Untitled R E I N W A R D T I A Published by Herbarium -Bogoriense, Kebun Raya Indonesia Volume 4, Part. 2, pp. 193 - 256 (1957) LAURACEAE* A. J. G. H. KOSTERMANS ** SUMMARY An attempt has been made at a classification of the Lauraceae, into taxa down to the rank of subgenus. The characters and their taxonomic value are discussed amply. A system of the family and keys are given. Two subfamilies, viz Lauroideae and Cassythoideae, are recognized; several new tribes, subtribes and subgenera have been described. The accepted genera, 31 in number, are surveyed, with synonyms, typification, description, and particulars. Additional notes to "The genera of Lauraceae", which appeared in 1952, are given. INTRODUCTION In a former paper1 I gave a historical survey of the genera of Laura- ceae, together with their typification and synonymy. In the present paper I shall endeavour to elaborate a tentative classification of those genera which I consider to be acceptable. I stressed in the first paper that the establishment of generic limits and the classification of genera should come after complete monographic revision. In as large a family as the Lauraceae this cannot be accomplished in a lifetime. Since I only have completely revised the subfamily Lauroi- deae2 (American species), the subfamily Persoideae remains unsatis- factorily classified. All African and Madagascar Lauraceae have recently been revised, some (part of the genus Beilschmiedia) by Robyns and Wilczek, the remainder by myself. During the last ten years I have had the opportunity to obtain first hand field knowledge of the Malaysian Lauraceae. * First published as Comm. of the Forest Research Institute, Indonesia No. 57, issued March 22, 1957. NOTA BENE. The present reprint is verbatim, with the following restrictions: a) names in the original paper indicated as new and provided with a Latin description are here not indicated so and provided with an English description, in the index, however, they are printed in bold face type; b) in the genera 5, 19, and 28 the name of the type-including subgenus has been conformed to that of the genus, the original name being there given in syn. The division of the text over the pages has remained the same from p. 206 onwards; the original page-number is found by subtracting 192 of the present number. There is not referred in the text to the above-mentioned publication. ED. ** D. Sc, Forest Service of Indonesia; Head of the Botanical Division: Forest Research Institute, Bogor; Cooperator of the Herbarium Bogoriense. 'A historial survey of Lauraceae in J. sci. Research Indonesia 1: 83-95; 113- 127; 141-159. 1952., 2 Sub-families as adopted by Pax. 194 R E I N W A R D T I A [VOL. 4 1957] A. J. G. H. KOSTERMANS : Lauraceae 195 The revisions have already yielded interesting results; genera formerly considered to be limited to certain continents proved to have a world wide distribution: Hufelandia, Tylostemon and Beilschmiedia, respectively from America, Africa and Asia, could be combined into one genus Beilschmie- dia; Potameia from Madagascar and Syndiclis from Asia could be com- bined; American Persea proved to be congeneric with Asiatic Machilus and it is likely that American Sassafras is not different from Asiatic Actinodaphne, whereas Ocotea and Cinnamomum are very close to each other; American Phoebe is likely not different from Asiatic Cinnamomum. Several views on the relationship between genera have been confirm- ed by anatomical research on the wood (cooperation in this field with W. F. Stern of the Yale School of Forestry, is much appreciated). I am Well aware that the system outlined below will be far from conclusive and perhaps never can be made conclusive, but I hope that it will bring some clarification. I disagree with those who have expressed the belief that a classification of Lauraceae "in se" would be impossible. A classification is especially necessary for workers in the field of palae- ontology. In recent years fragments have been described as belonging to certain genera of Lauraceae; but it is — even in contemporary species — impossible to refer specimens to their proper genus without flowers or fruit at hand. Our insight in the relationships and development of Laura- ceae will never be advanced by unreliable identification of fossil material. I wish to express here my gratitude to Ir. Soesilo H. Prakoso, Head of the Forest Service of Indonesia, who with a broad view for pure and applied science in botany, has given his support and enabled me to carry on with taxonomic work in the Indonesian Forest Service. Furthermore I have to thank Prof. Dr. C. G. G. J. van Steenis, who kindly went through the manuscript and suggested many emendations and alterations, and Mr. M. Jacobs for proofreading. I wish to thank Dr. A. C. Smith (National Science Foundation, New York), who pointed out to me that the names of a few type-including sub- genera were not in accordance with Art. 22 of the International Code (which I could not consult). These names have been brought up to date in this reprint. PRINCIPAL LITERATURE A. L. de Jussieu, Genera PI. 89-90. 1789 (Lauri); Lamarck-Poiret, Encycl. meth. 1783-1817; Ventenat, Tabl. Regne veg. 2: 245. 1799 (Laurinae) ; Willdenow, Spec. PI. 2: 477. 1800; Lindley, Nat. Syst. 201. 1836 (Lauraceae); C. G. Nees von Esenbeck in Wallich, PL Asiat. rar. 2: 57-76. 1831; Plant. Laur. Expos., Breslau 1833; Systema Laurina'rum 1836; Endlicher, Gen. PI. 315-323. 1837; Enchir. 196-205. 1841; Meissner in DC, Prodr. 15 (1): 1-260. 1864; Baillon, Hist. PI. 2: 429-486. 1870; Bentham in Bentham & Hooker f., Gen. PI. 3: 146-164. 1880; Pax in Engler & Prantl, Nat. Pfl.Fam. 3 (2) : 106-126. 1889; Mez in Jahrb. kgl. bot. Gart. Berlin 5: 1-556. 1889; Hutchinson, Fam. fl. PL 1: 90. 1926; Kostermans in Rec. Trav. bot. neerl. 33: 719-757. 1936; 34: 500-575. 1937; 35: 56-129 et 831-931. 1938; in Rev. Univ. Chilena 24: 201-232. 1939; in Humbert, Not. syst. Paris 8: 67-128. 1939; in Bull. Jard. bot. Bruxelles 15: 73-108. 1938; in Humbert, Fl. Madagascar, 8le Famille Lauraceae: 1-90. 1950; in Bol. techn. Inst. Agron. Norte (Belem-Para, Brazil) 28: 49-76. 1955; Robyns & Wilczek in Bull. Jard. bot. Bruxelles 19: 459-506. 1949; Lawrence, Taxon. vase. PI. 512-513. 1951. FAMILY CHARACTERS GENERAL Evergreen trees or shrubs (rarely parasitic climbers: Cassytha) with alternate (rarely opposite or subopposite: Beilschmiedia, Endiandra, Cryp- tocarya, etc. or whorled: Actinodaphne; occasionally in species of other genera) usually entire, rarely lobed (Sassafras), usually leathery leaves, without stipules, with usually numerous aromatic oil and slime cells (also in bark), pinnately or subpalmately veined (often triplinerved: Aiouea, Litsea, Neolitsea, Lindera, Cryptocarya, Cinnamomum, etc.) and usually densely reticulate; the reticulation as a rule not visible in the fresh leaves which often have a waxy appearance and often have a glaucous lower sur- face. Hairs — if present — simple, as a rule one-celled. Buds perulate, inno- vation flush-wise. Bark often aromatic. Timber usually not very durable, sometimes very durable (Eusideroxylon, Ocotea. irodioei Mez, Dehaasia caesia BL), finely grained, often yellow and with cigar-box wood smell. Inflorescences definite, (rarely indefinite: Cassytha) paniculate, race- mose or capitellate; before anthesis completely enveloped in large bracts (Actinodaphne, Sassafras, species of Beilschmiedia, Cryptocarya, etc.) or almost naked; ultimate flowers 3 or more together in axils of bracteoles or the end-flowers in pseudo-umbels, surrounded by persistent decussate bracts (Lindera, Laurus, Litsea) or irregular bracts (Umbellularia). Flow- ers usually small (the largest 2 cm in diameter; but usually less than 5 mm in diameter, the smallest: Potameia, often less than 1 mm in diameter), usually white or greenish white, sometimes yellow, sometimes reddish or turning red after anthesis (Persea subgen. AIseodaphne), usually aromatic; tepals either explanate or remaining almost closed (Persea subgen- Alseo- daphne, Licaria, Beilschmiedia). Flowers bisexual or by abortion monoecious, actinomorphic, usually trimerous (in the genera Laurus, Neolitsea, Potameia dimerous). Perianth gamosepalous, free, in bud valvate rotate, infundibuliform or urceolate with 6 or 4 tepals in two whorls, rarely 9 tepals in 3 whorls (Phyllostemono- daphne and Dicypellium) ; tepals equal or (Persea, species of Persea subgen. Alseodaphne) outer ones smaller, alternate, deciduous or persistent, some- times indurate; tube either deciduous or altered into a cup which surrounds the basal part of the fruit, or the fruit completely included in the tube, or the ovary adnate to the tube (hypogynous: Hypodaphnis). Stamens alter- nate, attached to the throat of the tube, perigynous or epigynous, definite 196 R E I N W A R D T I A [VOL. 4 1957] A. J. G. H. KOSTERMANS: Lauraceae 197 in number (rarely sub-definite, in Litsea) in 4 whorls (or rarely more whorls in Litsea), usually the central whorl abortive and reduced to more or less conspicuous staminodes; more rarely the 2 outer whorls also abortive; the third whorl always present (very rarely sterile: species of Cryptocarya, one species of Aniba etc.), as a rule with two more or less stalked glands at either side of the filament or the stalks of the glands connate with the filaments; more rarely all stamens with basal glands (Urbanodendron, one species of Endlicheria; the subgenus Pleurothyrium in Ocotea; species of Litsea) ; filaments present or anthers sessile; outer 2 whorls of anthers as a rule introrse (some exceptions in Licaria) ; all extrorse in Litsea; the 3rd whorl of stamens as a rule extrorse, sometimes with (all or partly) apical or lateral cells. Anthers 4-, or 2-celled by abortion (very rarely 1- celled: Potameia species) ; the connective (of the 2-celled anthers) usually produced beyond the anthers (ablastic part) ; the cells placed in pairs above each other or in an arc; usually all species in one genus with the same number of anther-cells, rarely one or two whorls of anthers with different (half or double) number of cells. Cells opening by valves from base to apex, very rarely (in Mezilaurus) from outside to inside. Pollen simple, globose, poreless, granulate. Staminodes (if present) of the outer whorls tepal-like or ligulate, of the fourth whorl as a rule sagittate or cordate-sagittate, stalked, very rarely provided with glands, sometimes staminodes minute or none. If more than 4 whorls of stamens are present (Litsea), the 4th and inner whorls may have glands. The glands are either small or large (filling the entire space between the stamens) or altogether lacking. Carpels probably 3, forming a single, one-celled ovary, which is usually superior, rarely sub-inferior or (in Hypodaphnis) inferior; ovule single, pendulous, anatropous; style distinct, rarely inconspicuous; stigma discoid, often with one shallow lateral incision, often depressed, some- times (Beilschmiedia, Litsea) inconspicuous, but decurrent at one side of the style as differently coloured tissue. Fruit baccate, sometimes enveloped (adnate or free) by the accrescent flower tube which is often ligneous {Cryptocarya, Ravensara, Eusideroxy- lon) or completely inferior (Hypodaphnis), sometimes on a naked pedicel, sometimes with its base surrounded by the indurate perianth {Phoebe, Apollonias) ; sometimes with its base (or a larger part) embedded in a cup; sometimes (Mezilaurus) with a small, flat disc at base. Where a cup is present, the perianth may be either more or less persistent (unaltered), or the base of the perianth persists (Cinnamomum) or the base of the stamens persists, which makes the cup double-rimmed (Licaria) ; the margin is either entire or may be wavy (base of tepals) ; the cup is always fleshy, often provided with large, flat, round warts (lenticels) ; the fruit- pedicel is either cylindrical or may be fleshy and sometimes highly colour- ed (Dehaasia; species of Persea subgen. Alseodaphne). Exocarp fleshy, very thin or thick (edible in Persea species), usually bitter, aromatic and astringent. Fruit either small (5 mm diameter in Litsea cubeba Pers.) to large (15 cm and more long in Persea americana Mill, and Eusideroxylon zivageri T. & B.). Sometimes a kind of aril (Potameia) is present which envelopes the embryo, but for its apical part. Seed without albumen with thin (rarely tough: Cassytha) testa; cotyledons large, flat-convex, pressed against each other (only in one species: Beilschmiedia variabilis Rob. & Wilcz., the embryo is transverse) ; corculum included, subpeltate; plumule well developed (2—8-leaved), often pilose. Rarely (Ravensara) ovary divided incompletely (in its lower half) into 6—12 compartments; the dividing dissepiments ruminate into the cotyledons. BOLE, CROWN, BARK, ROOT With the exception of Cassytha, which comprises parasitical, leafless twiners, all Lauraceae are woody plants. They range from small treelets or shrublike treelets of less than one metre height to trees of 40 and more metres, although they never become emergent trees in the tropical rain forest, but are confined to the second story, only some species belong to the first story. In some species the branches are weak and need the sup- port of other trees. Originally pyramidal, the canopy becomes often irreg- ular in mature trees, although some retain their regular appearance. The branching is either mono- or sympodial. In the latter case the tree remains pyramidal and has the typical whorls of leaves congested near the top of the branchlets, which is often thicker than the older part. (Phoebe, Mezilaurus, Persea, etc.). The main branches are horizontal, more or less erect, or irregular. Sometimes they are dorsiventral (differently coloured on upper and lower j surface) ; the leaves are then often in one plane. Myrmecophily is found [exceptionally (South America, New Guinea) ; in that case the ants eat away the pith of the branchlets (usually swollen). Shedding of branches by means of an abscission joint has been described by Van der Pijl (in Indon. J. nat. Sci. 1, 2, 3: 20. 