracked or fissured, scaling, the inner surface dirty brown-greenish; under bark dark red; inner bark reddish brown to yellow; latex clear yellow (white—SAWN 33639), very sticky. Twigs slightly flattened, 1.2-3 mm. across, 4-angled to 4-alate, drying blackish brown, glabrous or sparsely farinose when young; axillary innovations lacking basal scars; internodes 0.5-3(-5) cm. long, terminal internode of innovation usually 1/2 or less length of others; uppermost pair of axillary buds rounded, ca. 0.5 mm. long, obscured by petiole bases; terminal bud plump, 1.5-2.5 mm. long, with rufous to pale brown, puberulent to subad- pressed indumentum (hairs, Ficure 41, k-m), underdeveloped internode not apparent. Petiole (0.3-)1-2.3 cm. long, flat to concave above, convex below, glabrous; lamina elliptic to subobovate, (3.8-)5.7-14.5 by (1-)1.2-6.5 cm., acute tc acuminate and recurved at apex, decurrent to cuneate (rarely rounded) at base, slightly undulate and not recurved at margin, coriaceous, drying umber to olivaceous above, umber to fulvous below, glabrous or very nearly so, the midrib above narrowing gradually from base, margins slightly raised, adjacent midrib often slightly raised, center slightly depressed, 0.1-0. wide at midpoint, usually disappearing ca. 1 cm. short of apex, below not much raised, rounded or + striate, the venation above and below subobscure, raised, 9 to 19 veins /5 mm., angle of divergence 30-S0°. Inflorescences terminal and from adjacent foliate axils (rarely 2 per axil), with 7 to 13 flowers, usually unbranched, the axis 3.5-9 cm. long, glabrous, or puberulent near base when young, lowest internode 1.7-3.5 cm. long; bracts 1.5-3(-8) mm. long, soon deciduous; pedicels 0.5-1.9 cm. long, to 2.5 cm. long in young fruit, glabrous. Flower (?)hermaphroditic; tepals 8 or 10, the outer pair suborbicular, (2.3 (when 10 tepals)-)3-3.7 mm. long and across, the next pair, when 10 tepals, + ovate, ca. 4.5 by 3.2 mm., the inner ones obovate 1980] STEVENS, CALOPHYLLUM 645 to elliptic, 5-8.5 by 2.5-4.5 mm. (rarely ca. 6 by 1.5 mm.); stamens 55 to 125, the filaments to 4 mm. long, connate for up to 0.5 mm., the anthers suboblong, 0.5-1.5 mm. long, truncate to retuse at apex; ovary I-1.5 mm. long, the style 2-4 mm. long, the stigma excentrically peltate, 0.5-0.8 mm. across, infundibular. Fruit spherical to ellipsoid, 0.9-1.3 by 0.75-1.1 cm., apiculate or not, drying dark brown, + smooth when ripe, strongly wrinkled when young; outer layer not detaching cleanly from stone, less than 0.5 mm. thick, air spaces developing; stone + spherical to ellipsoid, 6-8.5 by 5.5-7.5mm., rounded at apex, the walls 0.2-0.3 mm. thick, smooth, unmarked; spongy layer thin. DistripuTion. Northeastern Borneo, never far inland, and the northern and western Philippine Islands (Map 48). SELECTED SPECIMENS SEEN. Borneo and adjacent islands. Brunei: Tutong, Kg. Danau, van Niel 3453 (L); Bukit Pasir Puteh, BRUN 5020 (Bo, BRI, K, KEP, io sinc); Belait, mouth R. Lumut, van Niel 4046 (L); Lumapas, KEP 3 (A, K, KEP, SING); Berakas, KEP 37212 (k, KEP, SING). SABAH. Sipitang: A NE. of Sipitang, 6 m., SAN 15191 (A, Bo, K, KEP, SAN, SING); Kuala Mengalong F.R., 30 m., SAN 73151 (K, L, SAN, SING); Sungei Buntoh, 3 m., SAWN 27978 (L, SAN, SAR); Marantaman, SAN 2492 (A, BO, K). Papar: Kawang F.R., SAN 30271 (kK, KEP, L, SAN, SAR, SING); Kimanis F.R., 6 m., A 30] . KEP, SING); Kg. Padawan, SAN 3031/1 (kK, KEP, L, SAN, SAR, SING); Bongawan 4.5 m., SAN 2316 (a, Bo, kK). Kota Kinabalu: Pulau Gaya F.R., m., ‘SAN 41303 (kK, L, SAN, SING); Putatan, 1 m., SAN 33639 (KEP, L, SAN, SAR, SING); Penampang, SAN 2813 (Bo, k); Bukit Baer SAN 19940 (san). Tuaran: Lumas F.R., 180 m., SAN 33837(k, L, SAN, SAR). Kota Belud: Kelawat F.R., 120 m., SAN 33644 (i KEP, L, SAN, SAR, SING). Kudat: Marudu, Kitaku L, P, SING, US); Loro F.R., 150 m., SAN 51054 (BisH, NY, SAN); Balembangan ,6m., SAN 69604 (SAN). Labuk & Sugut: Jembongan Is., SAN 18956 ce BO, K, KEP, L, SAN, SAR, SING). Sandakan: Leila F.R., Water Works, SAN 48174 (K, KEP, L, SAN, SAR, SING); Sepilok Trig. Hill, 90- 150 m., SAN 24904 (K, L, SAN); Bukit Luku, Buli, SAWN 20553 Lo L, SAR, SING). Philippine Islands. Patawan: Aborlan, Malasgao R., PNH 14132 (a, Bo, uc, us); Tapul Mt., vicinity of Puerto Princesa, Ebalo 534 (A, BISH, NY, UC); Silanga, BS 9585 (2) (k, us). CALAMIAN. Culion: sine loco, Herre 1087(a, Ny, uc). Luzon, Camarines Norte: Paracale, FB 27096 (a, k, P, us). Albay: sine loco, FB 20108 (Bo). Samar: sine loco, Philippine Plants 1630 (BM, BO, G, GH, MO, NSW, NY, P, SING). Ecotocy. Rather variable in Borneo, although favoring poorer (acid) soils (in kerangas vegetation and similar places), by seashore, in swamps (at Kimanis), in primary forest with nibong (Oncosperma Blume), in secondary forest, in mixed dipterocarp forests on hillsides; to 180 m. alt. In Philippine Islands also with wide ecological range (Merrill, 1923, loc. cit.). Flowering December, February, and March (flower scented, petals often notably reddish or purplish—e.g., BRUN 5020); fruiting March to May, July, and December (fruit bluish, or reddish black). GERMINATION AND YOUNG PLANT. The radicle probably breaks through the stone to one side of the base. The seedling has two pairs of leaves separated 646 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 by a well-developed internode. Subsequent growth is rather slow, but the young plants seen were growing on a dry ridge, which may have made their growth slower than normal. The young plants are erect, and the terminal bud is functional. Young plants 30 cm. tall lacked the characteristic short terminal internode on each innovation, but this was present on innovations of plants about 1.6 meters tall. (Stevens 702, from Sabah.) Locat use. The timber is used for building. Calophyllum obliquinervium is a distinctive species. Its twigs dry dark brown, the terminal internode of each innovation is notably shorter than the others, its leaf blades have steeply ascending venation, its inflorescences are terminal and usually unbranched, and its fruits are fairly small. The epithet obliquinervium (‘‘oblique nerves’’) was suggested by the characteristic venation of this species. Anderson felt that Calophyllum benjamina and C. obliquinervium could be separated on bark characteristics: the bark of specimens from kerangas vegetation is gray and fissured (see his notes on S 27681 at Kew), while that of specimens from peat swamps is yellow. Although his later reduction of C, benjamina to synonymy under C. obliquinervium (Ande 5, 1OC. CIE.) is correct, he included specimens of C. ardens in C. he i Ca ophyl- lum ardens is a characteristic peat swamp species that apparently always has yellow bark; the numerous differences between it and C. obliquinervium, a species usually of drier ground, are discussed under C. ardens. The similarity between Calophyllum obliquinervium and C. sakarium is only superficial, as the discussion after the latter species indicates. Specimens of Calophyllum obliquinervium from Borneo are similar to those from the Philippine Islands, although the Bornean specimens have slightly smaller flowers. Herre 1087, from Culion Island, in the Philippines, has pedicels only 3-5 mm. long and a leaf blade that is rounded at the base: it is not otherwise different from specimens collected in the Philippine Islands. 161. Calophyllum sakarium P. F. Stevens Figure 42, d-f. A speciebus aliis Calophylli in lamina mediocra obovata nervis lateralibus 3 usque ad 7 per 5 mm. (in pagina superiore ut videtur 6 usque ad 14 per 5 mm.), inflorescentiis terminalibus, floribus cum 8 tepalis, et fructu 1.1—1.3 cm. longo, differt. Tree 12-24 meters tall, d.b.h. to 65 cm.; outer bark yellowish green to yellowish white, scaly or fissured; inner bark pale red or red-brown: latex yellow. Twigs flattened, 2-3(-3.5) mm. across, + strongly 4-angled, drying brown, soon glabrescent; axillary innovations lacking basal scars; internodes 1.5-3 (—4.5) cm. long, becoming shorter toward apex (uppermost internode less than V2 length of others); uppermost pair of axillary buds rounded, ca. | mm. long, spreading; terminal bud bluntly conoid, 1.4-2 mm. long, with short, brown, tomentose indumentum (hairs, Ficure 41, h-j), underdeveloped inter- node 0.5-1.5 mm. long. Petiole 0.6-1.7 cm. long, shallowly concave above, 1980] STEVENS, CALOPHYLLUM 647 Figure 42. a-c, Calophyllum ardens (BRUN 297). a, habit, x 0.5. b,c, fruit x 0.75: b, longitudinal section; c, from outside. d-f, C. sakarium. d, SAWN 57404, terminal bud, x 3. e, f, SAN 21622, fruit, x 3: e, longitudinal section; f, from outside. convex below, glabrescent; lamina obovate, 4-9 by 1.7-4.7 cm., rounded to shallowly retuse at apex, cuneate to acute at base, distantly undulate but not recurved at margin, coriaceous, drying olivaceous above and sabelline below, often with grayish covering, soon glabrescent or sparsely and persis- tently puberulent on midrib below, the midrib above narrowed gradually from base, flat to slightly raised, 0.3-0.7 mm. wide at midpoint, below slightly raised, striate or not, the venation subobscure above and below, latex canals as prominent as veins above, 3 to 7 veins/5 mm., angle of divergence 40-55°. only 0.3 mm. in terminal inflorescences; bracts unknown; pedicels 0.7—2.5 648 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 cm. long, glabrous. Flower (?)hermaphroditic; tepals 8 (or 9), the outer pair suborbicular, ca. 3 by 3 mm., when 8 the inner ones obovate to suborbicular, 5-11 by 4-7 mm., when 9 innermost one ca. 6.5 by 3 mm.; stamens 105 to 160, the filaments to 4 mm. long, the anthers suboblong, 1.2-1.7 mm. across, retuse at apex; ovary ca. 1.5 mm. long; style ca. 3 mm. long; stigma peltate, (?)1-1.5 mm. across, 2-radiate. Fruit ellipsoid, ca. 1 by 0.8 cm., apiculate, drying vinaceous-brown, wrinkled; outer layer detaching from stone only with difficulty, 0.2-0.3 mm. thick, compact, (?)air spaces developing at maturity; stone ellipsoid, ca. 0.8 by 0.7 cm., rounded at apex, the walls 0.2-0.3 mm. thick, smooth, unmarked; spongy layer initially thick. . Sabah, Lahad Datu, V.J.R. Silam, 240 m., 12 Dec. 1966, SAN 57404 an ‘Sihonseo! (holotype, L; isotypes, K, SAN). DiIsTRIBUTION. Borneo (southeastern Sabah), local (Map 48). ADDITIONAL SPECIMENS SEEN. Borneo. SAasan. Lahad Datu: Pulau Sakar, 60 m., SAN 21612 (Kk, KEP, L, SAN), 75 m., SAN 21613 (kK, KEP, L, SAN, SAR, SING), 6 m., SAN 21622 (A, K, KEP, L, SAN, SAR, SING), SAN 24569 (kK, KEP, L, SAN, SING), 36 m., SAN 26993 (a, K, KEP, L, SAN, SAR, SING); Silam, ca. 36 m., SAN 29384 (kK, KEP, L, SAN, SAR, SING); Mile 1, Silam Road, 90 m., SAN 47686 (A, K, L, SAN); Mile 13, base of Mt. Silam, 75 m., SAN 73422 (SAN). Eco.Locy. Well-drained ridges or slopes, once in soil derived from ultramafic rock (SAN 21612); 6-240 m. alt. Flowering December, and February to April; submature fruit in April. Calophyllum sakarium can be readily distinguished by its obovate leaf blades that are rounded at the apices; its steeply ascending, rather distant venation that appears to be twice as dense on the upper surface of the blade the lower; its terminal inflorescences; and its small, ellipsoid fruits. The specific epithet is taken from the name of an island, Pulau Sakar, where this species has been collected. There is considerable variation in internode length along both vegetative and inflorescences axes. Although the terminal two pairs of leaves are never pseudoverticillate, as in Calophyllum obliquinervium, the uppermost internode may be less than half the length of the penultimate one. In addition, there is a tendency for the first internode of the axillary innovations to be notably longer than the rest. However, the basal internode of terminal inflorescences varies greatly in length and is frequently much shorter than subsequent ones. The relationships of Calophyllum sakarium are obscure. Although both C. ardens and C. obliquinervium have medium-sized leaves with steeply ascending venation, and C. obliquinervium has terminal inflorescences, the two species are readily distinguished by the characters given above. In addition, both species have denser venation (apparently equally dense on both surfaces of the lamina); C. obliquinervium has leaf blades that are usually acuminate at the apices; and C. ardens has much larger fruit. An anatomical feature that readily distinguishes C. sakarium is the presence of a large latex canal replacing most of the vascular tissue in alternate vascular bundles; each 1980] STEVENS, CALOPHYLLUM 649 normal vascular bundle is separated by three latex canals. This feature has not been observed in any other species. Calophyllum sakarium is possibly closer to C. pentapetalum, which also often has terminal inflorescences and small fruits. However, the latter species has an often smaller lamina that is rarely obovate in shape, denser venation (the same density on both surfaces), and hairs that are usually strongly papillate or even branched at the base. 162. Calophyllum ardens P. F. Stevens, sp. nov. Ficure 42, a-—c. C. obliquinervium auct., non Merr.; J. Anderson, Gard. Bull. Singapore 20: 154. 1963, pro majore parte: Pukol & Ashton, Checklist Brunei Trees, 83. 1964, pro parte; Smythies, Common Sarawak Trees, 61. p/. 21. 1965. Calophyllum sp. nov. J. Utero Trees Peat Swamp Forest Sarawak, 86. pl. 27c. 1972. A speciebus aliis Calophylli in foliis patentibus, lamina elliptica valde coriacea margine plana nervis lateralibus sub angulo 25—35° e costa abeuntibus, et fructu magno circa 3 cm. longo putamine parietibus circa 1.2 mm. crassis, iffert. Tree 15-30 m. tall, d.b.h. to 45 cm.; trunk without buttresses, numerous knee roots present; outer bark ochraceous, smooth, hard, with thin, curved scales; inner bark reddish brown, lamellate, or in transverse section bright yellow with dark brown lamellae; latex bright yellow, opaque, or brown, very Viscous. Twigs flattened, 2.5-3 mm. across, not angled or obscurely so, drying brown to blackish, glabrous at maturity; axillary innovations lacking basal scars, such scars perhaps present in young plants; internodes 1.5-4.5 cm. long; uppermost pair of axillary buds rounded, to 1 mm. long, spreading; terminal bud plump, 4.5-6 mm. long, with crustaceous, brown indumentum (hairs, Figure 41, aa), underdeveloped internode absent. Petiole 2-2.8 cm. long, broadly concave above, convex below, glabrous; lamina elliptic or cuneiform to obovate, 4.5-8.5(-10.7) by 1.7-4.6 cm., retuse or rounded to acute at apex, narrowly cuneate at base, quite flat at margin, very coriaceous, drying honey above and sabelline below, glabrous, midrib above narrowing gradually from base, flat to slightly depressed, surrounding lamina obscurely raised, (0.25-)0.4-0.6 mm. wide at midpoint, disappearing ca. 1.2 cm. below apex, below + flat, not conspicuous, the venation inconspicuous on both surfaces, slightly raised, 5 to 10 veins/5 mm., angle of divergence 25-35°. Infructescences from foliate axils, with scars of 7 to 14 flowers, sometimes with 3-flowered branches to |.6 cm. long, the axis 3.5-6 cm. long, glabrous, lowest internode 0.7-2 cm. long; bracts unknown; pedicels in fruit (0.2—)1.5— cm. long, glabrous. Flower not known. Fruit ellipsoid, 2.7-3 by 2-2.2 cm., subrounded at apex, drying brown, inconspicuously striate; outer layer detaching cleanly from stone, ca. 0.8mm. thick, compact, hard; stone ellipsoid, ca. 2.6 by 1.95 cm., obtuse at apex, the walls ca. 1.2 mm. thick, smooth, unmarked, paler area ca. 6 mm. across at base; spongy layer thin. 650 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Tyre: Brunei, Tutong, Bukit Pasir Puteh, 1.5 m., 25 July 1957, BRUN 297 coll. Ashton (holotype, L; isotypes, BO, BRI, KEP, SAR, SING). DistRiBUTION. Northern and western Borneo, scattered (Map 48). ADDITIONAL SPECIMENS SEEN. Borneo. Sarawak. 3rd Division: Loba Kabang South Protected Forest, S 659 coll. Anderson, 12 Jan. 1954 (KEP, sar, SING), KEP 79327 (a (frag.), KEP). Brunet: Telamba, KEP 37081 (KEP); Badas, KEP 48498 (KEP). KALIMANTAN. Barat: Kenepai, Teysmann, HB 8050 (po); Sambas, Paloh, 5 m., bb 13885 (Bo). EcoLocy. Low-altitude peat swamps; rare in alan forest (Shorea albida- Gonostylus—Stemonurus association), more common in Shorea albida—domi- nated forest, locally abundant in Shorea albida—Litsea—Parastemon association (Anderson, 1963, op. cit.); below 5 m. alt. Fruiting in July Local NAME AND USE. An Iban name, ‘‘kayu api,’’ means ‘‘firewood.”’ ‘‘The yellow exudate of the tree is richly secreted into the outer bark, imparting to it its color, and the flakes infused with dry sap ignite easily, burning with a smoky yellow flame. Hence its use in firelighting’’ (BRUN 297). Calophyllum ardens is a very distinctive species, immediately recognizable even when sterile by its spreading leaves and by its very coriaceous leaf blades that dry quite flat and have steeply ascending venation. The epithet ardens (“‘burning’’) is appropriate since the bark is used to start fires. Although both Calophyllum ardens and C. obliquinervium have been collected at Bukit Pasir Puteh in Brunei (the latter as BRUN 5020) and have been much confused in the literature, they are easily separable. The leaf blades of C. obliquinervium have steeply ascending veins similar to those of C. ardens, but they are less coriaceous, and the leaves themselves are more or less ascending on the twigs. Calophyllum obliquinervium has terminal, not axillary, inflorescences; its fruits are about one-third the size of those of C. ardens and have a thin-walled stone. Calophyllum ardens is a characteris- tic tree of peat swamps, but C. obliquinervium is very rare there; references to the latter occurring in peat swamps (e.g., Whitmore, 1975, p. 153) usually refer to C. ardens. There are anatomical differences, the most obvious being the thickness of the cuticle /epidermal cell wall complex; 25-35 jm. on both leaf surfaces in C. ardens, and less than 10 pm. in C. obliquinervium. The leaves from apparently young plants are narrowly elliptic and measure up to 12.5 by 2 cm. Axillary innovations in such specimens appear to have basal scars. (Teysmann, HB 8050.) 163. Calophyllum sp. Ficure 40, m-o. Tree 4.5—-25 meters tall, d.b.h. to 48 cm.; outer bark shallowly fissured and flaked. Twigs flattened, 1.8-2.5 mm. across, 4-angled, drying dark brown, transiently subfarinose; axillary innovations lacking basal scars; internodes (0.5—)1.2-3.5 cm. long; uppermost pair of axillary buds + rounded, ca. | mm. long, spreading; terminal bud plump to conical, 3-4.5 mm. long, with grayish brown, subcrusta- 1980] STEVENS, CALOPHYLLUM 651 ceous indumentum (hairs, Ficure 43, v—-x), underdeveloped internode to 3(-7) mm. long. Petiole 0.8-1.3 cm. long, deeply concave above, convex below, glabrous; lamina elliptic to suboblong, 4.8-12.5 by 1.8-4 cm., acute to subacuminate at apex, acute to attenuate at base, undulate and slightly recurved at margin, coriaceous, drying hazel to olive above and fulvous below, sparsely farinose-puberulent on midrib below when young, the midrib above narrowing gradually from base, raised, 0.12-0.2 mm. wide at midpoint, below raised, + angled to substriate, the venation subapparent above and below, raised, latex canals sometimes subimpressed on both surfaces, 6 to Il veins/5 mm., angle of divergence 70-80°. Infructescences from foliate axils, with scars of 7 to Il flowers, unbranched, the axis 2—4.7 cm. long, glabrous, lowest internode (0.2-)0.4-1.2 cm. long; bracts unknown; pedicels 0.7-1.8 by 1.2-1.5 mm., glabrous. Flower unknown. Fruit spherical, ca. 1.2 by 1.2 cm., apiculate or not, drying kaiser brown, sharply wrinkled; outer layer detaching cleanly from stone, 1.3-2 mm. thick, compact; stone sub- spherical, ca. 10 by 9 mm., rounded at apex, the walls ca. 0.15 mm. thick, smooth, unmarked; spongy layer thick at first. DistripuTIoN. Northeastern Sarawak, (?)Brunei (Map 48). SPECIMENS SEEN. Borneo. Sarawak. 3rd Division: Balleh, Ulu Mujong, Temiai/Temalad watershed, 1100 m., S 13399 (a, K, KEP, SAN, SAR, SING); Carapa Pila, Ulu Mujong, Balleh, 900 m., SAN 19958 (a, BO, K, KEP, SAN, SAR, SING); Ulu Pila/Mujong watershed, 850 m., S 19612 (A, K, KEP, SAN, SAR, SING); Kapit, Melinau, Bukit Mabong area, 1006 m.. S 25796 (SAR). BRUNEI: G. Pagon Periok, 1500 m., BRUN 1372 (?) (sar). Sine loco: S 10634 (sar). EcoLocy. Locally common in lower montane forest on basalt or dacite, 900-1500 m. alt. Fruiting in April; fruit green. Calophyllum sp. 163 can be recognized by its short, subconoid terminal bud often with an obvious underdeveloped internode; its inconspicuous indumentum; its more or less elliptic leaf blades that dry bicolored (hazel to olivaceous above, fulvous below); and its spherical, wrinkled fruit with a compact outer layer |.3-2 mm. thick. Because of the way it dries, Calophyllum sp. 163 is superficially similar to some members of the C. blancoi complex, but its short terminal bud and unbranched hairs readily distinguish it. Calophyllum incumbens also tends to dry a similar color to Calophyllum sp. 163, but that species has a different terminal bud and fruit (cf. Ficures 40, m-o, and 32, a, b). Calophyllum tetrapterum var. obovale may have a terminal bud somewhat similar to that of Calophyllum sp. 163, but its obovate leaves have an obvious thickened margin, and the fruits are usually smaller and have a much thinner outer layer. BRUN 1372 possibly does not belong here; it is a sterile specimen, and Oo handwriting, but the adjacent numbers were collected by Smythies on Mt. Kinabalu, Sabah 652 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 q. rt, y, Williams 639, large hairs; s, PNH 39394, from inflorescence axis. t, u, C. rigidulum (BS 45686). V-X, Calophyllum sp. 163: v, x, S 13999, Scale = 60 um. (in c, d, scale = 120 pm.) 1980] STEVENS, CALOPHYLLUM 653 164. Calophyllum sp. Tree ca. 10.5 meters tall, d.b.h. ca. 28 cm.; trunk and bark unknown. Twigs flattened, |-1.3 mm. across, 4-angled, drying brown, farinose when young; axillary innovations lacking basal scars; internodes 0.5-3 cm. long; uppermost pair of axillary buds rounded, ca. 0.5 mm. long, spreading; terminal bud plump to narrowly conical, 1.3-2 mm. long, with subcrustose to subpu- berulent, brown indumentum, underdeveloped internode 1-6 mm. long. Petiole 4-6.5mm. long, deeply concave above, convex below, glabrous; lamina elliptic, 4.1-6.7 by 1.3-2.7 cm., acuminate at apex, cuneate at base, slightly undulate and barely recurved at margin, thinly coriaceous, drying + sabelline above and below, glabrous when mature, the midrib above gradually narrowed from base, raised, strongly sulcate at first, ca. 0.2 mm. wide at midpoint, below raised, rather inconspicuous, obscurely angled, the venation apparent above, subobscure below, raised, 13 to 18 veins/5 mm., angle of divergence 65-70°. Inflorescences from foliate axils, with 5 to 7 flowers, unbranched, the axis 0.3-1.2 cm. long, glabrous, lowest internode 1.5-7 mm. long; bracts unknown; pedicels 7-8 mm. long, glabrous. Flower (?)hermaphroditic; tepals 8, glabrous, the outer pair suborbicular to broadly ovate, 4-4.5 mm. long and across, only slightly concave, inner ones obovate, 6-7 by 2.5-3.5 mm.; stamens ca. 230, the filaments to 5 mm. long, the anthers elliptic, 0.25-0.3 mm. long, subapiculate to subretuse at apex; ovary ca. | mm. long, the style ca. 3.7 mm. long, the stigma peltate, ca. 0.7 mm. across, (?)radiate. Immature fruit ovoid, 7 mm. by 4 mm., apiculate. DistripuTion. Sarawak (Map 47); known only from a single specimen. SPECIMEN SEEN. Borneo. Sarawak. 3rd Division: Anap, Bukit Mersing, 800 m., S 22125 (K, SAR). Eco.ocy. Mixed dipterocarp forest on basalt hillside, ca. 800 m. alt. Flowering in September. Calophyllum sp. 164 can be recognized by its small, slender terminal bud, which may be nonfunctional, and its almost equally long and slender under- developed internode. Its elliptic leaves have fairly dense venation more prominent above than below, and its flowers have very small anthers. 165. Calophyllum gracilipes Merr. Philip. Jour. Sci. C. 5: 197. 1910, Enum. Philip. Fl. Pl. 3: 79. 1923. Type: Philippine Islands, Mindanao, Zam- boanga Province, Port Banga, 600 m., 3 Feb. 1908, FB 9405 coll. Whitford & Hutchinson (isotypes, kK, Ly, US). C. tetragonum Merr. Univ. Calif. Publ. Bot. 15: 199. 1929; Masamune, num. Phanerog. Born. 476. 1942; Keith, N. Borneo Forest Rec. ed. 2. 2: 315. 1952 (C. sp. cf. tetragonum’’); Meijer, Bot. Bull. Herb. Forest Dept. Sabah 7: 15. 1967. Type: Borneo [Sabah], near Sandakan, Sapajaya River, 10 m., 21 March 1923, D. D. Wood 1312 (holotype, UC; Isotypes, A, BO). C. tetragonum Merr. var. parvifolium Merr. Univ. Calif. Publ. Bot. 15: 199. 1929; Masamune, Enum. Phanerog. Born. 476. 1942. Tyre: Borneo 654 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 [Sabah], Elphinstone Province, Tawao, eae 21577 (lectotype, uc; isolectotypes, A, BISH, BM, C, F, G, GH, K, M, MO, NY, P, SING, U). C. cuspidatum Ridley, Kew Bull. 1938: 119. 1938; Masamune, Enum. Phanerog. Born. 475. 1942. Type: Sarawak Ds Division], Baram, March 1895, Hose 37 (holotype, k; isotypes, BM, C. pulcherrimum auct., non Wall. ex Choisy: Merr. Bibl. Enum. Born. PL 393, 1921 Masamune, Enum. Phanerog. Born. 476. 1942. Tree (rarely shrub), 3.724 meters tall, d.b.h. to 30(2-60) em.; trunk without buttresses (rarely with short, spreading spurs); outer bark yellowish or orange-brown (brown), smooth or finely cracking and scaling, hoop marked or not, the inner surface bright orange (rarely blackish or reddish (in Sabah)); under bark red; inner bark red to pale red; latex whitish- or grayish-brown to lemon yellow, not very sticky (rarely clear yellow, sticky). Twigs somewhat flattened, 0.6-2.5 mm. across, strongly 4-angled when young, soon becoming rounded (rounded), often with obscure horizontal lines at nodes, drying yellowish, greenish, or brown, glabrous or transiently farinose to puberulent; axillary innovations lacking basal scars; internodes 0.5—4 cm long; uppermost pair of axillary buds rounded to pointed, to | mm. long, erect to spreading, often inconspicuous; terminal bud narrowly conical, 1-6 mm. long, with brownish, crustaceous to adpressed (short-tomentose) indu- mentum (hairs, Figure 41, p-r, bb, cc), underdeveloped internode to 1 mm. long. Petiole (0.2—)0.35—1.4(-2.4) cm. long, deeply concave above, con- vex beneath, usually glabrous; lamina narrowly elliptic or elliptic to ovate (subobovate), (2.5—)4—14(-16) by (0.75-)1.3-5.4(-6.2) cm., acuminate to long- acuminate at apex, narrowly to broadly cuneate (rarely acute) at base, strongly and closely undulate and narrowly recurved at margin, thinly coriaceous, drying umber to sepia above and fulvous to grayish below, midrib often yellowish, glabrous or shortly and subpersistently brown-pubescent on midrib below, the midrib above narrowing gradually from base, raised, center often + sulcate, 0.15—0.3 mm. wide at midpoint, below raised to almost flat, rounded to striate, the venation subapparent above and obscure to subapparent below, raised, latex canals below raised or depressed (additional obliquely ascending latex canals near base), 6 to 12 veins/5 mm., angle of divergence 50-70(-75)°. Inflorescences from foliate axils (sometimes 2 together), with 3 (to 9) flowers, unbranched, the axis 0.5-3(-5) cm. long, farinose to subtomentose near base, lowest internode 0.3—1 cm. long; bracts not seen, apparently caducous; pedicels 0.5-1.5 em. long, glabrous, in fruit to 4.3 cm. by 1.5 mm. Flower (?)her- maphroditic; tepals (?4 to) 8 (to 10), glabrous or almost so, the outer pair ovate, 4.5-5.5 by 3-4.3 mm., the inner ones oblong, elliptic, or obovate, 6.5-8 by 3.5-6 mm.; stamens 60 to 110, the filaments to 5.5 mm. long, connate for up to 0.75 mm., the anthers suboblong, 0.4—0.9(-1.2) mm. long, shallowly retuse at apex; ovary 1.3-2 mm. long, the style 4.5-6 mm. long, the stigma subpeltate, apparently asymmetrical, 0.5-0.8 mm. across. Fruit subspherical, 1.3-2 cm. by 1.2-2 cm., apiculate or not, furfuraceous, drying rown, broadly wrinkled; outer layer detaching cleanly from stone, 0.6—1.5 mm. thick, air spaces developing under skin and next to stone, otherwise compact; stone subovoid to ellipsoid, 1-1.4 by 0.8-1.15 cm., rounded to 1980] STEVENS, CALOPHYLLUM 655 apiculate at apex, the walls less than 0.2 mm. thick, smooth, unmarked; spongy layer thin DistriBuTION. Borneo and Mindanao (Map 46). SELECTED SPECIMENS SEEN (for explanation of letters, see text). Borneo. Sarawak. Ist Division: G. Gading, 640 m., S 34645 (a, san, SAR); G. Buri, 75 m., S 36700 (sar); Sabal Tapang, 175 m., Stevens et al. 179 (a); G. Matang, 610 m., Stevens et al. 270 (a). 3rd Division: Kapit, Bukit Raya, 600 m., S 23910 (a, BO, K, SAR, SING); Kapit, Balleh, Sungei Mengiong, S 29682 (a, E, K, SAR, SING); Bukit Goran, 420 m., S 36259 (sar); Bukit Bukar, near Takalit, Katibas, 740 m., S 36398 (sar). 4th Division: Ulu Mayeng, Kakus, 60 m., S 21841 (y) (4, BO, KEP, P, SAN, SAR, SING); Bukit Mersing, Anap, 150 m. S 21903 (y) (A, K, KEP, SAN, SAR, SING). 5th Division: Limbang, Ulu Medamit, 305 m., S 32327 (Bo, kK, SAN, SAR, SING). Brunel: Labi Hills, F.R., KEP 48124 (KEE), SABAH. Sipitang: Mensapol, SAWN 65998 (L, san). Kota Kinabalu: Gaya Is. below trig., Jesselton side, 60 m., SAN 33559 (kK, KEP, SAN, SAR, sinc). Kota Belud: Bukit Matindok, 12.6 km. from Kota Belud, 310 m., SAN 32182 (L, SAN, SAR); Mt. Templer F.R., SW. of Sungei Talupit, 457 m., SAN 76245 (K, SAN, SAR). Tenom: G. Lumaku F.R., W. of Sapong Est, SAN 43154 (K, KEP, L, SAN, SAR, SING); Pangi, 8 km. WNW. of Tenom, 579 m., SAN 15115 (a, BO, KEP, SAN, SING); Mile 79, Pangi, 150 m., SAN 5047] (san). Ranau Mt. Kinabalu, E. shoulder, 762 m., RSNB 209 (a, B, BO, CANB, K, AN, sING), Lohan, 1372 m., SAN 44653 kK. L, SAN, SAR, SING); half way eenane Tambunan and Keninguan, 732 m., KEP 80422 (ker); along Mamut road, E. of Mamut camp, 900-1400 m. Rokawa & Hotta 5943 (san). Labuk & Sugut: Telupid, S. of Agricultural Station, 60 m., SAN 53625 (san); Ulu Sungei Segualid Lokan F.R., mile 41 Labuk Road, SAN 72628 (k, san). Tingod, Lagsikan F.R., 90 m., A 3409 (k, KEP, L, sinc); Dusun, 60 m., SAN 77366 ae Kinabatangan: Lamag, Bukit Garam, 36 m., SAN 36563 (k, L, SAN, SING); Sungei Pin V.J.R., SAN 5204] (san). Lahad Datu: Bukit Pintasan, Ulu Segama, SAWN 79109 Se near Kuala Sungei ea SAWN 71040 (San); Mostyn, Madai F.R., 45 m., SAN 67748 (san). Tawau: near Tawau, Elmer 21121 (A, BISH, BM, BO, C, F, G, GH, K, M, MO, NY, P, SING, U, UC, US); Kelumpang, Balong, 30 m., SAN 17342 (k, KEP, L, SAN, SAR, SING); Gemok Hill, 180 m., SAN 39219 (x, Bs ou Ulu Sungei _ SAN 32955 (k, L, SAN); Membalu F.R., 60 m., SAN 22770 (a, BO, K, KEP, SAN, SAR, SING); [Sungei] Umas-Umas, 6m., SAN 4575 (k, SING). noun Timur: Kabiran, Sungei Scan 100 m., bb 11657 (Bo). Barat: Liang Gagang, Hallier 2992 (?) (A, BO, K, L); Sekadau, Pait, 250 m., bb 8029 (y) (po), Tamang, 50 m., bb 8027 a (BO). Philippines. MinpDANao. Zamboanga: Sax R., Williams 2193 (GH, K, NY, US). Ecotocy. Mixed dipterocarp forest, to 1372 m. alt. Flowering February (Philippines), March, and July to September (flower (really stamens?) some- times yellowish or greenish); fruiting January, February, and May to No- vember (fruit yellow to olive or brownish). GERMINATION AND YOUNG PLANT. The radicle breaks the stone just to one side of the base. The seedling has two pairs of leaves separated by an internode to ca. 5 mm. long. Subsequently produced internodes are a little longer; 656 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 the terminal bud is functional; the stem is initially arched and only later straightens. (Stevens et al. 159, 179A, 360.) Calophyllum gracilipes can be recognized by its thinly coriaceous leaf blades that are more or less undulate ie slightly recurved at the margin, its few-flowered inflorescences, and its subspherical, furfuraceous fruits. The epithet gracilipes - cane foot’ “e was chosen because of the distinctive long, slender pedi alophyllum ies is closely related to C. aurantiacum and C. recurva- tum; for differences that separate C. gracilipes from these species, see C. aurantiacum and C. recurvatum. alophyllum gracilipes has been confused with C. tefrapterum var. tetrap- terum, but the two are not closely related. In C. tetrapterum var. tetrapterum the lamina is a darker color when dry, and the margin is less prominently undulate. The flowers of the two species are similar, although C. tetrapterum var. tetrapterum has more profuse inflorescences. In fruit the two are quite different: the fruits of C. tetrapterum var. tetrapterum are about half the size of those of C. gracilipes, they lack a furfuraceous surface, and they have a thinner outer layer. There is a considerable amount of vegetative variation within Calophyllum :; g species. Specimens in the list above that are denoted by ‘‘y’’ have a fairly large lamina (toward the upper limits of the measurements given), the midrib on the lower surface is rather strongly raised and yellowish in color, and the veins appear to be dense because of the prominence of the latex canals. However, these characters vary individually between other specimens. The indumentum on the terminal bud is quite variable in its development: although normally subcrustaceous to very short-tomentose, it is sometimes tomentose (e.g., S 36700, Stevens et al. 159; see also Calophyllum recurvatum). The specimens of Calophyllum gracilipes from Brunei and the 3rd Division of Sarawak have slender twigs 0.6-1 mm. across, petioles 2-5 mm. long, and narrowly ovate to elliptic leaf blades 2.7-6.6 by 0.75—2(-2.7) cm. that are long-acuminate at the apex. These specimens include the type of C. cuspidatum, but they cannot be clearly distinguished from other specimens with larger leaves and a coarser facies. Hallier 2992 is included only with hesitation. It has leaf blades that are rounded at the base and flowers with only four tepals. Another noteworthy specimen is SAN 33559, which has very small leaves less than 3.5 cm. long: this merely represents the lowermost extreme of great variation in leaf size within Calophyllum gracilipes. The specimens determined by E. D. Merrill as Calophyllum gracilipes and C. tetragonum are very similar; in his description of the latter species, Merrill noted that its alliance was clearly with C. gracilipes. When described, C. tetragonum var. parvifolium was distinguished from var. tetragonum by small differences in leaf size; it is clear from the preceding discussion that it is impossible to recognize variants in C. gracilipes based on leaf size. Calophyllum 1980] STEVENS, CALOPHYLLUM 657 tetragonum var. parvifolium has been lectotypified on the sheet of Elmer 21577 at uc. Elmer 21577 and the syntype, Elmer 21751, are almost identical; they both agree with the protolog and have been widely distributed. 166. Calophyllum recurvatum P. F. Stevens, sp. nov. Ficure 44, a, b. A Calophyllo gracilipes, quo aliter simile est, in gemma terminali indumento tomentoso praedita, ramulis subalatis, ramulis, foliis, floribus et fructibus grandioribus, et lamina margine valde recurvato pagina inferiore venulis lateralibus haud conspicuis, differt. Tree 3-20 meters tall, d.b.h. to 23 cm.; trunk without buttresses; outer bark reddish brown, brown-yellow, or grayish, sometimes mottled, with lines of lenticels, the inner surface bright orange to orange-red; under bark pale red to reddish; inner bark rose; latex yellow or clear yellow-brown, not viscous, or almost colorless and watery. Twigs slightly flattened, 2.5-3.7 mm. across, strongly 4-angled to subalate, often with obscure horizontal lines at nodes, drying brown, glabrous or sparsely tomentose when young; axillary innovations lacking basal scars; internodes erect, inconspicuous; terminal bud broadly conical, 3-6.5 mm. long, with brown, tomentose indumentum (hairs, Ficure 41, v, w), underdeveloped internode absent. Petiole 0.8-2.2 cm. long, narrowly concave above, convex below, sometimes transiently tomentose below; lamina elliptic to suboblong, (9-)12-24 by (2.6-)3.7-9.8 cm., gradually acuminate at apex, cuneate to acute at base, rather distantly undulate and strongly but narrowly recurved at margin, coriaceous, drying sepia to gray-olivaceous above and sepia to wood-brown below, glabrous or transiently tomentose on midrib below, the midrib above narrowing gradually from base, raised, 0.2-0.35 mm. wide at midpoint, below raised, angled, the venation apparent and slightly raised above, subobscure below, 5 to 9 veins/5 mm., angle of divergence 70-80°. Inflorescences from foliate axils, with 3 to 9 flowers, unbranched, flabellate, the axis (0.3-)2-6.2 cm. long, sparsely tomentose, at least at base, lowest internode 0.3-2(—4.3) cm. long (exceptionally terminal, with up to 13 flowers, branched, axis to 8.5 cm. long); bracts ovate to elliptic, 5-7 mm. long, deciduous; pedicels (0.5-)1-2.2 cm. long, glabrous, incrassate in fruit, to 3.5 mm. thick, flattened. Flower (?)hermaphroditic; tepals 8 to 12, the outer pair ovate, 5-9 by 5-7 mm., the next pair broadly elliptic, 11-12 by 8.5-9.5 mm., the inner ones narrowly elliptic to obovate, 10-12 mm. by 3.5-5 mm.; stamens ca. 125, the filaments up to 3.5 mm. long, + connate, the anthers oblong, 1.5-2.1 mm. long, eee to subretuse at apex; ovary ca. 2.5 mm. long, the style 45.7 mm. long, the stigma asymmetric, ca. | mm. across. Fruit subspherical, ca. 2.3 by 2 cm., apiculate, furfuraceous, drying brown, broadly wrinkled; outer layer detaching cleanly from stone, ca. 0.7 mm. thick, large air spaces developing; stone spherical, ca. 1.3 by 1.3 cm., rounded at apex, the walls ca. 0.25 mm. thick, smooth, unmarked; spongy layer thin. Type: Sarawak, Ist Division, Kuching District, 12th mile, Penrissen Road, 658 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 61 Ficure 44. a, b, ra aaa recurvatum (S 26205): a, terminal bud, x f, 3; b, habit, x 0.5. c-f, C. aurantiacum (Stevens et al. 195). c-e, fruit: c, from pte x |; d, ee section, X 1; e, surface, x = f, terminal bud, eG, rigidulum (BS 45686), el bud, x 6. h, 1, Calophyllum sp. 70 eure 38909), fruit, x I: h, longitudinal Aare hon outside. 19 Oct. 1966, S 26205 coll. ak Nudong & ak Bubong (holotype, a; isotypes, K, L, SAN, SAR). DistripuTION. Borneo (Map 47). 1980] STEVENS, CALOPHYLLUM 659 ADDITIONAL SPECIMENS SEEN. Borneo and adjacent islands. Sarawak. Ist Division: Semengoh Arboretum, S 24338 (k, L, SAN, SAR, SING), S' 25370 (K, KEP, L, SAN, SAR, SING), 90 m., S 7737 (Bo, kK, L, SAR, SING), Stevens et al. 143 (a); Lundu, Bukit Jebong, 60 m., Stevens et al. 212 (a), 90 m., Stevens et al. 218 (a), 120 m., S 15450 (bo, k, L, sar); G. Gading, 686 m., S 35069 (SAR). Sine loco: Native collector for Bur. Sci. Philip. 641 (a, p, Ge). Beccari, PB 2304 (rt). Kavimantan. Timur: Bukit Liang Karing, Jaheri 1224 (Bo); Lilibloelan Tepoetsy, Jaheri 882 (no). Barat: G. Sendjudjul, 80 m., Hamzah BO). Ecotocy. Mixed dipterocarp forest, sometimes over limestone; 40-686 m. alt. Flowering August and September; fruiting September and December (fruit greenish yellow or pale green). GERMINATION AND YOUNG PLANT. The radicle probably breaks through the stone to one side of the base. The seedling has two pairs of leaves separated by an internode less than 5 mm. long. Subsequent growth is slow, a young plant 22 cm. tall having 14 internodes in 12 cm., and although the terminal bud is functional, the stem is initially curved and straightens only after some time. (Stevens et al. 143.) Calophyllum recurvatum can be recognized by its tomentose indumentum, subalate twigs, fairly large leaf blades with sharply recurved margins, flowers with eight to twelve tepals, and subspherical, furfuraceous fruits. The filaments are connate for much of their length; as a result, the androecium may remain on the old flower after the tepals have fallen—an unusual condition in the genus. The epithet recurvatum (‘‘recurved’’) was chosen because of the sharply recurved edges of the leaf blades. Although Calophyllum recurvatum is a distinct taxon, it is not easy to decide at what rank it should be recognized. The similarity between C. recurvatum and C. gracilipes extends to the young plant (in both, the main shoot is initially curved, only later straightening) and to their somewhat similar yellowish bark. Moreover, there are three specimens more or less intermediate between the two. S 36700 (Sarawak, Ist Division, G. Buri, 75 m.) has a densely tomentose terminal bud and a sparsely tomentose stem and inflo- rescence axis, an elliptic leaf blade the size of that of C. gracilipes (up to 9 by 2.8 cm.) that is subpruinose below, and rather large flowers. Stevens et al. 159 is a sterile specimen from Gunong Matang (lst Division) with the vegetative characters of S 36700. Kostermans 6013 (Kalimantan, E. Kutei, Menubar R., NE. of Sangkulirang, 100-200 m.) has well-developed, tomentose indumentum, but in other vegetative characters (except its more or less oblong lamina) it is like C. gracilipes. Further collections from southwestern Sarawak and northwestern and northern Kalimantan are needed to enable a better understanding of the relationships between the two taxa 167. Calophyllum aurantiacum P. F. Stevens, sp. nov. Ficure 44, c-f. A Calophyllo gracilipes et C. recurvato, quibus fructibus furfuraceis similibus habent, in ramulis mox demum teretibus, et folio Parvo coriaceo costa supra leviter vel haud elevata et ante apicem evanescenti, differt. 660 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Sprawling shrub or tree ca. 4.5 meters tall; trunk without buttresses; outer bark yellowish (small plants) or grayish and pale brown mottled, smooth, the inner surface bright orange; under bark bright orange; inner bark pale red; latex sparse. Twigs flattened, 1.2-2.5 mm. across, slightly 4-angled or -striate, soon becoming rounded, drying brown, soon becoming dark brown or blackish, puberulent when young; axillary innovations lacking basal scars; internodes 0.4-3.5 cm. long; uppermost pair of axillary buds rounded, to | mm. long erect; terminal bud plump or conical, 1.8-3 mm. long, with short, tomentose, brown indumentum (hairs, Ficure 41, s, y), underdeveloped internode absent. Petiole 4-11 mm. long, deeply concave above, convex below, glabrous; lamina elliptic to obovate, (2.5-)3.4-6.5(-7.7) by (1.1-)1.8-3.5 cm., rounded to subretuse at apex, cuneate toward and shortly and abruptly rounded at base, slightly to broadly and distantly undulate and recurved at margin, coriaceous, drying fulvous to olivaceous above and fulvous below, glabrous, the midrib above gradually narrowed from base, not very evident, at first subdepressed, becoming elevated, 0.05—0.15 mm. wide at midpoint, disappearing 2-4 mm. before apex, below raised, + angled, the venation subevident (rarely obscure) on both surfaces, slightly raised, 5 to 9 veins/5 mm., angle of divergence 50—60°. Infructescences from foliate axils, with scars of 5 to 7 flowers, not branched, the axis 0.8-1.3 cm. long, sparsely puberulo-tomentose toward base, lowest internode (0.25—-)0.6-1.1 cm. long; bracts unknown; pedicels in fruit 0.6-1.1 cm. by 1.5 mm., glabrous. Flower unknown; old tepal persisting at base of fruit obovate, ca. 5mm. by 3.5 mm. Submature fruit subspherical, 1.2-1.6 by 1-1.6 cm., rounded at apex, furfuraceous, drying brown, smooth or broadly corrugated; outer layer detaching cleanly from stone, 0.7—1 mm. thick, subcompact; stone spherical, 0.9-1.3 by 0.8-1.2 cm., rounded at apex, the walls up to 0.2 mm. thick, smooth, unmarked; spongy layer at first well developed. Tyre: Sarawak, Ist Division, Bau, Tai Ton, ca. 105 m., 12 December 1975, Stevens et al. 195 (holotype, A; isotypes, sar and to be distributed). DistrRiBUTION. Sarawak, local (Map 48). ADDITIONAL SPECIMEN SEEN. Borneo. SARAWAK. Ist Division: Bau, Tai Ton, Bukit Numpang, 165 m., S 22784 (sar). Ecotocy. Small bush or tree of limestone hills, 105-165 m. alt. Submature fruit in December. Calophyllum aurantiacum can be recognized by its more or less terete twigs; its short terminal bud; its small, elliptic to obovate, rather coriaceous leaf blades; and its furfuraceous fruits. The midrib on the upper surface of the lamina is at most slightly elevated and disappears short of the apex. The epithet aurantiacum (‘‘orange’’) was suggested by the bright orange color of the inner surface of the outer and under bark. Calophyllum aurantiacum is related to C. gracilipes and C. recurvatum, two species that also have furfuraceous fruits; in all three the venation on 1980] STEVENS, CALOPHYLLUM 661 the underside of the fresh leaf is practically invisible. Calophyllum aurantiacum can be distinguished from these other species by the characters given above. The narrow but abrupt basal rounding of the lamina develops during the drying of the specimen—living leaves of Calophyllum aurantiacum are cuneate at the base. 168. Calophyllum griseum P. F. Stevens, sp. nov. Ficure 45, h-k. A speciebus aliis Calophylli in lamina mediocra coriacea in siccitate supra saepe cinerascenti venulis lateralibus manifestis, inflorescentia termi- nali indumento farinoso-puberulento praedita, et in fructu sphaerico in sicci- tate rugoso putamini parietibus minoribus quam 0.2 mm. crassis, differt. Tree 12-30 meters tall, d.b.h. to 40 cm.; trunk without buttresses; outer bark grayish brown, scaly, with scales yellowish brown and newly exposed bark dark brown (pale citrine), or deeply (shallowly—BRUN 670) fissured, or cracked, inner surface bright orange; under bark brownish red or dark red: inner bark brownish red or pale reddish; latex yellow, clear (yellow- green—BRUN 670). Twigs slightly flattened, 1.7-3(-3.5) mm. across, usually strongly 4-angled, drying brown to yellow, brown-farinose at first; axillary innovations lacking basal scars; internodes 1.5-5.5 cm. long; uppermost pair of axillary buds + pointed, 2-3.5 mm. long, spreading; terminal bud plump, 4.5-7 mm. long, with grayish, subcrustaceous indumentum (hairs, Ficure 43, m-o), underde- veloped internode absent to 4(-7) mm. long. Petiole 0.8-1 .6 cm. long, broadly concave above, convex below, glabrous; lamina ovate or suboblong to elliptic, 8.5-16 by 3.5-6.2 cm., subrounded or acute to acuminate at apex, acute to cuneate at base, broadly and deeply to barely undulate and slightly recurved at margin, coriaceous, drying sepia to gray-olivaceous above and sepia to hazel below, glabrous at maturity, the midrib above narrowing fairly quickly near base, raised, 0.2-0.4 mm. wide at midpoint, below raised, angled toward apex, striate toward base, venation + apparent on both surfaces, raised, 4 to 8 veins/5 mm., angle of divergence 70-80(-85)°. Inflorescences terminal and from adjacent foliate axils, with up to 20 flowers, branched, sometimes flabellate, the axis 2-4 (axillary inflorescences) or up to 9 (terminal inflores- cences) cm. long, puberulent, lowest internode 1-2 cm. long; bracts unknown; pedicels 2.5-10 mm. long, puberulent, stout. Flower (?)hermaphroditic; tepals 8, the outer pair broadly ovate, 5-5.5 by ca. 4.5 mm., short-puberulent on back, the next pair subelliptic, 7.5-10 by 4-7.5 mm., sometimes puberulent toward base, inner ones oblong to elliptic, 7.5-11 by 2.5-5.5 mm.; stamens 190 to 265, the filaments to 7 mm. long, connate basally for up to 1.2 mm., the anthers oblong, 0.6-1 mm. long, apiculate to retuse at apex, ovary 1.3-2 mm. long, the style ca. 4 mm. long, the stigma peltate, ca. 0.9 mm. across, (?)4-radiate. Fruit spherical to ellipsoid, 1.7-2.2 by 1.2-1.7 cm., rounded at apex, drying dark brown; broadly wrinkled; outer layer detaching cleanly from stone, 1.7-3.2 mm. across, thick, subcompact, air spaces developing under skin; stone subspherical, 1.4-1.8 by I-1.5 cm., rounded at apex, the walls less than 0.2 mm. thick, smooth, unmarked; spongy layer thin. [voL. 61 JOURNAL OF THE ARNOLD ARBORETUM Q GIOMIE 1980] STEVENS, CALOPHYLLUM 663 | : 6 . 100 ia lO 120] AP 49, Fela of Calophyllum poilanei (open star), Calophyllum sp. 170 Geld Star), subhorizontale (circles), C. exiticostatum (half-circles), C. griseum ee ), C. echinatum (star in solid circle), C. rigidulum (inverted triangle), Calophyllum sp. 178 (side-facing triangle), and C. brachyphyllum (erect triangle) in Southeast Asia—Malesia. Tyre: Sarawak, 4th Division, Bintulu, Nyabau catchment area, 100 feet [30 m.], 22 June 1966, S 24614 coll. ak Luang (holotype, a; isotypes, kK, KEP, L, SAN, SAR, SING). DistriBUTION. Northwestern Borneo (Map 49). ADDITIONAL SPECIMENS SEEN. Borneo. Sarawak. Ist Division: Semengoh Ar- vere ve m., Stevens et al. 131 (a); Lundu, Sungei Arok, S 9624 (Bo, nc); G. Santubong, 90 m., Stevens et al. 302 (a). 3rd Division: Ulu Rejaue FA 515 coll. Egon, 4 Aug. 1938 (kLU, sar). 4th Division: Miri, Ficure 45. a, b, ae poilanei (Poilane 6572): a, habit, terminal bud, x 3. c, C. echinatum oe /U-215), X 1.5 (axillary buds eee off). d-f, C. ee (bb 31756). d, e, fruit, x 3: d, longitudinal section; e, from outside. f, terminal bud, x 6. g, Calophyllum sp. 178 (FB Hee terminal bud, x 3. 1 C. griseum. h, S 15872, terminal bud, Ds S 15872, fruit, x 1: i, from outside; j, transverse section. k, S 25802, habit, 0.5: 064 JOURNAL OF THE ARNOLD ARBORETUM [voc. 61 near Sungei Dalau F.R., KEP 80069 (ker); Bintulu, Simalajau F.R., S 15872 (A, BO, K, L, SAN, SAR, SING); Nyabau catchment area, 120 m., S 24648 (a, K, KEP, L, NY, SAN, SAR). BRUNEI: Berakas F.R., 60 m., S 2035 (?) (sar); Anduki F.R., 3 m., S 2243 (sar); Labi Hills, Bukit Teraja, Mile 19!/4, 150 m., BRUN 670 (BO, BRI, K, KEP, L, SAR, SING); Labi, KEP 30474 (a, KEP, SAR). SABAH. Beaufort: Lumat, SAN 25802 (Bo, kK, KEP, L, SAN, SAR, SING). KALIMANTAN, Barat: Melawi Tjatit, B. Bansa, 400 m., bb 26445 (Bo, L); Melawi, B. Watas Emang nabij Ng. Betoeng, 175 m., bb 26869 (?) (L). Ecotocy. Well-drained mixed dipterocarp forest, 3-400 m. alt. Flowering March, June, July, and September (flower scented fruiting December, January, and August (fruit pale grayish brown). GERMINATION AND YOUNG PLANT. Young plants about | meter tall are erect and have a functional terminal bud; the internodes are well developed. The young plants are conspicuous because of their pale, yellowish white bark. Calophyllum griseum is a distinctive species. The upper surface of the lamina often has a grayish, waxy covering that is later rubbed off; this grayish covering suggested the specific epithet. The lamina is fairly coriaceous, with distinct and rather distant venation; the inflorescence is terminal and puberu- lent, and the pedicels and the backs of the outer tepals are also puberulent; the flower has eight tepals; and the broadly wrinkled fruits have a thick, subcompact outer layer and a thin-walled stone. 169. Calophyllum subhorizontale M. R. Henderson & Wyatt-Smith, Gard. Bull. Singapore 15: 325. pl. 11. 1956; T. C. Whitmore, Tree Fl. Malaya 2: 170. 1973. Type: Malaya, Pahang, Kuala Lipis, 500 feet [150 m.], KEP 658 coll. Carroll (holotype, sinc; isotype, KEP). Tree 6-8 meters tall, d.b.h. to 29 cm.; outer bark ochre, smooth, to purplish or yellowish, fissured; inner bark reddish to white; latex yellow, sticky. Twigs flattened, 3.5-7 mm. across, with (2 or) 4 (or 6) prominent rounded lines, drying shiny, brown when young (yellowish when older), at first puberulent; axillary innovations (?)lacking basal scars; internodes (0.5—)1—5 (-7.5) cm. long; uppermost pair of axillary buds rounded, ca. | mm. long, suberect; terminal bud plump to conical, 4-7 mm. long, with brown, short, tomentose indumentum (hairs, Figure 46, 1, m), underdeveloped internode absent. Petiole 0.5-1.6 cm. long, broadly concave above, convex below, glabrous when mature; lamina elliptic to suboblong or obovate, 10.5-27 by 5.5-9 cm., obtuse to acute at apex, acute at base, distantly undulate and slightly recurved at margin, coriaceous, drying sepia above and umber to sabelline below, + puberulent on midrib below when young, the midrib above narrowing gradually from or rather quickly near base, raised, 0.3-0.7 mm. wide at midpoint, below raised, angled, the venation above + apparent, finely raised, below + apparent to subobscure, broadly raised, 3 to 6 veins/5 mm., angle of divergence 65-85°. Inflorescences from foliate axils, with 7 to 9 flowers, unbranched, the axis (1-)2-4 cm. long, sparsely tomentose near base, lowest internode (0.3-)1-1.7(-2.5) cm. long; bracts ovate, to 4 mm. 1980] STEVENS, CALOPHYLLUM 665 amt “tel Ficure 46. Hairs (from terminal bud, unless otherwise noted). a—-d, Calo- phyllum insularum. a-c, bb 30965: a, b, apex and base of hair ca. 780 pm. long; c, base of hair ca. 420 ym. long, arrow marks abrupt transition in cell type. d, bb 30937. e-g, C. exiticostatum: e, g, bb 2299; f, Stevens et al. 959, slight birefringence. h, i, C. membranaceum (Wang 449). j, k, C. 0, Calophyllum sp. 170 eeu 38909), apex and base of hair ca. 780 um. long. Scale = 120 pm long, caducous; pedicels 1-3.5 cm. long, glabrous, in fruit to 4 cm. by 3 mm. Flower (?)hermaphroditic; tepals (8 to) 12, sometimes glabrous, the outer pair orbicular to broadly elliptic, 7-12 by 7-10 mm., the inner ones elliptic to oblong, 13-17 by 4.5-11 mm.; stamens (?)150 to 260, the filaments 4-7.5 mm. long, connate for up to 0.7 mm., the anthers oblong, |-1.9 mm. long, rounded to slightly retuse at apex; ovary ca. 2 mm. long, the style ca. 7 mm. long, the stigma peltate-infundibular, ca. 1.7 mm. across, obscurely radiate. Fruit ovoid, ca. 4.3 by 2.5 cm., acute at apex, drying vinaceous-brown, wrinkled; outer layer not detaching cleanly from stone, 1.5-3 mm. thick, compact; stone ellipsoid, ca. 2.4 by 1.9 cm., rounded at apex, the walls ca. 1 mm. thick, smooth, unmarked; spongy layer (?)thin. DistrRIBUTION. Malay Peninsula (Map 49). 666 JOURNAL OF THE ARNOLD ARBORETUM [voL. 6] SELECTED SPECIMENS SEEN. Malaya. SELANGor: Sungei Buloh F.R., KEP 5315 (KEP, SING). NEGRI SEMBILAN: G. Angsi F.R., KEP 23817 (Kep, SING). TRENGGANU: Sungei Tong ou KEP 94773 (a, KEP, L, SAN, SING); Kemaman, Bukit Kajang, 0 m., Corner s.n., 25 Nov. 1935 (sinc). PAHANG: cag Tembeling, near Kuala Keniyum, cee Redab, 240 m., FRI 8584 (k, KEP, L, sinc); Bukit Goh F.R., KEP 17316 (ker); Raub, Bilut Reserve, KEP 23381 (kep, siNG); Kuala Lipis, KEP 1267 (kep, sinc); Lesong F.R., 45 m., FRI 19825 (k, KEP); sine loco, KEP 1151 (KEP). EcoLocy. Sometimes in rocky stream beds (FRI 8584), 45-240 m. alt. Flowering March and August (flower scented); fruiting March and December. Calophyllum subhorizontale is a distinctive species that can be recognized by its relatively small, tomentose terminal bud; stout, more or less shining twigs; large, coriaceous leaf blades with distant venation; large flowers usually with more than eight tepals; and ovoid fruits. The tepals tend to persist at the base of the fruit. The fruit dries wrinkled and has a moderately thick outer layer and a well-developed stone. The epithet subhorizontale was suggested by the clear and widely ascending venation of this species. An incomplete specimen from Vietnam, Chevalier 38909, has fruits superfi- cially similar to those of Calophyllum subhorizontale, but the leaf blades, with their fine venation, are reminiscent of those of C. soulattri. At least the outer pair of tepals is densely puberulent on the back, and the outer layer of the fruit is only ca. 1.2 mm. thick and is notably fibrous (see Calophyllum sp. 170). The data for the type locality and collector were kindly confirmed by K. M. Kochummen; the collector’s name on the Singapore sheet is given as Phillips. 170. Calophyllum sp. Ficure 44, h, 1. (?)Tree; details of trunk and bark unknown. Twigs strongly flattened, 2-3 mm. across, persistently 4-subalate, drying brown, subpersistently tomentose; axillary innovations lacking basal scars; internodes (1.3-)2—7 cm. long; uppermost pair of axillary buds rounded, ca. 2 m. long, suberect; terminal bud bluntly conical, 3 to ca. (?)5 mm. long, with brown, tomentose indumentum (hairs, Ficure 46, n, 0), underdeveloped internode absent. Petiole |.1-2 cm. long, broadly concave above, convex below, glabrous when mature; lamina oblong to elliptic, 9-13 by 5.8-7 cm., retuse to shortly and abruptly acuminate at apex, acumen ca. 4 mm. long, acute at base, undulate and slightly recurved at margin, coriaceous, drying near sabelline above and below, subpersistently tomentose on midrib below, the midrib above narrowing quickly near base, raised, 0.3-0.4 mm. wide at midpoint, below raised, + angled, the venation above and below subapparent, raised, 10 to 16 veins/5 mm., angle of divergence 70-75°. Inflorescences unknown; pedicel in fruit 4-7 by ca. 2 mm., short-tomentose. Flower known only from remains at base of fruit; tepals ca. 16, the outer pair broadly ovate, 4.5-8 by 6-7.5 mm., puberulent on back, the next pair subelliptic, 7-12 by 7-11 mm., + puberulent on back, the inner ones obovate to elliptic, 1980] STEVENS, CALOPHYLLUM 667 to 18 by 11 mm., sometimes puberulent in strip down back; stamens numerous, the filaments to 10 mm. long, connate for up to 2.3 mm., the anthers ellipsoid, ca. | mm. long, retuse at apex; ovary, style, and stigma unknown. Fruit ovoid to ellipsoid, 2.5-3 by 2.1-2.5 cm., sharply pointed at apex, beak ca. 7 mm. long, drying brown, with rather distant and shallow wrinkles; outer layer detaching + cleanly from stone, ca. 1.2 mm. thick, compact except for air spaces under skin, fibrous; stone ellipsoid, 2-2.4 by 1.8-2.1 cm., rounded at apex, the walls 1.1-1.4 mm. thick, at base 2.2-3 mm. thick and slightly fibrous, smooth, obscurely triradiate at apex, basal (?)plug ca. 6.5 mm. across; spongy layer thin. Distripution. Vietnam (Map 49); known only from a single collection. SPECIMEN SEEN. Vietnam: Nhatrang, entre Suodi Ca et Sudi Giau, Chevalier 38909 (a, P). Calophyllum sp. 170 can be recognized by its bluntly conical, tomentose terminal bud, very strongly and persistently angled twigs, rather large leaf blades with close venation, tomentose pedicels, flowers with twelve tepals (outer ones tomentose on back), and large, ovoid, beaked fruits with thick stone walls. Calophyllum sp. 170 is similar in fruit to C. subhorizontale, although it differs in terminal bud, twig, and leaf. In leaves it resembles C. soulattri but differs in terminal bud, flower, and fruit. Although the fruits are not attached to the shoots, they nevertheless appear to belong to the specimen. 171. Calophyllum brachyphyllum Merr. Philip. Jour. Sci. 17: 290. 1920, Enum. Philip. Fl. Pl. 3: 78. 1923. Type: Philippine Islands, Mindanao, Surigao Province, Hegapit River, ca. 350 m., 25 April 1919, BS 34482 coll. Ramos & Pascasio (isotypes, A, K, P, ue): Small tree; trunk and bark not known. Twigs slightly flattened, 1-1.5 mm. across, obscurely 4-angled, drying brown when young, grayish when older, sparsely farinose when young; axillary innovations lacking basal scars; internodes 0.8-3.5 mm. long; uppermost pair of axillary buds pointed, ca. 1.5 mm. long, erect; terminal bud narrowly conical, 3-4 mm. long, with dense, adpressed, grayish indumentum (hairs, Ficure 41, x, z), underdeveloped internode absent. Petiole 1-2 mm. long, concave above, convex below, glabrous; lamina broadly ovate to elliptic, 1.3-2.7 by (0.85-)1-2 cm., retuse at apex, rounded to shallowly cordate at base, quite flat at margin, coriaceous, drying sepia to near sabelline above to sepia below, minutely puberulent on midrib above and below when young, the midrib above narrowing near base, becoming flat to slightly raised, margins at first + raised, center sulcate, less than 0.2 mm. wide at midpoint, below raised, striate, conspicuous, the venation above and below subobscure, very slightly raised, 13 to 17 veins/5 mm., angle of divergence 55-70°. Infructes- cences from uppermost or adjacent axils, with scars of 3 to 5 flowers, unbranched, the axis 0.5-1.8 cm. long, glabrous, lowest internode ca. 5 mm. 668 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 long; bracts unknown; pedicels |-2.5 cm. by 2 mm., glabrous. Flower unknown. Fruit spherical, to 1.3 by 1.3 cm., rounded at apex, drying dark brown, smooth; outer layer detaching + cleanly from stone, 1|.5-1.8 mm. thick, compact; stone spherical, ca. | by | cm., rounded at apex, the walls 0.5-0.6 mm. thick, smooth, unmarked; spongy layer at first thick. DistriBuTION. Philippine Islands; known only from a single collection from Mindanao (Map 49). EcoLocy. Stream-side forest, ca. 350 m. alt. (Merrill, 1920, loc. cit.). Calophyllum brachyphyllum can be recognized by its small, coriaceous leaf blades that are retuse at the apex and more or less cordate at the base; the venation is dense. The small fruit, with its relatively very thick outer layer and stone wall, is also distinctive. The epithet brachyphyllum (‘‘short leaf’’) alludes to the short leaves of this species. Calophyllum brachyphyllum is apparently a rather isolated species. It is not close to any of the cordate-leaved taxa centered on C. venulosum, nor leaves and dense venation like C. exiticostatum, but it differs from that species in its more coriaceous leaves that have a much stronger midrib, and in its fruit (that of C. exiticostatum is smaller, has a thin outer layer that is disorganized by air spaces, and has thin stone walls). 172. Calophyllum rigidulum P. F. Stevens, sp. nov. Ficure 44, g. A speciebus aliis Calophylli in lamina parva percoriacea marginibus planis costa supra subplana infra lata versus basem leviter elevata versus apicem (sub)depressa, differt. Shrub ca. | meter tall; trunk and bark not known. Twigs flattened, |-1.5 mm. across, 4-angled, drying dark brown when young, yellowish when older, sparsely farinose-puberulent; axillary innovations lack- ing basal scars; internodes 0.7-2.5 cm. long; uppermost pair of axillary buds subacute, 0.5-1.5 mm. long, erect; terminal bud plump, 3-4 mm. long, with brown, crustaceous indumentum (hairs, Ficure 43, t, u), underdeveloped internode absent. Petiole 2-5 mm. long, strongly concave above, convex below, glabrous; lamina elliptic to subobovate, 1.4—3 by 0.75-1.8 cm., rounded to slightly retuse at apex, cuneate at base, quite flat at margin, very coriaceous, drying dust color above and fulvous-umber below, when young subfarinose on and near midrib on both surfaces, the midrib above narrowed at base, at first depressed, becoming level, ca. 0.2 mm. wide at midpoint, below broad, + depressed toward apex, raised at base, the venation obscure on both surfaces, subdepressed below, 13 to 16 veins/5 mm., angle of divergence 55-60°. Infructescences from foliate axils along twig, with scars of 3 to 11 flowers, sometimes with 3-flowered branches ca. 4 mm. long, the axis 0.6-1.4 cm. long, sparsely puberulent, lowest internode 4-7 mm. long; bracts ovate, ca. 1.6 mm. long, deciduous; pedicels 3-6 mm. long, glabrous. Flower not known; outer pair of tepals persisting at base of immature fruit suborbicular, 1980] STEVENS, CALOPHYLLUM 669 ca. 2 by 2 mm., thick. Young fruit ovoid, ca. 8 mm. long, apiculate, drying brown; outer layer less than 0.3 mm. thick. Type: Philippine Islands, Luzon, Tayabas Province, Mt. Alzapan, 6000 feet {1829 m.], 7 June 1925, BS 45686 coll. Ramos & Edano (holotype, a; isotypes, B, K, UC, US). DistriBuTIon. Philippine Islands (Map 49); known only from the type locality. Ecotocy. Shrub in mossy forest, ca. 1830 m. alt. Calophyllum rigidulum is a distinctive, albeit poorly known, species. The leaf blades are small, very coriaceous, and rounded to shallowly retuse at the apex, and the venation is fairly dense. On the upper surface of the blade the midrib is rather narrow and indistinct, while on the lower surface it is broad but more or less depressed. The epithet rigidulum means ‘‘rather rigid’’—appropriate for this rigid-leaved species. 173. Calophyllum insularum P. F. Stevens, sp. nov. Ficure 40, d. Calophyllum sp. A, aff. C. caudatum Kanehira & Hatusima, P. F. Stevens, Austral. Jour. Bot. 22: 362. 1974. A speciebus aliis Calophylli in lamina chartacea 4.9-10 cm. longa et 1.1-2.5 cm. lata venatione densa, ramulo, gemma terminali, costa laminae, inflores- centia et tepalis exterioribus utrinque indumento tomentoso subpersistenti praeditis, et inflorescentiis axillaribus singulifloribus, differt. Tree to 33 meters tall, d.b.h. 40 cm.; trunk without buttresses; outer bark dark brown, with numerous shallow fissures; inner bark brown; latex yellow, clear. Twigs slightly flattened or not, 0.9-1.3 mm. across, not 4-angled to obscurely so, with obscure horizontal line at nodes, drying blackish, subpersistently tomentose; axillary innovations frequently with scars within 6 mm. of base; internodes 1-3.5(-7) cm. long; uppermost pair of axillary buds subacute, ca. 1.5 mm. long, erect; terminal bud narrowly conical, 5—8(-9.5) mm. long, shape obscured by dense, reddish brown, tomentose indumentum (hairs, Figure 46, a-d), underdeveloped internode to 1 mm. long. Petiole 0.7—-1.3 cm. long, deeply concave above, convex below, subpersistently tomentose; lamina narrowly ovate to elliptic, 4.9-10 by 1.1-2.5 cm., acuminate at apex, cuneate at base, undulate and slightly recurved at margin, coriaceous, drying olivaceous to umber above and sepia to gray-sepia below, + tomentose on midrib, margin, and lower surface when young, persistently so on midrib below (and margin), the midrib above abruptly narrowed at base, flat to slightly raised, 0.15-0.4 mm. wide at midpoint, sometimes sulcate, below raised, slightly angled or rounded, venation evident on both surfaces, raised, (13 to) 16 to 19 (to 22) veins/5 mm., angle of divergence 70-80°. Inflorescences from foliate axils near ends of twigs, with | flower, unbranched, the axis 3-5 mm. long, tomentose, lowest internode 3-5 mm. long; bracts not known; pedicels 5.5-8 mm. long, tomentose. Flower known only in late bud, (?)hermaphroditic; 670 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 tepals 8, the outer pair triangular, 7.5-8.5 by 4-5 mm., concave, tomentose outside and sericeous inside, the next pair elliptic to obovate, to 7.5 b 4 mm., thinner; stamens 80 to 100, the anthers oblong, 1.3-2.3 mm. long, subretuse to apiculate at apex; ovary |.5-2 mm. long, the style 1.5-3.5 mm. long, the stigma peltate, ca. 1.3 mm. across. Fruit not known. Tyre: Eil. Meos Noem [Irian Jaya], Seroei, 200 m., 5 Oct. 1939, bb 3094] (holotype, L; isotypes, A, NY, SING). DistriBuTION. Papuasia, known only from islands in Geelvink Bay (Map ADDITIONAL SPECIMENS SEEN. Papuasia. IRIAN Jaya. Geelvink Bay: Meos Noem, Seroei, 200 m., bb 30937 (a, L), bb 30944 (a, L, NY, SING), bb 30959 (a, L), bb 30965 a L), bb 30967 (L, sinc); Japen Is., Aisao, 200 m., BW 10569 (L, LAE), BW 10585 (L, LAE). Eco.Locy. Colline rainforest, ca. 200 m. alt. Late bud in October. Calophyllum insularum can be recognized by its tomentose indumentum, its narrowly ovate to elliptic leaf blades with dense venation, its single-flowered, axillary inflorescences, and its flowers. The flower buds are bilabiate, with the tips of the outer tepals separating early (perhaps due to the way that the specimens were dried). The outer pair of tepals has hairs on both surfaces. The epithet insularum (“‘of the islands’’) was chosen because this species has so far been collected only on islands. Calophyllum insularum can be distinguished from C. trachycaule, with which some of the specimens cited above had been confused, by its more delicate facies (thinner twigs, and narrower, thinner, more pointed leaves), its solitary flowers with eight tepals, the outer two of which are hairy on both surfaces (the inflorescences of C. trachycaule have five or more flowers, each with four subglabrous tepals), and its anthers, which are two to three times the size those of C. trachycaule. There are also anatomical differences between the two (Stevens, Joc. cit.). The indumentum on the petiole often consists of rather scattered long hairs mixed with short ones (Ficure 46, a-d). 174. Calophyllum poilanei Gagnep. ex P. F. Stevens, sp. nov. Ficure 45, a, b. C. poilanei Gagnep. in Humbert, Fl. Gen. Indo-Chine Suppl. 1: 269. 1943; Pha Ng guyen, Cay-Co6 Mién Nam Viét-Nam, 179. bang 62bis F. 1960; Pham, Cay-Co Mién Nam Viét-Nam. ed. 2. 2: 299. fig. 1970. Nomen invalidum. A speciebus aliis Calophylli in ramulis crassis nitidis 4-subalatis, nodis defoliatis prominentibus, et inflorescentiis terminalibus axillaribusque floribus pedicellis brevibus (usque ad 10 mm. longis) vel nullis, differt. Small tree 3-4 meters tall, d.b.h. to 7 cm.; bark not known. Twigs slightly flattened, 4-5 mm. across, strongly 4-angled to winged, drying 1980] STEVENS, CALOPHYLLUM 671 brown, shiny, Saar tomentose when young; axillary innovations with basal scars; internodes 0.7-3.5(-5) cm. long; upper pair of axillary buds rounded, ca. | mm. long, aaieadiee, not conspicuous; terminal bud conical, ca. long, with brown, tomentose indumentum (hairs, Ficure 46, j, k), erences internode absent. Petiole 2-5 mm. long, broadly concave above, convex below, glabrescent; lamina oblong to subobovate, 11.7-21 7-6 cm., acuminate at apex, broadly rounded to shallowly cordate at base, slightly undulate and strongly recurved at margin, marginal thickening mm. wide, coriaceous, drying cinnamon above and fulvous below, glabrous at emai the midrib above abruptly narrowed near base, becoming + raised, 0.2-0.5 mm. wide at midpoint, below strongly raised, angled toward apex, striate toward base, venation prominent on both surfaces, raised, 6 to 12 veins/5 mm., angle of divergence ca. 80°. Inflorescences terminal and from adjacent foliate axils, with numerous flowers, often flabellate, branched, branches to 2.5 cm. long, the axis to 10 cm. long, tomentose when young, 4-alate, lowest internode 1-2 cm. long; bracts ovate, ca. 7 Opiceuachicdiie tepals 10 (rarely 11), the outer pair ovate, ca. 8 by 5-6 mm., short-tomentose on back, the inner ones elliptic to oblong, 10-15 by 3.5-7 mm., outer two with strip of indumentum down back, next two with tuft of hairs near apex; stamens 130 to 180, the filaments to 9 mm. long, the anthers oblong, 1-1.3 mm. long, + rounded at apex; ovary 1.2-1.4 mm. long, the style 6-8 mm. long, the stigma subexcentrically peltate, ca. 1 mm. across, 3- or 4-lobed. Fruit not known. pe: Vietnam, Prov. Nhatrang, nord de Ninh-Hoa, massif de la Mere et l’Enfant, 1300 m., 18 May 1934, Poilane 6572 (holotype, P; isotype, P). DistripuTion. Central Vietnam (Map 49). ADDITIONAL SPECIMEN SEEN. Vietnam: Nhatrang, nord de Ninh-Hoa, massif de la Mere et l’Enfant, 1300 m., Poilane 6454 (P). Ecotocy. Stony ground, ca. 1300 m. alt. Flowering in May; flower scented. Calophyllum poilanei is one of the most distinctive species of Calophyllum, with its shiny, little-branched, more or less four-alate twigs that have very prominent leaf scars. The lamina is rounded to shallowly cordate at the base, the inflorescence is terminal, and the flowers usually have ten tepals. The flowers have short pedicels or are sessile; this gives the inflorescence a distinctive appearance. Although the twigs, leaves, and flowers are very robust, the plant itself is reported to be small. The epithet commemorates E. Poilane. Gagnepain (loc. cit.) seems to have been somewhat confused over the 4 mm. It is possible that Gagnepain’s ‘‘bracteoles’’ were bracts (these fall off very early—only a single one was seen) or the outer pair of tepals (in which case he may have confused their measurements with those of the next pair.of tepals). 672 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 61 The description given above is needed because Gagnepain’s description (foc. cit.) is in French and invalid. 175. Calophyllum membranaceum Gard. & Champ. Jour. Bot. Kew Misc. 1: 309. 1849; Choisy, Descr. Guttif. Inde, 43. 1849, Mém. Soc. Phys. Hist. Nat. Geneve 12: 423. 1851; Walp. Ann. Bot. Syst. 2: 191. 1851; Seem. Bot. Voy. H.M.S. Herald, 368. 1857; Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 290. 1862; Bentham, Fl. Hongkong. 25. 1861; Forbes & Hemsley, Jour. Linn. Soc. Bot. 23: 75. 1886; Vesque, Epharmosis 2: ¢. 4. 1889, in C. DC. Monogr. Phanerog. 8: 551. 1893; Gagnep. in Humbert, Fl. Gen. Indo-Chine Suppl. 1: 275. 1943; Auct. Ic. Corm. Sin. 2: 883. 1972. Type: Hongkong [Mt. Victoria], Champion s.n. (holotype, kK). C. bonii Pitard in Lecomte, Fl. Gen. Indo-Chine 1(4): 317. 1910; Gagnep. in Humbert, Fl. Gen. Indo-Chine Suppl. 1: 268. 1943. Tyee: Tonkin eee Ninh Thai, Muong Lang, 23 April 1889, Bon 4136 (lectotype, C. acne Pitard in Lecomte, Fl. Gén. Indo-Chine 1(4): 317. 1910. Tyre: Tonkin [Vietnam], Ninh Thai, Muong Lang, 17 March 1890, Bon 4301 (lectotype, p; isolectotype, P). C. spectabile auct., non Willd.; Hooker f. & Arnott, Bot. Beechey Voy. 174. 1883. Shrub or small tree 1-5 meters tall, d.b.h. to 10 cm.; details of bark unknown. Twigs slightly flattened, 1.3-2.5 mm. across, strongly 4-angled to subalate (with 2 additional raised lines decurrent from petiole), drying brown, glaucous when older, sparsely puberulent when young; axillary innovations lacking basal scars; internodes (0.5—)1.5-7 cm. long; uppermost pair of axillary buds pointed, to 1.5 mm. long, erect; terminal bud narrowly conical, 3.5-7 mm. long, with dense, brown, subadpressed indumentum (hairs, Ficure 46, h, i), underdeveloped internode 0.1 mm. long. Petiole 0.4—1.4 cm. long, very deeply concave above and convex below, sparsely puberulent when young; lamina oblong to elliptic, 4.2-13.1 by 1.4-4.6 cm., acuminate at apex, cuneate at base, slightly undulate and recurved at margin, coriaceous, drying umber to olivaceous above and below, glabrous or sparsely and subpersistently puberulent on midrib below, the midrib above gradually narrowing from base, sharply raised, 0.1-0.35 mm. wide at midpoint, below raised, angled to striate, the venation on both surfaces apparent, raised, (5 or) 6 to 11 (or 12) veins/5 mm., angle of divergence 70-80°. Inflorescences axillary or terminating short axillary shoots, with (1 to) 3 to 9 flowers, sometimes flabellate, unbranched, the axis (0.1-)0.35—2.7(-5) cm. long, sparsely to subdensely puberulent, lowest internode (0.1-)0.3-1.5(-3.4) cm. long; bracts elliptic, 1-8 mm. long, + deciduous, or foliaceous, to 5.2 by 2.3 cm., subpersistent; padieels 3-9 mm. long, sparsely puberulent. Flower (?)hermaphroditic; tepals 8 (to 12), t outer pair suborbicular, 3-4.5 by 3-4 mm., sometimes puberulent on a near base, the next pair broadly elliptic, 5.5-8 by 3.5-4.5 mm., the inner ones elliptic to obovate, 5.5-10 by 2-4.5 mm.; stamens 125 to 250, the paca 1.5—-4 mm. long, connate for up to 1.5 mm., the anthers suboblong to elliptic, 1980] STEVENS, CALOPHYLLUM 673 0.4-0.9 mm. long, + retuse at apex; ovary 1.2-1.8 mm. long, style 4.5-5.5 mm. long, stigma obscure. Fruit ellipsoid (rarely ovoid), 1.4-1.8 by 0.9-1.15 cm., minutely apiculate or not, drying pale brown, wrinkled; outer layer not detaching cleanly from stone, 0.4-1 mm. thick, compact; stone ellipsoid, 1.3-1.7 by 0.8-1 cm., rounded at apex, the walls ca. 0.1 mm. thick, smooth, (?)unmarked; spongy layer thin. Distrisution. Southern China (Kwangtung, Kwangsi; reported also from Yunnan (Auct., Ic. Corm. Sin., Joc. cit.)), Hong Kong, and northern Vietnam (Map 50) SELECTED SPECIMENS SEEN. China: Kwanctunc: Pok Lor, Fung Hom A-548 (Ny); Yam na Shan (Yit Nga Shan), Mei isan Distr.. W. T. Tsang 21525 (A, K, Ny, P); Ting Wu Shan, W. Y. Chun 6406 (a, £, K, US, w); San-on Distr., Ng-Tung Shan, 940 m., 7. M. Tsui ae O (F, K, Mo, Nsw); Lofoushan, Ying 1714 (SING). KWANGsI: Sehfengdarshan, S. Nanning, 762 m., R. C. Ching 8086 (A, NY, US); Shap Man Taai Shan, near Ping Hoh village, SE. of Shang-sze, Kwangtung Border, W. T. Tsang 22047 (a, 3m, Pp); Hainan, Mochung Mt., Tingkap, Tingan, S. P. Ko 52238 (ny), Yaichow, H. Y. Liang 61994 (kg, M, NY), Five Finger Mt., 500 m., CCC 8495 (a, BM, C, E, G, K, MO, P), Dung Ka to Wen Fa Shi, N. K. Chun & C. L. Tso 43684 (ny), Ka Chik Shan & vicinity, Ch’ang Kiang Distr., S. K. Lau 1633 (a, BM, Ny, P), Ng Chi Leng, Fan Yah, 914 m., N. K. Chun & C. L. Tso 44133 (a, E, F, NY), Kan-en Distr., Chim Fung Ling, near San Mo Watt village, S. K. Lau 3856 (a, P), Ngai Distr., Yeung Ling Shan, S. K. Lau 86 (a, B, BM, E, G, K, MO, NY, P, US, W), Po Ting, 365 m., F. C. How 71875 (a, pm). Hong Kong: Aberdeen, Y. S. Lau 47 (a); Kowloon, Wong Keng Tsai, Shatin, Y. W. Taam 2298 (G, NY, Us); Wong Kum Cha, Y. K. Chan 3043 (Ny); Pic Victoria, Bodinier 1450 (£, p); Ma On Shan, S. Y. Hu 9329 (a); ravine leading to Pok-foo-lum, Lamont 43 (bm); Kadoorie Farm, Kuan Yin Shan, S. Y. Hu 12014 (a); Tai Mo Shan, S. Y. Hu 11274 (a). Vietnam: Tam Dao, 850 m., Petelot 4241 (A, BO, NY, P, US); Nin Thai, Muon Long, Bon 3955 (e); Quombi, Balansa 1504 (c, Pp); station experimental de Phu Tho, Chevalier 41053 (rp); Ha Coi, Tuai Wong Mo Shan, near Chuk Phai, W. T. Tsang 29024 (a, Bo, Cc, E, G, K, P, SING). Map 50. Distribution of Calophyllum membranaceum in Asia (circles, loca- lized; squares, unlocalized). 674 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 EcoLocy. Locally common, lowland or (usually) colline forest, sometimes in rocky places; 50-915 m. alt. Flowering April and May, rarely in March; (flower scented); fruiting October to December, rarely September or January (fruit yellow to orange—those reported as black seem to be immature). LocaL uses. Used in Yunnan to reduce the inflammation around bruises and to kill pain (Auct., Ic. Corm. Sin., Joc. cit.) Calophyllum membranaceum can be easily recognized, even when sterile, by its strongly angled twigs, its suboblong, thinly coriaceous lamina that often dries greenish and has prominent veins leaving the midrib at 70-80°, and its deeply concave petiole. The inflorescence is somewhat variable. There are sometimes fully developed leaves in the lower part of the inflorescence axis so that the inflorescence appears to be terminal. The asymmetrical, nonpeltate stigma of the flower and the thin outer and woody layers of the fruit are also distinctive. The epithet membranaceum (‘‘membranaceous’’) was presumably suggested because of the texture and thinness of the dry leaf blades. There is relatively little variation in flower and fruit, but specimens of Calophyllum membranaceum from Hong Kong frequently have more than eight tepals. In all other respects they agree with specimens from other parts of the range of the species. Calophyllum bonii was based on specimens with very small inflores- cences—this was the key character used by Gagnepain (loc. cit.) to separate it from the other species of Calophyllum growing in Indochina. However, as mentioned above, inflorescences in C. membranaceum are variable in appearance, and C. bonii has been reduced to synonymy under C. membrana- ceum without hesitation. Bon 4136 is designated the lectotype of C. bonii; the original description was also based on Balansa 1504. Calophyllum tonkinense was based on three numbers collected by the Abbé Bon (Bon 3159, 3282, and 430/) that are indistinguishable from other specimens assigned to C. membranaceum. Calophyllum tonkinense has been lectotypified on Bon 4301, a specimen with flowers. 