1953). In some species (Aiouea) decurrent leaf spurs are present, which make the branches angular or ribbed. Buttressing is common in many species, although large buttresses are an exception. Pneumatophores occur (rarely) in some species of marshy habitat. Stilt roots are, thus far, not known. The bole is usually smooth (although thick, deeply fissured barks loccur too) and then often covered with numerous large, round, flat lenti- ceels. The dead bark is as a rule very thin and is shed in small fragments. In some Malaysian species (Persea subgen. Alseodaphne) the branchlets are pure white. In some South American species (Mezilaurus, Ocotea ver- ruculosa Mez) the branchlets are covered with a corky layer, apparently an adaptation to habitat. The living bark is either thin or may be very thick; its colour varies between white, yellow, pink and dark red; it is usually brittle but for an inner fibrous layer; often it has a more or less 198 REINWARDTIA [VOL. 4 1957] A. J. G. H. KOSTEHMANS: Lauraceae 199 pronounced aromatic smell; in some species (Cinnamomum iners Bl.) where the bark is scentless and has only mucilage cells, the roots are fragrant. The bark is almost always characterized by the presence of secretion cells containing essential oil or mucilage and of remarkable stone cells in the pericyclic region with onesided thickening; the secretion-cells are found in the cortex and especially in the bast. The stone cells usually contain starch grains and sometimes calcium oxalate crystals of different shapes. The number and length of the bast fibres is very variable. The medullary rays are from one to more cells wide. The oil and mucilage is formed in a resinogenous mass in the cells; according to Janssonius (in Blumea 6: 408-451. 1950) the mass is surrounded by a ligneous membrane (which is somewhat thicker in oil-cells), which shrinks together with the resinogenous mass, if soaked in alcohol. This does not fit with Tschirch's theory, according to which the resinogenous mass is formed in an resino- genous layer of the cell wall. The mucilage is sometimes so abundant that it has economic value (Litsea species in Indochina and Persea species in Indonesia, for making hair-fixatives and glues). Sometimes (Litsea species in, Indonesia) the mucilage cells are more common in the sapwood. Shirasawa found, that in Cinnamomum, camphora Nees & Eberm. the oil-cells developed imme- diately below the vegetation point. LEAF The leaves are usually leathery and (on the upper surface) of a waxy, glossy dark green colour; the lower surface is often glaucous (layer of wax) ; in this character they strongly resemble the leaves of Myristica- ceae and Monimiaceae. The dried leaves have a definite colour in different species. The lower leaf-surface is often provided with domatia in the nerve- axils; they appear to represent a constant character for certain species and not to be caused by insects. Pinnately veined leaves are a rule, but in several genera (Aiouea, Cinnamomum, Lindera, several species of Ocotea, Endlicheria, Cryptocarya, Ravensara, American Phoebe) tripli- nerved leaves occur, (that this character is of no generic value is import- ant for palaeontological identification). The venation, which becomes visible after drying, is either lax or very dense (areolate in Endiandra and in species of several other genera). The leaves are spirally arranged (phyllotaxis 2/5 and 3 / 8 ), sometimes subopposite to opposite (Beilschrfiiedia, Endiandra,, Cryptocarya, etc.), sometimes whorled (Actinodaphne, some species of Endlicheria, Aniba, etc.). Stipules are absent, but "Hochblaetter" are often present, the "Hochblaetter" are distinctive in Actinodaphne on the long internodes between the whorled leaves; they drop very soon, and hence are hard to study in herbarium specimens. Formerly it was supposed that only tem- perate representatives of the family had perulate leafbuds, but I found large numbers of this kind in species of the tropical rain forest where periodical leaf shedding is far from uncommon. Some species of P&rsea subgen. Alseodaphne in the wet rain forest e.g. are completely bare for a short period (a couple of days) ; in most species, however, the older leaves survive one or more flushes. The scales determine the shape of the leafbud, which may be a very useful vegetative character to recognize some genera (Beilschmiedi-a,, Cryptocarya). The leaves are entire (perhaps slightly wavy in Nectandra sinuata Mez), the only exceptions are found in the lobed leaves of Sassafras and some aberrant leaves of the cultivated "avocado" (Persea americana Mill.) as described by Huber. The leaf surface is flat and smooth; rare exceptions are found where the leaves become boat-shaped and bullate (Endlicheria bullata Ducke, Mezilaurus arassiramea Taub., Beilschmiedia bullata Kos- term.). The leaf margin is always strengthened by sclerenchym. Even if the species is entirely glabrous, hairs are often present on leafbud scales. The hairs are simple, silky, woolly or strigose; as a rule the pilosity is denser on the lower, than on the upper surface, although the reverse is also represented (Ocotea subgenus Nectandra); the hairs are usually silvery, but golden coloured hairs occur in several genera (Actinodaphne, Endiandra, Licaria, Ocotea, Persea, etc.). The hairs are mostly unicellular, frequently thick-walled, the lumen sometimes almost obliterated (Santos found differences in hair structure in Philippine Cinnamomum species). The leaf is generally dorsiventral, but palisade tissue is more strongly developed towards the abaxial than the adaxial surface of the leaf in species of Nectandra, Phoebe, Mezilaurus. The leaf cuticle is delicately punctate in some species of Ocotea and varies in thickness in different species of a single genus from the same habitat. The epiderm of both surfaces in leaves of Cinnamomum consists of a single layer of rectangular, thick-walled and highly cutinized cells. In the surface view the cells are polygonal in outline with sinuate, thick walls. The hypoderm is one-layered, but shows a tendency to form two layers (Cryptocarya, Endiandra). The lower epidermis is commonly papil- lose; in the Aniba rosaeodora-group these papillae are highly developed. Pappilae-like cells are usually connected with a waxy lower leaf surface. 200 R E I N W A R D T I A [VOL. 4 1967] A. J. G. H. KOSTEKMANS: L'auraceae 201 Stomata are on the lower surface, usually they are sunken, and over- arched by subsidiary cells, which are usually arranged parallel to the pore and of the rubiaceous type, but the subsidiary cells of the depressed stomata are not easily recognizable. The guard cells are provided with ridge-like processes in most genera. The mesophyll consists of 1—3 layers of palisade cells, which may be constant in a genus or varying. Individual cells of the palisade layer in some species of Actinodaphne, Ocotea, and Beilschmiedia are transformed into stone cells, visible as dots on the sur- face of the leaf. Sclerified cells, immediately within the lower epidermis are present in Ravensara; spongy parenchyma containing large lacunae filled with stellate tissue occurs in Aniba, Cryptocaryw, Licaria, Persea, Systemonodaphne and Urbanodendron, The midrib is often surrounded by a mixed sclerenchymatous sheath, containing stone cells with U-shaped thickenings. Medium sized and small veins are frequently vertically trans- current by sclerenchymatous elements; the sclerenchymatous sheaths in a few species of Persea are spread out beneath the upper epiderm. Oil cells in mesophyll and midrib are generally spherical with suberized walls and yellowish contents, frequently giving rise to transparent dots in the leaf; they are situated in the palisade or spongy tissue, rarely (Umbellu- laria) in the lower epidermis as well. Mucilage cells are found in almost all genera; they are similar in shape to oil cells and are confined to the palisade tissue or they occur also in other parts of the mesophyll. Crystals usually occur in the form of small needles or spindles. The petiole, as a rule, has the same diameter over its entire length; in rare cases it is thickened towards its base {Mezilaurus; some species of Persea subgen. Alseodaphne, etc.). The petiole in transverse section through the distal end exhibits a shallow crescentic vascular strand in species of several genera or a shallow arc of separate but closely placed bundles {Beilschmiedia roxburghiana Nees) ; there are no accessory bundles. Sometimes massive stone-cells are present in the petiolar ground tissue {Cinnamomum zeylanicum Bl.; Ra- vensara) . WOOD ANATOMY X Cork not formed until a relatively late stage; often arising in the epidermis or outer part of the cortex, but pericyclic cork said to occur in Cinnamomum zeylanicum Bl. Cork cells thin-walled in species of Litsea, Persea, Sassafras with inner tangential walls sclerosed in species of Cin- namomum, Dehaasia, Dicypellium, Litsea, Ocotea. A secondary ring of Extracted more or less verbatim from Metcalfe and Chalk, Anat. of Dicotyl. 2: 1147-11521950; W 1 ' . S t e r n i n Trop. Woods 100. 1954 and Record in Trop. woods 80: 15. 1944. stone cells reported to arise in the phelloderm in species of Actinodaphne, Cryptocarya, Dehaasia, Litsea, Ocotea. Primary cortex of relatively old branches sometimes containing stone cells. Pericycle characterized by isolated strands of fibers interspersed with stone cells to form a continuous or locally interrupted ring. Some of the stone cells usually provided with thin outer tangential walls, but scle- rosed on the inner tangential and radial walls. Pericyclic sclerenchyma rings of the above type, recorded in species of Licaria, Beilschmiedia, Cin- namomum, Cryptocarya, Dicypellium, Endiandra, Endlicheria, Laurus, Lin- dera, Litsea, Ocotea, Persea, Phoebe, Sassafras. Secondary phloem usually containing a few spindle-shaped fibers with narrow lumina, but fibers sometimes aggregated into bundles in Licaria, Beilschmiedia, Ocotea. Stone cells similar to those of the pericycle sometimes accompany the fibers in the secondary phloem, particularly in species of Cryptocarya and Litsea. Xylem in the form of a continuous cylinder traversed by narrow rays (1—6 cells wide) ; relatively broad rays occur in certain genera; rays uni- seriate (1—14 cells high) and multiseriate rays (3—6 cells high). Vessels usually with simple perforation; scalariform plates oecassional. Peripheral part of the pith in a few species consisting of amyliferous cells with thicker walls than those of the central part. Groups of stone cells some- times present in the pith, e.g. in species of Beilschmiedia and Ravensara. Secretory cells, containing oil or mucilage, present in the primary cortex, phloem, wood, and pith of the axis. Oil cells in the primary cortex and pith approximately isodiametric, but those in the phloem vertically elongated (barrel-shaped) ; recorded in the young stem of species of Lica- ria, Aniba, Beilschmiedia, Cinnamomum, Cryptocarya, Dicypellium, Lau- rus, Litsea, Ocotea subgen. Nectandra, Persea, Sassafras, Umbellularia. Mucilage cells, usually vertically elongated, most frequent in the phloem (especially in Cinnamomum and other barks, but also recorded in the primary cortex and wood. Cells with unidentified but probably tannini- ferous contents common and often abundant, particularly in the cortex and phloem, e.g. in species of Beilschmiedia, Cinnamomum, Laurus, Per- sea. Crystals common in the unlignif ied tissues, mostly acicular or spindle- shaped ; solitary prisms noted, e.g. in Beilschmiedia. WOOD —Growth rings as a rule present, although sometimes in a single species {Persea americana Mill.) present or absent. Vessels mostly medium-sized (100—200 µ mean tangential diameter; up to 650 µ,, according to Stern), but sometimes small (less than 100µ.), e.g. in species of Laurus and Lindera; sometimes very long (1224µ.), solitary and in numerous small multiples; multiples of 4 or more cells sometimes moderately common, e.g. in some species of Persea subgen. Alseodaphne, 202 R E I N W A R D T I A [VOL. 4 1957] A. J. G. H. KosHEKMANS: Lauraceae 203 Aniba, Cryptocarya, Eusideroxylon, and Laurus, and sometimes with a tendency to form loose oblique rows, e.g. in Lindera and Persea; 3—40, mostly 5—12, per sq. mm., fewer than 5 per sq. mm. in some species of Beilsehmiedia, Cinnamomum, Cryptocarya, Hypodaphnis and Licaria; ring porous in Sassafras. Perforations typically simple, but sporadic scaliform plates are of moderately common occurrence, particularly in the Persoi- deae; such plates usually with few bars but with many in Persea; accord- ing to Knoblauch, as quoted by Solereder, the perforation plates are entirely scalariform in Beilsehmiedia pendula Benth. (not confirmed). Intervascular pitting alternate, typically large, but occasionally small, e.g. in Ocotea rodiaei Mez and in Lindera species (not in material of L. bifaria Benth.) ; occasionally striated owing to coalescent apertures, e.g. in Laurus and Umbellularia; Solereder refers to spiral striation in Cinnamomum eamphora Nees & Eberm. and Litsea, pits to ray and wood parenchyma typically including many large, elongated and simple or only partially bordered pits, often almost scalariform, and sometimes unilaterally com- pound but with only small round pits similar to the intervascular pitting in Hypodaphnis zenkeri Stapf, Laurus nobilis L., Lindera erythrocarpa Mak., Neolitsea acuta-trinerva Kan. & Sas., and Ocotea rodiaei Mez. According to Bailey the sieve-like structures reported by Janssonius are not true vestured pits. Tyloses common, sclerotic in Eusideroxylon zivageri T. & B. and Ocotea rodiaei Mez and, according to Record and Hess and Stern, in some species of Aniba and Licaria; the walls of sclerotic tyl- oses are often laminated and they show ramiform pitting (Stern). Mean member length 0.35—0.8 mm. Parenchyma paratracheal, typically as an irregular and often incomplete sheath round each vessel (vasi-centric), sometimes tending to be aliform, e.g. in Beilsehmiedia, Cryptoca,rya, Eusi- deroxylon, Hypodaphnis, Lindera, and Mezilaurus, and occasionally locally confluent; sometimes very abundant in broad irregular bands in Eusider- oxylon zwageri T. & B.; irregularly spaced bands; that appear to be ter- minal and bended apotracheal present in some species of Beilsehmiedia, Cryptocarya, Endiandra, Hypodaphnis, Lindera, Ravensara and Mezilau- rus. Most authors refer also to diffuse parenchyma, but, though individual strands, separated from the irregular sheaths round