176. Calophyllum echinatum P. F. Stevens, sp. nov. Ficure 45, c. A speciebus aliis Calophylli in gemma terminali longa, pari superiore gemmarum axillarium conspicuo patentique, lamina plerumque saltem 9 cm. onga margine arcte undulata costa supra lamina elevata circumcincta venulis lateralibus 15 usque ad 22/5 mm., et fructu strato exteriore fibroso fibris putamine conflatentibus, differt. Tree to 26 meters tall, d.b.h. to 75 cm.; details of bark not known. Twigs slightly to strongly flattened, 2.5-3 mm. across, 2-, 4-, or 6-angled, soon becoming rounded, drying yellowish, farinose-puberulent when young; axillary innovations lacking basal scars; internodes (1—)2-6.5(-10) cm. long; uppermost pair of axillary buds + acute, |.5—7 mm. long, spreading to suberect; terminal bud narrowly conical, (0.7-)0.9-1.7 cm. long, with brown, subcrusta- ceous indumentum (hairs, Ficure 41, t, u), underdeveloped internode 3.5-9 1980] STEVENS, CALOPHYLLUM 675 mm. long. Petiole |.7-2.4 cm. long, strongly concave above, convex below, drying blackish, glabrous; lamina ovate or elliptic to suboblong, (5.2-)9-13.5 by (2.2-)3.2-5.2 cm., acute to short-acuminate at apex, acute to cuneate at base, closely and strongly undulate and slightly recurved at margin, coriaceous, gray- to green-olivaceous above and sepia-olivaceous-sabelline below, subpersistently puberulent on midrib below, the midrib above abruptly narrowed near base, raised, surrounding lamina also raised, 0.3-0.55 mm wide at midpoint, below strongly raised, angled, the venation subobscure on both surfaces (subapparent above), slightly raised, 15 to 22 veins/5 mm., eae of divergence 70-80°. Inflorescences from uppermost foliate axils, with 11 flowers, the branches 3-flowered, to 1.3 cm. long, the axis 4-6 cm. ae puberulent, lowest internode 1-3.5 cm. long; bracts not known; pedicels ca. 2.8 cm. long, puberulent. Insect-damaged flowers only known; tepals 4, the outer pair broadly ovate, ca. 6 by 5 mm., puberulent on back, the inner pair obovate, ca. 9.5 by 6 mm.; stamens and ovary not known. Fruit ellipsoid, ca. 5.3 by 3 cm., obtuse at apex, drying vinaceous-brown, deeply and distantly wrinkled; outer layer not clearly distinguishable from stone, fibrous, large air spaces developing; stone ellipsoid, ca. 2.7 by 1.7 cm., rounded at apex, the walls to 0.8 mm. thick, in places only 0.1 mm. thick, not marked, rough, with branched, anastomosing fibers arising from stone; spongy layer (?)thin Type: Celebes, Malili, Oesoe, Cel. /II-407 coll. Waturandang (holotype, DisTRIBUTION. Known only from the Celebes (Map 49). ADDITIONAL SPECIMENS SEEN. Celebes: Malili, Oesoe, 50 m., bb 32596 (Bo, L), 200 m., Cel. /JJ-215 (a, Bo, L); Thawatta, 200 m., Cel. /II-407 coll. Gonggrijp (A, BO, L); Lingkomomo, Timampoe, 300 m., bb 8566 (Bo). EcoLocy. Forest, 20-300 m. alt. The fruits of Cel. /I/-407 coll. Gonggrijp appear to be galled; i are flattened-ellipsoid structures ca. 6 by 2.3 cm. that are open on one si Calophyllum echinatum is a distinctive species that is easily recognizable, er 9m long. The fibers arising from e stone give it a spiny appearance, ene: the epithet echinatum (‘‘prickly’ Although some specimens of Calophyithin soulattri from the Celebes (e.g., Cel. / ITI-63) have a midrib similar to that of C. echinatum, they do not otherwise approach that species. In C. soulattri the uppermost pair of axillary buds 676 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 is usually erect (rather than spreading), the lamina is thinner, the lowest internode of the inflorescence is shorter, the pedicels and outer tepals are nearly always glabrous, and the tepals are smaller. The fruit of C. soulattri is quite different from that of C. echinatum: it is ee one-quarter the length and has a compact, slightly fibrous outer layer. 177. Calophyllum exiticostatum P. F. Stevens, sp. nov. Ficure 45, d-f. A speciebus aliis Calophylli in lamina parva 2-5(-7.5) cm. longa basi plerumque rotundata venulis lateralibus densis et costa infra apicem evanes- centi, petiolo brevi 1.5—4 mm. longo, et fructus circa 7 mm. longo strato exteriore forte proviso, differt. Large tree 30-45 meters tall, d.b.h. to 140 cm.; trunk without buttresses; outer bark yellowish, with shallow, diamond-shaped fissures, the inner surface orange; under bark dark red; inner bark pale red; latex at first yellow, clear, becoming opaque Twigs slightly flattened: 1-1.5 mm. across, strongly 4-angled (with obscure horizontal lines at nodes), drying blackish brown, puberulo-tomentose when young; axillary innovations lacking basal scars; internodes 0.5-3 cm. long; uppermost pair of axillary buds rounded, ca. 0.8 mm. long, erect; terminal bud plump to conical, 2.3-3 mm. long, with brown, subadpressed indumentum (hairs, Figure 46, e-g), underdeveloped internode absent. Petiole 1.5-4 mm long, broadly concave above and convex below, subpersistently puberulent; lamina ovate to elliptic, 2—S(-7.5) by 0.9-2.6(-3.1) cm., rounded (rarely subretuse or subacute) at apex, broadly to narrowly rounded or cuneate at base, not undulate or slightly so and not recurved at margin, coriaceous, drying umber to fulvous above and cinnamon to sabelline below, subpersis- tently puberulent on midrib below, the midrib above gradually narrowed from or abruptly narrowed at base, flat to slightly raised, subsulcate at base, 0.05-0.2 mm. wide at midpoint, below + level or slightly raised, striate, generally disappearing 2-3 mm. below apex, the venation obscure to subapparent on both surfaces, slightly raised, 14 to 23 veins/5 mm., angle of divergence 40-60(-70)°. Inflorescences axillary, with 5 to 7 flowers, unbranched, the axis 4-10 mm. long, densely pubescent, especially near base, lowest internode ca. 2 mm. long; bracts not known; pedicels 3.5-6.5 mm. long, pubescent. Flower (?)hermaphroditic; tepals 4, the outer pair ovate, 4-5 by ca. 2 mm., puberulent on back, the inner pair + elliptic, 3.5-4.5 by 1.5-3 mm., with strip of indumentum down back; stamens 40 to 50, the filaments to 2.5 mm long, the anthers oblong, 1-1.2 mm. long, rounded to shallowly retuse at apex; ovary ca. 1.5 mm. long, the style ca. 2.2 mm. long, the stigma peltate, ca. 0.6 mm. across. Fruit spherical, ca. 7 by 7 mm., rounded or minutely apiculate at apex, drying vinaceous-brown, smooth; outer layer not detaching cleanly from stone, ca. 0.8 mm. thick, with large air spaces developing beneath skin; stone spherical, ca. 5.5 by 5.5 mm., rounded at apex, the walls ca. 0.15 mm. thick, smooth, unmarked; spongy layer thin. ype: Sumatra, Palembang, Lematang Ilir, Semangoes Reserve, 75 m., 8 May 1940, bb 31756 coll. Buwalda (holotype, a; isotypes, Bo, K, L, NY, SING). 1980] STEVENS, CALOPHYLLUM 677 DisTRIBUTION. Southern Sumatra and northern Borneo (Map 49). ADDITIONAL SPECIMENS SEEN. Sumatra. SELATAN: Lematang Oeloe, 110 m., Grashoff 161 (po), 150 m., Grashoff 224 (Bo, L); Pasemahlanden, Djangkar, 900 m., bb 8118 (Bo); Afd. Redjang, bij desa Loeboek binjai, Rimba Ketjau, bb 2299 (po, L). Barat: [Barisan], Manindjau, Silajang, 500 m., bb 3971 (Bo). Borneo. Sasan. Ranau: Poring turn-off, Kota Kinabalu-Sandakan road, 640 m., Stevens et al. 559 (a), 563 (a). Ecotocy. Colline rainforest, 75-900 m. alt. Flowering in March; fruiting in GERMINATION AND YOUNG PLANT. The young plants grow erect, and the terminal bud is functional. The leaf blades are up to 11.5 by 3 cm Calophyllum exiticostatum is an easily distinguishable species. It has strongly four-angled twigs, a short terminal bud, and small, very densely veined leaves that are usually rounded at the base. The inflorescence axis is more or less puberulent, the flowers have four tepals, and the spherical, smooth fruit has large air spaces in the outer layer. The midrib, which disappears just below the apex of the blade, suggested the epithet exiticostatum (‘‘going-out— midrib The closest relative of Calophyllum exiticostatum may be C. calaba, although the latter species has somewhat larger leaf blades with usually less steeply ascending venation and a midrib that continues to the apex of the blade. In addition, the fruit of C. calaba lacks the strong epicarp of C. exiticostatum. Calophyllum brachyphyllum, from the Philippine Islands, also has small leaf blades with dense venation, but this species has a much more coriaceous blade that is retuse at the apex, and larger fruits with both the outer layer and the stone thicker. Despite the 1500 km. distance separating the localities of Calophyllum exiticostatum in Sabah (near Mt. Kinabalu) and Sumatra, the Bornean specimens are vegetatively similar to the Sumatran ones. The only differences seem to be that in the Sumatran specimens the midribis a little more prominent, the lamina is a little larger (measurements in parentheses in the description above), and the horizontal lines at the nodes are more obvious (cf. also C. gracilipes). However, material in flower and fruit is needed to confirm this similarity; the description of the bark is taken from Bornean material. 178. Calophyllum sp. Tree ca. 8 meters tall, d.b.h. to 6 cm.; trunk and bark unknown. Twigs flattened, 3-3.5 mm. across, 4-angled, drying dark brown, tomentose when young; axillary innovations lacking basal scars; internodes 2.5—4 cm long; uppermost pair of axillary buds subrounded, ca. 2 mm. long, erect; terminal bud bluntly conical, ca. 4 mm. long, tomentose, underdeveloped internode absent. Petiole 2.8-3.3 cm. long, rather narrowly concave above, convex below, glabrous when mature; lamina ovate-oblong, 16.5-18 by 6-6.5 cm., short-acuminate at apex, rounded to acute at base, distantly undulate and slightly recurved at margin, coriaceous, drying umber above and fulvous- 678 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 umber below, with sparse hairs persisting on midrib below, the midrib above gradually narrowing from base, raised, 0.3-0.4 mm. wide at midpoint, below raised, substriate to subangled, the venation above apparent, below subob- scure, raised, latex canals impressed below, 5 or 6 veins/5 mm., angle of divergence 70-80°. Inflorescences from foliate axils, with ca. 5 flowers, unbranched, the axis 3-3.5 cm. long, puberulo-tomentose, lowest internode 0.6-1.1 cm. long; bracts unknown; pedicels 0.5-1.4 cm. long, puberulo-tomen- tose. Flower (?)hermaphroditic; tepals ca. 12, the outer pair subovate, ca. 1.5 by | cm., puberulent at base on back, the next pair suborbicular, ca. 1.3 by 1.3 cm., the inner ones oblong-obovate, |.1-1.6 by 0.4-1 cm.; stamens ca. 410, the filaments to 8 mm. long, the anthers oblong, 1.2-1.6 mm. long, retuse at apex; ovary ca. 3 mm. long, the style ca. (?)6 mm. long, the stigma peltate, ca. 1.6 mm. across, 2-radiate. Fruit unknown. DistriBuTION. Philippine Islands (Map 49). SPECIMEN SEEN. Philippine Islands. Luzon. Tayabas: Malbog, 100 m., FB 30772 (Ny). Ecotocy. Forest, ca. 100 m. alt. Flowering in February; flower scented. Calophyllum sp. 178 can be recognized by its short, bluntly conical terminal bud, its long petioles, its relatively large, ovate-oblong leaf blades with the venation clearly raised on the upper surface but obscure and barely raised (with latex canals impressed) on the lower, and its few-flowered axillary inflorescences. The flowers are large and have about twelve tepals. It is premature to describe this taxon in the absence of fruits, but its conical, rather short terminal bud and its distant venation suggest that it is not related to the variable Calophyllum blancoi. 179, Calophyllum macrophyllum Scheffer, Natuurk. Tijdschr. Nederl.-Indié, VII. 2: 405. 1873; Vesque in C. DC. Monogr. Phanerog. 8: 609. 1893: Lauterb. Bot. Jahrb. 58: 14. 1922; T. C. Whitmore, Gard. Bull. Singapore 22: 12. 1967; P. F. Stevens, Austral. Jour. Bot. 22: 377. 1974. Type: Gebeh [Irian Jaya], Teysmann, HB 7574 (holotype, Bo; isotypes, c, K, L, MEL). (?)Tree; details of trunk and bark unknown. Twigs slightly flattened, 5-7 mm. across, strongly 4-angled to -alate, drying brown, glabrous at maturity; terminal bud + conical, ca. 5 mm. long, with brownish indumentum, hairs adaxially curved and very papillate; other details of twigs and buds unknown. Petiole 1.3-1.7 cm. long, shallowly concave above, angled below, glabrous at maturity; lamina elliptic to suboblong, 32-45 by cm., acute at apex, cuneate at base, barely undulate and slightly recurved at margin, coriaceous, drying olivaceous above and fulvous-umber below, glabrous at maturity, the midrib above rather quickly narrowed at base, raised, 0.3-0.4 mm. wide at midpoint, below strongly raised, angled, the venation obscure above, apparent below, raised, with 3 to 5 veins/5 mm., angle of divergence ca. 80°. Inflorescence, flower, and fruit not known. 1980] STEVENS, CALOPHYLLUM 679 DistTRIBUTION. Gebeh Island (Map 45); known only from the type collection. The very large leaf blades with distant venation and the relatively small terminal bud allow Calophyllum macrophyllum to be recognized readily, but it is still known only from the rather fragmentary sterile collection on whic the original description was based. The epithet macrophyllum (“‘large leaf’’) was presumably suggested because of the size of the leaves. Although it seems unlikely that Teysmann, HB 7574 was taken from a sapling (Stevens, Joc. cit.), the leaves would be large even for a young plant of Calophyllum. Fertile collections of C. macrophyllum would be of consider- able interest; it is possible that it is related to C. celebicum. Nomina Non Satis CoGNITa Calophyllum cerasiforme Teijsm. & Binn. ex Koord.-Schum. Syst. Verzeich. 10. Nomen I cannot identify the specimen on which this name is based (Koorders 10623, from Sumatra); see also Calophyllum sp. 9. Calophyllum kunstleri King var. longifolium Boerl. Catal. Pl. Phanerog. Horto Bogor 2: 80. 1901; Koord.-Schum. Syst. Verzeich. 2: 58. 1910. Nomen. I have been able to identify neither the specimens cited in the references above, nor that listed under Calophyllum kunstleri in Koorders-Schumacher (loc. cit.). The ‘‘Calophyllum kunstleri’’ currently grown at Bogor (plant VI C 237) is C. calaba var. bracteatum. my eae lonchophyllum O. Schwarz, Repert. Sp. Nov. 24: 89, 1927. Type: stralia, Northern Territory, Finniso River district, Bleeser A22. Tree. Branches terete, branchlets plicate-sulcate; buds with fuscous, adpressed, pubescent indumentum, the hairs contorted, unbranched. Petiole 5-9 mm. long; lamina long-lanceolate, 12-19 by 1.4-3.5 cm., long-acuminate at apex, narrowly acute (?and/or cuneate) at base, coriaceous, glabrous, the midrib above immersed-prominulous, below carinate, yellowish, the lateral nerves spreading horizontally, very close. The description above is based on the protolog. Bleeser A22, from which Schwarz took his description, was apparently a sterile specimen and has since been destroyed, having been held in Berlin. It is impossible to tell to what species the name Calophyllum lonchophyllum should be referred. Calophyllum minus Ridley, Dispersal Pl. World, 207. 1930. Nomen. As noted by Ridley (/oc. cit.), Guppy found that fruits of this species, which grew inland in the Solomon Islands, float. The name is not mentioned in Guppy, Solomon Is. 1867. 680 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Calophyllum vanoverberghii Merr. Philip. Jour. Sci. C. 9: 454. 1915. Type: Philippine Islands, Luzon, subprovince of Bontoc, Bauco, ca. 1700 m., 12 Sept. 1918, Vanoverbergh 1466 (pNu, destroyed). Small tree ca. 3 meters tall. Twigs drying grayish or olivaceous, sometimes slightly pubescent at tips; terminal bud ferrugineous-pubescent. Petiole ca. | cm. long, slightly pubescent at first, otherwise glabrous; lamina oblong, 5-8 by 2-3 cm., obtuse at apex, acute or slightly decurrent at base, thickly coriaceous, drying concolorous, nitid, glabrous when mature, nerves very close. Inflorescences axillary and terminating short lateral branchlets, branched, with 3 short branchlets, often also with additional flowers, the axis 1.5—2.5 cm. long, the lowest internode very short, usually somewhat ferrugineous-pubescent; pedicels 5—10 mm. long. Tepals 4, the outer pair broadly ovate, ca. 6 mm. long, sometimes glabrous, concave, the inner pair narrowly obovate, ca. 8 mm. long; stamens indefinite, the filaments to 3.5 mm. long, the anthers narrowly obovoid or oblong-obovoid, 1.5-2 mm. long; ovary glabrous, style ca. 2 mm. long. Fruit unknown. Although Merrill’s description of the inflorescence of Calophyllum vanover- berghii (translated above) reads somewhat like that of C. soulattri, a species which also has four-tepaled flowers, C. soulattri never has an oblong, very coriaceous lamina that dries concolorous and nitid. BS 75484 (from Catan- duanes, Luzon) may belong to C. vanoverberghii; the outer layer of the fruit of this specimen is not compact, unlike that of C. soulattri, although in inflorescence the two are similar. Calophyllum sp. Merr. Jour. Arnold Arb. 19: 355. 1938. Merrill (/oc. cit.) noted that Rheedia umbellata Houtt. (Nat. Hist. II. 3: 3. 1774) was a new name for R. javanica Burman f. (FI. Indica, 118. 1768) and that it might be a species of Calophyllum. I have not been able to identify R. javanica more precisely Nomina ExcLUDENDA Calophyllum acuminatum Lam. Encycl. Méth. Bot. 1: 553. 1785. Type: sine loco, anon. (lectotype, P). The specimen that Lamarck had on hand consists of a fruiting element, referable to Calophyllum tacamahaca, and a leafy element, referable to a species of Garcinia. The name C. acuminatum was lectotypified on this latter element (see P. F. Stevens, Jour. Ace Arb. 57: 181. 1976). Calophyllum akara Burman f. Fl. Indica, 121. 1768. Type: based on Rheede, Hortus Malabar. 5: 15. t. 8. 1685 (Akara patsjoti). According to Merrill (Philip. Jour. Sci. 19: 366. 1921), this is properly called Tetracera akara (Burman f.) Merr. (Dilleniaceae). Lhe angustifolium Roxb. Fl. Indica. ed. 2 (W. Carey, ed.). 2: 608. , Hortus Bengal. 41. 1814, nomen; Choisy, Descr. Guttif. Inde, 1980] STEVENS, CALOPHYLLUM 681 43. 1849, Mém. Soc. Phys. Hist. Nat. Genéve 12: 423. 1851; Planchon & Triana, Ann. Sci. Nat. Bot. IV. 15: 293. 1862; T. Anderson in Hooker f. Fl. Brit. India 1: 276. 1874; Vesque in C. DC. Monogr. Phanerog. 8: 609. 1893. Type: Malaya, Penang, Roxburgh s.n. (lectotype, BM, quoad Garcinia). The protolog to this species reads ‘‘Twigs cylindric. Leaves short- -petioled, lanceolate, with lengthened, somewhat obtuse points, lucid, ely veine Flowers in axillary fascicles; pedicels with a cyatheiform apex... .”’ The are two elements on a sheet presumably of the type at the British Museum: a vegetative shoot of a species of Calophyllum with ‘‘Calophyllum angustifo- lium’? written on one of the leaves, and a shoot of a species of Garcinia with pistillate flowers just past anthesis. The only part of the original description clearly referable to Calophyllum is the mention of the leaves being lucid and finely veined, although Roxburgh also says that ‘“‘peon’’ (poon) spars for the masts of ships come from this species—probably also referring to this element. The mention of the leaves having somewhat obtuse points best fits the shoot of Garcinia, as does the description of the inflorescence and the pedicel. Since most of the protolog of Calophyllum angustifolium seems to be based on the shoot of Garcinia, it is reasonable to lectotypify the name on this element. I do not know to which species of Garcinia this shoot belongs; the epithet angustifolia is already occupied in Garcinia (G. angustifolia A. C. Smith, from New Guinea). The shoot of Calophyllum is not complete enough for identification; the terminal bud is not properly developed. Calophyllum augia Steudel, Nomencl. ed. 2. 1: 260. 1840, nomen illegitimum for Augia sinensis Lour. Fl. Cochinch. 1: 337. 1790, ibid. ed. Willd. 1: 411. 1793. Merrill (Trans. Am. Philos. Soc. n.s. 24(2): 244. 1935) equated this name with Rhus succedana L. (Anacardiaceae). The only element in the original description that might refer to Calophyllum is the mention of the flower having 100 stamens. ‘“Calophyllum chinense Walp.? ex Seemann in Bonplandia’’; T. Anderson in Hooker f. Fl. Brit. India 1: 270. 1874. Nomen. Anderson (loc. cit.) nae this name with Ochrocarpus longifolius ees Bentham & Hooker f. (=~ Mammea longifolia (Wight) Planchon & T I have been unable to find any other reference to Calophyllum Pale included in Kosterman’s (1956, 1961) revisions of the Asiatic and Pacific species of Mammea, but from Seemann’s discussion it seems to be a synonym of Calysaccion longifolium Wight (= M. longifolia). Calophyllum dubium Steudel, Nomencl. ed. 2. 1: 261. 1840. Based on C. longifolium Wall. Catal. 4851. 1831. Nomen. 682 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 See Calophyllum longifolium, below. Calophyllum excelsum Zoll. & Moritzi, Nat. Geneesk. Arch. Neerl. Indié 2: 582. 1845 As Merrill (Jour. Arnold Arb. 26: 94. 1945) noted, this name is to be referred to Ochrocarpus odoratus (Raf.) Merr., properly called Mammea odorata (Raf.) Kosterm. Calophyllum hexapetalum Hooker f. Trans. Linn. Soc. 23: 163. 1860. Type: [In ora septentrionale insulae] Borneo, Low s.n. (holotype, k). This is certainly a species of Mesua, but was not included by Kostermans (1969) in his list of new combinations in the genus Calophyllum longifolium Wall. Catal. 4851. 1831. Nomen. Wight took up Wallich’s epithet when he described Calysaccion longifolium Wight. This is correctly to be called Mammea longifolia. oe nagassarium Burman f. Fl. Indica, 121. 1768. This name has usually been referred to Mesua ferrea L. (Steudel, Nomencl. ed. 2. i: 260. 1840); however, type material of M. ferrea apparently belongs to the species hitherto called M. thwaitesii (Kostermans, 1976). The species of Mesua commonly cultivated must be called Mesua nagassarium (Burman f.) Kosterm. Kostermans (loc. cit.) typified C. nagassarium on Rumphius, Herb. Amboin. 7: p/. 2. 1750. However, Burman may well have received material of C. nagassarium, since he stated ‘‘habitat in Amboina & Java, unde anno 1759. missa.’’ There are two sheets of C. nagassarium originally in Burman’s herbarium that are now in Geneva. Calophyllum neurophyllum Schlechter, Bot. Jahrb. 39: 193. 1906. Type: New Caledonia, Hugeln bei Noumea, 20 m., Oct. 1902, Schlechter 15091 (isotypes, G, P). This is correctly called Mammea neurophylla (Schlechter) Kosterm. Calophyllum pustulatum Ridley, Kew Bull. 1938: 118. 1938; Masamune, Enum. Phanerog. Born. 476. 1942. Type: Sarawak, near Kuching, 23 Jan. 1893, Haviland 2117 (holotype, K; isotype, L). name is to be referred to Mesua, but it is not included in that genus by Kostermans (1969). Calophyllum touriga C. T. White & Francis, Proc. Roy. Soc. Queensland 35: 63. 1923. Type: Australia, Queensland, Gourka pocket, Atherton Tableland, Jan. 1923, Merrotsy s.n. ean Se BRI, n.v.; see L Smith, Proc. Roy. Soc. Queensland 68: 48. 1957). This species is correctly called Mammea touriga (C. T. White & Francis) L. S. Smith. 1980] STEVENS, CALOPHYLLUM 683 Calophyllum cae F. Villar in Ceron, Catal. Pl. Herb. Manila, 229. plate. 1892. E: Philippine Islands, Luzon, Camarines Sur Province, Mt. 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Dassanayake, ed., A revised handbook to the flora of Ceylon. Vol. 1. Amerind, New Delhi). In this account, endosperm is said to be sometimes present as a thin, jellylike layer around the cotyledons; Thwaites (Jour. Bot. Kew Misc. 6: 72. 1854) noted that CP 2446 (C. trapezifolium) had a green embryo surrounded by abundant endosperm. These observations must be confirmed. Kostermans gives interesting information on the color and appearance of the young leaves. The species mentioned in his account are listed below; brief comments have been added so that Kostermans’s account and the present one can be integrated. 1. Calophyllum inophyllum. 2. Calophyllum tomentosum (see also Thwaites, Enum. Pl. Zeyl. 51. 1858). Kostermans 24848 is considered to be a close match with material previously identified as C. elatum; reports of that species from Sri Lanka are included without hesitation in the synonymy under C. tomentosum. 3. Calophyllum moonii. The flowers are reported to have only four tepals. 4. Calophyllum calaba. Material assigned to C. calaba var. worthingtonii above is believed to represent the young stage of ‘‘typical’’ C. calaba 5. Calophyllum bracteatum. The seedlings are noted as having very narrow, linear leaves; saplings 2 meters tall already show the leaf variation characteristic of this species. 6. Calophyllum trapezifolium. Galls are said not to occur on this species. 7. Calophyllum zeylanicum. This is included in C. trapezifolium in the present account. 8. Calophyllum cuneifolium. 9. Calophyllum walkeri. 10. Calophyllum thwaitesii. The conflicts with earlier discriptions of this species and its variation that Kostermans mentions are resolved by the description of C. vergens. 11. Calophyllum cordato-oblongum. 692 JOURNAL OF THE ARNOLD ARBORETUM [vot 61 INDEX+ Acacia, 540, 542 —*articulatum, 134, 157, 197, 542, Agathis, 529, 559, 594 550-552, 554 Anacardiaceae, 474 —augia, 681 Apoterium, 167 —*aurantiacum, 160, 165, 189, 633, —sulatri, 167, 277, 290 643, 656, 658, 659-661 Augia, —aureobrunnescens, 129, 179, 625, —sinensis, 681 636, 637, 638, 652 —aureum, 136, 179, 630, 637, 638, Balsamaria, 167 652 —inophyllum, 167, 325, 335 —auriculatum, 481, 486 Calaba, 167 ee 269, 270, 272, 275, Calophyllodendron, 149 —austrocoriaceum, 348, 358, 360, 364 —cuddalorense, 149 —*austroindicum, 148, 154, 157, 175, Bea ear a 204, 241-244, 250, 251, 255 —indicum, 149 —balansae, 164, 176, 209, 211, 215, Calophyllum, 119-140, 141, 142-166, 167, 168, 169-691 = 225-227, 228, 229, 233 sect. Apetalum, 120 , 506 eee —*b 192, 525, 528, 532, 53 —sect. Hypodermata, 120, 145 “banyengit 5 53 3, : —benjamina, 644, 646 —sect. Microphyllum, 120 —*bicolor, 165, 202, 535, 536-541, —acuminatum, 275, 279, 680 542 —acutiputamen, 202, 617, 618, 652 —biflorum, 123, 128, 132, 144, 158 —*aerarium, 201, 426, 432, 443-445 164-166, 181, 187, 190, 193, 442, —akara, 680 454, 463, 464, 471, 472-476 —*alboramulum, 123, 125, 131, 153, —*bifurcatum, 133, 137, 165, 196 158, 179, 191, 204-206, 209, 211, 199, 535, 536, 538, 540,545. 430, 445 —bintagor, 325, 334 —amblyphyllum, 536, 558, 561, 562, __blancoi, 123, 137-139, 142, 158, 185, —amoenum, 220, 225, 261, 262, 270, 195, 196, 206- 218, 219, 225, 529, 273, 274, 466, 507, 518 631, 651, 678 ——var. obtusifolium, 262, 274 —blumei, 325, 334 —amplexicaule, 496, —blumutense, 512, 513 —*andersonil, 188, 396, 426, 428, 429, —bonii, 672, 674 430, 478, 497 —borneense, 207, 371, 372, 375, 436 —angustifolium, 221, 680 —brachyphyllum, 194, 502, 503, 633, —antillanum, 2 3, 667, 668, 677 —apetalum, 119, 142, 149, 175, 244, __bracteatum, 131, 174, 221, 258, 253-256, 258, 273, 279 275-277, 479, 691 —*archipelagi, 156, 182, 536, 552, __brasiliense, 130, 143, 149, 153, 270, 574, 580, 581, 583, 584 7 —*ardens, 126, 160, 187, 633, 643, __brassii, 123, 124, 199, 264, 292-294, 646-648, 649, 650 612 tItalics indicate synonyms; asterisks, new names and combinations; boldface, main references to names discussed. 1980 ] STEVENS, CALOPHYLLUM 693 —burmannii, 158, 258, 270, 274, 279, 172, 258, 305, 308, 316-318, 319, 466, 562 320 —— var. brachiatum, 274 —chinense, 681 —— var. bracteatum, 261, 274 —cholobtaches, 278, 287, 291, 292, —— var. bractiatum, 261 401, 402 var. parvifolium, 258, 562 —*clemensorum, 145, 184, 253, 494, aeolian 131, 498, 499, 500 497, 503-505, 525, 635 —calaba, 119, 129, 149, 154, 156, 167, —*collinum, 127, 130, 132, 160, 165, 174, 177, 179, 182, 183, 189, 225, 200, 542, 551, 571, 572, 579, 586 248, 254, 256-275, 277, 367, 677, _—comorense, 172, 305, 337, 338 691 —*complanatum, 160, 184, 368, 383, ——var. *australianum, 156, 164, 386, 389, 394, 395 203, 244, 258, 264, 269, 270, 272, —*confertum, 186, 396, 416, 425-427, 579 428 —-—var. *bracteatum, 123, 135, 142, —confusum, 165, 197, 203, 536, 538, 155, 164, 230, 231, 244, 258, 261- 540, 547, 548 267, 270-273, 275, 466, 514, 679 —congestiflorum, 596, 603 — var. calaba, 137, 160, 164, 237, _—cordato-oblongum, 174, 255, 277, 244, 257, 258- 260, 270-272, 274 414, 478, 479, 691 ——var. *cuneatum, 164, 257, 258, —coriaceum, 131, 179, 350, 358, 360, 264, 267, 268, 272, 419 362-364, 371 —var. E, 164, 197, 258, 264, 268, —costatum, 160, 200, 542, 572-574, 269, 290.5273 , 593 ——var. *worthingtonii, 164, 237, —costulatum, 132, 138, 164, 165, 181, 244, 258, 260. 261, 270, 271, 691 454, 463, 465, 470, 471, 528 —calabiformis, 149 —cucullatum, 131, 132, 194, 362, —calaboides, 254, 256 382-384, 385, 396 —*calcicola, 160, 184, 414, 446, 488, | —cumingii, 497, 500 492, 493, 552 —cuneatum Henderson & Wyatt- —caledonicum, 127, 144, 198, 542, Smith, 267, 272, 412, 419 549, 566-569, 580 —cuneatum Vidal, —canum, 133, 139, 142, 158, 180, 185, —cuneifolium, 128, 164, 174, 240- 193, 207, 280, 364, 370, 371-375, 244, 247, 248-250, 691 376, 377, 417, 476, 626, 640 —curtisii, 262, 271, 275 —carrii, 198, 569-571, 593 —cuspidatum, 654, 656 ——var. carrii, 542, 552, 570 —cymosum, 277, 291, 434, 437 ——var. longigemmatum, 552, 570 —dasypodum, 141, 142, 156, 183, 193, —*castaneum, 126, 132, 138, 185, 195, 275, 358, 364-368, 422, 443, 297, 351, 353-355, 361, 362 Sle —*caudatum, 197, 535, 536, 538, 540, —decipiens, 240, 251, 253, 256 541, 542, 544, 545, 669 —depressinerve, 389 —celebicum Koord., 279, 582 —depressinervosum, 130, 144, 158, —*celebicum P. F. Stevens, 157, 200, 165; 176, 178 4182< 191, 368; 385, 396, 437, 445-447, 679 386, 389-393, 399, 412, 529 —cerasiferum, 150, 158, 160, 165, —diepenhorstii, 278, 291 166, 201, 542, 549, 556, 557, 562- —*dioscurii, 141, 156, 182, 192, 514, 565, 568, 600 525, 528, 530-532, 533, 538 —cerasiforme, 679 —*dispar, 164, 179, 183, 191, 192, —*ceriferum, 128, 176, 446, 494, 497, 494, 517, 520, 522, 524- 527 500-502 —dongnaiense, 175, 211, 224, 234, 235 —changii, 207, —drouhardii, 173, 258, 305, 311, 312 —chapelieri, ee 144, 164, —dryobalanoides, 176, 209, 224, 230, 694 JOURNAL OF THE ARNOLD ARBORETUM [vot 61 232, 233, 508 —gracile, 507 —dubium, 681 —pgracilipes, 123-125, 165, 192-194, —*echinatum, 197, 633, 663, 674-676 625, 633, 653-657, 659, 660, 677 —elatum, 217, 220, 225, 226, 238,691 —gracillimum, 182, 375, 531, 549, —elegans, 165, 186, 188, 404, 407, 624-626, 628, 652 409, 410, 411, 414, S11 —grandiflorum, 121, 123, 124, 156, —ellipticum, 153 —enervosum, 135, 177, 416, 423, 424, 432 —eputamen, 139, 154, 173, 321-323, 324, 337, 338 ——var. eputamen, 140, 322, 323 ——-var. grandis, 322-323, 326 —euryphyllum, 123, 129, 130, 133, 200, 574, 590-593, 598 —excelsum, 682 —*exiticostatum, 160, 181, 189, 663, 665, 668, 676, 677 —_ferrugineum Merr., 206, 214 164, 469- —ferrugineum Ridley, 132, 144, 181, 182, 348, 432, 458-467, 471, 476, 478 ——var. ferrugineum, 123, 165, 459, 460, 462-464, 471 ——var. neriifolium, 460, 467 I23, 432, 230, 5 463, ——var. *orientale, 164, 189, 432, 442, 459, 460, 462, 465, 466 —*fibrosum, 172, 305, 320, 321 —flavocorticum, 206, 21 oo 178, 348, 396, 421, 422, ee cas 232, 390, 399, 400, 507, 510, 512-514, 530, 531, 617 —foetidum, 395, 507, 513, 514 —fragrans, 429, 430, 476, 478 —fraseri, 136, 154, 158, 178, 183, 232, 272, 416, 417-420, 436, 441 — 357, 361 —gaimanum, 600, 601, 602 404, _Kearcinioides, 128, 135, 142, 145, 153, 154, 160, 188, 190, 412-417, 419, 421 —glabrum, 206, 217, 218 —glaucescens, 186, 368, 384, 385, 404 —globuliferum, 508, 511, 515 —goniocarpum, 123, 132, 140, 165, 199, 202, 294, 591, 598, 606-608, 610-612 165, 178, 414, 487-489, 492, 590 —grandifolium, 132 “eum 345, 363, 481, 507, 511, — i. 191, 652, 661-664 —harmandii, 262, 275 —hasskarlii, 262, 364, 367 —*havilandi, 187, 414, 446, 450-452, 469 —heterophyllum, 138, 165, 200, 201, 598, 606, 607, 608, 609 —hexapetalum, 682 —hibbardii, 278, 289, 291 —hirasimum, 160, 200, 551, 598, 600, 603, 6 —hirtellum, 277, 291 —horstii, 452, 457 —hosei, 131, 160, 178, 78 188, 430, 460, —*humbertii, 172, 300, 305, 306, 307 —izermannii, 481, 485, 486 —incrassatum, 345, 347, 348, 361 —*incumbens, 141, 160, 183, 192, —inophylloide, 432, 436, 439, 440 ——var. inophylloide, 436, 440 . Singapurense, 432, 440, 472 —inophyllum, 119, 123, 130, 135, 139, 141-144, 149, 150, 154, 156, 158, 161, 163, 167, 168, 170, 173, 174, 177, 181, 189, 196, 258, 280, 287, 316-318, 324-335, 337, 439, 447, 557, 558, 560, 568, 569, 691 ——var. blumei, 325 ——f. oblongata, 325, 334, 335 ——f. obovata, 325, 33 —insigne, 350 —*insularum, 130, 132, 137, 142, 165, 199, 542, 621, 665, 669, 670 —intramarginale, 432, 442 —Jjavanicum, 481, 485, 489 —kajewskii, 166, 584, 585 —karoense, 272 1980] STEVENS, CALOPHYLLUM 695 —kingianum, 385 —*milvum, 128, 129, 142, 160, 172, —kiong, 278, 291, 292 258, 300, 302-304, 305-307, 310, 324 —kunstleri, 263, 279, 287, 294, 297, —wminahassae, 279 460, 679 —mindanaense, 206, 216 —var. longifolium, 679 —wminus, 679 —lanceola, —miquelii, 432 —lanceolarium, 335 —molle, 130, 139, 179, 185, 355, 375, —lanceolatum Blume, 335, 555 378, 379, 382, 524, 527, 621, 626, —lanceolatum Teijsm. & Binn., 364, 633, 640, 641-644 367 —montanum, 566, 568, 580 —lanceolatum Warb., 278, 290 —moonii, 123, 129, 134, 143, 174, 237, —lancifolium, 278, 291 280, 299-302, 691 —lanigerum, 348, 350, 353, 356-361, | —morobense, 197, 203, 598, 616, 617 364 —motleyi, 518, 523 ——var. *austrocoriaceum, 135, 180 —*mukunense, 138, 160, 179, 184, 185, 350, 353, 355, 357, 358-360, 454, 488, 490-492 361 —muscigerum, 638 ——var. lanigerum, 190, 353, 357, —nagassarium, 682 358, 360, 361 —nathorstii, 148 —laticostatum, 156, 165, 195, 199, —neo-ebudicum, 123, 132, 141, 157, 552, 579-582, 583, 584 160, 165, 166, 198, 202, 289, 542, —laxiflorum, 173, 307-309, 310, 318 548, 549, 556, 557-562, 566, 580, —*leleanii, 129, 132, 144, 201, 288, 629 291, 573, 574, 587-589, 590, 621 —neriifolium, 460 —leptocladum, 132, 137, 197, 558, —neurophyllum, 682 560, 628, 629, 631, 652 —nodosum, 123, 125, 126, 130, 144, —leucocarpum, 137, 158, 165, 202, 145, 160, 163, 165, 178, 187, 368, 538, 548-550, 626 385-389, 391, 393, 395, 396, 399, —lineare, 154, 161, 164, 177, 494, 520, 412, 524 522, 523, 524 —novoguineense, 128, 165, 197, 533- —*lingulatum, 172, 300, 304-306 536, 538, 541, 544, 547, 548, 617 —lonchophyllum, 679 —obliquinervium, 123, 128, 142, 160, —longifolium Wall., 681, 682 186, 191, 194, 427, 633, 643, 644- —longifolium Willd., 153 646, 648-650 —lowei, 375, 377 —oblongifolium, 460 —lowii, 139, 160, 180, 185, 280, 347, —oblongum, 279, 289 368, 371, 374, 375-378 —obovale, 510, 515 —macrocarpum, 128, 135, 140, 160, _—obscurum, 201, 574-576, 593, 598 175, 182, 190, 432, 446, 452-457 —odoratissimum, 27 —macrophyllum, 196, 617, 678, 679 —oliganthum, 195, 383, 396, 411, 412 —marginatum, 436, 443 —ovatifolium, 325 —medium, 263 —paludosum, 279, 290 _—_megistanthum, 207, , 214 —palustre, 375, 378 —membranaceum, es i 177,665, —*paniculatum, 137, 144, 173, 300, 672-674 305, 312-316 —mesuaefolium, 395 —aff. paniculatum 1, 300, 305, 313, —microphyllum T. Anderson, 385, 314, 315 389 —aff. paniculatum 2, 300, 305, 314, —microphyllum Planchon & Triana, 315, 316 546 —aff. paniculatum 3, 305, 315, 316 —microphyllum Scheffer, 546, 547 —papuanum, 123, 124, 129, 133, 134, 696 JOURNAL OF THE ARNOLD ARBORETUM [vou 61 140-142, 144, 158, 160, 165, 196, 130, 133, 145, 158, 161, 165, 169, 574, 593-595, 597, 600, 602 178, 187, 260, 268, 279, 366, 383, 385, 386, 389, 391, 393, 395-400, —parkeri, 154, 176, 414, 416, 419, 420, 421 —parviflorum, 139, 144, 154, 173, 302, 303, 323, 324, 326 —parvifolium Choisy, 165, 197, 535, 536, 540, 546, 547 —parvifolium Vesque, 385 —pauciflorum, 123, 129, 142, , 196, 595-600, 602, 604 —peekelii, 123, 125, 140, 158, 160, 162, 166, 200, 574, 584-587, 590, 618 —pelewense, 193, 292, 383, 396, 401, 402, 413 ) 160, —pentapetalum, 131, 134, 195, 454, 485, 494-500, 502, 649, 668 ; ee 454, 485, 497, 498, 499, 500, 503 ——var. Serene 194, 454, 485, 494, 496, 497, 499, ——var eee 497, 498, 499, 500 163, 165, 620-624 172, 173, 258, 304, —*persimile, 129, 145, 198, 598, 617, —pervillei, 128, 08, 309, 310 —piluliferum, 165, 203, 606-608, 613, 614, 652 166, —pisiferum, 123, 124, 128, 154, 158, 161, 177, 181, 183, 191, 192, 367, 466, 470, 494, 512, 518-523, 524- 526, 53 —plicipes —pliocenicum, 149 —*poilanei, 138, 154, 175, 663, 665, 670-672 —polyanthum, 133, 138, 142, 154, 157, 160, 164, 174, 176, 215, 217, 219, 220-226, 227, 229, 238, 273, 274, 350, 466, 555 —*praetermissum, 128, 129, 160, 186, 396, 416, 426, 427, 428 —praineanum, 417, 507 —procerum, 577 —pseudotacamahaca, 131, 497, 499, 00, 644 —pseudovitiense, 158, 289, 557, 559, 560 —pseudowallichianum, 207 —pulcherrimum, 123-125, 127, 128, 443, 481, 507, 514, 617, 654 , 682 —*pyriforme, 128, 145, 165, 191, 192, 402-405, 406-409, 432, 457 216 —ramiflorum, 577 eri 139, 173, 305, 326, 338- 340, Sa 123, 124, 128, 138, 139, 165, 192, 630, 633, 656, 657- 659, 660 —retusum, 258, 263, 371, 395, 459, 466-469, 472, 478, 518 ——var. cambodgense, 518, 523 ——var. cochinchinense, 518, 523 ——var. parvifolium, 262, 274, 466 —revolutum, —rhizophorum, 432, 447, 450 —*rigidulum, 160, 194, 652, 658, 663, —rigidum, 121, 123-125, 128, 133, 135, 138, 161, 180, 183, 189, 192, 258, 280, 287, 294-298 —robustum, 123, 130, 132, 133, 161, 202, 258, 264, 298, 299 —*roseocostatum, 127, 165, 191, 386, 404, 405-407, 409 —rotundatum stoiundifoliui: 137, 154, 160, 178, 503, 505, 625, 633, 634-636 —rubiginosum, 130, 132, 137, 139, 179, 191, 375, 382, 405, 621, 626, 630, 633, 638-641, 643 —*rufigemmatum, 125, 130, 132, 139, 141, 163, 169, 180, 348, 349, 350- 356, 362, 640 —rufinerve, 137, 201, 202, 598, 606- 610 —*rugosum, 142, 154, 176, 621, 630, 631-634, 635 —rupicola, 133, 144, 158, 161, 178, 183, 494, 497, 512, 522, 529 ——var., 123, 183, 497, 516-518 ——var. elatum, 507, 514, 515, 517, 177, 515-518, 1980 ] STEVENS, CALOPHYLLUM 518 —saigonense, ae 274, 275 r. na , 268 —*sakarium, 42, 160, 186, 189, 633, 643, 646-649 —samoense, 557, 560-562 —sangkae, 518, —savannarum, 134, 163, 196, 552-554, 624 —schefferi, 546 —sclerophyllum, 121, 123, 124, 150, 160, 161, 165, 181, 186, 525 542, 127, 188, 414, 439, 440, 444, 445, 447-450, 452 —scriblitifolium, 123, 128, 180, 184, 347, 364, 368-371 —sil, 142, 158, 161, 165, 198, 572-574, 9 —solomonense, 589 —sorapa, 307, 3 —smilesianum, 221, 225 var. lutea, 221, 225 144, 278, 291, 08 —sorsogonense, 280, 289 —soulattri, 137, 121, 140-143, 160, 161, 166, 123-125, tT 292, I33;, 130- 160, 587, 134, —spectabile, 131, 150, 158, 207, 236, ZA os ceo Bs 335, 344, 346, 350, 363, 371, 292. 299, 202, 315; 318, 514, 56 —*stipitatum, 165, 178, 186, 386, 396, 404, 407, 408-410 —streimannil, 123, 203, 573, 598, 615, 616, 617 —suberosum, 123, 130, 140, 145, 160, 163, 165, 620, 622-624 166, 1973-598 .°6 17, 618- —subhorizontale, 180, 182, 439, 663, 664-666 —subluridum, 44 —subsessile, 131, d 165, 179, 454, 487, 488, 489, 490, 492 —*sundaicum, 123, 160, 164, 486, 165, 697 181, 190, 446, 454, 463, 465, 466, 47 467- —suriga, 277, 290, 291 —symingtonianum, 154, 160, 176, 179, 211, 222, 229-232, 233, 234, 287, 417, 463 —tacamahaca, 123, 124, 139, 141, 154, 158, 173, 292, 305, 325, 326, 335- 337, 338, 340, 498, 680 —tahanense, 346, 347, 348, 350 —tenuicrustosum, 555, 556, 557 —tenuivenium, 483, 485 —tetragonum, 653, 656 ——var. parvifolium, 653, 656, 657 —tetrapetalum, 277, 290, 344 —tetrapterum, 121, 127, 128, 156, 164, 177, 184, 232, 399, 400, 411, 443, 505-515, 517, 518, 522, 617 —— var. *blumutense, 506, 508, 512, 213,514 ——var. *obovale, 135, 169, 186, 187, 190, 506, 508, 510-512, 513-515, 651 ——var. tetrapterum, 123, 158, 169, 193, 400, 494, 506-510, 513-515, 531, 656 —teysmannii Miq., 119, 121, 124, 127, 130-132, 136, 144, 164, 165, 181, 188, 190, 367, 420, 430, 431-443, 505 —var. *bursiculum, 130, 134, 142, 186, 432, 434, 436, 440, 441 ——var. *inophylloide, 123, 126, 130, 135,..1365- 157.158, tol. 166, 190, 414, 420, 430, 432, 436-439, 440- 443, age 476 —var. teysmannii, 123, 130, 134, 136, 158, 426, 432-435, 439-442 —teysmannii Zoll. ex. Planchon & Triana, 262, 270, 275, 364, 367, 442, 443 —thorelii, 132, 142, 160, 176, 209, 211, 213; 215, 218-220, 221, 5225, 229, 235 ——var. oxycarpum, 218, 220 —thwaitesii (incl. a), 164, 175, 237, 240-244, 252, 691 ——f, 244, 251, 253, 505 —tomentosum, 123, 128, 131, 174, 221, 222, 226, 235-238, 269, 302, 366, 691 —*touranense, 164, 176, 204, 209, 211, 215, 225, 226-228, 229 698 —touriga, 682 trachycaule, 123, 141, 165, 201, 202, 598, 604- 608, 609, 610, 612, 614, 670 —trapezifolium, 135, 136, 142, 148, 154, 160, 164, 174, 222, 237, 240, 242, 243, 244-248, 250, 691 —treubii, 279 —*undulatum, 131, 132, 157, 160, 165, 197, 535, 536, 540, 542-545 —vanoverberghii, 291, 680 —venulosum, 121, 123, 124, 131, 132, 134, 141, 156, 158, 179, 272, 454, 479-487, 489, 490, 492, 493, 552, 590, 668 ——var. *tenuivenium, 134, 165, 182, 480, 483, 485, 614 —-—var. venulosum, 134, 142, 165, 184, 194, 454, 480, 481-483, 485, 486, 488, 492 —venustum, 507, 514, 637 —*vergens, 134, 145, 164, 174, 204, 237, 244, 251-253, 505, 691 —*vernicosum, 135, 139, 173, 300, 305, 326, 340-342 —versteegil, 278, 291 —*verticillatum, 128, 135, 137, 144, 164, 172, 258, 300, 305, 308, 309, 317, 318-320, 342 —vexans, 123, 130, 139, 158, 160, 165, 166, 196, 203, 574, 597, 600-603 —vidalii, 683 —vitiense, 141, 158, 160, 165, 166, 198, 554-557, 558, 560, 564, 568, 80 ——var. 556, 557 ——var. vitiense, 542, 549, 555, 556, wei *amblyphyllum, 543, 555, —wakamatsul, 325, 334 —*waliense, 121, 123, 199, 573, 574, 588, 589, aol —walkeri, 136, 138, 150, 160, 164, 175, 222, 237, 238-240, 242-244, 248, 249, 251, 691 —wallichianum, 180, 185, 207, 214, 263, 279, 326, 342-350, 352, 353, 355, 368, 375, 638, 640 ——var. *incrassatum, 123, 127, 130, 156, 165, 326, 343, 344, 345, 346, 349, 350, 353, 364 . *tahanense, 165, 326, 343, JOURNAL OF THE ARNOLD ARBORETUM [vot 61 346, 347, 350 —___var. wallichianum, 123, 165, 326, 343-345, 353 —warburgii, 278, 577 —warenense, 533, 536 —wawaroenti, 582 —whitfordii, 136, 137, 157, 217, 566, 629-631, 652 —wightianum, 253, 256 195, 207, 21, 225 —*woodii, 192, 351, 354, 368, 378- 380, 381 —zeylanicum, 244, 247, 248 —zschokkei, 278, 291 Calophyllum sp. 10, Henderson & Wyatt-Smith, 358 Calophyllum sp. 23, Wyatt-Smith, 204 Calophyllum sp. 44, Henderson & Wyatt-Smith, 434, 439, 441 Calophyllum sp. 45, Henderson & Wyatt-Smith, 530 Calophyllum sp. A, P. Henderson & F. Stevens, Calophyllum sp. B, P. F. Stevens, 609 Calophyllum sp. C, P. F. Stevens, 620, 624 Calophyllum sp. D, P. F. Stevens, 571 Calophyllum sp. 9, 184, 204, 211, 222, 233, 234, 679 Calophyllum sp. 53, 364, 380, 381, 382 Calophyllum 54, 368, 381, 382, 6 Calophyllum sp. 61, 396, , 401 190, 351, 354, 179, 351, 354, 128, 188, 383, Calophyllum sp..°65,, 165,. 169; 121, 403, 404, 407, 408, 409 Calophyllum sp. 73, 130, 178, 416, 422, 423 Peas sp. 95, 194, 488, 493, 494 Calophyllum sp. 98, 160, 189, 494, 502, 503, 520 Calophyllum sp. 121, 551, 552, 565, 566 Calophyllum sp. 129, 156, 165, 199, 551, 574, 580, 581, 582, aay. 153, 157, 195, 542, 185, 583, 584 Calophyllum sp. 143, 160, 165, 197, 1980 | STEVENS, CALOPHYLLUM 591, 598, 606-608, 610, 612, 613 Mammea, 149-151, 681 Calophyllum sp. 151, 158, 181, 182, —americana, 144, 149 621, 625, 626-628, 652 —longifolia, 290, 681 Calophyllum sp. 163, 193, 621, 643, —neurophylla, 682 650, 651, 652 —odorata, 682 Calophyllum sp. 164, 193, 630, 653 —suriga, 290 Calophyllum sp. 170, 175, 658, 663, —touriga, 682 665, 666, 667 Melaleuca, 540 Calophyllum sp. 178, 195, 663, 677, Mesua, 143, 149-151, 682 678 —ferrea, 682 Calophyllumpollenites rotundus, 149 —nagassarium, 125, 682 Calysaccion chinense, 681 —nuda, a —horstii, 452 —thwaitesii, 682 —longifolium, 681 Uiise sts la truncatum, 141 Castanopsis, 598 Casuarina, 462 Nothofagus, 598, 604 Ochrocarpus, 149 Dacrydium, 462 Derris, 374 Dimorphanthera, 294 Diospyros, 161, 164 Dipterocarpaceae, 233, 333 Drimys, 164, 166 Paramammea, 127, 149, 151 Dryobalanops, 233 Parastemon, 650 Podocarpus, 294 Poecilonet.on, 149, 151 —pentapetalus, 496 Oncosperma, 645 Endodesmia, 150, 151 —calophylloides, 151 } —indicum, Eugenia, 161 —pauciflorum, 151 Ponna, | Garcinia, 149, 680, 681 Psorospermum, 127 —angustifolia, 681 —alternifolium, 127 ie 4 . Raatamaa Quiina descaineana, 443 143 Gleichenia teen 529 Gonostylus, 65 Guttiferae, 117, 119, 143, 150 —subfam. a lal 117, 119, Rheedia javanica, 680 —umbellata, 680 Rhus succedana, 681 Rinorea, 161 144, 149, ——tribe mic iviieas 14? Schuurmansia vidalii, 683 ——tribe Endodesmieae, 150 Shorea, —subfam. Clusioideae, 14 —albida, 650 ——tribe Garcinieae, 143 Stemonurus, 650 Guttiferoxylon fareghense, 149 Syzygium, 161 Heliciopsis, 121 Tetracera akara, 680 Tovomita pentapeialn: 494, 550 Kayea, 149 Vaccinium, 294 Lebrunia, 150, 151 Vateria, 333 Lithocarpus, 294 Litsea, 650 Whampoa sinensium, 443 NOTICE Because of continually increasing production costs, the Editorial Committee of the Journal of the Arnold Arboretum is requesting authors to help defray publication expenses. Beginning with manu- scripts received and accepted for publication as of March, 1980, a page charge will be levied. Acceptance of manuscripts for the Journal will, however, continue to be based solely on appropriate- ness and scientific merit, not on an author’s ability to meet page costs. Information on page charges and general editorial policies can be obtained from the Managing Editor. sagt of ane onal ——_ : E = = ae, 1980 ~ CONTENTS OF VOLUME 61, ‘NUD BERS A Revise of the Old World Species of Calo (Gautier) Pichi STEVENS ers a es ee SYSTEMATIC TREATMENT, ‘CONCLUDED . eater, aot TLiveRATURE CITED sO So ADDENDUM .... < = ae = etic sheet ee INDEX OF Scientiric Nawes. ea ee Volume 61, Number 2, Races pages 117-424, was issued December 19, 1980. JOURNAL oF tHe ARNOLD ARBORETUM HARVARD UNIVERSITY VOLUME 61 NUMBER 4 wate ‘US ISSN 0004- 2625 3S of ae aE: Arboretum Published quarterly in January, Bue ‘July, and eee by the Arnold Arboretum, Harvard University, = ‘Subscription price $25. 00 per year. : Subscriptions and remittances should be sent to ous E. B. Schmidt, ame Arboretum, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. Claims will not be accepted after SIx months from the date of issue. a POSTMASTER: send address changes to Ms. E: B. Schmidt, Arnold Arboretum, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U. S. A. Volumes 1-51, ‘reprinted, and some back numbers of volumes 52-56 are _ available from the Kraus “aR Corporation, Route 400, Millwood, New York 10546, U. S.A EDITORIAL COMMITTEE 8. AS Spongberg, Editor __E. B, Schmidt, Managing Editor P-S. Ashton. -K. S. Bawa -P. F. Stevens Sone or Wood, Jr. : Printed at Edwards Brothers, Inc., Ann ae Michigan OVER: rE rg bs rs i eee Wet pee : epee Sree 2s neatly 160° times) of a Fijian seach Calophy tum Ieucocarpum —C: Smi ith, for e basis of this year’s stylized cover design. A row of trichomes is’ used in the tnaioe on the back cover, while a cluster of eae appears on the offprints. __As in recent years, the designs, based on holotype material in the herbarium of the Amold Arboretum, were drawn by Karen Stoutsenberger. st Calophyllum oles (Guttiferae) is a wide-ranging genus of oe forest trees in bo th the New and Old Worlds, and some species are of considerable economic portance as sources of valuable timber. Its use on the cover of this volume BOF the Journal of the Arnold Arboretum is appropriate since the April and July _ numbers are devoted to P. F. Stevens’ Ss revision of thie Old World representatives of this genus. —S.A.S. Second-class postage paid at Boston, Massachusetts, and additional offices. JOURNAL OF THE ARNOLD ARBORETUM VoLuME 61 OcToBER 1980 NuMBER 4 TAXONOMY OF THE WEST INDIAN CYCADS JAMES E. ECKENWALDER TEN GENERA of Cycadales exist in world floras. Four genera, all members of the Zamiaceae, are natives of the Western Hemisphere, and two of these are found in the West Indian region. The monotypic Microcycas (Miq.) A. DC. is endemic to Cuba, while Zamia L. is found from Mexico south to Bolivia and Brazil, as well as in the West Indies (Schuster, 1932). Within the West Indian region, cycads are indigenous to all of the Greater Antilles, the Bahamas, Florida, and the southeastern coast of Georgia but are absent from the Lesser Antilles and the Virgin Islands. Because of its restricted distribution and seeming morphological isolation, Microcycas has hereto- fore caused little dispute, but the taxonomic history of West Indian zamias, like that of many other West Indian plant groups (Howard, 1977), has comprised a confusing array of divergent circumscriptions and delimitations, resulting in widely different species tallies (from one (present study) to 14 or more (de Candolle, 1868)) for the region. Altogether, some 35 entities have been described, accounting for about 40 percent of the names proposed in the Diverse taxonomic treatments can 2 oe hes to complex variation patterns and political fractionation of the region, as well as to differences in philosophy among taxonomists. Most previous workers have had a very narrow species concept and have acknowledged neither extensive variation within populations nor continuity between populations. Often species were described on the basis of single specimens— sometimes cultivated plants for which the original sources were unknown. Other than Schuster’s (1932) synonymies, which are tortuous and partly invalid under the present International Code of Botanical Nomenclature, there has been little effort to consolidate nomenclature in different subregions, and nomenclatural and taxonomic chaos persist. This paper proposes a new taxonomy of West Indian cycads, based on field, herbarium, and garden studies; explains the contrast between this class- ification and those of previous workers; and accounts for most published names in the complex. peony © President and Fellows of Harvard College, 1980. Journal of the Arnold Arboretum 61: 701-722. October, 1980. 702 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 TAXONOMIC AND NOMENCLATURAL HISTORY West Indian cycads were described by pre-Linnaean authors, usually as species of **Palma,”’ and Linnaeus (1763, 1764) founded his genus Zamia, with its sole species, Z. pumila L., on their descriptions and illustrations. Smith (1961) raised an important question concerning the typification of Z. pumila. He correctly contended that the Linnaean protologue contained references to two currently recognized species of Zamia, and that correct application of the name depended upon lectotypification by subsequent authors. Most authors have either rejected Z. pumila because of this mixture (e.g., Schuster, 1932) or have used it for the West Indian cycad described and illustrated by Commelijn (de Candolle, 1868; Carabia, 1941). Smith, in contrast, argued that Miller (1768) had first lectotypified the name using the other references. Thus, he contended that Z. pumila L. should be applied to the Mexican species currently known as Z. furfuracea Aiton, and that the West Indian species that had almost universally sani the Linnaean epithet should be known as Z. debilis Aiton. This argument, which has largely been ignored, can be set safely aside within the framework of the ICBN. Miller’s account need not be taken as a lectotypification of Z. pumila L. because it describes Palma pumila Miller and contains no reference to Linnaeus’s combination. Nomenclaturally, it can be considered an independent publication of the epithet. The first post-Linnaean use of Z. pumila may be by Aiton (1789), who applied it (as a synonym) to a West Indian cycad. Zamia pumila L. ma thus be retained for the common West Indian cycad that has traditionally been known (in part) by that name, and Z. furfuracea Aiton remains the correct name for a Mexican species. Soon after the original description of Zamia, new West Indian species were added to the genus. Aiton (1789) contributed Z. debilis Aiton and Z. integrifolia Aiton, the atter a renaming of Z. pumila L. With Z. media Jacq. and Z. angustifolia Jacq., Jacquin (1791, 1798) created the narrow species concept, based on leaflet widths, in West Indian Zamia. Aiton and Jacquin set trends followed by many later authors dealing with the taxonomy of Zamia in the West Indies—purposeful or inadvertent renaming of previously described entities, and proliferation of described taxa based on slight differences in leaflet width. As an example of the former trend, cycads with narrow leaflets from the south coast of the former Oriente Province of Cuba have successively been named Z. stricta Miq., Z. vatesii Mig., Z. angustissima Miq., Z. multifoliolata A. DC., and Z. guggenheimiana Carabia. Many plants attributable to these taxa are virtually indistinguishable from cycads from Eleuthera in the Bahamas named Z. angustifolia Jacg., and from others from Mayagtiez Province, Puerto Rico, named Z. portoricensis Urban. The second trend is reflected in such epithets as Z. angustissima Mig., Z. angustifolia Jacg., Z. media Jacq., and Z. latifoliolata Prenl. The tendency toward a narrow species concept defined on leaflet width is even reflected in treatments of fossils of Zamia from the West Indies (Hollick, 1932). Additional names proposed in the last 200 years are included later in the synonymy of Z. pumila subsp. pumila. warf cycads from western Cuba were first distinguished as Zamia pygmaea Sims (1815). Since these diminutive cycads also display variations in leaflet size compara- — ble to those of the more widespread larger plants, they too have engendered numer- ous specific epithets based on local variants. These names are included in_ the 1980] ECKENWALDER, WEST INDIAN CYCADS 703 synonymy of Z. pumila subsp. pygmaea (Sims) Eckenw. in the taxonomic treatment presented below. The other Cuban endemic, Microcycas calocoma (Miq.) A. DC., was first described as a species of Zamia in 1851. De Candolle (1868) transferred it to a new monotypic genus because of its striking differences from the other species of Zamia then known. Its rarity and distinctiveness have inhibited recognition of segregates, and little controversy has attended taxonomic accounts of this species since its identity was firmly established (Caldwell & Baker, 1907). Modern treatments of West Indian cycads adopt one of four differing points of view on the taxonomy of the Zamia pumila complex. 1) Each subregion has several species, which may or may not be shared between subregions (Schuster, 1932; Chamberlain, n. d.). This viewpoint is particularly attractive to botanists working within a single subregion and has characterized most treatments to date of West Indian cycads, including accounts for Florida (Small, 1933), the Bahamas (Britton & Millspaugh, 1920), Puerto Rico (Britton & Wilson, 1926), and Cuba (Leon, 1946; Liogier, 1969). It is less attractive when viewed against overall variation in the West Indian region. 2) Each subregion has a single endemic species (Read, | 67). This view ignores parallel variation in cycads of different subregions, particularly in Cuba, Puerto Rico, and the Bahamas. 3) The West Indian region has a single endemic species of Zamia. This is the viewpoint adopted here and rests on evidence presented below. It is more conservative than treatments such as that of Carabia (1941), which accept two or three species and are intermediate between viewpoints | and 2. 4) West Indian cycads are conspecific with those of the Central and South American main- land. W. C. Steward (pers. comm.) contends that Zamia is monotypic. This concept of Z. pumila ignores apparent discontinuities in variation to include a wide array of diversity in morphology, ecology, and chromosome number under one specific epithet. Since Steward’s arguments are not yet published, they can not be examined in detail here. Of the four taxonomic viewpoints enumerated above, numbers 2 and 4 do not seem to accord with the facts of variation in West Indian zamias, as seen here. Based on information presented below, a treatment reflecting viewpoint | must embody great arbitrariness in delimitation of taxa. I favor a conservative approach (number 3) as most workable and most in accordance with present variation in these plants. MATERIALS AND METHODS Approximately 1500 herbarium specimens (including 33 holotypes and isotypes) of Microcycas and Zamia from throughout the West Indian region were examined in the following herbaria: 4. AAH. BM. DAO. DAV. DUKE. F. FAU, FLAS, FSU, FTG. GA, GH. MICH, MO, MSU, NCU, NY. TEX. UC. UNC. US, USF. and the personal herbarium of Dr. J. de Js. Jimenez (Santiago, Dominican Republic). Over 500 living plants at Fairchild Tropical Gar- den, representing 34 accessions from known localities in all West Indian subregions and including a few interpopulational hybrids, were also studied. Field work in the Bahamas and throughout Florida complemented the herbarium and garden studies and provided information concerning the association of morphological variations with habitat differences. Occurrences were mapped, and leaflet length, width, anc vein number (characteristics that dominated previous species Ea eee were 704 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61] recorded. Variation in these characters is analyzed below. Additional morphological features, both vegetative and reproductive, were recorded; they also influenced taxonomic decisions. They are not specifically analyzed below but are summarized in the taxonomic descriptions presented. The short lists of representative specimens come from the full range of each taxon and emphasize morphological variation within those subregions in which each occurs. e 60 - / Oo oO 55 - oO / (e) y , fe) / e 2 50 + e / e ,0 Oo / @) : ; fe) fe) 45- ) 9 0 oO fe) eo , oe, fe) oO oO , - oe , o 405 eocee ,° ° / = @e@eoo 7 fe) © ge = e ® ofee 35 5 eo r axe) fo) aa fo) © eee’ fe) eo. /e rr) e o cee c e oe /0 @ > 30-4 eno oo e@ > cae cee‘e @ coe 60 © e - © @000CRDee oO 2 © ce esome oe ~ 254 © come oOo © CoO ecoaqmnce ee To aoca0D @ = © 00sss0’6O @0 o 3 oe fe) = 20 4 e fe) 0090080 @ 15 O aonnasae ee coocoe a0 ocqcacse cagas0 10-7 ooo focoeys) fone) (000 @ 4 GO 5 7 CO@ mene) foe) ] if T T T T T ] T 5 1O 15 20 Co 30 29 40 45 Leaflet width (mm) Ficure 1. Regression of vein number on leaflet width of West Indian zamias: circles, Zamia ae subsp. pumila; =. subsp. ee a; Bo ele — subspe- ).8 11] cies; regression line dashed, y = 1.66x + 3.29, = ( 1980] ECKENWALDER, WEST INDIAN CYCADS 705 RESULTS AND DISCUSSION Previous workers, often lacking fertile material, emphasized leaflet characteristics in their specific delimitations of West Indian zamias. Because leaflet length, width, and vein number differ between populations, these (especially the latter two) are the features that were most often used to distinguish new species. Furthermore, since early workers had few specimens available to them, their descriptions included narrow ranges of variation in these characters, and later authors continued this tradition (Read, 1967). A scatter diagram of the two most frequently used distinguish- ing characters (leaflet width and vein number), based on a representative sample of ). The correlation — 500 specimens, shows them to be closely associated (FIGURE coefficient for these characters for all 1043 specimens measured is 0.94, and similar high correlations characterize plants from cach of the West Indian subregions (TABLE 1). Thus, 88 percent of the variation in vein number in these cycad leaflets is accounted for by corresponding differences in leaflet width. It is not surprising, therefore, that authors could always find differences in vein number to reinforce differences in leaflet width. In fact, the features are structurally related: no matter Tape 1. Correlation coefficients of leaflet width with vein number and leaflet length for Zamia in the West Indian region. REGION VEIN NUMBER n LEAFLET LENGTH n West Indies 1043 0.07* 1083 Florida 401 0). 16** 345 Bahamas 69 0.45*** 106 Cuba 364 0.10% 427 Jamaica 11 0.24 n.s 19 Dominican Republic 3] os" 46 Puerto Rico 167 0.21* 130 + Correlation ceefficients significant at: *p<.05, **p<.01, ***ps.001. how wide or narrow the leaflets are, there are always about two veins per mm. of width. These veins arise by numerous dichotomies of two original traces from the rachis within the first 4 mm. after they enter the petiolule. Rarely, dichotomies may occur before the traces enter the leaflet, so that three or four traces enter, but this is as frequent in narrow leaflets as in wide ones. Thus. two traces enter most leaflets, and these ramify to maintain a relatively constant relation of vascular tissue to other tissues of the blade. Cycad taxonomists were incorrect in using differences in these features as independent differences between West Indian Zamia species, and leaflet width will be emphasized in subsequent analyses in this paper In contrast to leaflet width and vein number, a scatter pee of leaflet width vs. leaflet length (Ficure 2) shows no obvious association, and the low, but mostly significant, correlation coefficients (Taste 1) argue for the independence of these features within each subregion, as well as within the entire West Indian area. Leaflet length seems, instead, to reflect the general robustness of the plant. For instance, the correlation coefficient of leaflet length with total leaf length is 0.70, and other features are also associated in a loose group of ‘‘size’’ characters that are indepen- — 706 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61] fo) width Leaflet 2:5 5:0 7-5 lOO 12:5 15-0 17-5 20:0 22-5 25-0 27-5 Leaflet length (cm) Ficurr 2. Scatter diagram of leaflet width vs. leaflet length of West Indian zamias: circles, — ae subsp. pumila; spots, subsp. pygmaea; mixed symbols, both subspecies; 1 500. dent of a series of characters associated with leaflet width (Eckenwalder, in prep.). Therefore, leaflet length is used in addition to leaflet width to represent these two sets of vegetative characters in the analyses that follow. When leaflet widths of individuals are plotted in frequency histograms for the West Indies and for each of its subregions (Ficurr 3), it is apparent that these frequencies follow smooth distributions within areas and sum to a smooth, unimodal distribution for the entire region. These frequency distributions provide little support for recognition of many taxa in the region, a point emphasized by superimposing ranges of leaflet width of species accepted for the West Indies by the most recent comprehensive monographs (Schuster, 1932; Chamberlain, n. d.) onto the overall West Indian histogram (Ficure 4). Morphological boundaries between taxa accepted by these authors are clearly arbitrary, and the modal width for cycads of the region (6.5—7.5 mm.) is not even included in values for any of Chamberlain's species. If species boundaries are not recognizable in these data, Schuster’s further segregation of these cycads among several subsections of the genus seems even less warranted. Data on leaflet length show a similar trend of smooth distributions within subre- (Omen 60 50 pense) : 1 ‘O Ps E \ ML — a nu S) 10 ° LE nol lipo om o, — —_, ) 10 1S 20 25 30 Leaflet width (O-Smm) Ficure 3. Histograms of leaflet oe of zamias of the West Indian region: A, Florida (n = 754, % = 7. 8); B, Bahamas (n = 100, x = 9.1): C, Dominican Republic (n = 31, X = 12.7); D, Cuba (n = 364, x = 13.1); E, Puerto Rico (n = 167, x = 15.4); F, Jamaica (n = 11, X = 22.4); G, entire West Indian ae (n = 1427, X = 10.5): triangles mark locations of means. SAVOAO NVIGNI LSAM YWACTVWMNA AOA [0861 LOL e d cl ‘a C f 904 b) g BO = h kK iB , = T0- ia : > 604 noi 2 so 504 = .~ 40-7 te) » Boos 0) a E pa | 40 45 Leaflet width (0-5 mm) Ficurr 4. Ranges of leaflet width given by Chamberlain (n. d.) for West Indian species of Zamia superimposed on frequency ue of leaflet widths for the entire West Indian region: a, Z. angustissima; b, Z. angustifolia; c, Z. floridana; d, Z. pygmaea; e, Z. silice media; g, Z. latifoliolata; h, Z. portoricensis: pygmaea, all others of subsp. pumila). 1, Z. kickxii; j, Z. pumila; k, Z. ottonis (d. e, i, and k are synonyms of Z. ere ao 80L WOLAYOPTNV ATIONYV AHL JO TVWNUNOL 19 ‘10a ] 1980] ECKENWALDER, WEST INDIAN CYCADS 709 804A ) individuals ne) oO Number of ea ie T t aw wa T T I T t ea aS 2-5 5-0 7-5 10-0 12-5 15-0 17-5 20-0 22-5 25-0 27:5 30-0 32:5 35-0 Leaflet length (O-5cm) Figure 5 pete of leaflet lengths of zamias . the West Indian region: A, peas = 754, x = 11.3); B, Bahamas (n = 100, x = 13.5); C, Cuba (n = 364 = 7.6); D, Jamaica (n = 11, X = 14.4): E, peeies: Republic (n = 31, x = 15, 5); F, Puerto Rico (n = 167, % = 15.9); G, entire West Indian region (n = 1427, X = 11.1); triangles mark locations of means. gions and overall for the West Indies (Ficurr 5). In Cuba, however, cycads I have distinguished as Zamia pumila subsp. pygmaea contribute to a bimodal distribution of leaflet length. In their extreme form in westernmost Cuba, these smaller plants appear strikingly different from individuals of subsp. pumila from eastern Cuba, but there is an uninterrupted cline of intergradation in robustness across the length of the island (Ficure 6). This cline steepens at the border between the two subspecies in eastern Cuba (Map 1), so that most individuals can be readily identified to subspe- cies. With this exception, leaflet length, like leaflet width, provides little evidence 710 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 A = —— = a - I I I | O S) fe) 15 ral) ren 30 Leaflet length (cm) Ficure 6. Dice-grams of leaflet lengths of Cuban zamias from west to east across island by _ Prowniees A, Isla de Pinos (n = 61, x 5.2); B, Pinar del Rio (n = 127, 5.4); C, Habana (n = 34, X = 5.4): D, Matanzas (n = 16, k = 6.2); E. Las 2D = 130, x = 7.0); F, Camagtiey (n = 37, k = 9.0); G, Oriente (n = 112, % = 13.1 wrizontal line = range: vertical line = mean: large rectangle = twice the standard deviation; small rectangle = four times the standard error, delimiting 98% confidence interval of mean. — favoring segregation of taxonomic entities among West Indian zamias. Other characters examined, both vegetative and reproductive, also show continu- ous variation patterns that discredit the traditional recognition of numerous taxonomic segregates, but I have concentrated on those emphasized by previous workers. Particular combinations of features characterize many distinctive local populations that have been described as species in previous taxonomic treatments. For example, individual islands of the Bahamas have populations of Zamia differing markedly in appearance (Figures 7, 8; Eckenwalder, in press), and these populations were divided among four species by Britton and Millspaugh (1920). Similarly, Small (1933) recognized four species of Zamia in Florida, where plants of coastal hammocks in the northeast (Z. umbrosa Small) seem sharply distinct from those of the remainder of the peninsula (Z. floridana A. DC.). But distinctive local races that appear to maintain their integrity in particular subregions seem much less distinct in the context of total variation throughout the West Indies. Formal recognition of these variants would be difficult to achieve consistently without further multiplying de- om SS an) tee ° °o & oO @ ee oo [J . 200 km fod e ee a > F e @ 80° 78° 76° 74° 72° 70° 68° 66° Map |. Distribution of native West Indian cycads: spots, Zamia pumila subsp. pumila; circles, subsp. pygmaea (Cuba only); mixed symbols, intermediates (Cuba only); triangles, Microcycas calocoma (Cuba only). [0861 SAVOAOD NVICNI LSAUM “WAC TVMNA MOF 712 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 Ficure 7. Outlines of representative leaflets of Bahamian zamias: A, Eleuthera (Correll & Hill 45240, mo); B, Long Island (Britton & Millspaugh 6271, GH—1sotype of Zamia lucayana); C, Grand Bahama (Britton & Millspaugh 2576, ¥): D, Great Abaco (Correll et al. 42713, vrG); E. New Providence (Britton & Brace 704, Ny); F, Andros (Brace 6990, scribed entities and would not clarify the biological situation. The distribution of variants is complicated by effective dispersal by both birds and seawater (Eckenwalder, 1980), which has confused what may once have been a more straightforward situation. Now, however, the variation pattern within Z. pumila seems taxonomically irresolv- able (making the species an ochlospecies, as described by White, 1962, and by Whitmore, 1975), and the course adopted here is to describe and accept variabil- ity without formalizing it in a necessarily arbitrary infraspecific hierarchy like that of Schuster (1932) TAXONOMIC TREATMENT Key to INDIGENOUS West INDIAN Cycaps Stems erect, to 20 m. tall: leaves numerous, more than 10 per crown, 10 dm. or more long, with petiole ca. Y% total length of leaf; leaflets drooping, lanceolate, pointed, entire, articulated to rachis by police Oo petiolule 5-8 mm. wide; male and female cones with tawny pubescence, massive, 4 dm. or more long, peduncle less than 5 length of cone: scales of male and ‘male cones with rounded, raised apex; fleshy layer of seed pink or peach. ............ icrocycas calocoma. Stems largely subterranean: leaves few 2 per crown, I—8(-15) dm. long with petiole ca. ¥ total length of leaf: leaflets horizontally spreading or ascend- ing, linear, oblong, oblanceolate to obovate, rarely lanceolate, usually blunt, usually with few to many callose teeth apically, articulated to rachis by callose ie) — 1980] ECKENWALDER, WEST INDIAN CYCADS 713 petiolule 2-5 mm. wide; male and female cones with purple or rusty pubescence, compact, 0.2—2.5 dm. long, peduncle at least 4% length of cone; scales of male and female cones with flat, truncate apex; fleshy layer of seed orange or red. pea ew Mae 4 a ree woe ede ein geen ee bea ae he Zamia pumila. 2. Plants robust, stems 2.5 cm. or more in diameter; leaves 4—8(—15) dm. lon leaflets 5—25(—35) cm. long, usually linear or oblong, usually revolute with thickened margin, thick and leathery; cones 0.5—2.5 dm. long. ........... eae e Beg gre BA Gels Sakata Z. pumila subsp. pumila. . Plants dwarfed, stems 2.0 cm. or less in diameter; leaves 0.3—6(—10) dm. long; leaflets 2.5-7(-10) cm. long, often obovate, margin thin, thin and fragile; cones 0.2-1.0 dm. long. ............. Z. pumila subsp. pygmaea. N Microcycas calocoma (Mig.) A. DC. Prodr. 16(2): 538. 1868. Zamia calocoma Miq. in Van Houtte, Fl. Serres Jard. Eur. 7: 141. 1851. Type: Cuba, cult. Hort. Amstelod., Miquel s.n. (holotype, 1). ILLustRations: Caldwell, Bot. Gaz. (Crawfordsville) 44: figs. /0-/4. 1907; Caldwell & Baker, Bot. Gaz. (Crawfordsville) 43: fig. /. 1907; Schuster, Pflanzenr. 99: figs. OP, 8K; pl. 5. 1932 (0-5 mm) = . 3 = 10-7 ) 4 ; = } =—9— | oS — ® a) 5 3 7 b T T T T T T T T T T T T T ] T q lO 15 20 Leaflet length (0O-l cm) x) for zamias from — Ficure 8. Plot of means of leaflet width ) VS. ee length different Bahamian iene a, Andros (n = 27, .4, y = 7.3); b, Eleuthera ( 24, x 3.6); c, New Boies Ci = 9,x = 12.0, y = 8.8); d, Great Abaco (n = ee = 12.4, y = 15.4); e, Grand Bahama (n = 18, x = 12.5, y¥ = 16.1); f, Long Island (n = 17, X = 17.3, ¥ = 19.3): horizontal and vertical lines represent twice the standard error on each side of mean. 714 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Arborescent, sparingly branched cycads up to 20 m. tall; crown formed by 10 to 50 pinnate leaves; leaves I-2 m. long, emerging in annual flushes, covered by transient, mealy, tawny pubescence; leaflets 50 to 80 pairs, 8-20 cm. long, lanceolate, reflexed, caducous after I—3 years from more persistent rachis, the rachis falling after 4 or 5 years, leaving an armor of persistent leaf bases that eventually abscise, the older portions of trunk developing shallowly furrowed, thin bark; male and female cones annual, | or 2, emerging from among scale leaves of crown, massive, 4-7 dm. long, with 500 to 1500 peltate hexagonal scales in ca. 20 straight rows, the external face tawny-pubescent, with raised, rounded knob; outer fleshy layer of seed coat light pink, turning peach at maturity, seed 2.5-3 cm. long. 1 = 13 (Sax & Beal, 1934). Distripution. Endemic to woodlands of canyons and foothills of the Sierra del Rosario in Pinar del Rio Province, western Cuba (Map 1), where it is apparently rare (Caldwell, 1926). Although the species is in little immediate danger of extinction because it is not commercially exploited and because the slopes on which it grows are unsuitable for cultivation, it is listed in Appendix | of the International Conven- tion on Trade in Threatened and Endangered Species (Ayensu & DeFilipps, 1978). REPRESENTATIVE SPECIMENS. Cuba. Pinar pet Rio: Britton et al. 9637 (Ny), Ekman 10845 (us), 18186 (Ny), Shafer 13750 (xy. us), 13882 (Ny. us), Van Hermann 3322 (NY. US Microcycas is scarcely differentiable from Zamia. When this Cuban endemic was first elevated to generic rank by de Candolle (1868), the majority of known species of Zamia were low, West Indian or Mexican plants that were superficially very different in stem, foliage, and cone from the massive Z. calocoma. Later, chromo- some counts of 2 = 8 for species retained in Zamia seemed to contirm their isolation from the n = 13 Microcycas (Sax & Beal, 1934). Embryological features, as well as the greater number of both spermatozoids and archegonia, were unique to Microcycas among all cycads studied and also reinforced separation of the two genera (Chamber- lain, 1935), The only character used to separate the two genera in generic keys to the cycads, however, has been the flattened knob of the male cone scales of Microcycas, compared to the truncate microsporophyll of Zamia (de Candolle, 1868; Schuster, 1932). In recent years, increasing access to arborescent species of Zamia from forests of Central and South America have eliminated many apparent differences between Zamia and Microcycas. Living plants of these formerly little-known species are becoming available for study at Fairchild Tropical Garden and other institutions. The taxonomy of these species is unresolved, but some of them have individual leaf and leaflet or cone characteristics of M. calocoma. None duplicates the combination of characters found in the Cuban cycad, but collectively they possess all of the gross morphological features formerly thought to distinguish Microcycas from Zamia. Zamia, tormerly considered to be chromosomally uniform with 2 = 8, has been 3 formerly shown to have a diversity of chromosome numbers up to the 2 = thought to be confined among cycads to Microcycas (Norstog, 1974, and in press). Recent anatomical work has further shown that Zamia and Microcycas share an unusual branched hair type not found in other cycads, including the related genus Ceratozamia Brongn. (D. W. Stevenson, pers. comm.). The only features now 1980] ECKENWALDER, WEST INDIAN CYCADS 715 remaining that distinguish the two genera are the extreme proliferation of both archegonia and spermatozoids in Microcycas. Embryology of the arborescent zamias of Tropical America is virtually unknown (Norstog, 1977), and a decision to return M. calocoma to the genus Zamia, although likely in the future, should await embryological investigation of Z. chigua Seem. and its allies. Microcycas calocoma was probably isolated in the West Indies by the same events that isolated Z. pumila. It has probably diverged from Central American arborescent zamias Just as Z. pumila diverged from its closest relatives in the Central American Z. furfuracea complex. Zamia pumila L. subsp. pumila Zamia pumila L. Sp. Pl. ed. 2. 2: 1659. 1763. Lecrorype: Palma prunifera humilis non spinosa, insulae hispaniolae, fructui jujubino similis, ossiculo triangulo, J. Commelijn, Horti Med. Amstelod. I: ¢. 5S. 1697 (lectotype, Ams; imprints, Hunt Inst.!, ny!). Palmifolium pumilum (L.) Kuntze, Rev. Gen. PI. 2: 803. 1891. Zamia debilis Aiton, Hortus Kew. 3: 478. 1789, nomen illegit. Palmifolium debile (Aiton) Kuntze, Rev. Gen. Pl. 2: 803. 1891. Zamia humilis Salisb. Prodr. 400. 1796, sphalm. Zamia media var. commeliniana Schuster, Pflanzenr. 99: 155-157. 1932. Zamia integrifolia Aiton, Hortus Kew. 3: 478. 1789. Type: Florida, cult. Kew, Aiton s.n. (holotype, Bm; tracing, Ny!). Palmifolium integrifolium (Aiton) Kuntze, Rev. Gen. Pl. 2: 803. 1891. Zamia floridana var. purshiana Schuster, Pflanzenr. 9 lobe We. Zamia angustifolia Jacq. Ic. Pl. Rar. 3: t. 636. 1791. Type: Bahamas, cult. Hort. Vindob. (lectotype, pl. 636). Palmifolium angustifolium (Jacq.) Kuntze, Rev. Gen. Pl. 2: 803. 1891. Zamia media Jacq. Pl. Rar. Horti Caes. Schoenbr. 3: 77, 78. tt. 397, 398. 1798. Type: West Indies, cult. Hort. Schoenbr. (lectotype, pl. 398). Palmifolium medium (Jacg.) Kuntze, Rev. Gen. Pl. 2: 803. 1891. Zamia media var. jac- quiniana Schuster, Pflanzenr. 99: 157, 158. 1932, nomen illegit. Zamia tenuis Willd. Sp. Pl. ed. 5. 4: 846. 1806. Type: Bahamas, cult. Hort. r illdenow s.n. (holotype, B: isotype (frag.), Ny!). Palmifolium tenue (Willd.) Kuntze, Rev. Gen. Pl. 2: 803. 1891. Zamia media var. tenuis (Willd.) Schuster, Pflanzenr. 99: 158. 1932 Zamia dentata Voigt, Syll. Pl. Nov. 2: 52, 1826, Zamia angustissima Miq. Nieuwe Verh. Eerste KI. Kon. Ned. Inst. Wetensch. Amsterdam, ser. 3. 4: 184. 1851. Type. trop. Amer. [Cuba], cult. Hort. Amstelod., Miguel s.n. (holotype, 1). Zamia angustifolia var. angustissima (Miq.) Regel, Gartenflora 27: 13. 1878. Palmifolium angustissimum (Miq.) Kuntze, Rev. Gen. Pl. 2: 803. 1891. Zamia stricta Mig. Nieuwe Verh. Eerste KI. Kon. Ned. Inst. Wetensch. Amsterdam, ser. 3. 4: 183. 1851. Type: trop. Amer. [Cuba], cult. Hort. Amstelod., Miquel s.n. (holotype, L). Zamia angustifolia var. stricta (Mig.) Regel, Gartenflora 27: 131578: cine strictum (Miq.) Kuntze, Rev. Gen. PI. 2: 803. 1891 Zamia yatesii Mig. Nicuwe Verh. Eerste KI. Kon. Ned. Inst. Wetensch. Amster- dam, ser. 3. 4: 18, 1851. Type: trop. Amer. [Cuba], cult. Hort. Amstelod., Miquel s.n. (holotype, L). Zamia angustifolia var. vatesti (Miq. ) Regel, Garten- _~ — 716 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 61 flora 27: 13. 1878. Palmifolium yatesii (Mig.) Kuntze, Rev. Gen. Pl. 2: Zamia floridana A. DC. Prodr. 16(2): 544. 1868. Lectorype. E. Florida, Fort Brooke, Hulse s.n. (holotype, G-pc; isotype, Ny!). Zamia angustifolia var. flor- idana (A. DC.) Regel, Gartenflora 27: 13. 1878. Palmifolium floridanum (A. DC.) untze, Rev. Gen. Pl. 2: 803. 1891. Zamia multifoliolata A. DC. Prode: 16(2): 545. 1868. Tyrer: Cuba [Oriente], San Juan de Buenavista, Wright 597 (holotype. G-pc: isotypes, . mo!, Ny!, us!). Palmifolium multifoliolatum (A. DC.) Kuntze, Rev. Gen. Pl. 2: 803. 1891. Zamia latifoliolata Prenl. Bull. Soc. Vaud. Sci. Nat. 11: 278, re Type: Republica Dominicana, road to Punto ¢ LAU). Zamia concinna Hort. Booth ex Regel, Gartenflora 27: 9. 1878, nomen inval. (pro sya. Z. pumila L.). Zama cylindrica Hort. Booth ex Regel, Gartenflora 27: 9. 1878, nomen inval. (pro syn. Z. pumila L.). Zamia linifolia Hort. Pawl. ex Regel, Gartenflora 27: 13. 1878, nomen inval. (pro syn. Z. angustifolia Jacq.). Zama linearifolia Linden, Il. Hort. 28: 32. 1881, nomen nudum. Zamia portoricensis Urban, Symb. Antill. 1: 291. 1899. Type. Puerto Rico, Mayagtiez, prope Guanica in sylva ad Ensenada, Sintenis 3782 (holotype, B isotype (frag.), Ny!). Zamia media var. portoricensis (Urban) Schuster, Pflanzenr, OO 158. 1932; Zamia allison-armourti Millsp. Publ. Field Columbian Mus., Bot. Ser. 2: 23. 1900. Tyre: Republica Dominicana, Distrito Nacional, near river flowing into Santo Domingo, Millspaugh 817 (holotype, &!; isotype, nv!). Zama erosa Cook & Collins, Contr. U.S. Natl. Herb. 8: 267. 1903. Lecrorypr (here designated): Puerto Rico, Ponce, Coamo Springs, Cook s.n. (lectotype, Ny!; 1solectotype, Ny!). Zamia lucayana Britton, Bull. New York Bot. Gard. 5: 311. 