the vessels and solitary oil cells, may appear to be scattered among the fibers, it appears to be somewhat misleading to classify these as diffuse. Strands usually of 2—4 cells. Tendency to be storeyed observed in Mezilaurus lindaviana Schw. & Mez and distinct storeys reported in 2 species of Cryptocarya from Madagascar. Oil cells present in more than half the species examined, often very abundant and sometimes very large, e.g. in Beilsehmiedia man- nii Benth. Rays typically 2—3 cells wide and up to 4—8 cells wide in some species or specimens of Beilsehmiedia, Cinnamomum, Cryptocarya, Endiandra, Eusideroxylon, Persea and Ravensara; less than l m m . high; uniseriates typically very few and low and composed of mixed upright and procumbent cells, but more numerous and composed entirely of upright cells in some species of Beilsehmiedia and Cryptocarya; occasional 'aggre- gate' rays noted by Dadswell and Eckersley in Cryptocarya glaucescens R. Br. and C. corrugata C. T. White, such rays, according to Francis, associated with the deep indentations characteristic of the mature stem; 4—11, mostly 5—7, rays per mm.; typically rather weakly heterogeneous, with marginal row of square cells, sometimes homogeneous, e.g. in Hypo- daphnis zenkeri Stapf and Umbellularia californica Nutt. Janssonius and Dadswell and Eckersley refer to occasional sheath cells in some species. Cells often with dark solid contents; crystals not observed. Occasionally with a distinct tendency to arrangement in echelon, particularly in some species of Beilsehmiedia and in Mezilaurus lindaviana Schw. & Mez. Com- monly containing oil cells. Gonggrijp notes the presence of silica in Endi- andra and Litsea. Fibers typical with simple pits that are more numerous in the radial than in the tangential walls, but the pits occasionally with small, rather indistinct borders in some of the species lacking septate fibres, e.g. some species of Actinodaphne, Beilsehmiedia; Endiandra, and Ocotea rodiaei Mez. Septate in Licaria p.p.; Actinodaphne p.p., Aiouea, Persea subgen. Alseodaphne, Aniba, Cinnamomum p.p., Dehaasia, Dicy- pellium, Endlicheria, Licaria, Lindera (one species only), Litsea p.p., Ocotea p.p., Persea, Phoebe, Ocotea subgen. Pleurothyrium, Mezilaurus and Umbellularia; occasional septate fibers may occur in some of the other genera (Kanehira, Stern). Septate fibers are not limited to the neigh- bourhood of the vessels, as in the Myristicaceae. Walls varying from very thin to very thick (the lumen is practically obliterated in certain species) sometimes gelatinous. Mean length 0.7—1.6. mostly 1.0—1.4 mm. Vasi- centric tracheids recorded by Janssonius in Phoebe dedinata Bl., Lindera subumbelliflora Kosterm., and several species of Litsea. Oil or mucilage cells are characteristic of most of the woods of this family, occurring usually in either the wood parenchyma or the rays and relatively rarely in both, e.g. in some species of Licaria, Aiouea, Cinnamo- mum, Cryptocarya, Persea and Phoebe; more commonly present in the wood parenchyma than in the rays and then occurring round the vessels and in the bands of parenchyma, if these are present. Isolated cells some- times appear to be scattered among the fibers in transverse sections, but at least some of these are due to the presence on the margins of the rays of occasional high oil cells which may project into a transverse section that misses the rest of the ray. According to Janssonius (1934) oil and mucilage cells are indistinguishable except by their contents and are to some extent interchangeable; he found oil cells to be more numerous than mucilage cells, except in Cinnamomum iners Bl., C. javanicum BL, and C. burmannii Bl.; to these may be added Persea thunbergii Kosterm. Oil or mucilage cells not observed or very rare in some species or specimens of Actinodaphne, Beilsehmiedia, Cryptocarya, Endiandra, Hypodaphnis, Lin- dera, Neolitsea and Ocotea rodiaei Mez, and Kanehira notes the following genera of which individual specimens may or may not posses these cells: Actinodaphne, Cryptocarya konishii Hay., Lindera, Litsea, Machilus (=' Persea), Phoebe, and Neolitsea. 204 R E I N W A R D T I A [VOL. 4 INFLORESCENCE The inflorescences are always axillary (in some American Persea species terminal, according to Mez), even in species with sympodial rami- fication. With the exception of Cassytha the dichasial or racemose inflo- rescences are limited. The inflorescences originate either from a mixed bud (but this is not very common) or from a flower bud. They are as a rule panicles, each ramification subtended by a bract; the ramifications repeat the phyllo- taxis, but often the phyllotaxis becomes of a lower order (% or even Vi in Persea americana Mill.) ; this is common in small (reduced) inflorescences. Sometimes the inflorescence is a raceme; its cymose nature may be deduced from the 2 bracts which subtend the flower (e.g. Eusideroxylon melagangai Sym.). A pseudo-spike occurs in some species (Nectandra leucantha Nees). When the ultimate ramifications are shortened, the flowers may be arrang- ed in a head {Endlicheria glomerata Mez; Litsea) ; pleiochasia are some- times present {Licaria gvianensis Aubl.) ; in Licaria capitata Kosterm. the entire inflorescence is reduced to a head on a long peduncle. Sometimes the inflorescences occur on shortened branchlets (Litsea, Lindera, Ocotea aniboides Mez). As a rule the inflorescences are erect (in Endlicheria arunciflora Mez they are perhaps pendulous) ; the infructescence, by the weight of the fruit, is often pendulous. The inflorescence bracts usually drop at an early stage, although more or less persistent bracts occur in species of several genera. Male inflorescences are as a rule more-flowered, more branched and bigger than female ones {Endlicheria). In Litsea and Lindera the flowers, arranged in a pseudo-umbel, are surrounded by decussate, large, persistent bracts; usually this involucre drops after anthesis, but (especially in male flowers: Litsea) it persists and the inflorescence drops as a whole. The lower pair of the involucral bracts is usually transversal, the third one is just above the umbel-bearing bract. The pseudo-umbel is actually a shortened spiral; each flower is subtended by a small bract (sometimes, however, reduced). Rarely the number of involucral bracts is reduced to two. The perianth of the flower is often reduced, as its function is taken over by the involucrum. The inflorescence of Sassafras should be (according to Mez) of the same type; the bracts at the base of the inflorescence are supposed to be the involucral bracts; the bud scales replace the involucral bracts to protect the flowers. The bracts, however, are not decussate and do not persist (they drop before anthesis) and hence are not exactly comparable 1957] A. J. G. H. KOSTERMANS : Lauraceae 205 to those of Litsea and Lindera, although they have probably originated in the same way. In Asiatic species of Beilschmiedia and in Actinodaphne the same phenomenon occurs, and it is assumed that Umbellularia has the same kind of enveloping bracts, although in this genus the spirally arranged bracts occur at the top of the inflorescence. In Cassytha the ramification of the inflorescence continues indefini- tely; each branch is subtended by a reduced bract; the spiral follows the number %. FLOWER The flowers are actinomorphic, usually bisexual, but dicliny is rather common. In dioecious species the female flowers are usually bigger, the tube especially is larger. In Endlicheria all species are dioecious, in Litsea and Lindera several, in Ocotea (including the subgen. Nectandra) the greater part of the species. In monosexual flowers remnants of the reproductive organs are as a rule present in the opposite sex, but in the male flower the reduced ovary may be completely suppressed, although style and stigma are always better preserved. It is assumed that the "primitive" flower has two outer whorls of fertile, glandless stamens, a third whorl of gland-bearing fertile ones and a fourth whorl of staminodes. These staminodes are either relatively large and heart- or spear-shaped (Persea, Beilschmiedia, etc.) or minute (Ocotea, etc.); this character has been considered of importance to segregate genera. The unilocular ovary has a single ovule, opposite the first leaf of the second perianth whorl; it is pendulous and anatropous and has 2 inte- guments. According to anatomical investigation the two perianth whorls appear simultaneously and hence there is no reason to consider the inner whorl of corolla nature. In some genera (Persea) the outer tepals are smaller than the inner ones; the reverse is very rare (Endlicheria paradoxa Mez). Sometimes the outer whorl of stamens is petaloid (Phyllostemono- daphne and Dicypellium). In Potameia, Neolitsea, and Laurus the perianth consists of two whorls of two decussate tepals each. In (abnormal) flowers of Persea americana Mill, it has often been observed that the perianth becomes dimerous, although the number of stamens in each whorl remains three. Abnormal numbers of tepals are often found in Litsea. Mez, who observed flowers with 5 tepals, concluded alliance with Berberis. Octo- litsea, as established by Liou Ho, with 8 tepals, should be considered an 206 REINWARDTIA [VOL. 4 1957] A. J. G. H. KOSTERMANS: Lauraceae aberrancy. In Litsea sometimes the reverse happens and tepals are chang- ed into stamens. Mez stressed the apetalous nature of the flower by pointing at Cassytha, where the 1/3 -spiral is continuous through the (reduced) leaves, the inflorescence bracts and the perianth. STAMENS. During its development the anther has usually already been completed before the basal part of the meristem develops into a filament. The anther consists of parenchymous tissue with numerous oil and slime ducts, the remnants of the not resorbed tapetum layer, the pollen cells, a layer of fibers that only partly covers the peripheral part, and an only slightly thickened epiderm. The anthers are originally four-celled; this is clear from two-celled ones where the ablastic upper part often contains a single undeveloped or partly developed cell. In Potameia chinensis Kosterm. and P. velutina Kosterm. the two remaining cells become confluent, although the flap is still two-lobed; some species of Mezilaurus show the same tendency. Occasionally sterile stamens are present and as a rule they are reduced to foliar staminodes; in some cases, and often in the third stam- inal whorl, the anthers are normal in outline but lack cells (Aniba kappleri Mez, Cryptocarya species from Madagascar, etc.). If 4 anther-cells are present, the "primitive" case is represented by 2 pairs above each other. In several genera the 4 cells are arranged in one arc-like row; this may be due to enlargement of the base of the anther; intermediate stages between superimposed anthers and those in one row are common. The anthers are extrorse, introrse, lateral or apical; if there are two pairs of anther-cells, one pair (especially of the third whorl) may open in a direction different from that of the other pair. Mez tried to correlate extrorsity or introrsity with the position of the floral glands. Although the theory is ingenious, it cannot explain all cases. In some species of Persea subgen. Alseodaphne and Licaria the anthers are very thick and fill the available space completely; they have small glands; the cells are apical, which perhaps may be explained by pointing out that it is only by this arrangement that pollinating insects can easily reach the pollen. When lateral cells are present, it also seems to me that this makes the pollen better accessible to insects. In flowers with spreading perianth the outer two whorls of stamens are as a rule also spreading and hence their introrse cells are well exposed; in these flowers the third whorl is erect and thus has its extrorse cells well exposed. Extrorsity and introrsity have little generic value. In Litsea usually all anthers are extrorse, which also is easily understood for this kind of umbel-like inflorescence. The velum of the anther cells contracts considerably after dehiscence; it is torn off at its base along a line where strengthening elements are lacking. As I pointed out elsewhere, the valves in Mezilaurus seem to open from top to base; actually they open in the normal way from base to top, but the bases of the anthers are enlarged and have moved the bases of the cells outwards; the cells are apparently transverse. The filaments are either long and slender (Beilschmiedia species, Persea species, Litsea, etc.) or as broad as, or broader than the anthers, or completely lacking. The shape of the anther is rarely constant in a genus (constant in Aniba, Endlicheria). The third whorl of filaments is often connate (Systemonodaphne) or closely pressed together (Madagascar species of Cryptocarya). Irregularities in the number of stamens were found in Beilschmiedia roxburghiana Nees (LauromeirrilMa was created on account of these irreg- ularities). The number of anther whorls is generally used for delimitation of genera. It is, however, a character to be handled with care. If it is not correlated with other characters, it has no generic value. In Aiouea 1, 2, or 3 whorls may be present; in Beilschmiedia 2 or 3 whorls; in Endiandra 1 or 2 whorls, etc. FLOWER GLANDS are as a rule represented in all genera. Anatomical evidence (epiderm not cutinized; according to Mez cuticle occurring some- times; small-celled parenchyma) shows that they are no staminodes, as suggested previously. Oil cells and a rudimentary vascular bundle are present. They may be minute or so large that they fill the space between the filaments completely. They are either sessile or stalked and the stalks may be partly grown together with the filaments. As a rule each filament is flanked by two glands. Sometimes there is a considerable distance between the filaments and their glands. In the regular type of flowers with 3 whorls of fertile stamens usually the third whorl only is provided with glands, but exceptions in a genus are not rare. Rarely all filaments are accompanied by glands (Ocotea subgen. Pleurothyr'ium, Endlicheria species, Urbanodendron). The number of glands, their presence or absence, has no generic value. R E I N W A R D T I A [VOL. 4 1957] A. J. G. H. KOSTERMANS : Lauraceae 209 The glands are usually globular; exceptionally they may be flat or heart-shaped; in the latter case they strongly resemble staminodes. STAMINODES. There are two trends of reduction of stamens; in Lica/ria the two outer staminal whorls are reduced to thin, foliate staminodes, without any difference between antherial part and filament. In Phylloste- monodaphne and Dicypeilium the outer whorl has become completely tepaloid (in Eusideroxylon the 4 minute cells are still present, but the stamens are tepaloid) ; in Licuria subgen. Misantheca the two outer staminal whorls are entirely lacking. This mode of reduction, starting with the outer whorls, is also present in Endiandra and Mezilaurus. If, on the contrary, the two outer staminal whorls (or one) are fertile and the third whorl (and/or the second), the reduction of the stamens is usually less complete; they are not strap-like and show a distinct antherial part (Aiouea). The third whorl tends to be- come staminodial in some species of different genera (Aniba kwppleri Mez, A. canelilla Mez, Endlichewia paradoxa Mez, Cryptocarya perrieri Danguy, Alseodaphne coriacea Kosterm., etc.); in this case the reduction may be restricted to the anther being sterile only, or the stamens may be reduced to a stipe. The fourth staminal whorl in most genera is staminodial. The staminodes are either heart-shaped and stalked (Beilschmiedia, Crypto- carya, Persea), or minute and ligulate (Ocotea, Cinnamomum). In some genera not all species show the same degree of staminal reduction. Some- times they still show the ablastic anther. According to Mez the staminodial tissue is similar to that of the floral glands and perhaps they may act as nectaries. I could not confirm this from living material, in which the slimy, glossy floral glands contrast with the dry staminodes. In Aiouea (and also in African Beilschmiedia species) the staminodes of whorl 4 are inserted lower than the other stamens. The staminodes may be either sessile or stalked, pilose or glabrous (floral glands are never pilose); these characters have only specific value. In Beilschmiedia roxburghiana Nees flowers with 6 or 9 fertile stamens may be found on the same plant; sometimes the missing stamens are represented by an equal (sometimes unequal) number of staminodes. GYNAECIUM. Although the ovary is one-celled and contains only one ovule, there is enough evidence that it is composed of 3 (or perhaps 6) carpels (pseudo-monomerous). This assumption is based on the following considerations: the stigma is sometimes 3-cleft (in the rudimentary ovary of the male flower) ; the fruit of Ravensara is divided (at the base) in 6 compartments; Mez found 6 primary vascular bundles in the ovary of Cinnamomum sericeum Sieb. The style is either long, short, or absent; the stigma is variable in shape; it is either inconspicuous (Beilschmiedia) to relatively large and peltate (Ocotea species) ; in the latter case there is often one lateral indent, which would point to a single carpel. The ovary is epigynous, perigynous, or hypogynous (Hypodaphnis). The single ovule is pendulous and ana- tropous and gives rise to a one-celled fruit. The alleged exception in Beilschmiedia roxburghiana Nees is due to an error in the original de- scription. FRUIT Meissner (1864) stressed the importance of the presence or absence of a fruit cup for generic delimitation. Starting from the assumption that a superior ovary is more primitive than an inferior one we may arrange the genera according to this char- acter, which is correlated with the character of the fruit. In flowers with a superior ovary the fruit is seated on a bare pedicel, with none or hardly a trace of the receptacle; in perigynous flowers the fruit is at the base partly covered by a cup, which represents the deep flower tube. In the most differentiated genera the ovary is included in the flower tube (adnate or not) ; the enlarged tube completely covers the fruit. Sometimes, how- ever, the flower tube enlarges more quickly than the ovary, which results in a fruit at first completely enclosed in a cup (Aniba, Ocotea, Litsea), although at a later stage it may be much larger than the cup. During this transitional stage it is very difficult to differentiate the fruit from those whose ovary is at first completely included in the tube and whose fruit is completely enclosed in the accrescent tube (Cryptocarya, Raven- sara) . In Eusideroxylon the flower tube is shallow, although it completely envelopes the mature fruit. Meissner combined genera showing transitional inclusion of fruit with genera like Cryptoearya; this is not advisable as the species showing transitional stages are not limited to definite genera. In genera in which the perianth drops completely and in which no trace of a cup is to be found (Beilschmiedia, Endiandra, Persea), the abscission line is very sharp; the perianth generally drops as a closed ring consisting of a small basal collar and the tepals. In rare cases the abscission line is somewhat above the base of the flower tube. In that case a flat, small disc will be found below the fruit (Mezilawrus). In Phoebe and Apollonias tube and tepals together enlarge slightly; the tepals harden and are pressed against the fruit. Although this is used as a generic character, the persistent, indurate perianth is also found in 210 REIN W A R D T I A [VOL. 4 1957] A. J. G. H. KOSTEBMANS: Lauraeeae 211 some species of other genera (Aiouea, Ocotea, Persea subgen. Alseo- daphne), although as a rule to a lesser degree. In Cinnamomum (but also in Phoebe amoena Mez and other American species of Phoebe) only the basal half of the tepals with the tube persists under the fruit. The abscission line here goes half way through the tepals and results in a cup crowned with 6 truncate lobes. In perigynous flowers the resulting cup varies considerably as to depth and size and in a single genus, like Litsea, an almost flat, inconspi- cuous cup may be found, or else a cup that covers the fruit completely (although in a different way than in Cryptocarya) ; as a rule, however, a normal hemispherical cup is present. The cup is generally more or less fleshy, although in a dried condition it may appear woody; it is either smooth, or (Aniba) covered with round, corky warts (lenticels). In a fresh condition it is usually green, yellowish green or red. The cup is either sharply separated from the stalk, or gradually merges into it. In the latter case the fleshy stalk and the cup may be highly coloured (Endlicheria). The rim of the cup is either simple, double or triple. Mez ascribed this phenomenon to a mechanical process by which the growing fruit carries the inner margin upwards. I cannot confirm this view. The second rim actually represent the grown-out tissue of the base of the fruit and second whorl of stamens together, and the third rim that of the third whorl of stamens. These double margins are constant in some genera: Licaria, Systemonodaphne, Urbanodendron, but occur also occasionally in species of other genera {Ocotea). Sometimes the rim is wavy, because the broadened bases of the perianth are persistent (Ocotea). The fruit pedicel is either cylindrical, or may (as in Endlicheria) be fleshy and conical, or (in Dehaasia) greatly swollen, club-shaped and highly coloured (also in Ocotea clavigera Mez), in Litsea species and in some species of Persea subgen. AJtseodaphne). The usually bright red stalks form a striking contrast with the black fruit. The fruit is a one-seeded berry. The exocarp is ordinarily glossy and smooth, although in some cases it becomes woody (Licaria camdra Eos- term., Alseodaphne paludosa King, Eusideroxylon, Cryptocarya). As' a rule it is black in colour, sometimes red; yellow fruits are rare (Actinb- daphne). In an immature stage the fruit is generally white or greenish white, later red; some fruit are pure porcelain-white (Litsea). The mejocarp is usually succulent; it is mostly thin, but may reach considerable thickness in the cultivated Persea americana Mill. According to Mez the mesocarp has its origin in the outer integument. The taste is usually bitter, aromatic and astringent. In most cases the endocarp is thin and smooth. In some Potameia species a kind of aril is present, which covers the embryo completely but for a small apical area. . In fruits completely covered by the accrescent flower tube the latter assumes the function of the exocarp and mesocarp; the exocarp of the fruit becomes woody (Ravensara, Cryptocarya, Eusideroxylon) and is often ribbed (Cryptocarya, Ravensara) or furrowed (Eusideroxylon). In Eusideroxylon, Cryptocarya, and Ravensara the accrescent flower tube becomes entirely adnate to the fruit, although the ovary in the flower is still free from the tube. In the flowers of these genera the style is often constricted by pressure where it passes the narrow apical aperture of the flower tube; in Eusideroxylon, however, the tube has a very v/ide aperture and nothing indicates at this stage, that the receptacle will later enclose the ovary. In the monotypical genus Hypodaphnis the ovary is inferior from the beginning; here the exocarp —- contrary to that of Cryptocarya, Ravensara, and Etisideroxylon — remains thin. The shape of the fruit it usually ellipsoid, sometimes globose, some- times slightly oblique (Persea subgen. Alseodaphne) and often (in genera without cup) provided with a neck at the base. The ripe fruit has either no trace of a style and may even be a little depressed or it is mucronulate (base of style). All tissue usually contain plenty of oil and mucilage cells. In young fruit a clear, jelly-like substance is often present at one side between the cotyledons; this may be residuary endosperm. The seed is composed of two large, flat-convex cotyledons that are easily separable (in Ravensara and Cassytha they are more or less fused) ; as a rule the cotyledons are white, rarely pink; they contain fatty oil (Persea), carbohydrates and proteins. The flattened apical (lateral in Beil- schmiedia variabilis Robyns & Wilcz.) corculum is completely covered by the cotyledons, which are attached to it by a large area of tissue; the first leaves of the plumule, which are often pilose, are well developed, usually one is larger than the other and covers it slightly. The outside of the cotyledons is either smooth or irregularly sulcate. In Ravensara (in the basal part of the fruit) the endocarp penetrates the seed with 6 septa (rarely 12 septa). The radicle is as a rule rather small, rarely swollen (species of Beil- schmiedia). The testa is thin and smooth (in Cassytha it is tough, probably because the seeds are dispersed birds that swallow them). DISPERSAL OF SEEDS. The fruit is dispersed mainly by birds, probably also by squirrels and monkeys. These are attracted by the glossy black, 212 R E I N WARDTIA [VOL. 4 1957] A. J. G. H. KOSTEBMANS: Lauraceae 213 red or yellow berries, often seated on a (swollen) red stalk. As there is no protecting coat except in Cryptocarya, Ravensara and Cassytha, the seeds only survive when they are not swallowed or too much damaged. The production of fruits is often enormous. The taste is as a rule bitter or pungent and I could observe that animals usually bite off a little of the tissue and then drop the fruit. The heavy fruits of Eusideroxylon are carried around by porcupines on the ground and sometimes by monkeys. Persea tonkinensis Kosterm. is dispersed by water. The species usually grows on alluvials along rivulets where periodical inundation occurs. They float by means of the air-filled space between seed coat and endocarp. Riverine species are apparently also dispersed by water. POLLEN AND POLLINATION According to Erdtman (Pollen Morphology and Plant Anat. 221. 1952) the pollen grains are non-aperturate, more or less sphaeroidal, 24—40(—70)µ in diameter, tenui-exinous, usually provided with spinules or spinuloid projections. The exine stratification is obscure. In general the pollen grains are similar to those in Gomortegaceae and Hernandiaceae. The flower glands point to insect-pollination; furthermore the flowers are often very fragrant. I never saw butterflies near the flowers and only occasionally found small beetles crawling around in them. Apparently self-pollination is frequent. The colour of the flowers is white or greenish, rarely yellow or red. In Persea subgen. Nothaphoebe they turn from yellow to red after anthesis. In some genera (Liearia, Persea subgen. Alseodaphne) the flowers hardly open and the anthers are so near the stigma that self-pollination seems inevitable. The existence of dioecious flowers points to the development of cross- pollination. Pollination has been investigated in Persea americana Mill, by Stout (in J. New York Bot. Gard. 25: 1-7. 1924) ; he discovered dichogamy. The flowers of one specimen open and close in two separate periods; during the first the pistil becomes ready for pollination, and fertilisation takes place during the second phase when the pollen is shed. Abnormal weather conditions may intervene and even cause overlapping of periods. Stout discovered two groups of varieties, of which one opens its flowers for the first period in the forenoon, the other in the afternoon. The pollination is effected by insects. Mez contents that the volatile oil prevents damage to the leaves by animals; in Indonesia, however, Persea americana Mill, is completely stripped of its leaves every year by caterpillars (Cricula trifenestrata). MYRMECOPHILY Mez gives examples of myrmecophily in American Pleurothyrium species. From New Guinea thus far two cases have become known {Crypto- carya caloneitra Kosterm. and Beilschmiedia myrmecaphila Kosterm.). The pith of the (secondary) terminal branches (usually swollen) is eaten away; there are oval or round entrance holes. CHROMOSOME NUMBERS The following figures are copied from Darlington — Janaki Ammal, Chromosome Atlas of cultivated plants (1945). Laurus nobilis x = 7 . . Cinnamomum- x — 12 Cinnamomum camphora ^i C. japonicum (== pedunculatum) 24 C. linearifolium C. obtusifolium C. sieboldii {=burmannii) C. zeylanicum Lindera x = 12 L . glauca . . . . . ; . . . . L. benzoin (= aestivale) Persea x = 12 P. americana P. pubescens (= caxolinensis) Sassafras x = 12 . S. albidum (= officinale) 42 24 24 24' 24 24 24 24 24 24 48 PALAEONTOLOGY The oldest fossil Lauraceae are of Tertiary age (Palaeocene). The pretended Cretaceous species (Lesquerreux) from N. W. America is appa- rently not lauraceous. Miocene and Pliocene fruit and flowers are known from Siberia, which makes it evident that Lauraceae occurred far North. In the Pliocene period Lauraceae disappear from Europe, except for the genus Laurus. 