1907. Type: Baha- mas, Long Island, Clarence Town and vicinity, Britton & Millspaugh 6271 (holo- type, ny!; isotypes, F!, Gu!, Ny!). Zamia umbrosa Small, J. New York Bot. Gard. 22: 136. 1921. Lecrorypr (here designated): Florida, Volusia, Daytona Beach, Small 8679 (lectotype, Ny!; iso- lectotypes, DUKE!, FLAS!, GH!). Zamia silvicola Small, J. New York Bot. Gard. 27: 128. figs. 1, 2. 1926. Type: e la Torrecilla, Prenleloup s.n. (holotype, Florida, Citrus, **Spanish Mound”’ near Crystal River, Small, Small, & DeWinkeler 10060 (holotype, xv!). Zamia floridana var. purshiana f. silvicola (Small) Zamia brachyphylla Hort. ex Schuster, Pflanzenr. 99: 159. 1932, nomen inval. (pro syn. Z. angustifolia Jacq.). Zamia linearis Mig. ex Schuster, Pflanzenr. 99: 159. 1932, nomen inval. (pro syn. Z, angustifolia var. yatesti (Mig.) Regel). Zamia media var. gutierrezii {. calcicola Schuster, Pflanzenr. 99: 155. 1932. Type Cuba, Pinar del Rio, Banos de San Vicente, Britton, Britton, & Gager 7392 (holotype, B. isotypes, Gu!, Nv!). Zamia calcicola Britton ex Schuster, Pflanzenr. 1980] ECKENWALDER, WEST INDIAN CYCADS Te 99: 155. 1932, nomen inval. (pro syn. Z. media var. gutierrezit f. calcicola Schuster). Zamia media var. jacquiniana f. brevipinnata Schuster, Pflanzen. 99: 158. 1932. Lectotype (here designated): Cuba, Oriente, Holguin to Myabe, Shafer 1406 (lectotype, B: isolectotypes, F!, NY!). Zamia subcoriacea Wendl. ex Schuster, Pflanzenr. 99: 155. 1932, nomen inval. (pro syn. Z. media var. commeliniana Schuster). Zamia verbruggiana Hort. ex Schuster, Pflanzenr. 99: 159. 1932, nomen inval. (pro syn. Z. angustifolia var. yatesii (Miq.) Regel). Zamia guggenheimiana Carabia, Caribbean Forest. 2: 89. 1941. Type: Cuba, Pilon. Ensenada de Mora, Manzanillo, Carabia 1403a (holotype, Ny!; isotype, F!). ILLUSTRATIONS: Small, J. New York Bot. Gard. 27: figs. 7, 2. 1926; Victorin & Leon, Itin. Bot. Cuba, figs. 33-37, 283, 254. 1942. Cycads with short, preponderantly underground stems, 2.5-6 cm. in diameter, these often highly branched in apparent dichotomies: crown formed by (1 to) 4 to8 (to 12) pinnate leaves; leaves (I—)3—7(—16) dm. long, emerging in annual or irregular flushes, covered by transient, rusty, filamentous pubescence; the leaflets (2 to) 10 to 30 (to 45) pairs, (4-)8-20(-35) cm. long, linear, lanceolate, oblong, or oblanceolate, horizontally spreading to secund, caducous after | or 2 years, only shortly before rachis, rachis abscising with leaf bases to leave smooth stem: male and female cones annual, long peduncled, with peltate, hexagonal scales in vertical files, the external face of scales rusty or dark purple pubescent, faceted, with flat center, the male cones | to 8 (to 63), emerging from among inner leaves of crown, 4—6.5(-13) cm. long, with 9 to 15 scales in each of 6 to 11 rows, the female cones solitary, terminal, more massive than males, 6—-10(-20) cm. long, with 3 to 5 scales in each of 5 or 6 rows; outer fleshy layer of ovular seed coat pink, turning bright orange or red at maturity, seed 1-2.5 cm. long. n = 8 (Sax & Beal, 1934) Distripution. Found sporadically throughout the native range of cycads in the West Indian region, but absent from Haiti and many of the Bahamas and rare in western Cuba, where it is replaced by subsp. pygmaea (see Map 1). It occurs in a variety of habitats ranging from open sea bluffs and sand dunes, through pine and oak savan- nas, to closed-canopy oak hammocks and tropical forests. It is most common near sea level on limestone and sand but also occurs on other substrates. Morphological features of leaflet width, attitude, and thickness are associated with habitat differ- ences, apparently affected by both genctic and environmental components (Eckenwalder, unpubl. data). The plants are frequently cultivated for ornament in their native regions and were formerly the basis of a starch-extraction industry (Small, 1921). Habitat destruction is the major current threat to them, especially in Florida, where residential development is leading to a drastic decline in number. The subspecies 1s listed as threatened in Florida (Ward & Pritchard, 1979). REPRESENTATIVE SPECIMENS. Florida. Avacuua: Small S450 (Mo. NCU, NY), D. & S. Ward 1838 (PLAS, FSU, NCU. US, USF), Weber s.n. (F Ny. US). BROWARD: Moldenke 504 718 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 (DUKE, MO, NY), Sandler s.n. (rau), Seibert 1168 (a. Mo). Crrrus: ples 4724 (FLAS), Lakela 26820 (usr), Small 10387 (xy). Dave: Beckner 1775 (ras. FSU. USE), Curtiss 2676 (A. GA. GH. MO, NY, US, USF), eas 10237 (MO. UC. US), rae "9265 ( (F, GH, MO.NY, TEX, US). Dixie: Harbison ay (aA. NcU, US). Gitcurist: Ward & Ford 3588 (LAs, NCU, USF). Grapes: Ward et al. 2423 (Frias. GH. USE). HERNANDO: Small et al. 10067 (DUKE, MO, NCU, NY). HittsporouGH: Blanton 6927 (b. Mo, us). Levy: Evers s.n. (plas), Godfrey Redfearn 52834 (puKE, Psu. GH. NY. USF), Miller 339 (us), Murrill s.n. (Mo). a ather M-172 (rLas. GA), Norstog s.n. (NY. USF), Schuster A-258-a (DUKE, FSU). ST. ae Smuth 522 (¢ GH. us), Ward & Moore 2362 (PLAs. Psu. NCU. USF). SARASOTA: Simpson 394 (p.GH. NY. US). SEMINOLE: Beckner 884 (FLAS. FSU. GA). TAyLor: Godfrey & Redfearn 52817 (DUKE, FSU, GH. NCU. NY, Uc. USF). Wotusia: Brown s.n. (Ncsc), Curtiss 2676b (¥. GA, MO. NY, US. USF), Read 1046 (a. era), Small 8679 (DUKE. PLAS. GH. NY). Georgia. Campen: Proctor s.n. (GA). GLyn: Duncan 23659 (Ga). Bahamas. Great ABaco: Brace 1746 (¢. Ny), Correll et al. 42577 (rrc). ANbrRos: Brace 6990 (fF. NY), Hill 3183 (F, FTG, NY). ELeEUTHERA: Britton & Millspaugh 5418 (re. Ny. uc. us), Correll & Hill 45240 (F. FrG. LL. MO, NY). GRAND BAHAMA: Brace 3693 (Fr. GH. MO. Ny. US), Britton & el Noe 2576 (F, NY, US). LonG Istanp: Hill 828 (r. GH. MO, NY, US), 2334 (FTG.NY). Ww ProvibENce: Britton e Brace 704 (vr ny). Cuba. Cee. Shafer 679 (PNY. “ 793 (PNY. US), 2659 (PONY. US). [sta DE Pinos: Carabia 1190 (¥). . VILLAS: Britton & Wilson 5443 (F. NY), on zales 381 (A), Leon & Carabia 19074 ( a FELASA ny. us). Matanzas: Carabia 1992 (re). Oriente: Britton & Cowell ee Ny, US), Carabia 3523 (r), Hioram 4924 tea GH, NY), Leon o Alain 18934 (pr, Pollard & Palmer 141 (e.Gu. MO. NY. US), 382 (PF. GH. MO. NY. US), Shafer 1406 (F, NY. US), Victorin 60037 (GH. Mo, Ny. US), Wright 1463 (F. Gu. Mo). PINAR DEL Rio: Baker 28944 ny), Carabia 3189 (r), Van Hermann 7153 (v. Gu). Jamaica. St. ANN: Clift s.n. (Frc), Howard & Proctor 15114 | Trecawny: Proctor 31556 (Ff. GH. LL). WESTMORELAND: Britton & Hollick 2070 hae: Clift s.n. (etc), Harris 10229 (ny. us Cayman Islands. Granp Cayman: Kimbe s.n. (TG) . Osment sn. (FTG). Dominican Republic. Avtacracia: R. & E. Howard 9774 (Gu. Ny. us), Liogier 12341 (Gu. Ny. us). Distrito Nacionac: Allard 13572 (us), Ekman 5800 (Gu. ye Rose et al. 3783 (Ny.us). Romana: Crafts 115 (bay), Miller 1007 (us), Taylor 352 (vp. ny), SAMANA: Abbott 1158 (us), Liogier 14383 (ny. us). SAN PEDRO bE Macoris ae 69] a ; Howard 9494 (GH), Rose et al. 3707 (vp. NY. us). Seipo: Jiménez 5052 (ny), Miller 1014 (us), Taylor 1/2 (xy). Puerto Rico. AGuaputa N. & E. Britton 9282 . ). ARECIBO: Blomquist 11984 (Dav. DUKE. MO. TEX), Stimson 3816 (DUKE. MICH, MO. NY), Underwood Griggs S47 (Pp. MicH. NY, Us). Mayacuez: Britton & Shafer 1832 (F. Mo. NY. er a sn. (F), Sintents 763 (Gu.us), Stimson 3272 (puKE). Ponce: Britton et i, is), Britton 8S57 (¥. Gu. Mo. Ny. us), Heller s.n. (us). SAN Juan: Britton etal. 2855 ea Ny. us), Goll 857 (Ny. us), A. & G. Heller 397 (Ny. us). _~ — > — — — — Zamia pumila is distinguished from other species of the genus by its low stature, underground stems, unarmed petioles, sparsely toothed, often rounded leaflets, small, long-peduncled cones with rusty or purple pubescence, and low-faceted cone scales. Although all of these features are found individually in other species of the genus, the combination is unique to the West Indian plants. This species is most closely related to the Mexican and Central American Z. furfuracea, which differs in being more robust and in having larger, spiny-petioled leaves, tougher, more serrate leaflets, longer-peduncled cones with tawny pubescence, and smaller red seeds. As noted above, Zamia pumila subsp. pumila is very variable in leaflet form both within and between West Indian subregions. It was initially expected that varieties 1980] ECKENWALDER, WEST INDIAN CYCADS 719 would be recognized within the subspecies. This would give some taxonomic recog- nition to the diversity, which is particularly troublesome in an area such as the Bahamas, where individual islands have distinctive, relatively homogeneous popula- tions (Ficures 7, 8; Eckenwalder, in press). This expectation was abandoned because of lack of morphological discontinuities in specimens of the subspecies from throughout its range. No coherent system of varieties could be devised that was not as arbitrary and as typological (in the perjorative sense) as the earlier systems of species here replaced with the present broadened concept. Local botanists are thus left with the somewhat unsatisfactory circumstance of not being able to give taxo- nomic recognition to distinctive variants that occur in their region and that intergrade only in some less familiar ground. The capacity for long-distance dispersal by seawater has evidently aggravated this situation (Eckenwalder, 1980). Zamia pumila L. subsp. pygmaea (Sims) Eckenwalder, stat. et comb. nov. Basionym Zama pygmaed Sims, Bot. Mag. 43: ¢. 1/747. 1815. Type: W. Indies [Cuba], cult. Hort. Loddiges, Saliebar Sa, i BM!). Palmifolium pygmaeum (Sims) Kuntze, Rev. Gen. Pl. 2: 803. I891. Zamia kickxii Mig. Monogr. se 71. t. 8, fig. A. 1842. Type: Cuba, cult. Hort. Ghent, Kickx s.n. (holotype, L). Palmifolium kic keii (Miq.) Kuntze, Rev. Gen. Pl. 2: 803. 1891. Zamia pygmaea var. kickxii (Mig.) Schuster, Pflanzenr. 99: 52,1932. Zamia ottonis Mig. Linnaea 17: 740. 1843. Type: as eye San Antonio Fundador de Caminar, Otto 36 (holotype, L: isotypes ¥ Crags), Rane ottonis (Miq.) 3 Kuntze, Rev. Gen. Pl. 2: 803. 1891. Pamia pyemaea var. ofttonis (Mig.) Schuster, Pflanzenr. 99: 153. 1932 a pugmaed var. wrightil A. DC. Prodr: 16(2): 543. 1868. Type: Cuba, abanas Chirigati, Wright 3192 (holotype, G-pe: = isotypes. BM!, GH!, Mo!). Zamia chamber lainii Schuster, Pflanzenr. 99: 153. Zamia silicea Britton, Bull. Torrey Bot. Cle . a 462. 1916. Type: Cuba, Isla de Pinos, vicinity of Los Indios, Britton, Britton, & Wilson 14166 (holotype, ny!; isotypes, F!, GH!, us!). Zamia media var. eae f. silicea (Britton) Schuster, WN Pfil cape potundifolin ‘Hort. ex e Sehasien acne 99: 152. 1932, nomen inval. (pro _Z. pygmaea var. kickxti (Miq.) Schu ILLustRATIONS: Victorin & Le6n, Itin. Bot. Cuba. figs. //3, 1/4, 173, 181, 189. Cycads differing from subsp. pumila primarily in smaller stature, with few differ- ences of proportion; stems 0.5—2 cm. in diameter; leaves (0.5—)I—4(—7) dm. long; leaflets 1-7(—10) cm. long, narrowly oblong to obovate, typically shorter and more numerous with respect to total leaf length than in subsp. pumila, much thinner in texture and more fragile; male cones 1.5—5 cm. long; female cones 2.5—7 cm. long; seeds 1-1.5 cm. long. n = 8 (Norstog, in press). DistripuTion. Dwarf cycads are endemic to western Cuba and Isla de Pinos (Map 1) and intergrade with subsp. pumila eastward across the main island (Ficure 6). These cycads typically occur in open limestone and serpentine communities, but the most robust individuals grow in moist forests of ravines on isolated mogotes (limestone 720 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 hills) from Pinar del Rio Province to Las Villas Province. Here they may be replaced subsp. pumila. They are uncommon in cultivation and are apparently relatively little affected by human activities. REPRESENTATIVE SPECIMENS. Cuba. Camacuey: Britton 2386 (ny), Britton et al. 13166 (Ny), Carabia 3405 ("). HaBana: Britton et al. 638 (b. NY), 6267 (ny), Carabia 1569 (r). Ista dE Pinos: Britton & Wilson 14877 (xy.us), Carabia 983 (vr), Killip 42674 (Gu, us). Las Vittas: Britton et al. 10233 (xy. us), Combs 647 (Gu), Howard et al. 290 (A. LL. MICH. NY. UC), Luna S85 (envy). Matanzas: Britton & Wilson 420 (ny), Carabia 1591 (re), Van Hermann 913 (ep Ny). Pinar pet Rio: Britton et al. 9664 (Ny. us), Shafer & Leon 13713 (e. Gu. NY), Van Hermann 2052 (¥. Ny.uC). Zamia pumila subsp. pygmaea is the smallest living ceycad. Although the largest individuals overlap subsp. pumila in total leaf length, the leaflets are proportionately shorter and broader than on corresponding plants of the more widespread subspecies, the stems are more slender, and the male and female cones and seeds are somewhat smaller. Chamberlain apparently crossed the two subspecies and obtained hybrids, some of which reached Fairchild Tropical Garden during the dispersion of his living collection after his death. These uncertainly documented plants appear intermediate, normal, and fertile. In all respects, plants of subsp. pygmaea resemble miniature versions of subsp. pumila. The differences are preserved after twenty years of cultivation at Fairchild Tropical Garden, so they are presumably genetically based. The evident similarity of the dwarf plants to subsp. pula and their intergradation with the more robust plants argue against their recognition at a higher taxonomic rank INTRODUCED CYCADS Zamia furfuracea Aiton, Hortus Kew. 3: 477. 1789. Zamia Lhe Sauvalle, Anales Acad. Ci. Méd. Habana 5: 54. 1868. Type. Ss ui del Rio, cult. Hort. Blain, Rangel (holotype, tac: isotypes, F!, NY!). ud mec dias var. gutierrezil (Sauvalle) Schuster, oe snr. 99: 154. 1932. a pumila Miller, Gard. Dict. ed. 8. no. 9. 1768. ECTOTYPE Palma fructu clavato polypreno, C. J. Trew, Pl. Selec. ft. 26. 1752 As noted both by Carabia (1941) and in correspondence associated with herbar- ium specimens at GH. this Mexican and Central American species has been cultivated for more than a century in the West Indian region. Plants growing in what was once a botanic garden in Cuba were renamed by Sauvalle and were considered as a West Indian endemic. Schuster (1932) incorrectly believed that these represented the widespread, particularly robust phase of Zamia pumila subsp. pumila (called Z. latifoliolata Prenl. by many authors) and made a combination appropriate to that view. Zamia furfuracea is contrasted with Z. piamila in the discussion of subsp. pumila. The two species may hybridize when brought together, and putative hybrids of this parentage can be found at Fairchild Tropical Garden. Cycas circinalis L. Sp. Pl. 2: 1188. 1753. 1980] ECKENWALDER, WEST INDIAN CYCADS 721 Cycas revoluta Thunb. Fl. Japon. 229. 1784. Cycas rumphii Miq. Bull. Sci. Phys. Nat. Neéerl. 2: 45, 1839. The familiar Asiatic sago palms are widely cultivated in the West Indies, as in all tropical and subtropical regions. They are arborescent cycads readily distinguished from the native plants by their leaflets with a midrib, and by their large seeds borne on elongate sporophylls not aggregated into compact cones but appearing in flushes alternating with flushes of foliage leaves. These most-isolated cycads are often retained as the sole extant genus of Cycadaceae, while other genera with distinct, pedunculate, megasporangiate strobili (Johnson, 1959) are segregated to the Zamiaceae (and Stangeriaceac). ACKNOWLEDGMENTS | thank the curators and directors of institutions whose living and preserved specimens I examined. | also thank my former colleagues at Fairchild Tropical Garden for numerous and varied contributions. A National Science Foundation grant to D. S. Correll supported some field work in the Bahamas. Financial support from Fairchild Tropical Garden contributed to extensive field work in Florida. Jack, Spencer, Susan, and others cracked the whip. LITERATURE CITED Arron. W. 1789. Hortus Kewensis. Vol. 3. 547 pp. G. Nicol, London. Ayensu. E. S., & R. A. DrFiuipps. 1978. Endangered and threatened plants of the United States. xv + 403 pp. Smithsonian Institution, Washington, D. C. Britton. N. L.. & C. F. Muuspaucu. 1920. The Bahama flora. vill + 695 pp. Privately published, New York. & Wiison. 1926. Cycadales. Sci. Survey Porto Rico Virgin Is. 6: 329, 330. CaLpwe.t,O. W. 1907. Microcycas calocoma. Bot. Gaz. (Crawfordsville) 44: 118-141. tt. 10-13 & C. F. Baker. 1907. The identity of Microcycas calocoma, Bot. Gaz. (Crawfordsville) 43: 330-335. CANDOLLE, A. bE. 1868. Cycadaceae. Prodr. 16(2): 522-548. Carapia. J. P. 1941. Contribuciones al estudio de la flora Cubana. Gymnospermac. Caribbean Forest. 2: 83-99. Cuamperain, C. J. 1935. Gymnosperms, structure and evolution, xi + 484 pp. University of Chicago Press, Chicago. _n. d. A taxonomic monograph of the cycads. Unpubl. ms. University of California, Los Angeles. Ecxenwatper. J. E. 1980. Divergent dispersal methods in the West Indian cycad, Zamia pumila L. Biotropica 12: 79, 80. n press. Zamiaceae. /n D. S. Correll, ed., Flora of the Bahamas Archipelago. Stanford University Press, Stanford. Houck, A. 1932. Descriptions of new species of Tertiary cycads, with a review of those previously recorded. Bull. Torrey Bot. Club 59: 169-189. Howarp. R. A. 1977. Conservation and the endangered species of plants in the fae JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Caribbean islands. Pp. 105-114 in G. T. Prance & T. S. Ettas. eds., Extinction is forever. vi + 437 pp. New York Botanical Garden, New York Jacquin, N. 1791. Icones plantarum rariorum. Vol. 3. 24 pp. + 194 it, Wappler, Vienna. . 1798. Plantarum rariorum horti caesarei schoenbrunnensis descriptiones et cones. Vol. 3. 80 pp. + /50 tt. Wappler, Vienna. ie A. S. 1959. The families of cycads and the Zamiaceae of Australia. Proc. Linn. Soc. New South Wales 84: 64-117. Leon. J. S. S. 1946. Flora de Cuba. Vol. 1. 441 pp. Contr. Ocas. Mus. Hist. Nat. Colegio **De La Salle.”’ No. 8. Cultural, S. A., Havana. Linnazus. C. 1763. Species plantarum. ed. 2. Vol. 2. Pp. 785-1684. L. Salvius, Stockholm. . 1764. Genera plantarum. ed. 6. 580 pp. L. Salvius, Stockholm. Liocier. A. H. 1969. Flora de Cuba, suplemento. 150 pp. Sucre, Caracas. Mitter, P. 1768. Gardener's dictionary. ed. 8. Rivington, London NorstoG, K. 1974. Zamiaceae. Jn A. Love. ed., [OPB chromosome number reports, 45. Taxon 23: 622. . 1977. The spermatozoid of Zamia chigua Seem. Bot. Gaz. (Crawfordsville) 38: 409-412. ———.. In press. Chromosome numbers in eee (Cycadales). Caryologia. Reap, R. W. 1967. Taxonomy of the cycads of the West Indies and Florida. Pp. IH-1-ll-6 in Fifth Conference on Cycad eee Department of Biology, University of Miami, Miami. Sax. K., ae - Beat. 1934. Chromosomes of the Cycadales. J. Arnold Arbor. 15: 255-258, tt. 107, 108 Scuuster, J. oe Cycadaceae. Pflanzenr. 99: 1-168. Sims, J. 1815. Zamia pygmaea. Bot. Mag. 43: t. 174/. SMALL. J. K. 1921. Seminole bread—the conti. J. New York Bot. Gard. 22: 121-137. 1926. Cycads. Ibid. 27: 121-129 - 1933, Manual of the southeastern flora. xxii + 1554 pp. Privately published, New York. Smith. L. B. 1961. The application of Zamia pumila L. Phytologia 7: 417. Vicrorin, F. M. & J. S. S. Leon, 1942. Itinéraires botaniques dans Vile de Cuba (premicre série). Contr. Inst. Bot. Univ. Montréal 41: i—viii, Warp. D. B., & P. C. Pritcuarp, eds. 1979. Rare and endangered biota of Florida. Vol. 5. Plants. xxix + 175 pp. University Presses of Florida, Gainesville. Wurre, F. 1962. Geographic variation and speciation in Africa with particular refer- ence to Diospyros. Publ. Syst. Assoc. 4: 71-10: Wuitmore, T. C. 1975. Natural variation and its taxonotuie oe within tropical tree species as seen in the Far East. Linn. Soc. Symp. Ser. 2: 25-34. DEPARTMENT OF BOTANY UNiversitry oF Toronto Toronto. Ontario M5S 1A] CANADA 1980] PAGE, DICOTYLEDONOUS WOOD, II 723 DICOTYLEDONOUS WOOD FROM THE UPPER CRETACEOUS OF CENTRAL CALIFORNIA, I VIRGINIA M. PAGE THE PRESENT REPORT is the second in a series describing the results of an investigation of an assemblage of woods from the Upper Cretaceous of central California. As reported in the first paper (Page, 1979), most of the fossil specimens were collected from three localities, considered to be Maastrichtian in age, situated in the foothills of the Mt. Diablo Range, west of the town of Patterson in Stanislaus County: locality 1 (Del Puerto Creek); locality 2a (Black Gulch); and locality 2b (an unnamed site near locality 2a). Pertinent information about these sites was presented in the earlier paper. Chmura (1973) places locality 1 stratigraphically above locality 2a; localities 2a and 2b are essentially time equivalent (Stein, pers. comm. ). In the present report, two additional localities are represented: — Locality 3. Cache Creek; R4W, T12N, Sec. 2, SEY, Morgan Valley 15’ quadrangle. Locality 4. % mile north of Pigeon Point, San Mateo County. Locality 3 is an exposure of the Sites Formation along Cache Creek, in Yolo County on the western side of the Sacramento Valley. Douglas (1969) and Passagno (1974) consider the Sites Formation as Coniacian in age, but correlations based on Goudkoff’s (1945) foraminiferal zones place the locality in the G-2 zone, or middle to late Turonian (Popenoe ef al., 1960). Locality 4 lies west of the Coast Ranges in an outcrop of the Pigeon Point Formation along the San Mateo County coast one-half mile north of Pigeon Point. Although numerous faults hamper stratigraphic work in this area, Hall er al. (1959) and Wentworth (1960 and pers. comm.) agree that the age of the formation is Campanian. All thin sections described here are deposited in the California Academy of Sciences Geological Collections in San Francisco (cas), and reference to these specimens is by accession number. The term fe is used in reference to the vessel as it appears in transverse section. The term “‘fiber’’ is used in its generic sense as defined in the Multilingual Glossary of Terms Used in Wood Anatomy (1964). Omission of an important feature in a description indicates that it was not possible to observe it due to poor preservation of the specimen. Growth rings are mentioned only if they are present, and pore frequency has been omitted because distortion of the grain in most specimens did not allow statistically significant measurements. In a general way the relative frequency can be inferred from the illustrations. The specimens are classified according to a system presented in the form of a synoptic key in the first paper of this series. The system is artificial and is merely a practical way to handle the structural patterns found among the fossil woods © President and Fellows of Harvard College, 1980 Journal of the Arnold Arboretum 61: 723-748. October, 1980. 724 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 under study that can not, for reasons discussed at length, be assigned to modern genera or families. The characters used in the classification are those most likely to be preserved: hence, groupings are defined according to distribution of pores and of axial parenchyma, and to type of perforation plate and of intervessel pitting. With reference to distribution of axial parenchyma, the intent was to separate specimens that have paratracheal parenchyma from those that do not. It is, however, necessary to reserve a category for those in which no axial parenchyma could be observed. The notion of frequency was inadvertently interjected into the classification by the desig- nation of an alternative (“‘sparse or absent’’) to the categories **parenchyma apotra- cheal only’? or *‘paratracheal parenchyma present.”’ For the sake of consistency, the word *‘sparse’’ should be removed from the synoptic key. Axial parenchyma cells, whatever their frequency, either are or are not associated with vessels. If the paren- chyma cells are sparse and are consistently associated with vessels, they are con- sidered paratracheal; if they are sparse and are not consistently associated with ves- sels, they are considered apotracheal. It should be mentioned that axial parenchyma cells are often difficult to recognize in sectioned (particularly in transversely sec- tioned) material. In fossilized woods the problems are more severe: such cells are often obscured by grain distortion; in transverse section they can be confused with fibers in which the secondary walls have been destroyed. Furthermore, mineral deposits within a cell may simulate the secondary thickening of fibers. Because of the difficulties in recognizing axial parenchyma in transverse section, it is important not to rely solely on such sections for determining its presence SYSTEMATIC DESCRIPTIONS The first four specimens—casc 60133, casc 60420, casc 60421, and casc 60422—are classified in Group HI. This group is characterized by mostly solitary pores and by perforation plates with fewer than 50 bars. Several specimens belonging to this group were described earlier (Page, 1979). casc 60133 represents section B2 (parenchyma apotracheal, intervessel pits opposite), and casc 60420, casc 60421, and casc 60422 represent section C (paratracheal parenchyma present). casc 60133 Figures I-4. Materia. Branch 2.5 cm. in diameter. Secondary xylem. Diffuse porous, the pores solitary, numerous, average radial diameter 86 wm. (70-99), average tangential diameter 96 jm. (86-116). Perforation plates scalariform, slightly oblique, with less than 10 widely spaced bars. Intervessel pits small, round, opposite; pits to ray cells oval to almost scalariform: pits to parenchyma oval: average vessel element length 681 wm. (S80—812). Parenchyma Figures I-11. 1-4, casc 60133: |, transverse section (note growth ring), x 53; 2 iransverse section, X 75; 3, tangential section, < 75; 4, radial section show vie enlarged ray cell, X 75. 5—8, casc 60420: 5, perforation plate, X 80; 6, transverse section, X 40; 7, tangential Saas es broad rays a 7 intervessel pits), 70; 8, transverse section, x —11, casc 60422, x 75: 9, 10, transverse sections Ll ne section. showing pore distribution variants; 125 PAGE, DICOTYLEDONOUS WOOD, II 1980] ts Gs enadeenad A Ste ata — “e, Sh ae Te 726 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 apotracheal diffuse. Rays 2 to 6 cells wide, numerous, the cells variable in size and shape, some much enlarged; uniseriates | to 6 cells high, heterocellular. Fiber pits on all walls small, bordered. Pith. Large, | cm. in diameter; primary xylem points numerous, separated by multiseriate rays. Locauity. Del Puerto Creek, locality |. Poor preservation obscured intervessel pits in all but a few vessels. Because of distortion, the uniseriate rays could not be counted; however, they do not seem to be numerous. Multiseriate rays are in various stages of dissection, and the lower rays have only one marginal row of square cells. Many are joined vertically by their uniseriate margins. The enlarged ray cells appear to have been crystalliferous (FiGure 4). The pith is filled with convoluted, thick-walled tubules resembling hyphae of certain fungal pathogens. Type [[B2 modern woods most similar to casc 60133 include certain members of the Ericaceae, Escalloniaceae, Cunoniaceac, and Myricaceae. Solitary crystals have been reported in the Myricaceac (Metcalfe & Chalk, 1950). casc 60420 Ficures 5-8. MarTeRIAL. Fragment of mature secondary xylem. Secondary xylem. Pores solitary, angular, some in radial pairs, average radial diameter 120 pm. (87-145), average tangential diameter 102 jum. (81-127). Perforation plates scalariform, 10 to 15 bars. Intervessel pits numerous, small to minute, opposite; pits to parenchyma similar, some elongated horizontally; average vessel element length 725 wm. (580-880); tyloses common. Parenchyma sparse, apotracheal diffuse and paratracheal as uniseriate sheath around some vessels. Multiseriate rays frequent, up to 15 cells wide and 2.7 mm. high, some dissected, the cells varying greatly in shape and size in no particular pattern, those in center mostly procumbent;: uniseriates infrequent, 3 to 5 cells high. Fibers thick walled. Most elements containing dark, amorphous inclusions. Locatiry, Cache Creek, locality 3. The amorphous inclusions and the tyloses obscure the end walls of many vessels: therefore, it is not certain whether all end walls are multipertorate. Similar wood patterns can be observed in genera of the Araliaceae and Dilleniaceae. According to Dickison (1967), apotracheal parenchyma predominates in t Dilleniaceae, and paratracheal parenchyma is of a diffuse nature. Fibers are septate in most species of the Araliaceae. Although fiber septae were not observed in the fossil, their presence could be obscured by the amorphous inclusions in the very narrow cell lumens. casG 60421 Figures 12-14. MareriaL. Stem or root with what appears to be only secondary xylem, in one radial section with some evidence of primary xylem. Grain much distorted. Speci- men probably from fork of a branch. PAGE, DICOTYLEDONOUS WOOD, II Pe | 1980] saan a pp mo ee La a. Ce Ss Ea a 30980 0 ES 5h oh oh on veel OE Wa =| a = Ge ee Toads oi lento! Tar n ef tkpP= Ow al =D) wt Wt mx x «kK & x ells, transverse section, transverse § tr ection, ction erse se , transverse section ansv 12 Ly. 9 | and axial parenchyma c 60421: f ray » CASG 20 ; 6042 . CASG 60424: 2-20. 12-14 wall pittin 5, 16 Wee: 19, 20 Figures | section showing , xX 70. section 728 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Secondary xylem. Pores solitary, angular, average radial diameter 139 wm. (102-203), average tangential diameter 171 fm. (87-232). Perforation plates scalariform, 40 to 50 bars. Intervessel pitting not observed; pits to ray cells and axial parenchyma scalariform or large and oval; average vessel element length 1413 wm. (1160-1624); tyloses common. Parenchyma paratracheal, apotracheal diffuse, and in uniseriate tangential lines. Rays up to I5 cells wide, averaging 1.7 mm. (12.7) high, heterocellular, the cells of central portion of large rays long procumbent, others short procumbent or square; uniseriate margins | to 5 cells high; sheath cells present: uniseriates numerous, up to 20 cells high, cells upright: rhomboidal crystals com- mon. Fibers thick walled, the pits large, bordered, apertures small. Locauiry. Del Puerto Creek, locality 1. Similar wood patterns occur in Dillenia (Dilleniaceac), Saurauia (Saurautaceae), and Kaunbachia (Cunoniaceae). CASG 60422 Ficures 9-11. Mareriac. Piece of secondary wood. Mostly compressed: grain distorted and cells crushed. Secondary xylem. Observable pores solitary, differing in size and frequency in various areas of undamaged section where not compressed (difference possibly represents either various portions of growth ring or early- and later-formed wood), average radial diameter 86 um. (S8—-104), average tangential diameter 78 zm. (58-93). Perforation plates scalariform, with ca. 15 coarse. widely spaced bars. Intervessel pits appear to be opposite, pits to ray cells and parenchyma scalariform: vessel elements ca. 770 wm. long: tyloses thick walled. Parenchyma appears to be mostly paratracheal. Rays 2 or 3 cells wide, the cells large, occasional larger ones may have been crystalliferous; sheath cells occasional; uniseriates appear to be infrequent. Fibers very thick walled, pit apertures small and slitlike. Phloem. Patches of fibers alternating radially with enlarged thin-walled cells and crushed cells. Fibers small in diameter, thick walled. Large, thin-walled cells becoming stone cells in outer layers. Cells of rays not preserved, but no perceptible broadening of ray tissue in outer layers. Locauiry. Del Puerto Creek, locality 1. The next two specimens, casG 60423 and casc 60424, are classified in Group VA. Members of this group have scalariform perforation plates with fewer than 50 bars, both solitary and multiple pores, and little or no parenchyma. Examples of modern woods of this type with opposite and/or scalariform intervessel pits and with vessel clements that average more than 900 wm. in length include Meliosma (Sabiaceae), Paypayrola and Leonia (Violaceae), and several members of the Flacourtiaceae and Araliaceae. Those with opposite and/or scalariform intervessel pits and with vessel elements averaging less than 900 wm. in length include certain genera of the Staphyleaceae, Dichapetalaceae, Rhizophoraceae. and Araliaceae. casG 60423 Figures 15, 16. 1980] PAGE, DICOTYLEDONOUS WOOD, II 729 MateriaL. Small branch 4.5 cm. in diameter. Grain distorted. Secondary xylem. Diffuse porous, pores solitary and in clusters and radial chains of 3 to 5, average radial diameter 98 xm. (58-128), average tangential diameter 81 jum. (46-104). Perforation plates scalariform, up to [8 coarse bars. Intervessel pits opposite, at least in part; pits to axial parenchyma large, oval: average vessel element length ca. 580 jum.; tyloses abundant. Parenchyma sparse, paratracheal. Rays 3 to 5 cells wide, over | mm. high, the cells large, mostly square or short procumbent; uniseriate margins | or 2 cells high, cells upright; uniseriates not numerous. Fibers septate in part. Pith. Small, round; cells rounded, thin walled. Locaury. Black Gulch, locality 2a. The secondary xylem pattern is similar to some observed in the Araliaceae and the Rhizophoraceae. Fiber septae are difficult to find in the fossil, but some can be observed; since there is no evidence of fungal hyphae, it is reasonably certain that they are truly septae. The nature of the perforation plates is also difficult to discern in many vessels: many are unquestionably multiperforate, but it is possible that simple ones are also present. This specimen is somewhat similar to casc 60420, except that in casc 60423 the rays are much narrower, no apotracheal parenchyma was observed, and pore multiples are present. casG 60424 Ficures 19, 20. MateriaL. Small portion of mature secondary xylem. Secondary xylem. Pores solitary and in radial rows of 3 to 5, average radial diameter 97 wm. (70-139), average tangential diameter 100 pm, (70-145). Perforation plates scalariform, bars averaging 35 (28 to 46). Intervessel pits small, opposite to laterally elongated; average vessel element length 1310 jum. (1160-1740). Parenchyma absent. Rays heterocellular, the cells mostly square, short procumbent, and upright, with some low procumbents; uniseriate margins | to 10 cells high, cells upright: many multiseriates connected vertically by margins; sheath cells occasional: uniseriates up to 14 cells high, cells upright. Fibers septate. Locatity. Black Gulch, locality 2a. Type VA woods, characterized by long vessel elements, opposite intervessel pits, and high heterocellular rays, occur in some of the more primitive members of the Violales. End walls in the vessels of the fossil have more bars than is characteris- tic of even the most primitive members of the Flacourtiaceae, and the intervessel pits are much smaller. There are numerous bars in the perforation plates in woods of the Violaceae, but intervessel pitting is scalariform or transitional. The family Lacistemaceae has small, opposite intervessel pits, but parenchyma in the four species examined ts — abundant. Pollen attributable to the Lacistemaceae has been described from locality 1 (Chmura, 730 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 The next two specimens, casc 60425 and casc 60426, are classified in Group VB, which comprises woods with scalariform perforation plates with fewer than 50 bars, and pores both solitary and in multiples. Woods of this type occur in at least 23 families. CASG 60425 Figures 17, 18. Materia. Stem or root 4 cm. in diameter. Secondary xylem. Pores solitary and radially aligned in groups of 3 or 4, with some clusters, average radial diameter 93 um. (72-114). average tangential diameter 81 wm. (64-107). Perforation plates scalariform, bars 10 to 15. Intervessel pits appear to be scalariform, average vessel element length 760 wm. (870-1100). Parenchyma abundant, apotracheal diffuse. Rays 2 to 4 cells wide. cells variable in size but mostly large and procumbent; uniseriate margins | or 2 cells high, cells square: uniseriates 3 to 10 cells high, the cells variable in size, mostly square Locauity. Del Puerto Creek, locality I. The wood anatomy of casG 60425 is much like that of Magnolioxvlon panochensis, a specimen described from locality 2a (Page, 1970), except that all the cells are larger, and the vessel elements are significantly shorter. casG 60425 may represent a root or a stem of the same species or, since it was collected from a slightly younger locality, a different species altogether. Few Group V woods have scalariform intervessel pits. Among those that do are Hibbertia (Dilleniaceae), and certain members of the Hamamelidaceae and the Magnoliaceae. CASG 60426 Ficures 21-23. MarteriAL. Small stem 3 cm. in diameter. Secondary xylem. Growth rings indistinct. Diffuse porous, pores solitary and in short chains and clusters, average radial diameter 58 am. (46-80), average tangential diameter 46 zm. (40-58). Perforation plates scalariform, with up to 50 or more fine bars. Intervessel pits small, opposite transitional; pits to ray cells small, numerous; average vessel element length 748 wm. (580-870). Parenchyma abundant, apotracheal, diffuse and in tangential pairs. Rays numerous, uniseriate, up to 23 cells high, heterocellular. Fibers thick walled. Pith. Small; cells small, rounded in transverse section, Phloem. About 3 mm. thick, cells in no particular pattern, walls of parenchyma Figures 21-29. 21-23, casGc 60426: 21, transverse section, X 62: 22, transverse 24-76 section, xX 200; tangential section, X 60. 24-26, casc 60427: 24, transverse section, X 67; 25, longitudinal alli showing bordered pits on vessel wall, x 250; 26, t ees section, x 60. 27, 28, casc 60428: 27, transverse section, x 28; 28, tangential section, x 80. 29, « cASG 60429, longitudinal section showing vessel wall pitting, = 7 =~ ~ 731 PAGE, DICOTYLEDONOUS WOOD, II 1980] ee Rone aa — 4 Se SP BF Seed ean ia : ae nena lo Sj : a re pons, wt pray pret S; whee Ue ares mo : ‘ ty, ea = a aes ee os To2 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 cells and ray cells highly thickened and tangentially elongated in outer layers. Rays continuous from xylem. Locauiry, Black Gulch, locality 2a. Because preservation Is poor, an accurate assessment of the number of end plate bars was not possible. It appears that most plates contain fewer than 50 bars; therefore, the fossil is placed in Group V. Type VB woods, with vessel elements averaging less than 900 om. in length, abundant parenchyma, and transitional to opposite intervessel pits, occur in such genera as /lex (Aquifoliaceae), Strombosia (Olacaceae), and Clethra (Clethraceae). Except for its more numerous end plate bars, the fossil is similar to the wood of Strombosia. Pollen attributable to both the Aquifoliaceae and the Olacaceae has been described from Black Gulch and Del Puerto Creek (Chmura, 1973) The following specimen, casG 60428, is classified in Group VIB. Members of this group have both simple and scalariform perforation plates, mostly solitary pores, and apotracheal parenchyma. CASG 60428 Figures 27, 28. MareriaL. Small stem 3 cm. in diameter. Secondary xylem. Pores ovoid, solitary, in slightly oblique arrangement, very small pores mixed with large ones, average radial diameter 121 fom. (70-153), average tangential diameter 104 wm. (87-157). Perforation plates simple and scalari- form with up to 30 bars; intervessel pits alternate or slightly elongated horizontally, occasionally almost scalariform near pith; pits to ray cells similar, apertures coalesc- ing obliquely in some cells: pits to parenchyma large; average vessel element length 551 wm. (406-696); tyloses occasional. Parenchyma abundant, apotracheal diffuse. Rays numerous, mostly uniseriate, some with biseriate centers, homogeneous to partly heterogeneous, up to 40 cells high, cells mostly procumbent, occasional marginal cells containing a single crystal. Fiber pits bordered: vasicentric tracheids occasional. Pith. Cells large, thin walled. Phloem. Alternating zones of fibers and stone cells separated by tangential rows of crushed thin-walled cells: cuboidal crystals occasional in parenchyma. Locauity, Black Gulch, locality 2a. Possibly because of poor preservation, vasicentric tracheids are discernible in only a few small areas in the two specimens with this pattern. All cells (except vessels) contain brown inclusions. Patterns similar to the above occur in Castanopsis (Fagaceae), Casuarina (Casuarinaceae), and Engelhardtia (Juglandaceae). Abundant apotracheal parenchyma and an oblique arrangement of solitary pores form a common pattern in these families. According to Moseley (1948), no species of Casuarina has only diffuse axial 1980] PAGE, DICOTYLEDONOUS WOOD, II 733 parenchyma. Although metatracheal bands of parenchyma are characteristic of most species of Juglandaceae and Fagaceae, there are some genera in which parenchyma is only diffuse. Vasicentric tracheids are characteristic of some genera of Fagaceae and at least one species of Casuarina. Scalariform perforation plates with few bars occur in species of some genera of the Juglandaceae including Engelhardtia, occasionally in small vessels in species of Casuarina, and in species of Nothofagus and Fagus. It is interesting to note that the three orders represented by these families are depicted by Cronquist (1968, p. 159) as closely related. Pollen attributable to the Fagaceae and the Juglandaceae has been described from both Black Gulch and Del Puerto Creek (Chmura, 1973). Group VIIC, which includes woods with both simple and scalariform perforation lates, pores in multiples as well as solitary, and some paratracheal parenchyma, is represented by specimen casc 60427. casG 60427 Ficures 24—26, MatERIAL. Stem 2 cm. in diameter. Secondary xylem. Semi-ring porous, the pores small, isodiametric, angular, soli- tary with some clusters and radial groups of 3, average radial diameter 49 wm. (43-58), average tangential diameter 52 um. (35-70). Perforation plates simple, a few scalariform, usually with 3 or 4 bars. Intervessel pits scalariform and transitional to alternate, large, bordered; pits to axial parenchyma large, round, almost scalariform; average vessel element length 464 wm. (406-520); tyloses abundant. Parenchyma terminal and sparse paratracheal. Multiseriate rays infrequent, 3 cells wide, the cells upright, walls thick; uniseriates low, the cells upright, square. Fibers septate near vessels; long, narrow, scalariform-pitted, tracheidlike elements associated with some vessels. Pith. Large, 7 mm. in diameter; cells small, more or less isodiametric, round in transverse section. Locauity. Del Puerto Creek, locality I. Loss of material during preparation of the thin section destroyed portions of the single growth ring, and grain distortion obscures much of what remains. Most of the axial parenchyma is associated with the larger vessels at the inner edge of the outer ring. Because the ring is incomplete and distorted, it is impossible to be certain that what can be observed represents the normal structure of the stem. The scalariform pitting and abundant tyloses make it difficult to determine the nature of the end walls, although most appear to be simple. There is a progression from scalariform intervessel pitting to transitional alternate between the primary xylem and the outer portion of the stem, where a mixture of the two types occurs. The rays are of the kind generally considered juvenile. These factors indicate that the mature secondary xylem of the plant may have had wider multiseriate rays and a mixture of scalariform or transitional and alternate pits. Since the wood was in a state of transition, comparison with modern mature woods Is valueless. All that can be said is that the fossil may be a transitional stage of mature patterns such as can be observed in some [voL. 61 JOURNAL OF THE ARNOLD ARBORETUM 734 Seaen- ae ea. s— Che ts . it Sis . aS tT ‘ =. @ ad eT ~~ ee Re & ential 31, tang x 60: : 30, transv 37. 30, 31, casc 60429 RES 30Q— Ficu ° erse section, x 75: 34, CASG 60432: tangential section, section, x 9S, 5: g, vessel wall sculpturin c transverse section near periph ction showing x 73; 37, transverse section near pith, 35, longitudinal se 36, x 7 x 75, ery of stem, co 1980] PAGE, DICOTYLEDONOUS WOOD, II 735 Araliaceae and in Leea (Leeaceae): scalariform tracheids are characteristic of Leea; terminal or initial parenchyma is uncharacteristic of both Leea and the Araliaceae. Group VIII is represented by specimens casc 60429 and casc 60430. Members of this group have simple perforations and scalariform, opposite, or transitional intervessel pits. This combination of characters is uncommon among modern woods but occurs in some Araliaceae, Cunoniaceae, Elaeocarpaceae, Fagaceae, Guttiferae, Leeaceae, Melastomataceae, Olacaceae, and Schisandraceae casG 60429 Figures 29-31. MateriAL. Mature secondary wood. Secondary xylem. Diffuse porous, the pores in multiples of 2 to 4, numerous, average radial diameter 143 wm. (100-174), average tangential diameter 110 pm. (75-156). Perforation plates simple, slightly oblique. Intervessel pits opposite, horizontally elongated to almost scalariform; pits to ray cells scalariform; average vessel element length 591 um. gene Parenchyma absent. Rays up to 3 cells wide, the cells mostly procumbent, varying in size; uniseriate margins | to 3 cells high, cells upright or square; uniseriates low, cells square or upright Locatity. Black Gulch, locality 2a. Type VIII woods with little or no parenchyma have been observed in the Vitaceae, Melastomataceae, and Fagaceae (Nothofagus). According to Metcalfe and Chalk (1950), re parenchyma is always present in the wood of Melastomataceae and itaceae. Some Magnoliaceae have wood patterns similar to that of the fossil, but at least some ees in these woods have scalariform perforations, and all have apotracheal parenchyma (mostly terminal). casc 60430 Figures 32—34. MateriaL. Secondary wood 2 cm. in diameter. Secondary wood. Pores solitary and in radial rows of 2 to 4, rounded, but flattened in areas of contact, average radial diameter 69 wm. (52-81), average tangential diameter 64 tm. (46-75). Perforation plates simple. Intervessel pits scalariform; average vessel element length 350 wm. (286-394). Parenchyma apotracheal. Rays mostly 2 or 3 cells wide, fe eis less than 500 pm. high, cells mostly procumbent; uniseriate margins | or 2 cells high; uniseriates infrequent, | to 9 cells high, cells square or upright. Locauity. Locality 2b. It is not certain that intervessel pits are scalariform in all vessels. Axial paren- chyma is present, but it is not possible to determine its amount or distribution, except that no paratracheal parenchyma was noted. A very similar stem, cAsG Ls consisting of a stacaie pith nearly | cm. in diameter surrounded by a layer of secondary xylem 13 mm. thick, was also collected at locality 2b. Near the pith there are a few scalariform ae plates with four or fewer bars. The rays are higher 736 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 and more heterogeneous than are those in casc 60430. The differences observed in the secondary wood of the two specimens are of the kinds that exist between early- and later-formed increments in the same plant. Type VIIIB woods with diffuse apotracheal parenchyma occur in species of the Cunoniaceae, Fagaceae (Nothofagus), and Olacaceae Group IX, which includes woods with simple perforations, alternate intervessel pits, and no parenchyma, is represented by two specimens, casc 60432 and CASG 60433. This pattern occurs in many families. casG 60432 Figures 35-37. MarteriaL. Stem 4.5 cm. in diameter. Secondary xylem. Pores solitary, in clusters, and in radial multiples of 3 to 6, average radial diameter 86 wm. (69-104), average tangential diameter 87 wm. (70-110), smallest near pith. Perforation plates simple, oblique. Intervessel pits alternate, crowded, minute; pits to ray cells similar; average vessel clement length 301 pm. (261-406). Parenchyma absent. Rays numerous, 3 to 7 cells wide, heterocellular, the cells variable in size and shape, large, square, or short procumbent, very irregularly organized; sheath cells common; uniseriates infrequent, | to 4 cells high. Small elements resembling tracheids present (no perforations observed). Fibers very short. Locauiry. Del Puerto Creek, locality 1. It is difficult to determine whether the irregular ray structure was characteristic of the plant as a whole since the only specimen collected represents the fork of a branch. Because this irregularity is consistent in the many sections cut from the sizeable specimen, however, | assume that it is typical of the plant. It appears that many rays that are uniseriate in tangential section may be radial uniseriate extensions of multiseriate rays in the process of reduction in width. Preservation is poor, and the grain is often distorted; therefore, the total absence of parenchyma is not certain. It could not have been abundant, for it would surely have appeared in one of the sections. Some members of the Compositae and Solanaceae have similar woods. Pollen attributable to the Solanaceae has been described from locality | (Chmura, 1973). — cAsG 60433 Ficures 38, 39. MareriaL. Small stem 17 mm. in diameter. Secondary xylem. Pores small, angular, mostly in radial multiples of 2 to 4, Ficures 38-47. 38, 39, casc 60433, x 63: 38. transverse section; 39, tangential section. 40-42, casc 60434: 40, transverse section, x 72:41, tangential section, x 70: 42, radial section, X 70. 43, 44, casc 60435: 43, transverse section, x 42; 44, 45, transverse section, X 45; 46, tangential section, x 7% 45-47, CASG 60419: transverse section, X radial section showing pitting on wall of ray cell, x 300; 47, 80. PAGE, DICOTYLEDONOUS WOOD, II et) ope ie) ba 1ghs pe Ses Peat a : ‘ ee u rj Mi wider [ea wk a a abated Ase Taq 738 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 clusters occasional, average radial diameter 69 wm. (34-87), average tangential diame- ter 50 pm. (29-58). Perforation plates simple, angle acute. Intervessel pits alternate, crowded, minute; average vessel element length 398 jzm. (377-464). No parenchyma observed. Rays numerous, low, uniseriate, some biseriate, the cells small, square or short procumbent, some upright. Pith. Almost 2 mm. in diameter; cells large, thin walled. Locauiry. Del Puerto Creek, locality | Wood patterns similar to that of the fossil can be observed in certain members of the Euphorbiaceae, Moraceae, Punicaceae, Sonneratiaceae, and Rubiaceae. ey Group XA, which includes woods with simple perforations, alternate intervesscl pits, and diffuse apotracheal parenchyma, is represented by specimens casc 60434, casa 60435, and casc 60419, Genera of many modern families are included in this group, ‘AsG 60434 Figures 40-42. MareriAL. Stem 2 cm. in diameter. Secondary xylem. Growth rings apparent with magnification. Pores small, round- ed, numerous, solitary and in radial pairs or triplets (occasional), average radial diameter 63 pm. (41-87), average tangential diameter 70 um. (46-87). Perforation plates simple. Intervessel pits small, mostly alternate, some opposite, apertures sometimes horizontally elongated; vessel element length ca. 412 fxm. Parenchyma sparse, apotracheal. Rays mostly low, average height 335 jum. (203-580), | to 3 cells wide, the cells square and short procumbent; uniseriate margins absent or up to 5 cells high, cells square or upright. Pith. Small, round. Phloem. Poorly preserved except for scattered stone cells and nests of fibers. Locauiry. Del Puerto Creek, locality 1. Some fibers appear to be septate, but there is some doubt about this interpretation. In many fibers extremely fine fungal hyphae follow the contours of the inner wall, at intervals crossing the lumen and continuing along the wall in the opposite direction. Unless the cells are carefully examined, the hyphae can easily be mistaken for septae. It is possible that passage along the full length of the fiber was obstructed by septae, and that hyphae crossings thus actually indicate the presence of septae. casG 60435 Ficures 43, 44. MareriaL. Stem 1.5 cm. in diameter. Secondary xylem. Growth rings distinct, demarcated by tangential pore clusters. Pores mostly solitary, occasionally in clusters or radial files, average radial diameter 65 um. (41-98), average tangential diameter 63 pm. (35-98). Perforation plates simple. Intervessel pits alternate, minute, the apertures narrow, extending beyond 1980] PAGE, DICOTYLEDONOUS WOOD, II 739 borders; average vessel element length 423 jm. (3 measurements). Parenchyma apotracheal diffuse. Rays mostly biseriate, some 3 cells wide, up to 12 cells high, the cells small, square and short procumbent. Ground tissue mostly fibers, some with conspicuously bordered pits, mainly in vicinity of vessels. Pith. Cells small, thick walled, round in transverse section. Phloem. Groups of stone cells alternating with thin-walled cells; multiseriate rays flaring out and forming conspicuous wedges in outer portion; vertical files of ‘‘beaded’’ cells resembling chambered parenchyma, containing what appear to be remnants of polyhedral crystals. Locatiry. Del Puerto Creek, locality 1. Due to poor preservation, the characteristics of the fibrous elements in the vicinity of vessels are not always clear. Those with conspicuous pits are not consistently associated with vessels. The specimen is closely similar to Carpinoxylon ostryopsoides (Page, 1970). There are a few multiperforate vessels in C. ostryopsoides, the rays are wider, and the parenchyma is more abundant. Aggregate rays are also present. Among modern forms with aggregate rays, there is wide variation in the age at which these rays are initiated. Since casc 60435 is half the diameter (3.0 cm.) of the specimen of C. ostryopsoides (mistakenly described as 1.5 cm.), and aggregate rays were observed in only the outer portion of the section, it is possible that these rays did not normally develop in early-formed wood. A specimen (cas 60436) collected at an earlier date but from the same locality (2a) as C. ostryopsoides is quite similar, although it lacks aggregate rays. Close similarities were also observed in pith and bark among the three specimens, although only casc 60435 has both tissues. Carpinoxylon ostryopsoides has a pith but no bark; casc 60436 has bark but no pith. These two specimens are probably from the same kind of plant, while casc 60435 may be a different species of either the same genus or a closely related one. Although all three specimens compare favorably with the Coryleae of the Betulaceae (except for the smaller pits in the fossils), there are no modern genera that bear a close resemblance. There is some similarity to Castanopsis of the Fagaceae, but here the intervessel pits are also large, the vessel-ray pits are even larger, and the pores jen are exclusively solitary. casc 60419 Figures 45-47. MareriaL. Fragment of mature wood. Secondary xylem. Pores solitary, rounded, ovoid, widely spaced in oblique radial arrangement, tangential clusters occasional, small pores mixed with large, average radial diameter 150 wm. (87—232), average tangential diameter 116 fxm. (69-174). Perforation plates simple, horizontal or slightly oblique. Intervessel pits small, alternate, the apertures narrow, sometimes coalescent; pits to ray cells similar, apertures sometimes coalescing obliquely; average vessel element length 494 wm. (435-696); tyloses occasional. Parenchyma abundant, mostly apotracheal diffuse, but some paratracheal. Rays numerous, mostly uniseriate, up to 20 cells high, the cells 740 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Square, upright, or short procumbent; multiseriates infrequent, up to 4 cells wide, cells mostly procumbent; all ray cells containing dark brown substance. Fiber pits round, bordered, crowded. Locatiry. Y2 mile north of Pigeon Point, locality 4. Axial parenchyma cells vary in diameter. It is sometimes difficult to distinguish between axial and ray parenchyma in radial section because the cells associated with vessels are small and frequently square or rhomboidal as in the rays. In transverse section, however, many large pores appear to have at least a partial sheath of parenchyma. Both rays and axial parenchyma are abundant; therefore, the association of either with vessels may be fortuitous. There is a general similarity between this specimen and casc 60428, but the cells in the rays of the latter are mostly procumbent, occasional cells contain crystals, and there is some evidence of vasicentric tracheids. Although scalariform perforation plates are present in casG 60428, this may not be a significant difference since a stem collected at Black Gulch (casc 60437) with secondary xylem identical to that of casG 60419 has a few vessels with scalariform perforations in the early-formed wood. casG 60419 has many characteristics of some woods of the Myrtaceae and 1s also similar to Licania and Parinaria (Rosaceae), and Monetes and Marquesia (Diptero- carpaceac); however, some paratracheal parenchyma is typical of the latter two genera. Group XIA is represented by 12 specimens consecutively numbered from casG 60438 through casc 60449. This group is distinguished by simple perforations, alternate intervessel pits, and vasicentric parenchyma—a pattern common among modern woods. A discussion of the relationship between the fossils and modern woods 1s presented following the descriptions of the specimens. casG 60438 Ficurrs 48, 49. MareriaL. An axis 2 cm. in diameter. The center is not preserved; therefore, it 1s not possible to determine whether the specimen is a stem or a root. Secondary xylem. Pores large, solitary and in radial or tangential multiples of 3, average radial diameter 202 wm. (174-232), average tangential diameter 161 pm. (145-191). Perforation plates simple. Intervessel pits alternate, crowded, apertures narrow; pits to parenchyma large; average vessel element length 383 wm. (261-522). Parenchyma paratracheal. Rays 3 to 9 cells wide, up to 2 mm. high, the cells large, square or short procumbent; uniseriate margins lacking; uniseriates low, heterocellular; occasional cells with opaque, amber-colored inclusions. Fibers short, average length similar to that of vessel elements. Phloem. Thick in proportion to amount of xylem; rays flaring out in cross section due to tangential expansion of cells; stone cells or other sclereids absent; many cells containing amber inclusions. Locatiry. Del Puerto Creek, locality 1. 1980] PAGE, DICOTYLEDONOUS WOOD, II 741 Ee _ Be My . Devesesshele * Ficures 48-56. 48, 49, casc 60438: 48, transverse section, x 38: 49, tangential section, X 54. 50, 51, casc 60440, 80: 50, tangential section; 51, transverse section. 52, 53, casc 60439: 52, transverse section, < 38; 53, tangential section, 4. 54-56, casc 60442: 54, transverse section, < 70; 55, longitudinal section th showing nests of sclereids, X 6; 56, radial section showing enlarged through pl secretory cell, 742 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 The large, numerous vessels and the high rays indicate that the plant may have been a liana. Inclusions such as those in the ray cells and bark may have been secretions of oil. The cells, however, are not especially enlarged. A’ proper radial section was not obtained. oy) casG 60439 Ficures 52, 53. MareriaAL. Stem or root. Secondary xylem | cm. in diameter with bark 2.5 mm. thick. Secondary xylem. There appear to be 13 or 14 faint growth rings. Pores large, solitary and in radial multiples, average radial diameter 232 wm. (128-365), average tangential diameter 174 pm. (116-232). Perforation plates simple. Intervessel pits small, alternate, apertures narrow; pits to rays variable, mostly large: pits to. axial parenchyma large, oval: average vessel element length 548 wm. (464-638); tyloses abundant. Parenchyma paratracheal. Rays variable, some very high, 2 to 4 cells wide, heterocellular; uniseriates up to 6 cells high, cells variable; many cells containing amber-colored inclusions. Fibers short. Phloem. Rays numerous, the cells enlarged tangentially, particularly at periphery of axis; scattered cells containing amber-colored inclusions; all cells thin walled, some parenchyma cells enlarged but lacking inclusions. Locauiry. Del Puerto Creek, locality This specimen is somewhat similar to casc 60438, although in that specimen the pores are more numerous, there are far fewer pore multiples, and tyloses are lacking. ggests that if it is a root, it may oe The presence of growth rings in casG 60439 st not have been subterranean. cAsG 60440 Ficurres 50, 51. MareriAL. Small stem 1.5 cm. in diameter. Secondary xylem. Pores mostly solitary, some radial multiples of 2 to 4, slightly angular, average radial diameter 100 wm. (58-121), average tangential diameter 80 wm. (58-98). Perforation plates simple, slightly oblique. Intervessel pits large, crowded, alternate; pits to ray cells large, oval, irregularly oriented: average vessel element length 399 um. (343-445). Parenchyma paratracheal, sheaths | or 2 cells thick. Rays mostly uniseriate, occasionally bi- or triseriate, the cells mostly square or upright, with numerous small pits on tangential and horizontal walls, occasionally slightly inflated and containing amber-colored inclusions. Some fibers septate. Pith. 0.5 cm. in diameter; nests of sclereids numerous: protoxylem points numer- led, other cells large. c ous; pith cells in vicinity of primary xylem small, thick wal Locatiry. Del Puerto Creek, locality 1. A specimen (casG 60441) collected at locality 2b and nearly identical in every way contains numerous inflated cells, which are interpreted to be secretory cells. 1980] PAGE, DICOTYLEDONOUS WOOD, II 743 casc 60442 Figures 54-56. MatEeRIAL. Stem 22 mm. in diameter. Secondary xvlem. Pores solitary with occasional radial files of 2 to 5, average radial diameter 106 pm. (70-128), average tangential diameter 85 jm. (52-104). Perforation plates simple. Intervessel pits small, alternate, crowded, pits to ray cells large, oval; average vessel element length 365 wm. (290-435). Parenchyma sparse, paratracheal. Rays | to 3 cells wide, the cells short procumbent or upright, often enlarged and containing amber-colored inclusions. Pith. Large, round; cells very large except in vicinity of primary xylem where small and thick walled. Nests of cells resembling potential sclereids common. Locauity. Black Gulch, locality 2a. casG 60443 FiGures 57-59. MarteriAL. Mature secondary wood. Secondary xylem, Pores solitary and in occasional radial multiples of 2 to 4, rounded, average radial diameter 118 wm. (93-139), average tangential diameter 92 wm. (81-104). Perforation plates simple. Intervessel pits small, alternate: pits to ray cells large; length of vessel elements ca. 765 wm.; tyloses occasional. Parenchyma sparse, paratracheal. Multiseriate rays 4 or 5 cells wide, cells mostly procumbent: uniseriate margins | to 3 cells high, cells upright; uniseriates infrequent, 2 to 4 cells high, cells upright; rays widely spaced; some enlarged cells may have been crystal- liferous. Fibers frequently septate. Locauity. Black Gulch, locality 2a. casG 60444 MaterIAL. Fragment of stem with pith and a small amount of secondary xylem. — Secondary xvlem. Pores mostly solitary and widely spaced with occasional radial multiples of 3, average radial diameter 129 wm. (104-174), average tangential diame- ter 115 wm. (93-145). Perforation plates simple, slightly oblique. Intervessel pits alternate, crowded; pits to parenchyma oval; length of vessel elements ca. 300 wm. Parenchyma paratracheal. Multiseriate rays mostly 3 cells wide, the cells mostly procumbent with | or 2 marginal rows of square cells, nearly homogeneous; uniseriates infrequent. Pith. | cm. in diameter, with large, mostly peripheral cavities. Primary xylem. Protoxylem points conspicuous, extending deep into pith, the adjacent pith cells very small, thick walled: average vessel diameter 79 zm. (63-93). Multiseriate rays extending into secondary xylem. — Locatity. Del Puerto Creek, locality |. The nature of the peripheral pith cavities is difficult to determine. There is no structural evidence to indicate whether they were a functional part of the stem or 744 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 eet eB Sen ee a Nees 7 ay ~ 2 0). eae = Ficures 57-65. 57-59, casc 60443: 57, tangential section, 78; 58, transverse section, X 52:59, radial section showing enlarged ray cell with contents resembling a crystal, x 80. 60, 61, casc 60445: 60, tangential section, x 64; 61, transverse section, X 57. 62, 63, casc 60448: 62, tangential section, X 78; 63, transverse section, * 70. 64, 65, casc 60447: 64, tangential section (note small intervessel pits), x 78; 65, transverse section, x 75. 1980] PAGE, DICOTYLEDONOUS WOOD, II 745 whether they were formed by some foreign agent. Vertical extent of the cavities 1s minimal. casG 60445 Ficures 60, 61. MateriaL. Fragment of mature secondary wood, diameter 2.2 cm. Secondary xylem. Diffuse porous, pores solitary with some radial pairs, radial files of 3, and occasional clusters of 3 or 4, average radial diameter 112 pm. (87-145), average tangential diameter 82 wm. (70-99). Perforation plates simple. Intervessel pits large, alternate; pits to ray cells large: vessel element length ca. 464 wm.; tyloses occasional. Parenchyma sparse, paratracheal. Rays up to 4 cells wide, irregular in height (0.58-3.1 mm.), some in process of dissection, the cells square and short procumbent, with tangential walls often inclined from vertical in radial section: uniseriate margins absent or up to 3 cells high, cells upright: uniseriates | to 4 cells high, cells variable in size and shape. Fibers thick walled. Locauity. Locality 2b. casG 60446 Materia. Small stem 1.5 cm. in diameter. Secondary xylem. One distinct growth ring present. Pores mostly solitary, radial pairs numerous, clusters of 3 occasional, average radial diameter 88 jum. (73-102), average tangential diameter 80 wm. (58-102). Perforation plates simple. Intervessel pits alternate, fairly large, particularly in ligule; pits to ray cells large, oval; average vessel element length 406 wm. (348-464): tyloses abundant. Parenchyma sparse, paratracheal. Rays mostly 2 or 3 cells wide, cells of multiseriate portion mostly procumbent; uniseriate margins | to 5 cells high, the cells square or upright, uprights more common toward center of stem; uniseriates mostly 5 cells high, cells square: some cells containing dark globules. Pith. Small. Phloem. Appears to have checkered pattern of groups of thin- and thick-walled Locauity. Locality 2b. casG 60447 Ficures 64, 65. MateriAL. Mature secondary xylem. Secondary xylem. Pores mostly solitary with some radial multiples, average radial diameter 154 wm. (116-191), average tangential diameter 116 wm. (87-145). Perfora- tion plates simple. Intervessel pits very small, alternate, apertures coalescent; pits to parenchyma oval; average vessel element length 407 wm. (314-515). Parenchyma paratracheal, sheaths | to 3 cells thick. Rays numerous, uniseriate or biseriate, 3 to 18 cells high, cells procumbent, some with one row of square cells at margins. Locaurry. Del Puerto Creek, locality |. 7146 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 cAsSG 60448 Figures 62, 63. MateriaL. Mature secondary wood. Secondary xylem. Growth rings indistinct, defined by few rows of radially flattened fibers and somewhat smaller pores. Diffuse porous, pores mostly solitary or in radial pairs, average radial diameter 103 jm. (75-127), average tangential diameter 81 jum. (58-104). Perforation plates simple, angle acute. Intervessel pits alternate, large, apertures small; average vessel element length 412 fm. (406-580); tyloses thin walled. Parenchyma sparse, paratracheal. Multiseriate rays widely spaced, variable in height from 174 wm. to | mm., mostly 3 or 4 cells wide, cells low procumbent; uniseriate margins | to 3 cells high, cells square; uniseriates infrequent; all cells containing dark globules. Fibers large in diameter, average 29 pm Locauity. Locality 2b. casG 60449 MareriAL. Stem 2 cm. in diameter. Secondary xylem. Pores solitary, in short radial files and in clusters, average radial diameter 52 wm. (29-75), average tangential diameter 53 pm. (29-81). Perforation plates simple. Intervessel pits not observed: pits to parenchyma large; average vessel element length ca. 500 jxm.; tyloses abundant. Parenchyma paratracheal. Rays low (about 12 cells high), 2 (or 3) cells wide, cells square or short procumbent;: uniseriates about 10 cells high, heterocellular. Fibers large in diameter, some septate. Pith. 3 mm. in diameter, prominently 6-lobed in transverse section; cells small. Locauity. Del Puerto Creek, locality 1. Except for its smaller pores and lower rays, casG 60449 is very similar to CASG 60448. Since one would not expect to find higher rays in mature wood than in the juvenile wood of the same plant (see Bailey & Howard, 1941), it is unlikely that the two specimens represent the same natural species. The mature wood was collected from locality 2b, and the small stem from locality 1, a stratigraphic distance representing an estimated time interval of approximately one million years. Although there is a general similarity between these specimens and casc 60430 and casc 60434, no paratracheal parenchyma was observed in the latter two. In addition, intervessel pits in cAsG 60430 appear to be scalariform, whereas those in casc 60434 are Opposite or alternate. Among the many families with type XIA wood patterns, woods with little or no apotracheal parenchyma occur in at least eight. Special features, when present, serve to differentiate the various families. For example, secretory cells are characteristic of the Lauraceae, while radial canals and crystalliferous ray cells are found in the 5 9) Anacardiaceae and Burseraceae. Canals also occur in the Araliaceae, and vestured intervessel pits are characteristic of the Leguminosae and the Combretaceae. Of the fifteen type XIA fossil woods described, casc 60438, casc 60439, casc 60440, Cas 60442, and two specimens described earlier (Udminiwn pattersonensis and U, mulleri (Page, 1967)) contain a few parenchyma cells that appear to have been oil or 1980] PAGE, DICOTYLEDONOUS WOOD, Il 747 secretory cells of some sort. In some specimens these cells are slightly inflated (Ficure 56), but in others they are distinguished only by the presence of amber- colored inclusions. Pith sclereids such as those described in casc 60442 and casc 60440 occur in several members of the Lauraceae. Infrequent rays, mostly solitary pores, and large vessel—ray pits as described in casc 60443, case 60444, casc 60445, and casc 60446 are characteristic of families such as the Anacardiaceae, Burseraceae, and some genera of the Araliaceae. Crystal- liferous cells (Figure 59) in the rays of casc 60443 indicate a possible alliance with the Anacardiaceae and/or Burseraceae. Pore multiples and low, numerous, mostly uniseriate rays composed of procum- bent cells as described in cas 60447 occur in some Combretaceae and Leguminosae. The presence of the vestured intervessel pits characteristic of these families is almost impossible to establish in fossil material. There are no distinguishing features in casc 60448 and casc 60449; however, the six-lobed pith of the former is uncharacteristic of the Lauraceae. CONCLUSIONS Remains of leaves and pollen of Cretaceous angiosperms are sufficiently abundant to provide an adequate source of information for the study of pre-Cenozoic angio- sperm history. By contrast, angiosperm wood is sparsely represented. Until the discovery of the suite of dicotyledonous woods from central California, only a few isolated specimens had been described. Very little, therefore, could be learned of the evolutionary history of the dicot stem. The number of specimens is still small, but we now have some evidence of the diversity of wood patterns that existed in the Upper Cretaceous, as well as some bases for observations with respect to the level of specialization displayed by these woods as compared with those described from later periods. Of special interest is the lack of highly specialized characters in the specimens of the collection studied. Such features have not been convincingly described in any other pre-Cenozoic woods of unquestioned provenance. There is, for example, no evidence of complex pore arrangements, storied elements, resin canals, or tile cells, and while axial parenchyma occurs in a variety of conditions, there is no evidence of multiseriate bands or aliform sheaths. Furthermore, there 1s a high proportion of specimens with scalariform perforation plates and medium or long vessel elements. The most highly specialized pattern is that of casc 60432, which has infrequent uniseriate rays, highly disorganized multiseriate rays with the cells vari- able in size and shape, very short fibers, and no observable parenchyma. The wood, in fact, resembles the secondary xylem of certain members of the Compositae. Although a discussion of some of the above observations was presented in an earlier paper (Wolfe et al., 1975), a more extensive treatment will be presented in a forthcoming report. ACKNOWLEDGMENTS I wish to thank John Sims, of the United States Geological Survey at Menlo Park, California, for the specimen from the Cache Creek locality; Carl Wentworth, also of 748 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61] the U.S.G.S. at Menlo Park, for the specimen from the Pigeon Point Formation; and Elisabeth Wheeler, North Carolina State University, Raleigh, North Carolina, for reading the manuscript and for her helpful comments. BIBLIOGRAPHY Battery. 1. W., & R. A. Howarp. 1941. The comparative oa of the Icacinaceae. Rays 7 a secondary xylem. Jour. Arnold Arb. 22: Cumura.C. A. . Upper Cretaceous (Campanian— vetoes ) angiosperm pol- len from rh western San Joaquin Valley, California, U.S. A. Palaeontographica 141B: 89-171. a A. 1968. The evolution and classification of flowering plants. xi + 396 pp. aa Mifflin Co., Boston oa Ww. ee _ Comparative morphological studies in Dillentaceae, 1. Wood anatomy. Jour. acs Arb. 48: 1-2. DouGias. R. G. 1969. Upper Cretaceous planktonic foraminifera in northern Califor- nia. Part 1. Systematics. Micropaleontology 15: 151-209, Goupkorr, P. P. 1945. Stratigraphic relations of Upp er Cretaceous in Great Valley, California. Am. Assoc. Petr. aes Bull. 29: 956-1007. JR Hari.C. A., _L. Jones. & S.A. Brooks. 1959, Pigeon Point Formation of rate Cretaceous age, San aa aie California. Am. Assoc. Petr. Geol. Bull. 2855-2865 Mercatre.C. R., & L. Ciark. 1950. Anatomy of the dicotyledons. Vols. 1. 2. Ixiv + 1500 pp. Clarendon Press, Oxford, England. _— F. M. 1948. Com oe anatomy and phylogeny of the Casuarinaceae. . Gaz. 110: 231-28 PAGE, 7 M. 1967. eee wood from the Upper Cretaceous of central Califor- nia: Part I. Am. Jour. Bot. 54: 510-514. ——.. 1970 Ne wood trom the Upper Cretaceous of central California: Part II. Ibid. 57: —1144. 279. See eee po a the Upper Cretaceous of central Califor- nia. Jour. Arnold Arb. 60: 32 PessaGno. E. A., Jr. 1974. A ae eee zonation for the Upper Cretaceous portion of the Great Valley Sequenc p. 61-80 in Soc. Econ. Paleontol. Minerol. Preprints, Pacific Section Meine San Diego, California. Popenor, P. . Imeay. & M. A. Murpry 1960. Correlation of the Cretaceous formations of the Pacific Coast. Geol. Soc. Am. Bull. 71: 1491-1540. Wentwortu, C. M. 1960. Sedimentary structure and inferred turbidity current origin of the Cretaceous Pigeon Point Formation, San Mateo County, California. Unpubl. report on student research project. 66 pp. Stanford University Dept. Geol., Stanford, California. DEPARTMENT OF BIOLOGICAL SCIENCES STANFORD UNI\ STANFORD, CALIFORNIA 94305 1980] HOWARD ET AL., POST-ERUPTIVE VEGETATION 749 THE POST-ERUPTIVE VEGETATION OF LA SOUFRIERE, GUADELOUPE, 1977-1979 RicHARD A. Howarb, JACQUES PORTECOP, AND PIERRE DE MONTAIGNAC Between July 8, 1976, and March |, 1977, a series of phreatic eruptions occurred from three fracture zones at the summit (ca. 1400 m.) of La Soufriére, a volcano on the island of Guadeloupe in the West Indies. Forcible ejections of gas, water vapor, mud, dust, and boulders, but no lava, destroyed the vegetation of the southeastern, southern, and southwestern slopes of the mountain from the summit to beyond the Savane a Mulets and Piton Tarade, a distance of approximately one kilometer (FIGURE I). The eruptions were accompanied by increased fumarole activity, resulting in toxic gas emissions from other summit vents and from those in the Col de l’Echell (ca. 1200 m.). The present study concerns the nature of the destruction of the vegetation reports on the recovery of certain taxa and details the early stages of revegetation of > it the affected area. The kistory of seismic-volcanic activity of La Soufri¢re is incomplete. The age of the present pumice deposits near the current town of Basse-Terre is estimated at approximately 27,000 years. There are records of the ejection of incandescent material from the volcano in 1590 and of fumarole activity in 1635. In 1797-98 an important radial fracture to the north-northwest of the summit ejected vapor. dust, and rivers of mud; this was repeated in 1836-37. In 1892 fumaroles opened in the Col de I’Echelle., No major events on Guadeloupe were associated either with the classic eruptions in 1902 of Mt. Pelée, Martinique, and La Soufriere, St. Vincent, or with the activity of the latter in 1972 and 1979. On La Soufriére of Guadeloupe, a new radial fracture developed on the southeastern flank in 1956, with the projection of vapor and dust in a seismic crisis lasting about four months; after this the volcano was quiet until the tremors of late 1975. The fracture that formed in 1956 reopened on July 8, 1976, with significant projections of vapor and dust. the formation of a geyser at the Col de l’Echelle, and a river of mud and boulders that descended the Carbet Valley. Ejections of dust continued on July 25 and August 9 and 10. On August 30 the abyss of Tarrisan, a summit crater, ejected rocks and dust, while new fissures appeared on the south flank of the mountain. Another event on September 14 blew out the newly opened fissure of August 30 with the ejection of boulders and a oe of mud and rocks in the direction of the Gallion River valley. On September 22 a plume of gas and dust reached an estimated altitude of 1500 meters above the crater. Ejections of dust occurred again on October 2 and between the 10th and the 2 The summit fissure La Croix erupted on January 14 and 15, 1977, projecting boulders and producing mud rivers on several slopes. After a month of relative quiet, the last ejections of dust and boulders took place on February 28 and March |, 1977. © President and Fellows of Harvard College, 1980. Journal of the Arnold Arboretum 61: 749-764. October, 1980. 