214 R E I N W A R D T 1 A [VOL. 4 A large number of fossil Lauraceae have been described. Species represented by fruit only (Reid & Chandler, London Clay Flora* 1933) may be referred to half a dozen genera. In contemporary species it is im- possible, without flowers at hand, to identify most of the genera. Conse- quently generic identification of fossil material on the basis of fruits and/ or leaves is usually dubious. Species represented by leaves only can hardly be placed at all. Hollick (in Bull. New York Bot. Gard. 12: 298-300. 1924) distinguishes between Nectandra and Ocotea, genera which in contempory species are only differentiated by the position of anther cells and some minor (and not constant) characters of the flowers. The identifications in question should therefore be considered utterly unfounded guess work. Triplinerved leaves are by palaeontologists consistently referred to Cinnamomum, although in contempory Lauraceae a dozen genera have species with such leaves. Triplinerved leaves are furthermore so common in other families that even the identifications as to family are doubtful. Reid and Chandler, without sufficient knowledge of the variability of fruit in modern Lauraceae, often identify fossil material by the fruit. The results, if not completely wrong, are at any rate dubious (fruits of Ocotea species are apiculate, in Beilschmiedia usually not; the cup in Cinnamomum, only occasionally leaves a trace on the base of the fruit, but in numerous cases, where the cup is shallow it certainly does not, etc.). Bandulska (Eocene of Bournemouth) approached the problem of identi- fication of leaves by means of studying cuticles and stomata. This approach should also be handled with care, as is proved by Aniba ridleyana Mez, which has the Aniba type of stomata, but actually represents a species of Ocotea. She herself stresses, that in modern species of a single genus the variation in stomata and cuticle is astonishing. Of Trianthera Conwentz a complete flower is preserved in amber, which makes it possible to relate it to Eusideroxylon. Palaeontological results consequently have given us thus far only an incomplete idea of the distribution of Lauraceae in the Tertiary period, and have not yielded information on ancestral problems. Actinoda'phne Aniba Beilschmiedia Benzoin Described fossil genera (incomplete) : Cinnanwmoides Daphnogene Cinnamomum Daphnophyllum Crowella Endiandra Cylicodaphne Laurinium 1957] Laurinoxylon LauriphyUum Laurocalyx Laurocarpum Laurus Lindera A. J. G. H. KOSTERMANS :: .Lauraceae 215 Litsea Mespilodaphne ' Neolitsea Ocotea Oreodaphne Persea RELATIONSHIPS Perseoxylon Protoravensara Sassafras Tetranthera Trianthera On account of the similarity of the dehiscence of the anther cells in Lauraceae and in some genera of Berberidaceae, Monimiaceae and Hama- melidaceae, relationship with these families has been suggested. There are now two current views: 1) Lauraceae are related to Thy- melaeaceae and 2) they belong in Ranales, Of the two families Hernandiaceae and Gomortegaceae, nobody doubts that they are very close to Lauraceae. Lindley (1853) already recognized that Lauraceae are far removed from Berberidaceae because of their polypetalous flowers, hypogynous stamens and endospermous seeds. Still, Hallier f. (1912) derived the family (incorporated in the sub- order Laurineae of the order Annonales) from hypothetical Proberberi- deae; he suggested relationship with Monimiaceae, Calycanthaceae and Chlorantaceae. Alliance with Thymelaeaeeae was accepted by Lindley (1853), Baillon (1870), Bentham (1880), who placed the family in the Daphnales series, and Pax (1889), who considered Lauraceae to be a connecting link between Polycarpicae and Thymelaeaeeae. Pax pointed to the significant differences between Lauraceae and Monimiaceae (acyclic flowers, several apocarp carpels, usual presence of/ endosperm, different pollen, etc.). Warming (1895), Bessey (1915), Johnson (1931), Engler & Diels (1936), Wettstein (1935), Gundersen (1943) and Pulle (1950), following Eichler (1886) adhere to relationship with Magnoliaceae (Magnoliales) of Ranales (Polycarpicae). Hutchinson (1926) placed the family (with Monimiaceae, Hernandia- ceae, Gomortegaceae and Myristicaceae) in a separate order Laurales (next to Annonales), which he considered as reduced perhaps from winter- aceous ancestors of Magnoliales. Apparently Myristicaceae are close to Lauraceae (apetalous flowers, ovary, etc.) ; Garret (1933) could find wood-anatomical support for this view. 216 REINWARDTIA USES [VOL. 4 Persea americana Mill, produces the familiar alligator pear or avoeado with edible mesocarp. The seeds contain oil. Well-known timbers are greenheart (Ocotea rodioei Mez) and iron- wood (Eusideroxyion zwageri T. & B.). Many others are used locally and are highly esteemed for their durability and fine grain (Ocotea bullata E. Mey., Persea lingue Nees, Dehaasia eaesia Bl., etc.). Most timbers are liable to attack by borers and fungi; on account of the excellent grain it is worth while to apply preservation methods. Several Lauraceae yield commercial volatile oils, such as rose wood (Ardba rosaeodora. Ducke and Aniba duckei Kosterm.). Numerous barks have commercial value because of their oil content: Cinnamon (Cinnamomum zeylanicum Bl. and C. cassia Bl.), Massoy (Ciryp- tocarya massoy Kosterm.), Litsea odorifera Valet, (an extremely sweet- scented bark), Licaria cinnamomoides Kosterm. with the smell of nutmeg (the fruits are sold locally), Aniba caneiilla Mez, Licaria puchury-major Kosterm., Dicypellium canjophyttatum Nees and Cryptocarya moschata N. & M. smelling of nutmeg and/or clove (marketable fruit), Rawensara aromatica Sonn. (saleable fruit), Endlicheria longifolia Mez with the odour of aniseed, Ocotea foeniculacea Mez with the scent of Foeniculum, etc. — Cinnamomum camphora Nees & Eberm. yields the well-known Japan cam- phor. Cinnamomum porrectum Kosterm. has saffrol in its bark (for scenting soap). — Alcaloids could be extracted from several barks and seeds: Aniba coto Kosterm., Ocotea veraguensis Mez, Ocotea rodioei Mez, etc. (cf. Baillon for further particulars). DISTRIBUTION Lauraceae are typical for the tropical rain forest where they ascend to 4000 metres. Not only do they represent an integral part of the mon- tane forest in this area; they are just as abundant in the lowland forests. They occur in marshy places and on well-drained soils. In forests under seasonal climatic conditions they become rare. North and South of their main area they are found as far as Cali- fornia (Umbellularia) and other parts of the United States of America (Sassafras) up to 45—50° N. L. In Europe they do not reach farther than the Mediterranean area (Laurus nobilis L.) ; in India they are found in Nepal; in China they go as far as Korea (Litsea) ; they also occur in Japan (Litsea, Lindera), they reach Southern Chile (island of Chiloe: Persea. lingue Nees) and Argentine; in South Africa only a few species 1957] A. J. G. H. KOSTEEMANS: Laiiraccae 217 are found (Ocotea bullata E. Mey.), but Madagascar has very many, in- cluding endemic genera. Lauraceae reach New Zealand "with two species (Beilschmiedia tawa Benth., B. tarairi Benth.). Richest in species are perhaps South and Central America, followed by Malaysia. In Africa the family is represented rather poorly. This may be due to two factors: the scarcity of the pure tropical rain-forest and the desiccation of the African continent since the Tertiary period. In Australia its main centre is the rain forest area of Queensland and New South Wales. Several genera have pantropical distribution (Beilschmiedia, Crypto- carya, Persea, Phoebe, Litsea, Cassytha), other are restricted to Asia and (or) Australia (Actinodaphne, Persea subgen. Alseodaphne, Cinnamo- mum, Dehaasia, Endiandra, Eusideroxyion, Lindera, Hexapora, Neolit- sea). — Cinnamomum, Actiodaphne and Litsea are distributed over the whole area, with their main centre in Malaysia. Eusideroxyion is restricted to Borneo and parts of Sumatra; Dehaasia to Malaysia (very rare in New Guinea) and Hexapora to Malaya. Lindera and Litsea are (more so than the other genera) represented in China, Korea and Southern Japan. Endi- andra has its centre in the Eastern part of Malaysia (and in Australia). Madagascar harbours the endemic genus Ravensara. Potameia has a disjunct area, the bulk of the species occuring in Madagascar, one in Nepal and one in Southern China. Africa has the remarkable endemic genus Hypodaphnis. The only genus rich in species in tropical Africa is Beil- schmiedia. Apollonias has one species in India and one in Cape Verde. Typical American genera are Ocotea, including Nectanclra (with Pleuro- thyrium), Aiouea, Aniba, Systemonodaphne, Urbanodendron, Licaria, Mezilaurus, Dicypellium and Phyllostemonodaphne. — With other con- tinents America shares Beilschmiedia, Cryptocarya and Litsea as well as North American Lindera and Sassafras; the latter genus has a disjunct area, one species occurring in America and two in China and Formosa. Australia has the same genera as Malaysia, with a preponderance of Endiandra. This continent is the main centre for Cassytha. SIZE OF THE GENERA The figures after the generic names indicate the number of published binomials. From the number of species of revised genera it may be assu- med that the actual number of species will be about10% - 40 % lower. The accepted genera are printed in bold face type. 218 B E I N W A R D T I A [VOL. 4 1957] A. J. G. H. KOSTIERMANS : Lauraceae 219 Acatsjavalli 0 Acrodiclidium 62 Actinodaphne 128 Adaphtos 0 Adenodaphne 1 Adenostemwm 1 (Gomorteg.) Adenotraehelium 1 Afrodaphne 15 Agathophyllum 16 Agriodaphne 1 Aiouea 46 Alseodaphne 58 Ampelodaphne 4 Anaueria 1 Aniba 74 Apella 0 Aperiphracta 4 Aperula 18 Api'wa 0 Apollonias 12 Aydendron 45 Balanopsis 4 Beilsehmiedia 236 Bellota 5 Benzoin 89 Berniera 1 Berrija 1 Bihania 1 Brassiodendron 1 Bistama 0 BoWw 2 Boldus 3 Borbonia 19 Bryantkea 1 Calodium 1 Ccdosmon 3 Calycodaphne 3 Camphora 47 Camphorina 4 Camphor omoea 10 Canella 2 Cansiera 1 Caryodaphne 5 Caryodaphnopsis 3 Cassytha 50 Cecidodaphne 1 Cedrota 2 Ceramocarpium 10 Ceramophora 6 Chanekia 6 Chibaca 1 (non Laur.) Christmannia 1 (= Salacia) Cinnamomum 341 Clinostemon 1 Colomandra 0 Cryptocarya 318 Cubeba 1 Curondia, 1 (= Salacia) Cussuta 1 Cyanodaphne 3 Cylicodaphne 59 Dambumeya 2 Daphnidium 20 Darwinia 1 Decapenta 1 Dehaasia 33 Dendrodaphne 1 Dictyodaphne 7 Dicypellium 1 Dipliathus 1 Dodecadenia 4 Douglassia 2 Dovera, 0 (Salvador.) Drimophylltom 0 Ehrhardia 0 Endiandra 95 Endlicheria 55 Endocarpa 1 Euodia 0 Euosmus 6 Euphoebe 7 Eusideroxylon 2 Evelyna 2 Evodia 2 Evonymodaphne 1 Evosmus 1 Farnesia 0 Fwtfa. 11 Glabraria 7 Goeppertia 16 Gymnobalanus 10 Haasia 23 Heckaria 1 Hexanthiis 1 Hexapora 1 Huberodaphne 1 Hufelandia 16 Plypodaphnis 1 Icosandra 1 Iteadaphne 2 Jozosmene 0 Jozoste 52 Lauromerrillia 1 Laurus 334 Lepidadenia 30 Leptodaphne 3 Lethedon 1 (?) Licaria 52 Lindera 135 Linharea 2 Litsea 474 Machilus 100 Malapoenna 168 Mespilodaphne 65 Menestrata 1 Mezia 6 Mezilaurus 10 Micropora 2 Misanteca 20 Nectandra 248 Nemodaphne 1 Neocinnamomum 8 Neolitsea 93 Neosilvia 0 Nesodaphne 3 Nobeliodendron 1 Nothaphoebe 41 Nyctandra 0 Nyrophyllum 0 Ocotea 697 Oreodaphne 178 Ozanthes 1 Ozarthris 4 (?wm Laur.) Parabenzoin 2 Parthenoxylon 3 Persea 240 Petalanthera 1 Peumus 5 (3) Phoebe 174 Phyllostemonodaphne 1 Pipalia 1 Pleurothyriwm 18 Polydenia 18 Pomatium 1 Porostema 2 Potameia 22 Pseudocryptocarya 1 Pseudolitsea 1 Psewdosassafras 2 Purkayasthea 1 Quinquedula 0 Ravensara 27 Rombut 1 Rumputris 1 Salgada 1 Sassafras 16 Sassafridium 2 Sckauera 0 Sciadiodaphne 0 Sebifera 3 Senneberia 0 Septina 0 Silvia 12 Spironema 1 Stemmatodaphne 1 Strychnodaphne 6 Symphysodaphne 1 Synandrodaphne 3 Syndiclis 2 Systemonodaphne 3 Tamala 8 Teleiandra 1 Tetradenia 45 Tetranthara 220 Thouvenotia 1 Tomex 12 Triplomeia 1 Tylostemon 45 Umbellularia 2 Urbanodendron 1 Volutella 1 Wimmeria 1 Yiishunia 1 The total number of published binomials is 5462 (1956). The accepted genera are listed below, arranged according to their size. Ocotea 697 Litsea 474 Persea 339 Cinnamomum 341 Cryptocarya 318 Beilsehmiedia 236 Phoebe 174 Lindera 135 Actinodaphne 128 Endiandra 95 Neolitsea 93 Aniba 74 Endlicheria 55 Licaria 52 Cassytha 50 Aiouea 46 Dehaasia 33 Ravensara 27 Potameia 22 Sassafras 16 Mezilaurus 10 Systemonodaphne 3 Eusideroxylon 2 Apollonias 2 Laurus 2 Umbellularia 2 Dicypellium 1 Hexapora 1 Hypodaphnis 1 Urbanodendron 1 Phyllostemonodaphne 1 The total number of binomials of accepted genera is 3435; we may assume that Lauraceae comprise between 2000 and 2500 species. 220 R E I N W A R D T I A CLASSIFICATION [VOL. 4 Linnaeus recognized only two genera (Laurus and Cassytha). •— A. L. de Jussieu (1789), under his Order Lauri enumerated the genera Laurus, Ocotea and Aiouea and as allied genera Myristica and Virola, and Hernandia. Several lauraceous genera (Ravensara, Cassytha, Lindera, Tomex and Liearia) were relegated to the genera incertae sedis. The first general monograph appeared in 1836 (C.'G. Nees von Esenbeck, Systema Laurinarum), who divided the family into 13 tribes. Nees created a considerable number of small genera and moreover reinstated older genera, which had formerly been incorporated in Laurus. Altogether he recognized 34 genera. The second monograph appeared in 1864 (Meissner in DC, Prodr. 