750 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 1300 900 900 ; j 1200 1000 -zZ> Ficurr |. Map of summit of La Soutricre, Guadeloupe. Area of complete destruc- tion of vegetation shown in dark dots: area of partial ade in lighter dots. a, summit or volcano and eae of phreatic emissions, 1456 m.: b, Savane a Mulets, 1142 m.; c, Piton Tarade; d, rai area: e, Col de TP’ Echelle: f. L’Echelle; g. La Citerne: h 1414 m Gallion River aics . Morne Amic, 1390 m.: j, Morne Carmichael, Since then all fissures and craters have been relatively quiet, with only casual emissions of steam from vents on the northern edge of the summit and near the Col de I’Echelle An ascent to the summit of La Soutriere is not a difficult climb. During the nineteenth century there were several trails from the southwest, one from the east, and another from the north. A cobblestone carriage track ascended from St. Claude to Bains Jaunes, where hot baths had been created using drainage from mountain streams. Although the area was easily accessible for over a century, collections of plant material remain inadequate, and nearly a dozen species reported from Bains Jaunes and the Soufri¢re have not been re-collected. Henri Stehlé (1935) published excellent, although brief, descriptions and enumerations of the vegetation around Bains Jaunes and the slopes and summit of La Soufricre. The illustrations presented in his paper, especially the double-faced pages between text pages 208 and 209, 224 and 225, and 240 and 241 (Ficurr 2), serve well for comparison with the recent and present conditions on the approach and summit of La Soufri¢re (Figure 3). In 1965 the government of Guadeloupe built a vehicular road to the area called Savane a Mulets and created a parking area in that location so that tourist buses could take groups of visitors to the area. An easy trail (Trace des Dames, Figure 7) ascends clockwise from the south to the north around the flank of La Soufri¢re, with a moderately steep climb to the summit. An undated (pre-1976) photograph (Ficure 4) in a recent paper by Sastre (1978) shows the undisturbed vegetation at the Savane a 1980] HOWARD ET AL., POST-ERUPTIVE VEGETATION 751 Mulets, currently the parking lot area. Neither the illustrations of Stehlé nor the photo by Sastre shows any plants killed by gases from fumaroles. When R. A. and E. S. Howard first climbed La Soufriére in 1950 (Howard, 1962, fig. 9), they found indications that fumarole emissions had had an effect on the woody vegetation even before the seismic activity of 1956. Stehlé (1935) described a plant association on the slopes of La Soufriére that he Ficures 2, 3. 2, view of ‘‘Clusietum guadelupense’’ and summit of La Soufriére from south (reproduced from Stehlé, 1935). 3, view of summit of La Soufriére from south (1979): complete destruction of Clusia formation evident in foreground. dae JOURNAL OF THE ARNOLD ARBORETUM [voc. 61 ’ called a **Clusietum guadelupense’’—a transition forest between the taller primary forest of more diverse composition in the area of Bains Jaunes and the stunted forests of the slopes at 1000 meters elevation. The size and dominance of Clusia is well shown in his plate reproduced here as Ficurr 2. Between 1150 and 1250 meters, a shrubby growth—a ‘‘Lobelietum guadelupense’’—dominates the area, although trees reduced in stature persist in depressions and valleys on the lee face of the mountain. Rocky areas near the summit are covered with bromeliads, forming a **Pitcairnietum guadelupense,”* and on the summit Stehlé recognized swampy areas as **Eleocharidetum guadelupense,”’ with the cliff faces having a **Sphagnetum guadelupense.”’ The southern and western flanks of La Soufri¢re affected by the recent volcanic activity are today a lahar, a volcanic desert of boulders, mud flats, and the debris of dead plants (Ficurrs 3, 6). The original vegetation of this area can be identified by comparison with areas on the northwestern flank of the volcano at comparable eleva- tions. Trees of Clusia mangle Rich. (Guttiferaec) dominate and form a tangle of spreading branches supported by adventitious or prop roots. Specimens of Cyrilla racemiflora L. (Cyrillaceae), Weinmannia pinnata L. (Cunoniaceae), [lex macfadyenit (Walp.) Rehder (Aquifoliaceae), Myrsine coriacea (Sw.) R. Br. (Myrsinaceae), Hedyosmum arborescens Sw. (Chloranthaceae), and Didymopanax atlenuatus (Sw.) Marchal (Araliaceae) are common, interspersed in the Clusia thickets. All are much- branched specimens, with the branching usually occurring from the base, Occasionally large, single-trunked trees of Richeria grandis Vahl (Euphorbiaceae) and Myrcia platyclada DC. (Myrtaceae) are found, and these are sparsely branched. Shrubs within the Clusia associations include Charianthus alpinus (Sw.) Howard (Melastomataceae), Freziera undulata (Sw.) Sw. (Theaceae), Besleria lutea L. (Gesneriaceae), and Palicourea crocea (Sw.) Roemer & Schultes (Rubiaceae). Philodendron giganteum Juss.) — Schott (Araceae) sprawls through the Clusia. Plants of Norantea spiciflora Krug & Urban (Marcgraviaceae) and Hillia parasitica Jacq. (Rubiaceae) may be woody at the base, and their scrambling branches appear vinelike from a distance. A few plants of Prestoea montana (Graham) Nicholson (Palmae) are found but rarely reach the canopy of Clusia. Two ferns, Dicranopteris pectinata (Willd.) Underw. and Histiopteris incisa (Yhunb.) J. Sm., are relatively common but not abundant in the dense growth. Between the swaths of the Clusia on slopes or flat areas is a shrub tangle that appears windswept and smooth. Although only 1-2 meters tall, the tangle is impene- trable from the ground and nonsupportive of any attempt to cross on top. Epiphytic masses of Guzmania plumieri (Griseb.) Mez, their rosettes filled with water, and soggy accumulations of Sphagnum and leafy liverworts cause a collector to become soaked while gathering specimens. The shrub association is of heterogeneous compo- sition; the principal plant species are Baccharis pedunculata (Miller) Cabrera (Compositae), Besleria lutea L. (Gesneriaceae), Charianthus alpinus (Sw.) Howard (Melastomataceae), Charianthus corymbosus (Rich.) Cogn., Clidemia guadalupensis —~ Griseb. (Melastomataceae), Freziera undulata (Sw.) Sw. (Theaceae), Miconia coriacea DC. (Melastomataceae), Miconia vulcanica Naudin, Psychotria aubletiana Steyerm, (Rubiaceae), Rondeletia parviflora Rich. (Rubiaceae), and Ternstroemia elliptica Rich. (Theaceae). Psychotria guadalupensis (DC.) Howard (Rubiaceae) is an epiphytic shrub, while Norantea spiciflora (Juss.) Krug & Urban (Maregraviaceae) and Symphysia racemosa Ficures 4-7. 4, Savane a Mulets area prior to 1965. 5S, parking area at Savane 4 Mulets, 1979: slope of La Soufriére on left, cascade of boulders evident in Col de l’Echelle; L’Echelle mountain in center. 6, dead shrub area on slopes of La Soufriére. 7, Trace des Dames in area of mud slides on western slope of mountain: unaffected Clusia formation evident in upper left, with Nez Cassé mountain shown in profile. [0861 NOILVLHOFA AAILANYA-LSOd “TV LA GUVMOH eSl 754 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 (Vahl) Stearn (Ericaceae) are scandent woody plants. The herbaceous species scattered throughout the shrub zone include Blechniwmn violaceum (Fée) C. Chr. (Polypodiaceae), Calolisianthus frigidus (Sw.) Gilg (Gentianaceae), Guzmania dussii Mez and G. plumieri (Griseb.) Mez (Bromeliaceae), Juncus guadeloupensis Buch. (Juncaceae), and Lobelia digitatifolia (Griseb.) Urban (Lobeliaceac). The single scandent herbaceous or subwoody vine present is Mikania ovalis Gilg (Compositae). On rock outcrops, cliff faces, or occasional boulders, there are woody and herba- ceous plants of various habits: Gaultheria swartzii Howard (Ericaceae), Lsachne rigidifolia (Poiret) Urban (Gramineae), Lycopodium clavatum L. (Lycopodiaceae), Machaerina restioides (Sw.) Vahl (Cyperaceae), Peperomia hernandifolia (Vahl) Dietr. and P. tenella (Sw.) Dietr. (Piperaceae), Pilea parietaria (L.) Blume (Urticaceae), Pitcairnia bifrons (Lindley) Read (Bromeliaceae), Pityrogramma chrysophylla (Sw.) Link (Polypodiaceae), Pterolepis glomerata (Rottb.) Mig. (Melastomataceae), Rhynchospora polyphylla (Wahl) Vahl (Cyperaceae), Sauvagesia erecta L. (Ochnaceae), Tibouchina ornata (Sw.) Baillon (Melastomataceae), and Viola stipularis Sw. Violaceac). When Calolisianthus frigidus (Sw.) Gilg (Gentianaceae) is found in this area, the plants are dwarfed and often flower when less than a foot tall. Following the eruption of July 8, 1976, Portecop made some observations on the initial effects of the volcanic activity. On August 15, 1976, Basse-Terre (the capital city) and surrounding areas were evacuated of 74,000 people; travel to the area was considered dangerous and was controlled. In October of 1976, Claude Sastre climbed La Soufri¢re. His observations are included in a brief appendix to his paper (1978). On December 8, 1976. Henri Stehlé ascended the volcano, and he has recently (1979) published the first part of his observations. Portecop returned to the mountain on December 20, 1976, and Howard and Portecop examined the slopes and summit in March, 1978, and again in April, 1979. The present report expands the brief published reports and extends the period of observation of lahars. The climate at the summit and on the upper slopes of La Soufriere is harsh. Under normal circumstances the mountaintop is cloud covered: it is estimated to be completely free of clouds only five or six days a year. The heavy morning cloud cover 1s generally dissipated briefly during part of the day, but it reassembles quickly, producing heavy showers and dangerously reduced visibility. By mid-afternoon the cloud cover is solid again. Rainfall is estimated at 10 meters annually, but mist accumulation is nearly continuous. Before the eruptions runoff of rainfall was controlled by the dense cover of vegetation, but both sheet and channel erosion are now apparent in the decimated areas. The original cover of volcanic dust on the vegetation has been washed off by rain, and the mud flows have been reduced in thickness on the flanks, resulting in deep accumulations of sediment in many valleys and depressions. The winds on the summit, classified as fresh (19-30 m.p.h.), reach gale force (30-65 m.p.h.) in gusts. Changes in temperature of 20°C. have been ee between the occasional full sun break in the clouds and the next passing _~ mis ae normal emission of acid gases from the summit fumaroles has had a cumula- tive effect in recent years on the flank vegetation on the western and southern slopes of La Soufriére. The current surface runoff and standing water show a pH of 4.5. g new leaves from apex; Dr. Portecop’s hand indicates broken end of large stem with apex developing new leaves of normal size. [0861 NOILVLH99UA FAILINUA-LSOd “TV LD GYUVMOH SSL 756 JOURNAL OF THE ARNOLD ARBORETUM [vot. 61 Drip from leaves, trunk water, and water held by the rosettes of leaves of bromeliads gave pH measurements of 4 to 5. R. M. Chevrier reported in 1976 that pH recordings taken at the level of Echelle during the eruption of July 14-16 went from 1.1 to 3.4. An unusually acid environment prevailed during the period of volcanic activity. The dust ejected during the volcanic activity was soon converted into mud, which coated the leaves of all plants and compacted and smothered the lesser vegetation. However, not one of the plants reported from the slope and summit vegetation has a completely deciduous habit. The vegetation is evergreen, and leaves persist for several years since apparently no abscission layer is developed. Portecop observed coatings 5-10 mm. thick on the leaves of Clusia (Ficuré 8) on July 26, 1977, after the first eruption. The acidity of this mud mass was certainly a factor in the defoliation and subsequent death of the plants. Heavy rains during the eruptions caused tremendous mud flows on the flanks of the volcano. At the present time many slopes of 30-45° have accumulations | meter thick, as revealed in profile in ravines where the old surface, compacted dead vegetation, and the new accumulations can be observed. The mud surface may havea rigid layer to | cm. thick, and transects of the flow show stratifications of silt and gravel. We observed that some plants of Pitcairnia, Anthurium, and Philodendron were exposed at the edges of ravines, where they were covered with layers of mud but were developing new growth on the ravine edge. A few plants of Machaerina that showed only the tips of leaves through the mud layer were excavated and found to be developing new rhizome offshoots that might in time penetrate the mud cover. However, ina flat area northwest of Savane & Mulets, Howard sank nearly three feet into a mire of mud when crossing a seemingly hard surface. In comparable open areas the current surface is depressed and is resilient underfoot as the mud-covered shrub layer, particularly the rhizomes of Philodendron and Guzmania, decays. Subsequent rainfall and surface runoff have sorted out areas of silt, gravel, and boulders. Hard-crusted areas resist colonization by plants, but silt and gravel zones now support seedlings that have germinated recently, There were no firsthand observations of the rain of boulders ejected during the eruption. In the Col de Echelle the former trail area is covered with piles of boulders and massive rocks, and the area is passable only by scrambling over and around the fallen rock (Ficure 5). To the south and west of the volcano summit, few new boulders can be observed, yet the effects of falling rocks are evident on the trees still standing. The tops of most woody plants are broken, and branches to 10 em. in diameter are seen broken at their junction with the trunk. Some horizontal branches Figures 12-15. 12, Didymopanax attenuatus, two adventitious shoots ee from marginal gone of injured stem, both shoots showing unusual bu devel- opment at base. 13, Charianthus alpinus, adventitious shoot development from base: upper branches have been killed, adventitious shoots have flowered and produced fruit. 14, Richeria grandis, trunks with multiple adventitious shoots of varying sizes developed from uninjured sections of trunk. 15, /lex macfadyenii, multiple adventi- tious shoots developed from base of unbranched plant: leaves of these shoots abnormal in shape and thickness. 1980] HOWARD ET AL., POST-ERUPTIVE VEGETATION Pew, are without bark on the upper surface. Bark damage is greatest on the side toward the mountain, indicating that the rain of rocks was directional. The bark may be stripped away for several decimeters. A few angular rocks, some to 24 cm. in diameter and a xX é BS 758 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 one estimated as weighing 4 kg., were found wedged in branches or in the crotches of standing trees. THE DEVELOPMENT OF ADVENTITIOUS SHOOTS From the combined effects of gas, acid, mud, and injury, the woody vegetation appeared to be completely dead, and the herbaceous vegetation eliminated. At present this is true of all plants of Clusta mangle: a number of other dead woody plants are unidentifiable. However, a few individual plants have had renewed growth after a period of leaflessness. Apical growth is infrequent and is restricted to plants of single stem habit (e.g., Philodendron). Rhizomatous plants or those forming clumps may have lost their original shoot apices when covered with mud and then developed new shoots from the base after being washed clean. Some trees and shrubs have developed adventitious shoots, but the variation in the patterns observed was not expected. Certain species developed adventitious shoots only near the apex of the broken shoots (Ficure 14). Others developed the shoots only at or near the base of the plant (Ficurr 15). The areas of injury where bark was removed (presumably by falling rocks) have, on some plants, begun to develop callus tissue although no adventitious shoot development has occurred. Other plants have developed adventitious shoots above the injury, below the injury (Figure 10), or on areas not associated with injury (Ficurrs 13, 14). Some woody species produce multiple adventitious shoots from a restricted area, suggesting the activity of multiple buds; others produce only solitary adventitious shoots. Regrettably, it is impossible to determine the age of the trunks or branches that develop adventitious shoots, and it is not Known if these are latent buds of considerable age that have been released. The multiple shoots may be of varying sizes. In some cases all shoots have developed equally in length and thickness, while in other cases a few of the multiple shoots have already exceeded the others in size. Examples were seen where the smaller or thinner shoots had begun to die. Horizontal branches or trunks that had extensive bark injury on the upper surface have often developed many adventitious shoots from the lower surface, opposite the injury, and a lesser number from the upper side in areas away from the injury. There seemed to be a tendency for the adventitious shoots from the lower surface (which had to curve around the trunk to assume an upright position) to branch from the base. This is in contrast to the unbranched adventitious shoots that developed on the upper surface. Species also varied in the nature of the growth and branching of the adventitious shoots. Didymopanax attenuatus was characterized by the development of swollen, almost subbulbous, bases to the adventitious shoots (Ficurr 12). Leaf scars were no more frequent in these swollen areas than farther up the stem. The lower internodes of some adventitious shoots were short, while in other species the first-formed internodes were contrastingly longer than later ones. Abnormal leaves with thickened petioles and blades of other than normal shape were found only on some specimens of Hex macfadyenii. However, there were contrasts in the type of leaf produced from the first or lower nodes. Smaller leaves (cataphylls with an early leaf fall) character- ized some species, while other species had normal leaves from the beginning of shoot development. 1980] HOWARD ET AL., POST-ERUPTIVE VEGETATION 759 In some species the growth of adventitious shoots is continuous, while in others it appears to be in units or flushes indicated either by sympodial apical growth below a dead bud or by longer internodes above a region of shorter internodal development with more proximate leaves (Howard, 1974). As many as three to five growth units were recorded for some adventitious shoots that had been produced within the past twelve months. REGROWTH OF SINGLE-STEMMED PLANTS The bromeliad Guzmania plumieri, the palm Prestoea montana, and the fern Blechnum violaceum all have tightly arranged leaves forming a rosette or crown. In the Guzmania this rosette normally holds water. During the volcanic eruption plants of all three species accumulated near the growing point quantities of acid mud that appears to have had lethal effects. Many plants are now seen without life and with the rosette of leaves well above the ground but still tightly packed with mud. Palms also have retained masses of mud in the apex and are dead, with only one seen in which the mud had been mostly washed away and new leaves were developing. A taxonomic problem remains regarding the bromeliad called Guzmania plumieri. Young epiphytic plants of this species do not develop a long stem system, but terrestrial plants of the same species seem to have been long lived and to have developed a leafy rhizome. In some plants this rhizome has been recorded over two meters in length; however, none of these plants was seen in flower, and no herbar- 1um record of G. plumieri refers to such growth forms. One long stem was washed clear of mud. Although the apex was dead, two adventitious shoots (small rosettes) had developed 17 and 32 cm. from the apex; these were green and were developing without evidence of adventitious roots. Climbing or rhizomatous bromeliad stems normally branch only immediately below a flower-producing rosette. This unusual example was marked and will be the subject of continued observation. Philodendron giganteum also produces a thick, climbing or scrambling stem several meters in length. Normally, it branches infrequently but develops feeding or holding roots along its entire length. Long stems of some plants of P. giganteum observed were broken as a result of the eruption; the upper detached end was observed to be living (Figure |1). Although the plant may have lost all its leaves, the renewal growth at its apex produced leaves of mature size. Many cases were seen where subapical fragments of stems, with roots, did develop adventitious shoots. These adventitious shoots were a fraction of the diameter of the original shoot and had produced leaves of reduced size, comparable to those of very young plants. No examples were found of new apical shoot development from rooted basal sections of an older plant. NOTES ON THE REGROWTH OF VARIOUS TAXA TREES Clusia mangle. The death of all individuals of this seemingly sturdy plant in the affected area was not expected. The standing trunks examined have no living bark, and no development of adventitious shoots is expected. The only example of survival 760 JOURNAL OF THE ARNOLD ARBORETUM [vov. 61 was a single shoot that had been depressed beneath a large stem of a Philodendron and protected by a mass of bromeliad rosettes. Green leaves attracted our attention, and upon excavation there appeared an example of natural layering. The stem system entering the ground was dead, but the apex had recently developed four pairs of leaves and emerged from the debris. Of interest was the development of adventitious roots immediately below the leaves as well as at the junction of the living and dead tissue, but with none for a distance of 30 cm. between (Ficurr 9). Cyrilla racemiflora. Trunks to 3 meters with a d.b.h. of 10.5 cm. were seen. Many adventitious shoots were observed; they developed from single buds from the base to a height of 2 meters on the trunk. Several adventitious shoots were producing flowers from the upper portions of a growth unit. Lateral branching had occurred from nodes immediately above the flowering area, while nonflowering shoots were unbranched. Didymopanax attenuatus. Standing trunks up to 3 meters tall with a d.b.h. of 8 em. were found. Stems of this plant seemed to be vulnerable to injury by falling rocks, suggesting a soft or succulent bark. Callus was developing rapidly and conspicuously around the injured areas, and the development of adventitious shoots was primarily adjacent to these areas. The shoots had conspicuously swollen bases and generally thin stems without growth units (Figure 12). One young inflorescence was developing. Ilex macfadyenii. This species, represented mostly by shrubs of less than 3 meters, is second to Clusia in frequency. However, two trees (4 and 6 meters tall) had diameters at breast height of 10 and 20 cm., respectively. Broken areas of bark were common, but callus formation was not conspicuous. Adventitious shoots were plentiful. pri- marily from the basal zone of even the largest trees (Ficurr 15). Many of the slender young shoots had retained cataphylls or reduced leaves at the first six or seven nodes. Abnormal leaves seen on several plants exhibited wider petioles, broader cuneate bases, and thicker blades than normal leaves. The shoots revealed three, four, and five growth units, the last in all cases being shorter than the others, with reduced leaf size near the apex and minute terminal buds. Adventitious shoots, which developed from the lower surface of damaged horizontal branches, were branched. Individual plants bore only staminate or pistillate flowers. but flowering on adventitious shoots was copious. The staminate flowers occurred in the axils of even the reduced leaves near the shoot apex. Pistillate flowers were not formed so close to the apex of the shoots. Only immature fruits were observed. Myrcia platyclada. Only one tree of this species was encountered. Truncated 6 meters above the ground and with a d.b.h. of 34 cm., it may have originally exceeded 10 meters in height. Adventitious shoots developed singly from supra- nodal areas at mid-height of the existing stem. The largest shoots seen were 14cm. long, without growth units, but with the younger leaves brightly colored, as_ is common in the Myrtaceae. The shoots were branched, but there was no evidence of flowering. Myrsine coriacea. Few trees have survived. The largest living tree was 2.5 meters in height; it had previously branched from the base and appeared shrublike. Advent- tious shoots were clustered, developing from multiple buds at the base to a height of 1980] HOWARD ET AL., POST-ERUPTIVE VEGETATION 761 30 cm. The lower nodes of the adventitious shoots had cataphylls, and the basal internodes were shorter than the upper ones. Sympodial branch development was observed on two shoots that had not yet flowered. Prestoea montana. Only one living plant of this palm was seen; it was | meter tall and had a flush of young green leaves. All other palms were dead, with the crowns and persistent leaf bases solidly packed with mud. Richeria grandis. Several large but badly broken trees remained standing: the largest remnant was 3 meters tall with a d.b.h. of 21 cm. Adventitious shoot development was abundant in the basal meter of the plants. Buds were multiple, but the shoots varied in size and smaller ones were dying (Ficure 14). Some adventi- tious shoots were produced singly near injured and callused areas. Weinmannia pinnata. This species is the third most common plant in the Clusia association. Trees generally were branched from the base, with the stems clustered. The largest plant still standing was 5 meters tall with a d.b.h. of 14 cm. Adventitious shoots were strongly developed from the current ground level but also occurred to a height of 3 meters. This was the only species that was developing adventitious shoots when we studied this area in 1978. The adventitious shoots were all clustered and of contrasting thickness. Areas of future shoot development along the stem were indicated by bright-colored multiple buds. Adventitious shoots showed three growth units; shoot growth is apparently halted by a resting terminal bud before development continues. The nodes at the immediate base of a growth unit were leafless, and the leaf scars were smaller than the bases of existing leaves. Several adventitious shoots were seen in flower, and young fruits were developing. SHRUBS Charianthus alpinus. One plant 3 meters tall with a d.b.h. of 6.8 cm. was seen. Others located were less than 2 meters tall and badly broken. Clustered adventitious shoots appeared to develop immediately below old and dead branches (Ficurr 10). No growth units were observed, although several shoots were in flower. Flowers were normally produced in cymes from the two apical nodes and also terminated the growth of the shoot. Subsequent branching developed from shoots produced in the two or three nodes below those that were flowering. The adventitious shoots had four or five short basal internodes, with the subsequent ones considerably longer. Palicourea crocea. Most existing plants were badly broken, suggesting that this is a plant of weak structure. Nevertheless, all plants of this species freely developed adventitious shoots, and all shoots were of one growth unit terminated by an inflorescence. Subsequent, seemingly dichotomous branching resulted from the growth of axillary buds at the base of the inflorescence or at an additional node immediately below SCANDENT PLANTS Hillia parasitica. Only one recognizable plant was encountered. It had a stem diameter of 2.6 cm. and was dead to near the base, where one adventitious shoot was developing. This shoot had a conspicuously swollen base similar to that of Didymopanax, and no growth units had been established. 762 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Norantea spiciflora. Plants of this semiscandent species seemed to have been buried in avalanches of mud but have recovered with remarkable frequency, One green shoot appeared to have developed adventitious roots above a dead area and was continuing growth. Other stems, 1-2 cm. in diameter, had sparse and scattered single adventitious shoots. The apices of all adventitious shoots showed the colorful flush growth of an elongated bud. The internodes of the adventitious shoots appeared to be shorter than those of undamaged plants, and the majority of adventitious shoots had developed clusters of adventitious roots from the lowest two nodes and internodes. It appears that the rapid shoot development awaits the establishment of this new root system. Philodendron giganteum. This is a very abundant plant with thick, scrambling or trailing stems. some to 15 cm. in diameter. The stems are apparently fragile, since most plants were broken. The apical meristem was not destroyed in the loss of leaves during the eruption, and new leaves have developed and produced blades of normal size (Figure 11). Sections of the stem without the terminal bud have developed slender axillary adventitious growth that produced leaves of smaller than normal size, No new root development was observed on the adventitious shoots. Symphysia racemosa. The majority of these scrambling plants had stems to 3.2 cm. diameter and were killed back to the current ground level. Adventitious shoots developed from the base of the plant from undamaged tissue. Scarred areas along the branches were developing callus but without indication of shoot production. The shoots had an average of six cataphyll units at the base, then abruptly large leaves, but ending with a series of much-reduced leaves. One shoot had three growth units recognizable by leafless areas presumably associated with cataphylls. Terminal growth was a flush of leaves, usually brightly colored, and normal stem extension was sympodial following the production of a terminal inflorescence. Several adventitious shoots had developed short lateral branches. SEEDLING INVADERS Volcanic activity of La Soufriére ceased in March, 1977. There are nonspecific reports of ‘‘seedlings present’’ for December, 1977. In March, 1978, we observed small but nonflowering plants of /schaemum, Machaerina, Pitvrogramma, Sauvagesia, and Viola. In April, 1979, we could recognize as invading plants mature, flowering, and fruiting specimens of Erechtites hieracifolia (L.) Raf., Ischaemum latifolium (Sprengel) Kunth, Phytolacca icosandra L., Sauvagesia erecta L., Ttbouchina ornata (Sw.) Baillon, and Viola stipularis Sw., as well as spore-producing plants of Histiopteris incisa (Thunb.) J. Sm. and Pityrogramma chrysophylla (Sw.) Link. We have no recollection from 1978 of Phytolacca or Erechtites in the area now devastat- ed, although both occurred occasionally as weedy plants near the parking area and along the trail. Phytolacca has multiseeded, fleshy fruits that might be transported by birds or on the shoes of visitors, while Erechtites, a composite, has plumed, wind-dispersed fruits. Eleocharis maculosa (Vahl) Roemer & Schultes (Cyperaceac), which was observed in fruiting condition, currently occurs in stream beds within the devastated area but probably existed there prior to the eruption. Seedling plants of Charianthus alpinus were seen flowering precociously with only two pairs of leaves and a total of four nodes present above the persistent cotyledons. Calolisianthus 1980] HOWARD ET AL., POST-ERUPTIVE VEGETATION 763 frigidus was also seen flowering as a very small plant. In the undisturbed shrub zone this species may reach 1.5 meters when in flower, while on rock cliffs the plants are known to flower when only 12 cm. tall. Didymopanax attenuatus seedlings occurred in clumps, due in some cases to the germination of several seeds in one fruit or the result of many fruits having been deposited in one place. Other species present as single plants—not clumps of seedlings—were Anthurium, Ilex, Palicourea, Sauvagesia, Viola, and Weinmannia. Seedlings of Miconia were infrequent but could not be determined to species. myriad of seedling rosettes of Guzmania could be distinguished from the less frequent seedlings of Pitcairnia. Only Guzmania has wind-dispersed seeds. Young sporophytes of Pityrogramma of both silver and gold color forms were more numer- ous than those of Blechnum violaceum. Young plants of Machaerina may develop only one distichous fan of leaves or proliferate early from the base, forming several fans of equal size. Of the nine taxa of woody plants normally associated in the area dominated by Clusia, only two were represented in the seedling populations. These—//lex an Didymopanax—have fleshy fruits. Although Clusia, Hedyosmum, Myrcia, and Myrsine have a fleshy fruit structure that would presumably aid distribution, they were not found as seedlings. Cyrilla and Weinmannia (both with small, dry seeds) and Richeria (with explosive fruits and large seeds) were not present. Clusia, Hedyosmum, and Richeria are dioecious taxa. The palm Prestoea montana has not been seen in fruit in the flank area of the volcano, and it is possible that existing plants are from past casual introductions of the fleshy fruits by birds. No seedlings have been seen. The shrubby species outside the Clusia associations include two species of Charianthus, two of Miconia, and one each of Besleria, Freziera, Norantea, Psychotria, Symphysia, and Ternstroemia, all of these taxa have fleshy and many- seeded fruits commonly red or blue in color. Only Miconia sp. and Charianthus alpinus are among the invading species of open areas. Rondeletia has small, light seeds, and Baccharis, a composite, has wind-dispersed fruits, but neither was found as a seedling. Of the epiphytes of the flank areas, Hillia parasitica has comose seeds, and Psychotria guadalupensis has black, fleshy fruits. Neither species was encountered as a Seedling. Of the herbs, only Guzmania (with plumose seeds) seems to have a specialized dispersal mechanism. Calolisianthus, [sachne, Machaerina, Sauvagesia, Tibouchina, and Viola, as well as Juncus, Lobelia, and Pterolepis, have numerous small, dry seeds. Viola seeds may be holoballistic, but for short distances only. The first six genera were represented by seedlings in mud flat areas open to colonization, often considerable distances from the nearest mature plants. e dominant plant of the ‘‘Pitcairnietum guadelupense’’ (Pitcairnia latifolia) has red-colored fruits that do not detach readily and that presumably are not distributed by animal vectors. Its abundance at the exposed higher elevations suggests that any seeds freed from the fruits could be carried by the wind to the mud flats where seedlings are now found. The dispersal of the very light spores of Blechnum, Histiopteris, and Pityrogramma is also easily understood. Gaultheria swartzii, found on rocky areas, has fleshy, black, many-seeded fruits but has not yet established itself as a colonizing plant. 764 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 The slopes of La Soutri¢re are within the Pare Naturel de Guadeloupe and are under the care of the Office National des Foréts. Additional study plots have been established, and future changes will be observed and reported. BIBLIOGRAPHY Howarp, R. A. 1962. Volcanism and vegetation in the Lesser Antilles. Jour. Arnold Arb. 43: 279-314. 1974. The stem-node-leaf continuum of the Dicotyledoneae. /bid. 55: 25- SASTRE, C. 1978. Plantes menacées de Guadeloupe et _ Sard I ees Bull. Mus. Hist. Nat. Paris II. 42: ¢ STEHLE, . Flore de la Guadeloupe et ene A ene botanique. 284 pp 1979 Especes . Bssai d’éecologie et de . Gouvernement de la Guadeloupe, Basse-Terre. Premicres sbservations sur la reconstitution du tapis végétal sur le volcan de la Soufriere de Guadeloupe apres les éruptions de juillet-aout 1976 et mars 1977. Bull. Soc. Bot. France 126: 349-359. R.A ARNOLD ARBORETUM Harvarb UNIVERSITY 22 Divinity AVENUE CAMBRIDGE, MASSACHUSETTS 02138 P.M. J. P. CENTRE UNIVERSITAIRE ANTILLES GUYANE, GUADELOUPE Orrick NATIONAL DES FORETS Basse-TERRE, GUADELOUPE 1980] HOWARD & CLAUSEN, SOUFRIERE PLANT 765 THE SOUFRIERE PLANT OF ST. VINCENT RICHARD A. HOWARD AND KRISTIN S. CLAUSEN Tre SOUFRIERE PLANT (Spachea elegans (Meyer) Juss.; originally S. perforata Juss.) was collected by Alexander Anderson along the Essequibo River in British Guiana in 1791 and introduced into cultivation in the botanical garden of St. Vincent. The following notes should end the saga of a species considered extinct in the wild but preserved in a botanical garden. How the story became a legend of the Caribbean area Is not clear. For many years the tale has been repeated that Spachea perforata had been a native of the slopes of the Soufri¢re on St. Vincent, and that all native plants had been destroyed by the eruption of that volcano in 1902. A single supposedly remained under cultivation in the botanical garden near Kingstow hen the senior author visited St. Vincent in 1950, the location of the plant in fie garden was not known, for most specimens in the garden were without labels. Two trees belonging to the Malpighiaceae were found in flower; herbarium specimens were made and determined, and the garden plants were again labeled. In a history of the St. Vincent Botanic Garden (Howard, 1954) the tale was published that, accord- ing to the records of the garden, these plants had been grown from seed obtained in the mountains. Extensive general collecting on St. Vincent by H. H. and G. W. Smith, John and Pamela Beard, Conrad Morton, George Proctor, R. A. Howard, and Keith and Elizabeth Laidler has not turned up a single native specimen of the Soufriére tree. More recently, Keith and Elizabeth Laidler (1975, 1978) became interested in propagating and reestablishing the species in the mountains of St. Vincent. These authors (1975) stated, *‘The eruption of Mt. Soufriére in the last century practically wiped out this arboreal species, which used to grow in relative abundance on the slopes of the volcano. The mature thirty-foot specimen now thriving in the Gardens is the only cutting that grew successfully from the single sapling which was discovered on the mountain after the catastrophe.’’ The Laidlers sent flowering specimens to Kew to verify that the plant was hermaphroditic. They also rooted cuttings of the tree, planted them in several places on the island of St. Vincent, and speculated on the possibility that the plant might still be discovered in the remote areas of the northwestern corner of the island. In the final report of their work it is admitted that, lacking care, the transplants all died. During the summer of 1979, Gavin Bridson, Librarian of the Linnean Society in London, called to the ane on of the senior author a large and unappreciated manuscript of a ‘‘Hortus’’ of the West Indies written by Alexander Anderson in approximately 1803. eee whose life story is being compiled by the senior author, was appointed director of the St. Vincent Botanic Garden in 1783 and served © President and Fellows of Harvard College, Journal of the Arnold Arboretum 61: 765-770. ee 1980. 766 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 in this capacity until ill health forced his resignation in 1811, a few months before his death. It was he who received and nurtured the plants, including several varieties of breadfruit, delivered from the South Seas by Capt. William Bligh. With William Lochhead, who was to succeed him as director of the Garden, Anderson made expeditions to Trinidad and Guiana to secure plants for the Botanic Garden. Anderson's manuscript described in detail the plants under cultivation in the garden and their sources, uses, and culture: it included many descriptions of plants new to science, as well as numerous exquisite drawings. The manuscript was sent to William Forsyth at Kensington Gardens and was intended for publication. Among the drawings was a colored plate of **Cullenia malpighioides,’*” supplemented in the text by a complete Latin description of the new genus and species. The name honored a Dr. Cullen of Edinburgh, who was the first in Scotland to use the Linnaean system in his teaching. Anderson reported that in 1791 he saw the plant some 200 miles up the Essequibo River, collected seeds, and secured some small plants to return to the garden on St. Vincent. When they flowered, he prepared the description and had the plant sketched. Although ‘‘Cullenia malpighioides’’ antedates the genus Spachea, the name was never validly published. In 1818 G. F. W. Meyer described Malpighia elegans as new, based on an E. K. Rodschied specimen collected on the shores of the island of Arowabisch in the Essequibo River. The original specimen is preserved at Gottingen. In 1824 de Candolle transferred the species to the genus Byrsonima, and in 1838 A. Jussieu created the new genus Spachea and published an illustration. Both de Candolle and Jussieu cited a collection of Leschenault from Surinam, now preserved in Paris and there indicated incorrectly as the type. In 1840 Jussieu added to the genus Spachea parviflora and S. ossana, trom Cuba; S. perforata, from St. Vincent; and S. tricarpa, from Brazil. All of the species were described, but without the citation of specimens. Longer descriptions were published in 1843 (Jussieu, 1843), and collections were cited. Most critical characteristics used by subsequent scholars seem to be derived from Jussieu’s descriptions. Spachea parviflora, based on a specimen in “*herb. DC."* was credited as “‘In insula Cubae juxta Habannam legit cl. de la Ossa.”’ Spachea ossana, based on specimens in ‘herb. DC, Hook.,’’ was described with the note ‘In Ins. Cubae juxta Habbannam legit cl. de la Ossa (herb. DC), et alterum specimen ex Ins. S. Vincentii exhibet herbarium Hookerianum cum S. perforata confusum.’* Spachea littoralis (‘herb. Mus. Par.’’) was attributed thus: ‘*Ad littora maris in Insula S. Thomae legit cl. Finlay.”’ Finally, S. perforata (herb. Hooker) was reported as ‘Specimen in Ins. S. Vincentiit lectum communicavit cl. Hooker.—Adsunt in herb. Lessert specimen ibidem acl. Caley lecta, foliis paulo majoribus, tenuioribus et omnino membranaceis, minus manifeste punctatis paulisper distincta.”’ Each of these species requires a To comment for clarification. José Antonio de la Ossa became the first director of the Jardin Botanico de la Habana on May 30, 1817. Leén (1918) referred to the fact that Ossa had prepared a work listing the uses of plants and their common names and had also included plants recently introduced from other countries and acclimatized there. The death of Ossa, which occurred shortly afterwards, frustrated the publication of his work. This manuscript has not been studied in detail, and it is not known if it exists in Cuba 1980] HOWARD & CLAUSEN, SOUFRIERE PLANT 767 today. It is clear that Eupatorium ossaeanum DC. (de Candolle, 1836), with a type ‘de la Ossa, Havana, Cuba,’’ is an endemic species of St. Vincent. It is probable that Spachea parviflora and S. ossana, attributed to Cuba, are plants obtained from the botanical garden of St. Vincent. A similar error was made by Jussieu in reporting Spachea littoralis was collected by Finlay from St. Thomas. Britton and Wils (1924) noted that *‘Spachea littoralis A. Juss., described by A. Jussieu as from St. Thomas, collected by Finlay, was really collected in Trinidad.’’ R. O. Williams (1947), who with the critical help of N. Y. Sandwith prepared the treatment of the Malpighiaceae for the Flora of Trinidad and Tobago, accepted only S. elegans and stated, *‘S. perforata Juss. recorded from St. Vincent and cultivated in the Royal Botanic Gardens, Trinidad, appears to be the same species.’’ In a later section Cheesman (1947) referred to Niedenzu’s work but noted, ‘‘A synonym of the latter |S. parviflora Juss. var. perforata (Juss.) Niedz.] is S. littoralis Juss., which was based on Finlay 198 from Trinidad, but this collection may have been from a cultivated plant.”’ The collectors of the material Jussieu cited for Spachea perforata were Guilding (for the specimen sent to Hooker and now in the herbarium at Kew) and Caley (for the specimen in the Lessert Herbarium (rp). Reverend Lansdown Guilding wrote a history (1825) of the botanical garden of St. Vincent that included Anderson’s list of the plants growing in the garden in 1806. **Cullenia malpighioides’’ is not included in this list, but neither is any species of Spachea. According to Guilding’s letters now in the archives at Kew, Guilding did send many specimens and drawings of plants in the St. Vincent Botanic Garden to Hooker for identification. George Caley, the director of the Garden from 1816 to 1819, commented in letters also preserved at Kew that Anderson neither numbered nor labeled the plants of the garden, making it necessary to send specimens to various people for identification. Grisebach (1859) recognized both Spachea elegans, from Trinidad (Purdie) and Guiana, and S$. perforata, from St. Vincent (Guilding). In the synonymy of S. perforata, Grisebach listed ‘‘Ossana Vincentiana, Juss.’’ with an exclamation mark, indicating that he had seen material. This name has not been publis Two monographs of the Malpighiaceae that treated the genus alae were published subsequently by Small (1910) and Niedenzu (1928). Small recognized four species: S. parviflora, from Cuba and St. Vincent: S. /ittoralis, from St. Thomas; S. perforata, from St. Vincent: and S. ossana, from Cuba and St. Vincent. No specimens are cited in treatments for North American Flora, and the lack of representative material in the herbarium of the New York Botanical Garden, where Small worked, suggests the descriptions and distinguishing characteristics were derived from the literature. Niedenzu treated these four species in a different manner. He accepted S. elegans, including his previously established forma ossana (Juss.) Niedz. (1914) (typified by the Ossa collections ‘‘ca. 1825: Havana’’), and S$ parviflora Juss. (typified by Ossa ‘‘ca. 1825: Havana’’), consisting of varieties typica and perforata (Juss.) Niedz. (1914). He cited S. /ittoralis in the synonym the species S. parviflora, but not under either variety typica or variety perforata. Niedenzu did cite Finlay 198, from Trinidad, along with material of Caley an Guilding from St. Vincent under his variety perforata. Recent collections from Trinidad and oties South America, as well as six different collections from the 768 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 trees in the botanical garden on St. Vincent, reveal that the distinctions between S. elegans and S. parviflora as used by previous workers are not reliable, and that only one species is represented. There remains a problem of the fertility or sex of the flowers of Spachea and particularly the ‘‘Spachea perforata’’ of the St. Vincent Botanic Garden. William Anderson, of the University of Michigan, a current specialist on the Malpighiaceae, states (pers. comm.), ‘Most Malpighiaceae have monotonously bisexual flowers, so it Was a great surprise to discover that Spachea does not. Some plants are function- ally pistillate, while the flowers on plants that produce pollen are morphologically perfect but perhaps functionally staminate, i.c., they do bear ovule-containing car- pels, but I suspect that those carpels do not set seed, since | have found no enlarging fruit on old inflorescences of those plants. These observations are based only on material of S. elegans and S. tricarpa.’> An examination of six collections of ‘‘Spachea perforata’’ made between 1825 and 1980 from the botanical garden in St. Vincent, as well as some collections from plants in Trinidad propagated from the St. Vincent plant, reveals anthers with pollen and carpels with ovules, but no specimen has fruit. The records of the St. Vincent Botanic Garden were examined by Dr. George Cooley in 1962. He noted the entry for 1912: °‘A large number of cuttings made of the old Spachea perforata, about a dozen appear to have rooted. These will be distributed when strong enough. No recent record of seed produced though it flowers profusely.’ An entry for 1915-16: “‘Spachea perforata showed signs of decay. A limited number of young plants have been raised, one of which was planted out in the Garden.”’ In 1918-19: ““Spachea perforata. Young specimen to replace old dying tree making good growth. No seeds produced in recent years.”* In 1929: “The old plant of Spachea perforata which was regarded as dead nine months ago produced a strong shoot from its base during the year. Probably planted by Anderson 130 years ago. In 1912 rooted cuttings from this tree were sent to various botanical establishments in the West Indies. One was planted in that part of the St. V. gardens known as Nutmeg ground. Now a healthy tree, 20 ft. in height.” These records certainly suggest that the St. Vincent plants are functionally staminate and perhaps date the cultivated plants in Trinidad from which specimens in various herbaria have co. — jos been taken Small (1910), in a key to the species, distinguished Spachea parviflora as having ‘stigmas truncate’’ and noted “fruits not seen.”’ According to Small, S$. liftoralis, S. perforata, and S. ossana have “stigmas slightly 2-lobed or reniform-capitate.”” He did not see the fruits of §. /itroralis but did record fruit size for S. perforata and S. ossana. The lobed stigma characteristic may well be correlated with functional carpels. As we have indicated, no existing specimen of S. perforata—considered to be endemic to St. Vincent and known only from the plants in the botanical garden—is known in fruit, and the source of Small’s descriptive detail of the fruit is a mystery. However, examination of the type material of S. ossana Juss. (DC.) reveals one sterile, leafless shoot, one leafless shoot with an inflorescence, and one leafless shoot with a small, unattached leaf trapped in a fruiting inflorescence. It is possible that Anderson sent to Ossa in Cuba some plants of the original 1791 introduction of his ‘*Cullenia malpighioides,’’ one of which was functionally pistillate, while retaining only functionally staminate plants on St. Vincent. The material of S. parviflora 1980] HOWARD & CLAUSEN, SOUFRIERE PLANT 769 Juss. (DC.) attributed to ‘“‘La Havanna, de la Ossa 1825°° is a leafy shoot with functionally staminate flowers, the anthers of which bear pollen. With regard to both S. parviflora and S. ossana, Leon and Alain (1953) stated, ““No observada por los botanicos modernos.”’ The legend of the Soufriére tree must be discounted, and the plant in the St. Vincent Botanic Garden must be recognized as Spachea elegans (Meyer) Juss., with the other taxa considered in this paper as synonyms. The synonymy of S. elegans is presented below. Spachea elegans (Meyer) Juss. in Delessert, Ic. Selec. PI. 19. ¢. 37. 1838. BasionyM: Malpighia elegans Meyer, Prim. Fl. Esseq. a. 18 Byrsonima elegans (Meyer) DC. Prodr. 1: 580. 1824. Spachea parviflora Juss. Ann. Sci. Nat. Bot. ser. 2. 13: 326. 1840; Monogr. Malpigh. 72. 1843. Type: Ossa s.n., Cuba (G-pc). Saute ossana Juss. Ann. Sci. Nat. Bot. setae “B: 326. 1840; Monogr. Malpigh. 184 PE: Ossa s.n., Cuba (G1 ee bers Juss. Ann. Sel. Hy Bot. Vincent (Herb. Hooker, k). Spachea littoralis Juss. Arch. Mus. Paris 3: 328. 1840. Type: Finlay s.n., ‘St. Thomas” (in error, really Trinidad) (P). Ossana vincentiana Juss. ex Griseb. in Griseb. Fl. Brit. W. Indian Is. 116. 1859, ser. 2. 13: 326. 1840. Tyee: St. — in synon. Spachea elegans f. ossana (Juss.) Niedz. Arbeit Bot. Inst. Lyc. Hosianum 5: 914. Spoehed parviflora var. typica Niedz. Ibid. 1914. Spachea parviflora var. perforata (Juss. ) oe Ibid. ACKNOWLEDGMENTS We are grateful to the curators of the herbaria mentioned for the privilege of cimens on site or by loan, to the officers of the Linnean Society for the “and to the library staff at Kew for Vincent Botanic examining spe opportunity to transcribe the Anderson **Hortus.’ their help in locating the letters of previous directors of the St. Garden LITERATURE CITED Britton, N. L., & P. Witson. 1924. Spachea littoralis. P. 448 in Scientific survey of Porto Rico and the Virgin Islands. Vol. 5. New York Academy of Sciences, New York. CANDOLLE, A. P. pe. 1824. eee Prodr. 1: 580. . 1836. Cae Ibid. 5: |: CueesMan. E. E. 1947. Spachea. P. 499 in Flora of Trinidad and Tobago, additions and corrections. Vol. |. Guardian Commercial Printery, Port-of-Spain Grisepacu, A. H. R. 1859. Flora of the British West Indian Islands. xvi + 789 pp. Lovell Reeve & Co., London. Guitpinc, L. 1825. An account of the botanic garden in the island of St. Vincent. 47 pp. Richard Griffin & Co., Glasgow 770 JOURNAL OF THE ARNOLD ARBORETUM [voL. 6] Howarp, R. A. 1954. A history of a botanic garden of St. Vincent, British West Indies. ee Rev. 44: 381- Jussieu. A. H. L. de. 1838. Malpghicese. P. 19, ¢. 3/ in J. P. B. Devessert, [cones selectae plantarum. Vol. 3. Fortin, Masson et Soc., Paris. ———. 1840. een synopsis. Ann. Sci. Nat. Bot. ser. 2. 13: 326. 1843. Monographie des Malpighiacées. Arch. Mus. Hist. Nat. Paris 3: 5-151, 265-616. Laipter. K., & E. Laipter. 1975. Synopsis of the results of the 1975 Durham Univer- sity expedition to St. Vincent. Carib. Conserv. News 1(3): 9, 10 1978. Synopsis of the results of the 1975 Durham University a to St. Vincent. /bid. 1(13): 19, 2 Leon, H. 1918. Las exploraciones botanicas de Cuba. Memorias de la Sociedad a 48 pp. Sociedad Editorial Cuba Contemporanea, Havana. — & H. Alain. 1953. Spachea. P. 16 in Flora de Cuba. Vol. 3. Contr. Ocas. Mus. Hist. Nat. No. 13 Meyer. G. W. I8I8. Malpighia Ailey Pp. 178, 179 in Primitiae florae Essequeboensis. H. Dieterich, Gottin Niepenzu, F. 1914. Malpighiaceae americanae. Arbeit Bot. Inst. Braunsberg 3: 18, 19, TT) ——. 192 eo In A. Encter & L. Diets, eds., Pflanzenr. IV. 141111): SMALL, J. K. ut Malpighiaceae. N. Am. FI. 25(2): 149, 150. Wituiams. R. O. 1947. Malpighiaceae. P. 449 in Flora of Trinidad and Tobago. Vol. 1. Government Printer, Port-of-Spain. ARNOLD ARBORETUM Harvarp UNIVERSITY 22 Diviniry AVENUE CampribGe, Massacuusetts 02138 1980] INDEX 771 INDEX Abella, Acronychia ornata, 24 — eats 89 Actinidia arguta, — uniflora, 89 — callosa, 83 Abies, 15. S51, 64. 65 — chinensis, 83 Abramsia, 95, 96, 101 — coriacea, 83 — trichotoma, [0 Actinidiaceae, 83 Acanthaceac. 88 aac ee 50 Acanthopanax, Eleutherococcus vs.. 107-111 = — cupularis Acanthopanax, 107-1 denocaulon adhaerescens, 90 — sect. Eleutherococcus, 109 Adiantaceae, 71 — divaricatus, 109 Adiantum Sin 71 — evodiifolius var. gracilis. 111 culus, 6 — giraldii var. pilosulus, 109 — wilsc — gracilistylus, 109 erred pilosa, 78 — ir. nodiflorus, 109 Ailanthus — — var. pubescens, 109 — altissima, 80 — — var. villosulus, 110 Ainsliaea fulvipes, 90 — henryi var. faberi, 109 Atrosperma Dea ers Notes on, with a New — lasiogyne, 109 Species from Fiji, 95-105 — nodiflorus, 109 Airosperma, 95—1( — pentaphylla var. variegatus, 110 — fuscum, 96, 99-101 — phanerophlebius, 10 — grandifolium, 96, 97, 99 finer l — psychotrioides, 95-101 — seoulensis, 109 — ramuense, 95-97, 100, 101 — sessiliflorus, 110 — trichotomum, 96, 97, LOL, 104 — — var. parviceps, 110 — vanuel 96, 97, 102-104 — setulosus, 110 Aizoaceae — sieboldianus f. variegatus, 110 Alangiaceae, 84 — spinifolia, 110 Alangium, 9, 65 — eee 107, 11 — chinense, 8 —— pubescens, 109 — handel, 49, 84 — — senopi f. angustissimus, [10 Albizia, 65 — f. dilatatus, 110 — kalkora, 80 — ternatus, I11 Aldrovanda a > =z trifoliatus, 110 Aleurites — villosulus, 110 — for dii — ward, 110 Alismatac 0 — yu, I1l Ainiahy tare: megaphyllum, 86 Acer, 50, 65 Alnus, 65 — amplum, 82 crematogyne, 50 cordatum, 82 Arcarapihncens, 51 — davidii, 82 Amelanchier, 65 — erianthum, 82 Ampelopsis chaffanjoni. 83 2 Amphicarpaea edgeworthii, 80 — ry, 82 Anacardiaceae, 81, 746, — longipes, 7 82 Anaphalis margaritacea, 90 — mono, 49, Anemone hupehen — a Nea 82 Angiospermae — sinense, &2 Antenoron neofiliforme. 74 — wilson, 82 Anthurium, 63 Aceraceae, 82 Apocynaceae, 87 Achlys, 56 nega 0 81. 732, 752 Acronychia. B52 Araceae, 56 772 JOURNAL OF THE ARNOLD ARBORETUM Aralia bodinieri, 84, 85 caisneana, 84 ete, 49, 69, 84, 726, 728, 729, 735, , 747, 752 dt ieen Arthromeris ei 71 pear ane Aspidiac Acidum ene TI Astilbe chinensis, 77 Athyriaceae, 7 Aucuba chinensis f. angustifolia, 85 oC ~~ Baccharts, 763 pedunculata, 752 Balsaminaceae, 82 egonia evansiana, 84 Belamcanda chinensis, 91 Betula, nee sinensis, 73 — alnoides, 73 fargesil, — luninifera, 73 wilsoniana, 73 Betulaceae, 50, - tribe Carlene: 739 Bicnoniscse: 8s Belchnum, 763 violaceum, 754, pean \ oT i “80 Botryopleuron nervosum, 88 Brandon Lignite, Fruits and Seeds of the. Rutaceae, 0 Bromeltaceae, 754 Broussonetia papyrifera, 74 Buddleta davidit Bulbostylis capil 9] Burseraceae, 747 Buxaceae, ¢ Buxus, 64 — microphylla, 81 ~J = = [voL. 61 Byrsonima, 766 elegans, 769 California, Dicotyledonous Wood from the Upper Cretaceous of Central, I], _ 748 Callicarpa bodiniert var. giraldi, 8 — rubella var. hemsleyana, 87 Calolisianthus, 763 — trigidus, 754, 763 Calophyllum (Guttiferae), Yd World Species of, 117—6% Calophyllum, 117-699 (for complete listing, see 692-699) v A Revision of the 7-699 — sinensis, 83 Campanulaceae Campanumoea javanica, 90 SLL Och acuminata, 50 Caprifoliaceac, 89 Cara giganteum, 91 Carex brunnea, 9 Caresium aeptacties 90 Carpinoxylon ostryopsoides, 739 Carpinus, 73 fargesit, 73 Carpolithus, 6 — nitidus, 5, 6 Carrierea calycina, 84 Carya, 65 Caryophyllaceae. 51 Castanea, 49, 65 henry, 73 mollissima, 73 Castanopsis, 65, 732, 739 chinensis, 73 —eyret, 73 — hystrix, 7 — sclerophylla, 73 — ubetana, 73 C asuarina, eee 733 0) 2 = ft ae = ne ios ie) ioe & — S i) oe on i y — labialis. 74 Cephalotaxaceae, 52, 72 1980] INDEX Cephalotaxus, 52, 53, 56-58, 65, 70 — fortunei, 56, 57, 72 = naprinetonid: oe — — var. drup , 30% 57 — — var. hing 56, 57 — — var. sine 7 — oliveri, 36-5 3 Ceratozamia, 714 Cercidiphyllaceae, 74 Cercidiphy! lum, 65 — japonicum, 74 es .8 Cestrum (Solanacea | from a Colombia. A New Spec Cestrum, | a 15 a ee 11S orae, 114 Geen 50 — cathayensis, 78 Chamabainia cuspid ata, 74 Ww 3 752, 755, 756, 761, 762 52. Chen nopadiaceae, 50, 51, 56 Chimonanthus praecox, 75 Chimonobambus, Chirita bicornuta, 88 Chloranthaceae, 72. 752 Chloranthus henryi, 72 oo 64 — axillaris, et base 68 — inunctum var. albosericeum, 76 6 COC — mairei, — platyphyllum. 76 oe erubescens, 84 Citr Clacvastis, 65 CLAUSEN, KRISTIN S., and RICHARD A. Howarb. The Soufriére Plant of St. Vincent. 765—770 Clematis, 65 ie coon 75 — meyeniana, — montana, 7¢ — spooneri, 75 — — var. spooneri, 75 subglabra, 74 Clerodendron bungei, 84 — mandarinorum, 87 — trichotomu eeeiGin 65 Clethra — fargesi, 85 ee 85, 732 i. 751-753, 756, nA ol. 763 — mangle, 752, 755, 758-766 Cocculus, 65 — see sare Colombia, aceae Combretac is 773 w Species of Cestrum (Solan- 5 te » Souther, 113-115 *) Comm ee. poate & 9] Commelinaceae ie Compositae, 49, 90, 736, 752, 754 Coniferae, a Coniferopsic 5, 02 Congr eee 70 _ 87 — kousa var. chinensis, 85 — henryi, — platypetala, — sinensis, Corylus farges 78 ii, 73 Cotoneaster eracilis, 78 65 Cretaceous of Central California. donous oe from the Upper Cudrania sta! 1 — ene 2? 78, 735, 736, phe — sempervirens, 56 osa, 56 Cuscuta chinensis. 50, 87 2 Cycads, Taxonomy of the West Indian, 701 2( Cycas circin tails — revoluta Dicotyle- . H, 723- 752 748 -722 774 JOURNAL OF THE ARNOLD ARBORETUM Cycas rumphii, 721 Cyclosorus aridus, 71 — parasiticus, 71 Cymbidium pasiioliint: QI Conencnum ete 87 54, Cyrilla, 763 dunes 752, 760 Cyrillaceae, 752 | in fortunel, 7 Cyperaceae, 49, 91 Cyperus orthostachys, 9] Cypripedium japonicum, 91 “ Dalbergia, - — dyeriana, — ee S. s Daphniphyllaceae ) Bae an g laucesce ns, 81 | ngisty 81 DARWIN. — VEN VP. Notes on Airosperma (Rubi aceae), with a New Species from Fiji, 95—105 Daucus carota, 85 : valien, 84 ease is as Dichroea febrifuga, 50 — — var. glabra, 77 — longipila, 7 Dicoty edoneae . 49, 72-90 Dicotyledonous Wood from the Upper Creta- s of Central California, bicranopter pectinata, 752 Dictamnus, Didissandra sesquifolia, 8 S = Se san ake bicornutus, 88 — hw 88 ee Stic _ 88 Diack, 761, — attenuatus Dillenta, 728 Dilleniaceae, 726, 728, 730 Dioscorea cirrhosa, 91 — kamaonensis var. fargesil, — japonicum = YY < = = = S = = Dryopteris fuscipes, 71 — 16: q52. a 758. 760, Ebenaceae, 86 Ec *KENWALDER JA [voL. 61 MES E. ao of the West Indian C eyes henryi, 49, — schnabeliana, viridis, 701- 84 Sec ag 69, 83, 735 Elaeocarpus Elatostema sessile, Eleocharis congesta subsp. chinensis, maculosa, 83 762 Japonica, 9 Eleutherococcus vs. oo inax, LO7-111 Eleutherococcus, divaricatus es — giraldil var. = pilosa, 09 yne, — nodiflorus, — rufinervis, — senticosus, gracilistylu = Xo 109 — phanerophlebius. 109 109 seoulensis, 1¢ — seca ee SUS, 107 9 109 — sicboldianus f. f: alates. 10 . parviceps, 110 10 varicgatus, 110 Emimenopterys henryl, 89 Engelhardt 733 E nkianthus c hinensts, 85 Ereaeeae 49, ~ serrulatus echtites, — cavalerei Epilobium ce ciietion: S4 Er 762 0 "85, 86, 726, 754 Eriocaulon buergerianum, 91 Escalloniaceae, Eucommia. 49 — ulmoides, 49, 50, 74 1980] Eucommiaceae, i dia . 17, 28, 29, 68 — me teas 4,9, 49 a ee 4, 80 Euonymus, — acanthocarpa, ar. eae. 82 — japonica var. pout 82 — myriantha, 82 — sanguinea var. oo 82 Eupatorium chinense, ee ( — ossacanum, 767 Sens 49, 68, 80, 81, 738, 752 Eurotia, 56 Eurya, 50, 69 — nitida var. aurescens, 83 Fee : — japonica, 82 Bvedicpata. 108 — scene 84 racilis, 111 ei 64, 65 Fagaceae. 49, 56, 73, 732, 733, 735. 736, 739 — longipetiolata, 73 — lucida, 73 Ficus, 68 — heteromorpha, 74 — lacor, 74 — sarmentosa, 74 — stapfii Fiji. nate on Airosperma (Rubiaceae), with a New Species from, 95-105 oe 129 Flora, The Metasequoia, and Its Phytogeogra- hi c Significance, 41—9¢ Fokienia, 65 ‘Fraxinu Freziera, 763 INDEX fit ps) Freziera undulata, 752 Fruits an _ ee of the Brandon Lignite, V. Rutace —40 Galium bungei, 89 Ginkgoaceae, 71 Gleditsia macracantha, 80 Glochidion pubescens, 81 Glyptostrobus, 64, 65 Gordonia, 64 The Post-Eruptive Vegetation of La Soufriére, —764 Guttiferae: Calophyllum, A Revision of the Old or sae 2 17-699 Guttiferae, 83, 132 pace a duss ~~ & i. 763 ee ae 754, 759 Gymnocladus, 65 — chinensis, 80 Gymnogrammaceae, Gymnospermae, 49, 3 12 Gymnostemma esa lum, 90 Hales ee 78, 73 Hedera nepalensis var. sinensis, 84 Hedyosmum, 763 — arborescens, Hedyotis lindlevaria var. elwingia. 5 ine) glabra, 89 — japonica, 8° Hema cuneata, 88 — procumbens, 89 eee um esquirolii, 85 Heteromeles —1 Houttuynia, 58 — cordata, 72 eeaa Beedle 82 Howarp, RIc A., and KRISTIN S. CLAUS sufrigre oa of St. Vincent, 765— 770 ARDA., JAC QUES fF rECOP, and PIERRE DE MONTAIGNAC. The P yst-Eruptive Vegetation of La Soufri¢re. Guadeloupe. 776 JOURNAL OF THE ARNOLD ARBORETUM 1977-1979, 749-764 Hu. Suu YING. Eleutherococcus vs. Acanthopa- nax, 107-111 Hu, Suiv YING. The Metasequota Flora and Its ea aes ie ea 41-94 HUNZIKER, ARMANDO T. A New Species of aan (Solanaceae) ae Southern Colom- bia, 113-115 Hydrangea, 64, 65 — patulum, 83 — petiolulatum, 83 Idesia, 65 —=“pOnyeatns 50, 84 ar. vestita, flex, 9. S50. 64, 65. 732. 763 = dealer var. cc a S| a . lichuane $i ~ elmerilliana. “a - 1, 81 — macfadyenii, 7 752, 756, 758, 760 —pernyl, $ i — amblyar ie 8 — pseud ae 80 — Jatifolium, 162 Itea, 65 Ixeris gracilis, 90 asminum, 69 — lanceolarium, - Juglandaceae, 50, 72, 73. 732. 733 Juglans, 64, 65 cathayensis, 72 — regia, 72 Juncaceae, 91, 754 Juncus, 763 — guadeloupensis, 754 — setchuensis, 9 eareos 51: t. Juniperus, 51, 54 nn pe + ~—l — formosana, 54, 72 — squamata, 49, 54, 72 Kadsura longipedunculata, 75 Kalopanax, — pictus, 8 — sciadophylloides, 10 © ow oe Kerria, 50 japonica, 78 Keteleeria, 52. S58. 61. 64. 65, 70 — pubescens Kyllinga ie oI Labiatae, 49, 87 Lacistemaceue, 729 Lagerstroemia indica, 84 L seegpe ee Larix, - potaninii, wv. ” 7I\ L (einthas hartir, 89 Lauraceae, 49, S50. 68, 75. 746, 747 2739 Leguminosae, 49. 68, 80, 746, 747 Leonia, 728 L cantanadnia: sinense. 90 Lepisorus Pees 71 a co n 87 Lillaceae, 49, 5( Lilium feucanthury var. chloraster, 91 [VoL. 6 l 1980] Lindera, 50, 67, 68, 76 — cercidifolia, 76 — communis, — subcaudata, 76 — umbellata, 76 — virdis, 49 Lindsaeaceae, 70 Liquidambar, 64, 65 — formosana, i INDEX Machilus chinensis, 76 Macleaya cordata, 77 Maesa, 69 — hupehensis, 86 Magnolia, 9, 15, 65 M agnoliaceae, 14. 68. 75, 730. 735 Magnolioxylon panochensis, 730 Malpighia elegans, 766, 769 Malpighiaceae, 765, 767, 768 Mahonia, 56, 65 Malus halliana, 78 — prunifolia, - Malvaceae. Mire cee 752 Marquesia, 740 ie naire 96 atteuccia orientalis, 71 777 Melastomataceae, 84, 735, 752, 754 C — cubeba, 7 eliaceae, — elongata, 76 Meliosma, 69, 728 — esquirolii, 76 beaniana, 82 — glutinosa, 76 — pende 2 — pungens —sl 82 — verticillata, 76 Menispermaceae, 75 Lloydia, 56 re te and ¥ Phytogeographic Sig- Lobelia, 90, 763 nificance ,41- io - Metasequ a —94 — dav idil — gly spoxroboies, 41-44, 46, 50, 51, 63. 65- 67 — ehabiank: 90) Michelin, 67. 68 — omiensis, 90 oo deat var. hupehensis, 75 — radicans, 90 — martini, 75 Lobeliaceae, 754 —— wilsonii 75 Loganiaceae, 87 Miconia, 763 iceré ryi, 89 — coriacea, 752 — » 12 Microcycas, 701, 703. 714. 715 — calocoma, 703. 711-715 Microsorium fortunei, 71 7 Olaningeede: ay sud cr be re) o Bo) © o = o G O Geran oe 754 — hyme — complanatum, 70 Minti ova valis 5 Lygodium been 70 Miscanthus sinensis, 90 ymia, SO Monetes, 740 — ovalifolia, 85 Monochoria vaginalis, 91 Lysichiton Monocotyledoneae, 49, 90. 91 8 MONTAIGNAC. PIERRE DE. RICHARD A. HOWARD. and JACQUES PoRTECOP. The Post-Eruptive Vegetation of La Soufri¢re, Guadeloupe, Lysimachia PoHGeL. ae pavendora; 88 Lythraceae 1977-1979, 749-764 Maackia chinensis, 80 Moraceae, 68, 74 Machaerina » 136, 762, 763 Morus cathayana, 74 — nee. 754 Mosla chinensis, 87 778 Mosla dianthera, 87 yrela, 70. — platyclada, 7 Myricaceae, 72 Myrsinaceae, 60. 86, Myrines 0 ) 752. 760 752 752, 760, 761 Myrtaceae, 740, 752, 760 Neolepisorus phyllomanes, 71] Neolitsea, 50, 77 — pinninervis, 77 — viridis, 76 f Cestrum (Solanaceae) from Southern aa A, 113-115 — spiciflora, 752, 762 Notes on Hi MEOSPS Ie (Rubiaceae), with a New Species from 95—105 eae, 73. 35, 736 inen Nyssaceae, 50, 84 Ochnaceae, 754 Olacaceae, 732, 735, 736 Old World Species of oo. (Gutuiferae) A Revision of the Oleaceae, 69, $6 Onagraceae, 84 Onocleaceae, 71 Onychium japonicum, 70 Orchidaceae, 91 Orixa, Japonica, & Ormosia puberula, 80 Orobanchaceae. 56 Osmanthus, 69 cooperi, 87 — fragrans, 87 Ossana vincentiana. we 767. 769 3 Ottella alismoides, 90 Oxalidaceae, § Oxalis griffithi, 80 Paederia chinensis, 89 — scandens, 89 JOURNAL OF THE ARNOLD ARBORETUM PAGE, [voL. 61 VIRGINIA M. Dicotyledonous Wood from the Up ey retaceous of Central California, I 72: a ne oo — crocea, 752, Palma. 702 — pumila, re 720 Palmae. 49, Palnaifolinin sal, — angustissimum, — debile, 715 — floridanum, — integrifolium — kickxu. 719 — medium, ~~ multitoliolatum, 715 716 715 716 = =~ > 715 — pygmaeum, 719 — strictum, 715 7 — subg. Acanthopanax, 108 — divaricatus. [09 Paris chinensis, Parnassia wrightiana var. Parrya, 56 P arthenocissus oo 83 Patrinia villosa, 89 Paulownia aa 88 Paypayrola, 728 Peperomia hernandifolia, 754 Sd flavida, 77 — tenella, 75 Perilla frutese Peristrophe rg ina, te Phellodendron, 205 -7¢9, 10, 36 — costatum, — elegans, : — curopacum. [3 — japonicum, 10, 13, 14 — lavallei, 10 — lusaticum, — molle, 10 — novae-angliae, — ornatum, 13-15 14-16, 35, 36 — sachalinense, 10 Philadelphus seat ee) Philodendron, . 758, 760 — viganteum, i 755, 759, 762 ]2=17, 29,5 1980] INDEX wai) Photinia, 67, 68, 78 Polygonum runcinatum. i — amphidoxa, 78 Polypodiaceae, 4, 9. 71, — beauverdiana, 78 Polypodium ee al — davidsoniae, 78 — lehmannii, 7 — franchetiana, 78 — mipponicum, 71 — parvifolia, 78 — phyllomanes, 71 Ph eee chinense, 88 Poly ate acutidens, 71 ee . 68 — tripteron, 71] — flexuosus, 81 — tsus-simense, 71 payilosticlie . 90 Pontederiaceae, 91 Pouch Significance, The Metasequota ae us, 6: Flora and Its, 41-94 o- adcagpedss 2 Phytolacca, i — tremula var. davidiana, 72 — icosandra, 762 pane OP. JACQUES, PIERRE DE MONTAIGNAC. Picea, 15, 51, 64, 65 and RICHARD A. Howarb. The Post-Eruptive Pieris, 5 Vegetation of La Soufriere, Guadeloupe. —- formosa, 85 1977-1979, 749-764 Pilea notata, 74 Post-Eruptive Vegetation of La Soufricre, Gua- — parietaria, 754 ie 1977-1979, The, 749-764 Pinaceae, 49, SO, 52, 53. 71, 72 Pratia, 90 Pinus, 51-53, 65 — fangiana, 90 — subg. Diploxylon, 51 — ania 90 — subg. Haploxylon, 51 Premna puberula, 87 — subg. Pinus, 51, 53 prenioeay montana, 752, 759, 761, 763 — subg. Strobus, 51, 53 Primulaceae, 86 armandii, 53, 71 protelinon « sini, 50, 91 — massoniana, 53, 71 Prunella vulgaris, 88 Piperaceae, 754 Prunus, 50, 65, 7 Pistacia, 65 — conradinae, 78 Pitcairnia, 756, 763 — ae 78 — bifrons, 754 — grayana, 7 — latifolia, 763 _ ae 78 Pitt asa 68, 78 — — var. obovata, 79 ao as — mume, 79 daphne 78 — padus, 79 — in — persica, 79 ar. a 78 — pseudocerasus, 79 — var. neritfolium, 78 — salicina, truncatum, 7 — spinulosa, 78, 79 i rostarhnn. 762, 763 — szechuanica, 79 — chrysophylla, 754, 762 — venosa, 79 Plagiogyria stenoptera, 70 — wallichit, 78 Se 0 — a 79 aginaceac, 89 — — var. leiobotrys, 79 pncee major, 89 Biter, 51-53, 58, 60, 64 Pl us, 65 — amabilis, 60, 62, 66, 71 Platycarya, 58 By ey 52. 53. 55, 56, 65 — strobilacea, 73 — forestit, 55 l e henryi, — gauseni, 55 Pogostemon glaber, 87 japonica, 55 Poliothyrsis sinensis, SO. 84 — menziesii, 55 Polygalaceae, 80 — sinensis, 55, -_ Polygala walters 80 — wilsoniana, 5 Polygonaceae, 74 Bees 98, a a Polygonum cuspidatum, 74 — aubletiana, 752 — multiflorum, 74 — anna 752, 763 — perfoliatum, 74 — insularum, 101 780 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Ptelea, - Rhododendron lutescens, 85 Pteridaceae, 70 — ovatum, 85 Pteridophyta, 49, 70, 71 — oxyphyllum, 85 Pteris henryt, 70 — simsil, 8 — vittata, 70 Rhus, 65 Pterocarya, 64, 65 — chi 8] Pterostyrax ck . 86 Pueraria LDA Punicaceé a Cason flavescens, 91 — sanguinolentus, 91 Pyracantha crenato-serrata, 79 S a Pyrrosia lingua, 71 — shearer Pyrus, 65 calleryana, 79 — pyrifolia, 79 Quercus, 9, 50, 64, 65, 73 — acuuissima, 7 — aliena, coccileroides. 73 2 5 TS pale Q nn ws) = ie Ranunculaceae. 74 Reineckea carne Revision of the Ol World ee of Calophyl- lum (Guttiferae), A. 117-699 Rhamnaceae, 69, 82 Rhamnus, 69 — esquirolii, 82 — paniculiflorus, 82 — rugulosus, 82 ~ “ Rhizophoraceas: 728, 729 Rhododendron, 50, 65 »phyllum, 85 fortunei, 85 oot ee dale i Gaal. 754 Riche ae — pranc ‘is. . 756, 761 Ricinus communis, 8 Robinia pscudo-acacia, 80 Rondeletia, 763 — parviflora, 752 Rosa, 65 — helenae, 79 roxburghil, 79 Rosaceae, 49, SO, 56, 68, Rubiaceae: Notes on nner ‘ill a New Species from Fiji, 95-105 Rubiaceae, 89, 95, 102, 104, 738, 752 — tribe Alberteae, 95 Rubus, 50, 65 — adenophorus, 79 — ampelinus, 79 — cavaleriel, 79 — ichangensis, 79 oe 79 Ruta, 3 Rutaceae, Fruits and Seeds of the Brandon Lignite, V., I-40 Riiraceaes 1-40, 68, 80 — tribe Toddalieae, 4, 7 Rutaspermum, 3, 20-22 - bognorense, 2 — minimum, 21, 22 ornatum, 21, 24 — rugosum ) — striatum, 21, 24, 27 Sablaceae. 69. 82, 728 Sageretia SS Henn: 2 Sagittaria pygmaea, 90 ear ae . Sagitt: St. Vincent, The Soufriere se a 765-770 eee 2 Salix. 50, 65 — biondiana, 72 &? i 1980] INDEX 781 Salix caprea, 72 Sorbus meee 79 — ernestil, 72 — folgn - — heterochroma, 72 nee Le — longiflora, 72 Soufriére, Gacbibs ees The Post- — rosthornii, 72 Eruptive ao of La, Sambucus, 65 Soufriére Plant of St. Vincent, a 765-770 racemosa, 89 Spachea, 766.-768 Sapium, 68 — elegans, Bees 770 — sebiferum, 81 — — f. ossana, 767, 769 Sarcopyramis nepalensis, 84 — littoralis, 766-769 Sassafras, 50, 65 — ossana, 766-769 — tzumu, 77 parviflora, 766-766 Sauraula, 728 — — var. eae: ei = Saurauiaceae, 728 — — var. typic 769 Saururaceae, 72 — peciorsta, 765. 76% Sauvagesia, 762, 763 — tricarpa, 766, ae recta, 754, 762 Spatheli Saxifragaceae, 77 Spermatophyta, 71 Schefflera, 69 Sphagnum, 752 — delavayi, &5 Sphenomeris chusana, 70 venulosa, 85 iraea, 65 Schima, 67, 69 — chinensis, 7 — parviflora, 83 — japonica var. fortunei, 79 — sinensis, 83 — prunifolia — superba, 83 Spondias, 64 Schisandra, 64 Stachyuraceae, 84 — enanthera, 75 Stachyurus, 58 Schisandraceae, 19; 135 — himalaicus, 84 Schizaeaceae, 70 Stangeriaceae, 721 Schoepfia, 65 Staphyleaceae, 82, 728 jasminodora, 65 Stenoloma chusana, 70 Scirpus wallichii, 91 Stephanan 5¢ Scrophulariaceae, 88 — chines 79 Seeds of the Brandon Lignite, Fruits and. V. Stevens. P. F. A Revision of the Old ue Rutaceae, 1-40 Species of es (Guttiferae), 117-696 Crane uncinata, 70 Stranvaesia, 5 Selaginellaceae, 70 — davidiana, % Serissa foetida, 50. 89 Strombosia, 732 Simaroubaceae, 80 Styracaceae, 69, 86 Sinarundinaria. 90 Styrax. 64, Sinopteridaceae, 70 oo 86 Skimmia, / — suberifolius, 86 Sloanea, 69 er id 87 — hemsleyana, 83 Sycopsis sinensis Smelowskia, 56 — 70. Smilax lancifolia, 91 52 megalantha, 91 “eee. 69. — myrtillus, 91 Symplocos, 50, 64, 69 Iparia, 9] anomala, 8 Solanaceae: A New He of Cestrum from > — botryantha. 86 Sou as ene. -115 — caudata, 86 Solanaceae, 56, 88, 736 — discolor, 86 Solanum ve 88 — lancifolia, 86 Sonneratiaceae, 738 — stellaris, 86 Sophora Sorbus, 65 Taiwania, 52, 58, 62, 64. 70 — aronioides, 79 — cryptomerioides, 62, 72 782 JOURNAL OF THE ARNOLD ARBORETUM [voL. 61 Taiwania flousiana, 62 Ulminium pattersonensis, 746 Tapiscia ulate 82 Ulmus, 65 Taxaceae, 50. 5 72 — bergmanniana, 74 Taxales, 52 — multinervis, ae SULCHBENENSIS: 50. 74 prunifolia, 74 Taxodiaceae, 52, 58, 72 Umbelliferae, 85 Taxodium, 66 Upper Cretaceous of Central California, Dicot- Jaxonomny of ve West Indian Cycads, 701-722 yledonous Wood from the, II, 723- sida. 52 Urticaceae, 74. 754 _54 — chinensis, 72 Vaccinium, 50 — wallichiana, 54 — bracteatum, 85 Ternstroemia. 50, 69, 763 — carlesii, 86 elliptica, 752 — hirtum, 86 — nitida, 8 83 — iteophyllum, 86 Tetracentraceae, 74 — japonicum, 86 Tetracentron sinense, 50, 74 — mandarinorum. 86 Tetrastigma serrulatum, 83 Valerianaceae, 89 Teucrium bidentatum, 88 Vegetation of ae Soutri¢re, Guadeloupe, 1977 The: ~ 49, 50, 69, 83. 752 1979. The Post-Eruptive, 749-764 Thely ane ee ie Tl Verbenaceae, 87 Thelypteris decursivo- pinnata, 71 Viburnum, 50, 65 Thuya, 65 — ichangense, 89 Tibouchina, 763 — ovalifolium, 89 — ornata, 754, 762 = propinquum, o Tipeney. BRucE H. Fruits and Seeds of the _— —— yar. mairei. 8 ae : ignite, V. Rutaceae, I-40 rhytidophyllum, Me Pilia, 65, — rosthornil var. xerocarpa, 89 — tuan i — seuigerum, & Villacea 3 — sympodiale, 89 Poddalia, 3, 68 — ternatum, 8 ; asiatica 80 theiferum, 89 Foona, 65 — tomentosum, 89 - sinensis, &O = ingu, 89 Torreya, 51, 52. 55 Viola, 762, 763 — californica, 55 — confusa, 83 — sepals: 53, 72 — inconspicua, 83 — jackil, 55 stipularis, 754, 762 — nucifera, 55 — triangulifolia, 83 — taxifolila, 55 Violaceae. 83, 728, 729, 754 Torricellia intermedia, 85 Wiseeie: 14. 8735 Toxicodendron, 65 Vitis. 65 Trachelospermum jasminoides. 87 Trachycarpus fortune!, Trem — nevi gata, 7 Follias: “yunnanensis, 75 Tae 52, 54, 64. 65. 70 sect. Hesperopeuce, 54, 55 — sect. Tsuga, 54 — caroliniana, 54 — chinensis. 54. 72 Zamia, 701-710, eed allison- eyOUh: Weigela japonica var. sinica, 89 Weinmannia, 76. — pinnata, 752, West Indian Cycads. Taxonomy of the, 701-722 Wood from the Upper Cretaceous = Central California, Dike eas ak I], 723-748 — mertensiana, 55 — angustifolia, 7 a 8, 715, 716 Tomita: 29 — — var. angustissima, 715 — var. floridana, 716 Ulmaceae. 68, 74 — — var, stricta, 715 Ulminium cullen: 746 — — var. yates, 715-717 1980] Zamia angustissima, 702, 708, 715 — brachyphylla, 6 ~~ ON — cylindrica, 716 — debilis, 702. 715 — dentata, 715 — floridana, 708, 710, 716 — furfuracea, 702, 715, 718, 720 a 702, 717 — gutierreziit, 720 — hun ‘lis Pegi) — integrifolia, 702, 715 — kickxil, 708, 719 — latifoliolata, 702, 708, 716, 720 linearifolia, 71 aris, 716 — linifolia, 716 lucayana . — media, 702, var. ae ee 715, 717 — — f. silicea, 71 — — var. _jacquini ana, 715 '. bres pa tH — — var. portoricens enuls, 715 — multifoliolata, 702, 716 — ottonis, 708, 719 — portoricensis, 702. 708, 716 eae ae 703, 708, Te, HB e TD IG, HB. 1 o a — ea 703, 704, 706, 708 709, TIL 73 a 720 — pygmaea, 702. a — rotundifolia, 719 — silicea, 708. 719 — silvicola, 716 — stricta. 702. 715 INDEX subs. aes ae 704. 706, 708. 709, , 715-72¢ 783 Zamia subcoriacea, 717 mamidcedes 701, 72 Zanthoxylum, 3, i 12, 15-26, 28, 29, 36, 49, 68 — acuminatum, 23, 24 — ailanthiforme, 22 — americanum, 18 oo POEDOICN ES 18 aeum, 23, 24, 36 — compre cum, pas — dissitoides, 20 — dissitum, 18, 8¢ echinospermum, 25-27, 36 5 — ft dum, 25 — yiganteum, 22 — hawaiiense. 22 — mone: 8 , 20, 26 8 ; ase, 18 — pimpinelloides, 18, 23, 24 — proceru — ee mum, — scandens, — schinifolium, 18 — simulans, 49, 8 — uffneyi, 22, 27-29, 36 — tomentellum. 26 — trifoliatum, 110 18, 22-27, 29, 36 = Qe — ee as 74 Zizyphus ARNOLD ARBORETUM JOURNAL OF THE 1980 VOLUME 61 HARVARD UNIVERSITY Dates of Issue No. | (pp. 1-116) issued 21 March, 1980. No. 2 (pp. 117-424) issued 19 December, 1980. No. 3 (pp. 425-700) issued 19 December, 1980. No. 4 (pp. 701-783) issued 24 April, 1981. Contents of Volume 61 1980 Fruits and Seeds of the Brandon Lignite, V. Rutaceae. BROGE ES MIMNEY sik. ccc e Ene EOT Ree eee eR ee I The Metasequoia Flora and Its Phytogeographic Significance. SEIU VING al wvn tanga kaa eee eee ones eee eae ace 4] Notes on Airosperma (Rubiaceae) with a New Species from Fiji. SUB VENI DARWIN: 25.05, gee beesecescseebe J32S 20 seked Pieeae es 9 Eleutherococcus vs. Acanthopanax. SEC NGO 235455 5d 54S dt aoe Vee eh ot ee ee 107 A New Species of Cestrum (Solanaceae) from Southern Colombia. PER NVEANIDOLL UNZIKER ~ 0.0 enacts oes oe ea he ae Bo ae es PS. SleMent OL OWNEISNIO- cays eee chee caw eus Buse eee e ee & 116 A Revision of the Old World Species of Calophyllum (Guttiferae). P : NS 5 Be SIEVENS. fis cececk ocigcpa atte ee Me Bk ig ee as Ae mE aes 117 Taxonomy of the West Indian Cycads. MES E. ECKENWALDER .......---0 0 ee eee eee eee eee ees 701 Dicotyledonous Wood from the Upper Cretaceous of Central California, II. NV IRGINUA: IM). PAGE. “oe o.f:5 wie een ee ea 28 9 a eae 722 The Post-Eruptive Vegetation of La Soufriere, Guadeloupe, 1977-1979. RICHARD A. HOWARD, JACQUES PORTECOP. and PIERRE DE MONTAIGNAC ..........00 2002 eee eee eee eee 749 The Soufriere Plant of St. Vincent. RICHARD A. HOWARD and KRISTIN S. CLAUSEN ..............00-- 765 UNGER 555, dite ecto esac ee Bett tte heats AS ae 3g 7T7\ —_ 1982 AETFAT CONGRESS Presented by the Department of Agriculture and Fisheries CSIR CONFERENCE CENTRE, PRETORIA, REPUBLIC OF SOUTH AFRICA, 19-22 January, 1982 Please address = meee, summaries of papers, and requests for further announcements The oe Secretariat $.229 CSIR PO Box 395 PRETORIA 0001 Republic of South Africa Telephone: National (012) 86-9211 International + 27 12 86-9211 extension 2077 (Elsie Coetzee) 2063 (Ann van Dyk) Telex: SA 3-630 - et of the ee perite October, 1980 _ CONTENTS OF VOLUME 6 61, , NUMBER Ra oe = “9 Taxonomy of the West Indian Cycads. JCKRENWALDER +: eros oe : oe 701 < Dicotyledonous Wood from the Upper eee a Central ~ Californi a, Il. — a : VIRGINIA M. PAGE. = = = 2 stan The Post-Eruptive Vegetation of es Soufriére, Guadeloupe es so ee 1979. : _ RICHARD A. ‘Howarp, Jacques | Portecor, AND PIERRE DE MONTAIGNAC . re ee ae 749 - The Soutriere: Plant of Sst. Vincent. ee ee Shine - -RicHarD A. HowarD AND Kaistin Ss. CLAUSEN . PS sure ee LOD "Volume « ae ‘Number 5, including pages 425-700, 7 19, 1380.