15 (1): 1-265). Although Meissner already combined several of Nees' genera, he still accepted 46 genera, which he divided into 4 tribes. Actually Lauraceae were divided into 3 suborders, of which the second (Gyroear- peae) is now included in the family Hernandiaceae; suborder 1 is Lauri- neae; suborder 3 Cassytheae. In our classification these two are considered subfamilies (according to modern rules). The suborder Laurineae was divided into 2 groups (without names) according to presence or absence of an involucrum of bracts under the subumbellate flowers. The group without involucrum comprised 3 tribes, mainly differentiated by the development of the flower tube in the fruit. The tribes were subdivided again, according to the number of anther cells and the number of flower parts. These subdivisions are not named. The group with involucrate flowers comprises only one tribus (Litsaeaceae) which was again subdivi- ded according .to the number of anther cells. Here the subtribus received names (Tetranthereae and Daphnidieae). Baillon (1870) recognized 8 tribus, of which 6-8 are now incorporated in Hernandiaceae. The 5 tribus of Lauraceae proper correspond more or less to those of Meissner (Oreo- daphneae Meissner are called here Ocoteae Baill.). Some genera of Meiss- ner's tribus Cryptocaryeae are moved to Baillon's tribus Cinnamomeae, which more or less covers Meissner's Perseaceae. Bentham (1880) divided the family into 4 tribus: Perseaceae without involucre, Litseaceae with involucre, Cassytheae and Hernandieae. He observed, that the fruit characters, as stressed by Meissner, were not sufficiently known. The subdivision of Perseaceae was made according to the number of anther cells and the development of the flower tube 1957] A. J. G. H. KOSTERMANS: Lauraceae 221 in the fruit, these subdivisions remained unnamed. Bentham recognized only 33 genera of the Lauraceae proper. Pax (1889) composed a very artificial classification based on the number of anther cells. In his system the aberrant genus Cassytha, often considered to belong to a separate family, was classified into the tribe Lauroideae, merely by its 2-celled anthers, and genera like Eusider- oxylon and Gryptocarya, which are closely related, were referred to widely separated tribes. Curiously enough wood-anatomical data (Dadswell & Eckersley 1940; Desch 1941) concur with regard to this artificial sub- division in Lauroideae and Persoideae. After Pax no general monograph has appeared. Mez' system of Ame- rican Lauraceae (1889) is worth to be mentioned here. He divided the family into two suborders: Laureae and Cassytheae. The Laureae were subdivided into two tribus: Perseae and Litseeae, differing by the ab- sence or presence of an involucrum; the next subdivision (unnamed) was made according to the number of anther cells. In the classification outlined below, I have adopted the following sequence of characters (according to their importance) : 1. The development of the flower tube in the fruit, which runs more or less parallel with inferior, intermediate and superior ovary. 2. The presence or absence of an involucrum of decussate persistent bracts surrounding and enveloping the pseudo-umbels. 3. The number of fertile stamens. 4. The number of anther cells. 5. The development of the 4th staminodial whorl. It should be stressed here that several genera are linked by one or a few intermediate species. Apparently the combination of characters is more or less indefinite and almost all combinations are represented. We may assume either that missing combinations are extinct or that the potentiality, that they will develope is still present. With such an assumption and without any indi- cation of the palaeontological succession I have refrained from trying to make a chronological family tree and have simply grouped related genera together. Even the position of the groups (tribus) has no proper phylogenetical base; nobody can tell whether Hypodaphnis with an infer- ior ovary has developed from Ocoteae, from Litseeae, or Perseeae, etc., although in our diagram it is placed at the end (top). The course of the phylogenesis is not clear; the only thing we know is, that the family must be rather old (ubiquist). 222 R E I N W A R D T I A [VOL. 4 SYSTEM OP THE FAMILY 1 a. Arborescent. Leaves normal subfam. A. Lauroideae 2 a. Inflorescences paniculate. Flower-umbels without involucre. Fruit without a cupula tribus I Perseeae 3 a. Anthers 4-celled subtrib. a. Perseineae : 1. Persea 2. Phoebe 3 b. A n t h e r s 2-celled . . . . . . . . . . s u b t r i b . b. B e i l s c h m i e d i i n e a e 3. Apollonias 4. Dehaasia 5. Beilsehmiedia 6. Endiandra 7. Mesilaurus 8. Hexapora 9. Potameia 2 b. Inflorescences paniculate. Flower-umbels without involucre. Fruit-base em- bedded in a cupula tribus II Cinnamomeae 4 a. Anthers 4-celled subtrib. a. Cinnamomineae 10. Ocotea .11. Cinnamomum 12. Actinodaphne 13. Sassafras H. Umbellularia . 15. Dicypellium i b . A n t h e r s 2-celled . . " . . . . . . . s u b t r i b . b . A n i b i n e a e 16. Aiouea 17. Aniba 18. Endlieheria 19. Licaria 20. Urbanodendron 21. Systemonodaphne 22. Phyllostemonodaphne 2 c. Flower-umbels surrounded by an involucre of decussate, large, persistent bracts. Fruit more or less embedded in a cupula tribus III Litseeae . . 5 a. Anthers 4-celled subtrib. a. Litseineae 23. Litsea 2J,.. Neolitsea 5 b. Anthers 2-celled ". . . . subtrib. b. Lauriineae 25. Lindera 26. Laurus 2 d. Inflorescence paniculate. Flower-umbels without involucre. Ovary superior. Fruit completely included in the accrescent flower tube. tribus IV Cryptocaryeae '6 a. Anthers 4-celled . . subtrib. a. Eusideroxylineae 27. Eusideroxylon, 1957] A. J. G. H. KOSTERMANS: Lauraceae 223 6 b. Anthers 2-celled . subtrib. b. Cryptocaryineae 28. Cryptocarya 29. Ravensara 2 e. Inflorescences paniculate. Flower-umbels without involucre. Ovary inferior. tribus V Hypodaphneae 30. Hypodaphnis 1 b . P a r a s i t i c a l c l i m b e r s w i t h o u t p r o p e r l e a v e s . . . . s u b f a m . B . C a s s y t h o i d e a e 3 1 . C a s s y t h a KETST TO THE GENERA la. Arborescent; leaves normal 2 b. Parasitical twiners with reduced leaves 31. Cassytha 2a. Flowers in pseudo-umbels (rarely single), surrounded by large, decussate, per- sistent bracts, forming an involucre, the bracts as a rule decussate . . . . 3 b. Flowers not surrounded by a persistent involucre 6 3a. Flowers dimerous 4 b. Flowers trimerous 5 4a. Flowers dioecious or bisexual; male ones with 3 whorls of 4 stamens, all bearing glands; anthers introrse, 2-celled. Female flowers with 4 large staminodes, all provided with glands 26. Laurus b. Flowers dioecious; male ones with 3 whorls of 2 stamens; inner whorl with glands; anthers 4-celled, introrse. Staminodes in female flowers as many as stamens in male ones 2U- Neolitsea 5a. Anthers 4-celled 23. Litsea b. Anthers 2-celled : . 25. Lindera 6a. Ovary inferior 30. Hypodaphnis b. Ovary superior or — if embedded in the flower tube — not adnate to the tube. 7 7a. Fruit completely included by the adnate, enlarged flower tube; usually only a small orifice at apex 8 b. Fruit seated on and partly covered by a shallow or deep cup developed from the flower tube 10 c. Fruit on a naked pedicel; or the perianth persistent, but no cup . . . . . . 22 8a. Anthers 4-celled 27. Eusideroxylon b. Anthers 2-celled 9 9a. Basal part of fruit septate; cotyledons ruminate . . . . . . 29. Ravensara b. Fruit 1-celled • . . 28. Cryptocarya 10a. Anthers 4-celled . . . . . . . . . 1 1 b . Anthers 2-celled . . . . . . . : . . . 1 6 l l a . Inflorescence before anthesis covered by long-persistent, large, non-decussate bracts (pseudo-involucrum) 12 b. Bracts of inflorescence small, soon deciduous 14 12a. Bracts at the end of a long peduncle . lit. Umbellularia b. Bracts at the base of the inflorescence . . . . . . , 13 13a. Leaves alternate, incised ' 13. Sassafras b. Leaves verticillate, rarely alternate, entire 12. Actinodaphne ^224 R E I N W A R D T I A [VOL. 4 14a. Tepals 9; fruit cup flat 15. Dicypellium b. Tepals 6; fruit cup flat or deep 15 15a. Staminodes of innermost whorl minute or none 10. Ocotea b. Staminodes of innermost whorl conspicuous, stipitate, heart- or arrow-shaped. 11. Cinuamomum 16a. Tepals 9 22. Phyllostemonodaphne b. Tepals 6 17 17a. Fertile stamens 9 18 b. Fertile stamens 3, 6, rarely 9 21 18a. All stamens provided with large glands. Fruit-cup with double rim. Flowers bisexual 20. Urbanodendron b. Only third whorl of stamens with glands, (in one Endlicheria species all stamens with glands, but the flowers dioecious) 19 19a. Flowers dioecious. Fruit cup fleshy, merging into the fleshy, slightly club-shaped pedicel 18. Endlicheria b. Flowers bisexual. Fruit-cup not fleshy, sharply demarcated from the not or hardly thickened pedicel 20 20a. The 3 inner stamens triangular, fleshy, connate; fruit cup with double rim, with persistent, not enlarged perianth 21. Systemonodaphne b. Inner stamens not connate, not fleshy; rim simple 17. Aniba 21a. The 2 outer whorls of stamens staminodial or none. Fruit cup double- or triple- rimmed, distinct from the pedicel 19. Licaria b. All anthers fertile, or those of the 2 outer whorls fertile, the inner whorl sterile, or the first whorl fertile, the inner two sterile. Fruit cup shallow, thickened, merging into the fleshy pedicel 16. Aiouea 22a. Perianth in the fruit indurate, clasping the base of the fruit 23 b. Perianth in the fruit deciduous, or, if persistent, not indurate and not clasping the fruit 24 23a. Anthers 4-celled 2. Phoebe b. Anthers 2-celled 3. Apollonias 24a. Fruit pedicel strongly thickened, fleshy, often highly coloured. Anthers 2-celled. U. Dehaasia b . F r u i t p e d i c e l h a r d l y o r n o t t h i c k e n e d (if t h i c k e n e d , a n t h e r s 4 - c e l l e d ) . . . . 2 5 2 5 a . A n t h e r s 4 - c e l l e d 1. Persea b. Anthers 2~celled 26 26a. Flowers dimerous ,. . . . 9. Potameia b. Flowers trimerous 27 27a. Leaves subverticillate; top of fruit-pedicel with a small disc. Anther-flaps opening from inside to outside 7. Mezilaunis b. Leaves alternate or subopposite; fruit-pedicel without disc. Anthers opening from base to top 28 28a. Fertiles stamens 3 (in one species 6). Leaves areolate . . . . 6". Endiandra b. Fertile stamens 6 or 9. Leave;? reticulate (in one species areolate). Outer 2 staminal whorls introrse; inner one extrorse 5. Beilschmiedia c.' Fertile stamens 9. Leaves reticulate. All anthers introrse. Fruit unknown. 8. Hexapora 1957] A. J. G. H. KOSHERMANS : Lauraceae L A U E A C E A E 225 Lauraceae Lindl., Nat. Syst. 200. 1838. SYNONYMS. — Lauri Juss., 1790. — Laurinas Vent., 1799. — Laurineae St. Hil., 1805. — Laureae Reichenb., 1828. — Laureacsae Lindl., 1833. * A . S u b f a m i l y L a u r o i d e a e K o s t e r m . , 1957. SYNON. — Laurineae verae Zoll. in Nat. en Geneesk. Arch. 1: 603. 1845; Miquel, Fl. Ind. Bat. 1 (1) : 891. 1855. — Laureae Mez, 1889. Arborescent. Leaves normal. Inflorescence definite. TYPE GENUS. — Lmirus L. I. Tribu3 P e r s e e a e Mez, 1889. Tribus Perseeae Mez in Jahrb. bot. Gart. Barlin 5: 3. 1889. SYNON. — Perssae Nees, 1836, p.p. — Perseacsae Maissn., 1834. p.p. — Persoideae Pax, 1889, p.p. Inflorescence paniculate, without involucrum, flower tube deciduous. a. Subtribus P e r s e i n e a e Kosterm., 1957. Anthers with 4 cells. 1. P E R S E A 1 [Plumier] Boehmer, 1760. Perssa [Plum.] Boshmer in Ludwig, Defin. 38. 1760. SYNON. — Famesia Heist, ex Fabric. 1763. — Nyrophyllum Necker 1790 (?). — Menestrate Vello^o 1825. — Alseodaphne Nees 1831. — Machilus Nees 1831. — Tamala Rafin. 1838. — Nothaphosbe Bl. 1851. — Euphosbe El. ex Meissn. 1864. — Stemmatodaphne Gamble 1910. — Caryodaphnopsis Airy-Shaw 1940. PRINCIPAL LITERATUEO). — Nees, Syst. 123. 1836; Endlieher, Gen. 317. 1836- 40; Msissner in DC. Prodr. 15 (1): 43. 1834; Baillon, Hist. PL 2: 469. 1870; Bsntham in B. & H., Gen. PL 3: 153, 157. 1880; Mez in Jahrb. bot. Gart. Berlin 5: 134—179. 1889; Pax in Engl. & Prantl, Pfl. Fam. 3 (2): 114. 18S9; Blake in J. Wash. Acad. Sci. 10: 9—21. 1920; Kostsrmans in J. sci. Ees. Indon. 1: 86 & 116. 1952. Trees or shrubs. Leaves alternate, chartaceous to rigidly coriaceous. Panicles axillary or sub-terminal- Flowers bisexual; tepals 6, the outer whorl as a rule smaHer than the inner one, deciduous or persistent (but not indurate and not clasping the fruit) ; tube very shallow. Fertile stamens 9 or 6 (two or three outer whorls) with, as a rule, long and slender filaments and introrse cells (in subgenera Nothaphoebe and Clavi- persea, stamens sessile); anthers of whorl three extrorse or lateral and 1 Antique Greek name of an Egyptian cauliflorous .tree, which has nothing in common with the modern Persea. 226 REINWARDTIA [VOL. 4 filaments flanked by glands. Fourth whorl consisting of conspicuous usually stipitate staminodes (small in subg-enera Nothaphoebe and Clavi- persea). Anthers as a rule 4-celled, rarely those of the third whorl 2- celled; cells usually large. Fruit on a naked pedicel, or the perianth (not enlarged) more or less persistent. Pedicel cylindrical or enlarged and fleshy. Pantropic. TYPE SPECIES. — Persea americana Miller. Number of binomials 239. It is not possible to estimate the exact number of "good" species. If we accept Mez' circumscription, most species are found in tropical America, Mez differentiated the genus from its nearest relative Phoebe by the position of the anther cells (in 2 pairs above each other in Phoebe and showing tendency to be placed in an arc in Persea) and the not enlarged perianth under the fruit. The way of separating the two genera is not satisfactory. The position of the anther cells has certainly no generic value. In Asiatic species of Phoebe the persistent, indurate and enlarged perianth is appressed to the fruit; the flower tube does not enlarge, and the pedicel of the fruit is not thickened. The genus Machilus has already been included by me in Persea, as the persistent, not enlarged, spreading or recurved perianth is also found in American Persea and the leaf- reticulation strongly resembles Persea. If the character of the indurate, appressed perianth can be main- tained in American species of Phoebe, it seems to be advisable to restrict only these species to Phoebe and to incorporate all other in Persea, and perhaps partly in Cinnamomum. Equally unsatisfactory is the delimitation versus Nothaphoebe and Alseodaphne. Nothaphoebe, as typified by N. umbelliflora BL, is char- acterized by sessile, thick anthers, minute staminodes and a more or less permanently semi-closed perianth with smaller outer tepals. The fruit is seated on a naked, not or hardly thickened pedicel. Alseodaphne, as typi- fied by A. semecarpifolia Nees, is more like Persea in its flower characters than Nothaphoebe by its more spreading perianth. Revision of all species is necessary to make certain whether Notha- phoebe and Alseodaphne may be kept separate from Persea or whether Bentham's view should be adopted to incorporate them, together with Phoebe, into Persea (as I have done here). Pax' system is unacceptable; it is partly based on wrong assumptions of floral and fruit characters of the genera concerned. Anatomical characters of leaves and wood yield no definite conclusion although at any rate they do not support separation of the genera ment- ioned above. 1957] A. J. G. H. KOSTEEMANS : Lauraceae 227 I agree with Airy-Shaw, that the character of 2- or 4-celled anthers is less important than stressed formerly in Pax' and Mez5 classification. In this paper I have incorporated Alseodaphne and Nothaphoebe in Persea, with the status of subgenera; moreover a new subgenus Clavi- persea is added, comprising species characterized by flowers as in Notha- phoebe, but with thickened, fleshy fruit pedicel with persistent tepals, Persea americana Miller yields the well-known avocado; P. lingue Nees from Chile is a well-known timber, like P. thunbergii (Sieb. & Zucc.) Kosterm., (basonym: Machilus thunbergii Sieb. & Zucc. in Munch. Abh, II Cl. Akad. Wiss. IV, 3. Abth. 202. 1943) from Japan. S u b g e n e r a five. — 1. Persea, syn. Eupersea Benth.: tepals deci- duous. 2. Machilus (Nees) Kosterm.: tepals persistent, patent. 3. Notha- pkoebe (Bl.) Benth.: tepals incurved, stamens sessile, staminodes small. 4. A7seodaphne (Nees) Benth.: tepals patent, deciduous, stamens with conspicuous filaments, staminodes large, fruit pedicel thickened. 5. Clavi- persea Kosterm.,: flowers as in Nothaphoebe, fruit peduncle swollen, fleshy, crowned by the persistent perianth. 2, P H O E B E 2 Nees, 1836. Phoebe Nees, Syst. 93. 1836. SYNON. — Persea Boehm. 1760. p.p. PKIHCIPAL LITERATURE. — Nees, Syst. 98. 1836; Baillon, Hist. PL 2: 468. 1870; Bentham in B. & H., Gen. PL 3: 157. 1880; Mez in Jahrb. bot. Gart. Berlin 5: 180. 1889; Pax in Engl. & Prantl., Pfl. Fam. 3 (2): 115. 1889 p.p.; Kostermans in J. scl. Res. Indon. 1: 122. 1952.' Floral characters as in Persea. Perianth persistent, indurate, enlarg- ed, clasping the basal part of the fruit. TYPE SPECIES. — Ph. lanceolata Nees. Number of binomials 174. The actual number of species depends on the delimitation of the genus. As characterized above, the genus is restrict- ed to Asia. The American species with cylindrical fruit-pedicel with or without persistent, not enlarged perianth, should be incorporated into I'ersea; the American species with thickened fruit-pedicel and disc-like cup, are perhaps better incorporated in Cinnamomum. Pax recognized two s e c t i o n s , of which Eupersea comprises the species of Phoebe proper. The section Gnesiopersea of Bentham, incor- porated by Pax with Phoebe, definitely belongs to Persea. 2 From the Greek cpoifios, phoibos = shining; the name of Apollon as sun god. 22S REINWARDTIA " [VOL. 4 b . Subtribus B e i l s c h m i e d i i n e a e Kosterm., 1957. Anthers with two cells. 3. A P o L L o N I A S3 Nees, 1833. Apollonias Nees, Frogr. grat. 10. 1833. PRINCIPAL LITERATURE1. — Nees, Progr. grat., Annexa pi. laur. expos. 10. 1833; Baillon, Hist. PL 2: 470. 1870; Bentham in B. & H., Gen. PI. 3: 152. 1880; Kostermans in Humbert, Fl. Madag., 81e Fam.: 2—10. 1950; in J. sci. Res. Indon. 1: 118. 1952. Flowers and fruit characters as in Phoebe, but anthers 2-ceiled- TYPE SPECIES. — A. barbusana (Cav.) A. Braun. Number of binomials 12; number of species 2. The Madagascar spe- cies, of which the fruit has become known better, were relegated to Beil- schniiedia. The genus has a disjunct area; one species in the Canary Islands, one in India. 4. D E H A A S U 4 Bluma, 1835. Dehaasia Blume. Rumphia 1: 161. 1835. SYNOS. •— Haasia Nees 1833. — Cyanodaphne Blume 1851. PRINCIPAL LITERATURE. — Bentham in B. & H., Gen. PL 3: 152. 1880; Kostermans in J. sci. Ees. Indon. 1: 120. 1952. Floral characters as in Persea subgen. Nothaphoebe, but anthers 2- celled. The fruit-pedicel is strongly enlarged, fleshy, and often highly coloured. TYPE SPECIES. •— D. incrassata (Jack) Kosterm. Number of binomials 33; the actual number of species will prove to be far smaller. The genu3 is closely related to Persea subgen. Nothaphoebe, but differs in the number of anther-cells and the swollen fruit-pedicel, which also occurs in Persea subgen. Clavipersea. Airy-Shaw advocates fusion of Alseodaphne and Dehaasia, which are very similar generally; because of the different pedicel and number of anther-cells, however, I think that Dehaasia merits generic status. The genus is restricted to Malaysia- 3 After the Greek god Apollon. 4 Named in honour of Dirk ds Haas, Head of the Dutch East India Company's establishment in Japan in 1677—1679, later (1687) governor of Amboina. 1957] A. J. G. H. KOSTERMANS : Lauraceae 5. B E I L S C H M I E D I A 5 Nees, 1831. 229 Beilschmiedia Nees in Wall. PL Asiat. rar. 2: 61, C9. 1831. SYNON. — Boldu (non Feuillee) Nees 1833. — Hufelandia Nees 1833. — Bail- schmidtia Reichenbach 1841. — Belloia Gay 1819. — Nesodaphne Hooker f. 1855. — Wimmer'a Nees ex Maissner 1834. — Bsrniera Baillon 1870. — Boldus O.K. (non Adans.) 1891. — Tylostemon Englsr 1898. — Afrodaphne Stapf 1905. — Thouvenotia Danguy 1920. — Ananeria Kostsrmans 1938. — Purkayasthaea Purkayastha 1938. — . jEsauromerrillia Allen 1942. PRINCIPAL LITERATURE. — Bentham in B. & H., Gen. PL 3: 152. 1880; Kostermans An Rec. Trav. bot. Neerl. 35: 837—868. 1938; Robijns & Wilczek in Bull. Jard. bot. jBruxelles 19: 459—-506 1951. Flowers bisexual. Tepals equal or subequal. Fertile stamens 9 or 6, 2-celled; two outer whorls introrse, glandless; third whorl extrorse, flank- led by glands; fourth whorl consisting of conspicuous, usually stipitate staminodes. Stigma inconspicuous, consisting of differently coloured tissue 1 decurrent at one side from the apex of the style. Fruit on a not (or hardly) swollen, bare pedicel. Leaves often suboppsite; reticulation lax, usually conspicuous. TYPE SPECIES. — B. roxburgkiana Nees. Number of binomials 238, the actual number of species will not be fles3 than 200. This pantropical genu.3 is well represented in Africa. The subdivision of the genus, as proposed by Robijns and Wilczek for the African species, is not applicable to Beilschmiedia in other con- 'tinents. S u b g e n e r a two. — 1. Beilschmiedia, syn. Ennearrhena, Stapf, ,1905; fertile stamens 9. 2. Hexaa^rhena Stapf, 1905: fertile stamens 6. 6. E N D I A N D R A 6 E. Brown, 1810. Endiandra R. Br., Prodr. Nov. Holl. 402. 1810. SYNON. — Dictyodaphne Blume 1850. — Brassiodcndron Allan 1942. PRINCIPAL, LITERATURE. — Baillon, Hist. PL 2: 474. 1870; Bentham in B. & H., Gen. PL 2 : 154. 1880. Floral characters as in Beilschmiedia, but (third whorl) only three often sessile stamens (one exception: E. fragrans Kosterm. with 6 fertile stamens). Staminodes minute or none. Leaves alternate or opposite with areolate reticulation. TYPE SPECIES. — E. glauca R. Brown. 5 Named in honour of K. T. Beilschmied, pharmacist and author of soms plant- gaographical works, born 1793 in Silesia, died 1843 in Herrnstadt near Breslau (Vra- tislava). 6 From the Greek evdeia, endeia = lack, and avijg, avdgog, aner, andros = man. A plant lacking stamens. 230 R E I N W A R D T I A [VOL. 4 Number of binomials 95; number of species perhaps 80. The genus is restricted to Malaysia, the Pacific region and Australia. It is closely related to Beilschmiedia, with which it is linked by E. fragrans Kosterm. with 6 fertile stamens, whereas B. endiandraefolia Kosterm. (ined.) has the areolate reticulation of Endiandra. It is also very similar to South American Mezilaurm, which has a small disc-shaped remnant of the flowertube at the top of the fruit pedicel. This occurs also in some Australian Endiandra species, but the anthers in Mezillaurus are different and the leaves are fascicled, near the apex of the branches. 7. M E Z I L A U S U S 7 Taubert, 1892, Mezilaurus Taub. in Bot. Zentralbl. 50: 21. 1892. SYNON. — Silvia Allemao 1854. — Silvaea Meissner 1834. — Endiandra Bentham 1880, p.p. — Mezia (nee Sehwacke) 0. Kuntze 1891. — Neosilvia Pax 1897. PRINCIPAL LITERATURE. — Kostermans in Rec. Trav. bot. neerl. 35: 109—12S. 1938. Floral characters as in Endiandra, but anthers often dehiscing later- ally. Leaves fascicled. TYPE SPECIES. -— M. na-valium (All.) Taub. Number of binomials 10; number of species 9. The genus is restricted to South America, It is closely related to Endiandra. Mezilaurus navalium Mez and M. itauha Mez furnish well-known commercial timbers, 8. H E x A P o R A8 Hooker f., 1886. Hexapora Hook, f., Fl. Brit. Ind. 5: 189. 1886. SYNON. — Micropora Hooker f. 1883. PRINCIPAL LITERATURE. — Hooker f. in Hook. Icon. t. 15k-7. 1886; Fl. Brit. Ind. 5: 862. 1886; Ridley, Fl. Mai. Pen. 3: 90. 1924; Kostermans in J. sci. Kes. Indon. 1: 144. 1952. Flowers bisexual; perianth of 6 equal, rotundate tepals. Fertile stamens 6, without glands, 2-celled, extrorse. Staminodes 3, thick. Stigma as in Beilschmiedia. Leaves alternate. Fruit unknown. TYPE SPECIES. — H- curtisii Hook. f. This monotypic genus is allied to Beilschmiedia and Endiandra. It is endemic in the Malay Peninsula. 7 Named in honour of C. Mez, born in 1836 at Freiburg; in 1900, professor in Halle, in 1910 in Konigsberg, a well-known botanist, author of monographs on Laura- ceae, Myrsinaceae, and Bromeliaceae. 8 Named for the 6 pores of the stamens. Actually there are 6 x 2 pores, and Hooker, when discovering his mistake, changed the name into Micropora. 1957] A. J. G. H. KOSTESMANS : Lauraceae 9. P O T A M E I A9 Th0U., 1808. Potameia Dupetit-Thouars, Nov. Gen. madag. 5: 16. 1803. SYNON. — Cansiera (non Juss.) Sprengal 1825. — Potwmica Poiret 1826. — Syndiclis Hooker f. 1836. PRINCIPAL LITERATURE. — Kostermans in Humbert, FL Madag., 81e Fam.: 10—18. 1950; in J. sci. Res. Indon. 1: 144. 1952; Communic. 55 Forest Research Instit. Bogor 3—35. 1957. Flowers bisexual; tepals 4, equal, in two opposite whorls. Fertile stamens 4, in two whorls, 2-celled (rarely one-celled). Staminodas 2, small, flanked with glands, or none; stigma inconspicuous. Fruit-pedicel with disc-like small remnant of tube with subpersistent perianth, or naked. TYPE SPECIES. — P. thouarsii R. & Sch. Number of binomials 22; number of species 21. The genus has a dis- junct area; 19 species occurring in Madagascar, one in Bhutan, one in Hainan. The latter two, formerly described under Syndiclis, show fusion of the 2 anther cells, although the valves are still bilobed, this is also the case in one Madagascar species (P. argentea Kosterm.). II. Tribus C i n n a m o m e a e Baill., 1870, emend. Kosterm., 1957. Cinnamomeae Baillon, Hist. PI. 2: 468. 1870, p.p. SYNON. — Oreodaphnsae Meissn. 1864. Inflorescences without involucrum, paniculate. Fruit-base embedded in a cupula. a. Subtribus C i n n a m o m i n e a e Kosterm., 1957. Anthers with 4 cells. 10. O C O T E A 1 0 AubL, 1775. Ocotea Aublet, Hist. PL Guyane fr. 2: 780. 1775. SYNON. — Borbonia [Plumier] Boehmer 1760. — Nectrandra Eolander ex Eott- bosll 1778. — Porostema Sehreber 1789. — Senneberia Necker 1790. — Perostema Raeuschel 1797. — Linharea Arr. de Camara ex Kostsr 1810. — Gymnobalanus Nees & Martius 1833. — Lsptodaphne Nees & Mart. 1833. — Mespilodaphne Nees & Mart. 1833. — Oreodaphne Nees & Mart. 1833. — Petalanthera Nees & Mart. 1833. — : Strychnodaphne Nees & Mart. 1833. — Teleiandra Nees & Mart. 1833. — Pleurothy- rium Nees ex Lindley 1836. — Pomatium Nees & Mart, ex Lindley 1836. — Calyco- daphne Bojer 1837. — Balanopsis Rafin. 1838, p.p. — Damburneya Rafin. 1838. — Agathophyllum (non Willd., nee Jussieu) Blume 1851, p.p. — Dendrodaphne Beurling 1854. — Adenotrachelium Nees ex Meissner 1834. — Agriodaphne Nees ex Meissner 1834. — Aperiphracta Nees ex Meissner 1884. — Camphoromoea Nees ex Meissner 1834. — Cannella Schott ex Meissner 1864. — Ceramocarpium Nees ex Meissner 1834. 9 From the Greek nora/AO?, potamos = river, and fieiov, meiou = smaller. 10 Local name Ocote in French Guiana. 232 REIN WARDTIA [VOL. 4 1957] A. J. G. H. KOSTEEMANS : Lauraceae 2S3 — Ceramophora Nees ex Meissnsr 1864. Nemodaphne Meissner 1834. — Sassafri- dium Meissner 1864. — Synandrodaphne Meissnsr 1834. — Adenotrachelima Baillon 1870. — Nyctandra Prior 1883—86. Sennebiera O. Kuntze 1904. PRINCIPAL LITEUATUEE. — Nees, Systema 277—340; 346—331; 380—470; 471—484. 1833; Meissner in DC, Prodr. 15 (1): 93—170. 1834; Baillon, Hist. Fl. 2: 473, 477. 1870; Bentham in B. & H., Gen. PI. 3: 157—158. 1880; Mez in Jahrb. bot. Gart. Berlin 5: 219—174. 1889; Kostermans in Bull. Jard. bot. Bruxelks 15: 73—88. 1938; in Humbert, Fl. Madagasc, 81e Fam.: 18—43. 1950; in J. sci. Res. Indon. 1: 1952; Communic. Forest Kessarch Inst., Bogor, 60: 1—44. June 1957; Allan in Ann. Missouri bot. Gard. 35: 16—62. 1948. Flowers in panicles, without involucre, bisexual or dioecious; flower tube small or conspicuous- Tepals equal, deciduous or persistent. Fertile stamens 9 in 3 whorls, the fourth whorl staminodial; staminodes minute or altogether lacking. Third whorl (only in one case all stamens) with glands. Anthers 4-celled, cells in pairs above each other or arranged in an ar"c. Cells of outer 2 whorls as a rule introrse, those of the third whorl generally extrorse. Stigma peltate. Fruit on a more or less developed cup with simple or double rim; perianth sometimes persistent. TYPE SPECIES. •— O. guianensis Aublet, Number of binomials (Ocotea) 449, (Nectandra) 248. The actual number of species will prove to be far smaller. This large genus is restrict- ed to tropical America. As Mez already indicated Ocotea and Neeiandra (and even Phoebe) are very difficult to separate. The position of the anther cells, the main and perhaps the sole difference between the two genera, is certainly not of generic value, as intermediate cases are not uncommon. The other characteristics (thin contra fleshy tepals, reflexed contra expanded tepals) are of even lsss generic value. Ocotea veraguensis Mez has the same number of glands as Pleuro- thyrium and as the number of glands is not stable in other genera, I have already incorporated Pleurothyrlum with Ocotea. Against Phoebe, Cinnamomum, and Persea, the genus Ocotea is mere- ly differentiated by the minute staminodes of whorl four. Ocotea and Nectandra are better combined and considered two sub- genera. Ocotea rodiaei Mez is the well-known greenheart of Guiana; O. bullata E. Mey. is a good timber from South Africa- S u b g e n e r a three. — 1. Ocotea: anther-cells in pairs above each other. 2. Nectandra (Rol.) Kosterm.: anther-cells in one row. 3. Pleurothy- rium (Nees) Kosterm.: all anthers with glands. 11. C I N N A M O M U M11 Boehmer, 1760. Cinnamomum Boshmer in Ludwig, Defin. 63. 1760. E^NON. — Camphora [Bauh.] Boshmer 1760. — Septina Noronha 1790 (?). —'• Camphorina Noronha 1790 (?). — Cecidodaphne Nees 1833. — Parthenoxylon Blume 1831. — Cynamonum Deniker 1883. — Neocinnamomum Liou-Ho 1932. PKINCINAL LITERATURE. — Nees, De Cinnamomo Disput. 1823; Syst. 1836; Meiss- ner in DC, Prodr. 15 (1); 10. 1834; Baillon, Hist. PL 2: 468. 1870; Bentham in B. & H., Gen. PL 3: 155. 1880; Perrot & Ebsrhardt, Les Canelliers de l'Indochine 1903; Cammtrloher in Bull. Jard. bot. Buitsnzorg 3, 7: 446—498. 1925; Lukmanoff, Nomencl. et Iconogr. Canell. et Camphr. 1—2: 1—28, pi. 1—16 (n.v.). Flowers bisexual, rarely polygamous; fertile stamens 9 (or 6) in 3 whorls; outer two whorls introrse, ^landless; inner whorl extrorse, flanked by glands. Anthers 4-c3lled, very rarely 2-celled. Fourth whorl consisting of conspicuous, stipitate, glandlezs staminodes. Stigma discoid or peltate. Flower tube accrescent, growing out into a cup which surrounds the basal part of the friut; often the basal part of (or the entire) perianth persistent on the rim of the cup. Leaves usually opposite and triplinerved. TYP3 SPECIES. — C. zeylaiucum Bl. Number of binomials 341; a revision will reduce the actual number of species considerably. The genus occurs from the Asiatic mainland to Formosa, the Pacific islands and Australia. The flower characters are similar to those of Persea and Phoebe, but the tube is usually deeper and grows out into a cup; sometimes this cup is very fleshy and shallow, merging into the pedicel. In the latter case the genus is very close to Aiouea, which also has triplinerved species and persistent tepals; in Aiouea the anthers are always 2-celled. In the few cases where in Cinnamomum the anthers are two-celled, the upper ablastic tissue is usually distinct. As in the genus Phoebe in the sense of Mez), there are species in Cinnamomum whose anther cells are arranged in an arc, these were considered to belong to a different genus (Neoc'nnamo- mum) by Liou-Ho. The arrangement of the anther cells, however, has no generic value. The alliance and delimitation between Cinnamomum and Aiouea remains to be cleared. Ocotea is apparently also related to Cinnamomum, but has smaller staminodes. Some American species of Phoebe should perhaps be included in Cinnamomum. 11 Latin transcription of the Greek: xivva/xcofiov, kinnamomon = cinnamon. 234 REINWARDTIA [tOL. 4 12. A C T I N O D A P H N E 1 2 Nees, 1831. Actinodaphne Nees in Wall., Fl. asiat. rar. 2: 61, 68. 1831. SYNON. — lozoste Nees 1831, p.p. — Jozoste Nees ex 0. Kuntze 1891. PRINCIPAL LITERATURE. — Bentham in B. & H., Gen. PL 3: 160. 1880; Hooker £., Fl. Brit. Ind. 5: 147—154. 1886; Koorders & Valet, in Meded. Lands PI. tuin 68: 110—123. 1904; Teschner in Engl. bot. Jahrb. 58: 385. 1923; Ridley, Fl. Mai. Pen: 3: 107—112. 1924; Liou Ho, Laur. Chine et Indoch. 155—162. 1932; Allen in Ann. Missouri Bot. Gard. 25: 440—414. 1938; Kostermans in J. sci. Res. Indon. 1: 115. 1952. Leaves usually verticillate. Flowers dioecious; tepals 6, subequal. Fertile stamens in male flower as a rule 9; the inner whorl biglandular. Anthers 4-celled, all introrse. Staminodes in female flower usually 9; the inner whorl biglandular. Stigma peltate. Fruit on a shallow or deep cup. TYPE SPECIES. — A. pruinosa Nees. Number of binomials 128; the number of species will be about 60—70. The genus is usually placed near Litsea, because of the introrse anther cells. It differs, however, by the lack of an involucre of decussate, per- sistent bracts, as found in Litsea and I therefore prefer to leave it near Ocotea and Cinnamomum. It is very close to Sassafras, with which it might be combined ultimately. The genus is restricted to the Asiatic main- land and Malaysia. 13. S A s S A F R A S13 [Kramer] Boehmer, 1780. Sassafras [Kramer] Boehmer in Ludwig, Defin. 36. 17S0. SYNON. — Euosmus Nuttall 1818, p.p. — Evosmus Rafin. 1838, p.p. •— Pseudo- sassafras Lecomte 1911. -— Yushunia Kamikoti 1933. PRINCIPAL LITERATURE. — Nees, Syst. 487. 1838; Nees & Ebsrm., in Handb. Med.- Pharm. Bot. 1: 448. 1830; Baillon, Hist. PI. 2: 439, 479. 1870; Bentham in B. & H., Gen. PI. 3: 160. 1880; Bantley & Trimen, Madic. PI. 3: 220. 1880; Mez in Jahrb. bot. Gart. Berlin 5: 484. 1889; Berg & Schm., Atl. off. PfL, ed. 2, 4: 79. 1899; Blake in Rhodora 20: 98. 1918; Render in J. Am. Arb. 1: 242—44. 1920; Record & Hess in Trop. Woods 69: 31—32. 1942; Kosterm. in J. sci. Res. Indon. 1: 88, 95, 147. 1952. Flowers dioecious. Tepals subequal. Stamens in male flowers and staminodes in female ones 9, all introrse, 4-celled; the inner whorl with glands. Flower tube enlarging into a fruit-cup. Flowers in shortened race- mes (pseudo-umbals), surrounded by deciduous, alternate bracts- Leaves alternate. TYPE SPECIES. — S. albidum (Nees) Nutt. 12 From the Greek: Aung, amivoQ, aktis, aktinos — ray, and dacpvrj, daphne = laurel. The leaves are arranged in star-shaped whorls. 'i* French, Italian and Spanish name, perhaps derived from latin: saxifraga. 1957] A. J. G. H. KOSTERMANS : Lauraceae 235 Number of binomials 4; number of species 2. The genus has a disjunct area, one species occurs in North* America, one in Formosa. The Formosan one is not constant in its number of anther cells and differs moreover by the presence of staminodes and bisexual flowers. I provisionally accept Render's view to consider Pseudosassafras as congeneric. The genus has always been put near Litsea, because all anthers are introrse. As I am not convinced that the in- or extrorsity of the anthers is more important than the involucre of decussate, persistent bracts, as found in Litsea, I prefer to place the genus near Actinodaphne. The genus is the only one that has lobed leaves. 14. U M B E L L U L A E I A 1 1 Nuttall, 1842. Dmbellularia Nuttall, N. Amer. Sylva 1: 103. 1842. SYNONYMS. — Drimophyllum Nuttall 1842. — Sciadiodaphne Reichenbach 1841. PRINCIPAL LITERATURE. — Bentham in B. & H., Gen. PI. 3 : 162. 1880; Mez in Jahrb. bot. Gart. Berlin 5: 482. 1889; Busss in Bsr. deut. pharm. Ges. 6 (2): 56—61. 1896; Jepson, The Silva of Calif. 2: 243. 1910; Bambacioni in Annali Bot. 22 (2): 99. 1941; Guenther, Ess. Oils 4: 207. 1950. Flowers bisexual; tepals 6, equal. Fertile stamens 9; the inner whorl with glands. Anthers 4-celled, first and second whorl introrse; third whorl extrorse. Fourth whorl staminodial. Flowers in a shortened raceme, sur- rounded by alternate, deciduous bracts. Fruit on a flat cup. Leaves alter- nate. TYPE SPECIES. — U. californica (Nees) Nuttall. Number of binomials 2; number of species 1. This monotypic genus occurs in California. It is usually considered to be related to Litsea, but in my opinion it is close to Actinodaphne. It differs more from Litsea than does Sassafras (in the position of the anther cells and the not decussate, non-persistent involucral bracts). It is possibly advantageous to incorporate it with Actinodaphne. 15. D I C Y P E L L I U M 1 5 Nees & Mart, 1833. Dicypellium Ness & Martins, Progr. grat. 14. 1833. PRINCIPAL LITERATURE. — Nees, Syst. 343. 1836 (excl. Licaria); Mez in Jahrb. bot. Gart. Berlin 5: 472—473. 1889; Kostermans in J. sci. Res. Indon. 1: 119. 1952. 14 Flowers in stalked umbels. is From Greek xvnelkov kupellon = cup; the cup is double-rimmed. 236 REINWARDTIA [VOL. 4 1957] A. J. G. H. KOSTEKMANS : Lauraceae 237 Flower and fruit characters as in Ocotea; tepals 9. Second and third whorl of -anthers foliaceous, ovate, 4-celled. Staminodes of the fourth whorl none. Tepals persistent, enlarged under the fruit. TYPE SPECIES. — D. caryophyllatum Nees. This monotypic genus from Brazil has a marketable fruit with clove fragrance. It is related to Ocotea, but also to Phyllostemonodaphne. b. Subtribu.3 A n i b i n e a e Kosterm., 1957. Anthers with two cells. 16. A.I o u E A16 Aublet, 1775. Aiouea Aublet, Hist. Guyane franc. 1: 310. 1775. SYNON. — Ehrhard'.a Seopoli 1777. — Douglassia Schreber 1783. — Colomandra Necker 1730. — Apivea Steudsl 1821. — Endocarpa, Eafinesqua 1838. PRINCIPAL LITERATURE. — Kostsrm. in REC. Trav. bot. neerl. 35: 57—104. 1938; in J. sci. Res. Indon. 1: 88. 1952; in Bol. tacn., Inst. Agron. Norts Brasil 28: 51—52. 1955. Flowers bisexual flower-tube not very deep. Tepals equal. Fertile stamens 9, 8 or 3; anthers 2-celied; those of the two outer whorls (with 3 exceptions) introrse. Sterile stamens represented by staminodes, which are often strap-like. Staminodes of the innermost whorl well developed. Stigma peltate. Fruit usually seated on a swollen obconical body with a slight concavity at top; sometimes perianth persistent and accrescent. TYPE SPECIES. — A. guianensis Aublet. Number of binomials 46, number of species 30. The genus is restricted to tropical America. The species are often difficult to distinguish from those of genera like Aniba and Endlicheria.' They frequently have a char- acteristic yellowish leaf colour and a thickened leaf margin. S u b g e n e r a three. 1. Aiouea: stamens of the outer two whorls fertile; of the inner two whorls sterile. 2. Endocarpa (Rafin.) Kosterm.: stamens of outer 3 whorls fertile; of fourth whorl sterile. 3. Trianthem Mez: stamens of first outer whorl fertile; of the other whorls sterile. 17. A N I B A 1 7 Aublet, 1775. Aniba Aublet, Hist. Guyane franc. 1: 327. 1775. SYNON. — Cedrola Schrebsr 1789. — Aydendron Nees & Mart. 1833. PRINCIPAL LITERATURE. — Kostsrm. in Rec. Trav. bot. neerl. 35: 866—1938; in J. sci. Res. Indon. 1: 83. 1952; in Bol. teen. Inst. Agron. Norta Brasil 28: 52—57. 1955. Flowers bisexual with conspicuous tube which increases in size after anthesis, becoming urceolate and temporarily enclosing the developing fruit; in the mature fruit appearing as a rather woody, mottled cup, sur- rounding the basal half of the fruit. Fertile anthers 9 (in one case the third whorl sterile), 2-cslled; outer two whorls introrse; third whorl ex- trorse and flanked by glands. Staminodes minute or none. Stigma as a rule inconspicuous, obtuse or truncate. The stamens in almost all species have the same characteristic shape. TYPE SPECIES. — A. guianensis Aublet. Number of binomials 74; number of species about 40. The genus is close to Endlicheria, from which it differs by its bisexual flowers, the different fruit cup and the differently shaped anthers. Aniba rosaeodora Ducke and A. duckei Kosterm. yield the well-known rose oil of commerce. The genus is restricted to tropical South and Central America. S u b g e n e r a two. 1. Aniba: stamens of the third whorl fertile, cells normal, extrorse. 2. Aioueopsis Mez: stamens of the third whorl sterile or with minute lateral cells. 18. E N D L I C H E R I A 1 8 Nees, 1833 (nom. eons.). Endlicheria Nees in Linnaea 8: 37. 1833. SYNOX. — Goeppsrtia Ness 1836. — Schaucra Nees 1838. — Sehaueria Nees ex Msissnsr 1834. — Ampelodaphne Meissnsr 1834. — Huberodaphne Ducke 1925. PRINCIPAL LITERATURE!. — Kostermans in Rec. Trav. bot. neerl. 34: 500—557. 1937; in J. sci. Res. Indon. 1: 17. 1952; in Bol. tzen. Inst. Agron. Norfce Brasil 28: 62—65. 1955; Allen in J. Arn. Arb. 26: 421. 1945. Flowers dioecious; tuba distinct. Male flowers with 9 fertile stamens; anthers as a rule 2-celled; the two outer whorls introrse; the inner whorl extrorse and usually flanked by glands. Staminodes none or minute. Ovary sterile, stipitiform. Female flowers as a rule slightly smaller and with broader tube; stamens smaller, sterila, but with same shape; stigma discoid or peltate, conspicuous. Fruit cup usually rather shallow, fleshy, merging into the usually fleshy, thick pedicel. TYPE SPECIES. — E. sericea Nees. Number of binomials 55; number of species about 40. The genus, which is restricted to tropical South and Central America, is related to Aniba by its floral characters and to Aiouea by its enlarged fruit pedicel. 16 Local name Ajoue in French Guiana. 17 Local name in French Guinea. 18 Named in honour of H. L. Endlicher, born 1804, Prssburg, died 1849, Vienna, Professor in Vienna and a well-known botanist. 238 REINWARDTIA [VOL. 4 1057] A. J. G. H. KOSTEKMANS : Laura-ceae 239 19. L I c A R I A19 Aublet, 1775. Licaria Aublet, Hist. Guyane franc.. 1: 313. 1775. SYNONYMS. — Acrodiclidium Nees 1833. — Evonymodaphne Nees ex Lindley 1838. — Dipliathus Rafin 1838. — Triplomeia Rafin. 1838. — Misanteca Cham. & Schdl. 1831. — Symphysodaphne A. Richard 1850. — Nobeliodendron 0. C. Schmidt 1930. — Chanekia Lundell 1937. — Clinostemon Kuhlm. & Sampaio 1928. PRINCIPAL LITERATURE. — Kostermans in Rec. Trav. bot. neerl. 34: 575—604. 1937; 35: 123—125. 1938; in J. sci. Res. Indon. 1: 89. 1952; in Bol. tacn. Inst. Agron. Norte Brasil 28: 65—71. 1955. Flowers bisexual; flower tube usually distinct. Stamens of the two outer whorls changed into small staminodes or wanting; stamens of the third whorl fertile, free or partly connate, basal glands present or none; cells two, introrse, extrorse or sub-apical. Fourth whorl of staminodes minute, usually absent. Stigma inconspicuous. Fruit cup large, with a double, rarely triple margin. TYPE SPECIES. •— L. guianensis Aublet. Number of binomials 58; number of species about 45, The genus is restricted to tropical South and Central America. It is related to Ardba, and Macbride suggested merging the two genera. Several species yield merchantable fruit with nutmeg or clove aroma, S u b g e n e r a two. 1. Licaria, syn. Acrodiclidium (Nees) Kosterm. 1957: staminodes of the two outer whorls present. 2. Misanteca (Cham. & Schdl.) Kosterm.; staminodes of the two outer whorls lacking. 20. U R B A N O D E N D R O N 2 0 'Mez, 1889. Urbanodendron Mez in Jahrb. bot. Gart. Berlin 5: 80. 1889. SYNON. •— Aydendron Nees & Mart. 1833, p.p. PRINCIPAL LITERATURE. — Kostermans in Rec. Trav. bot. neerl. 35: 106—109. 1938; in J. sci. Ees. Indon. 1: 146. 1952. Flower characters as in Licaria, but stamens 9; anthers 2-celled, those of the two outer whorls introrse; of the inner extrorse. All fila- ments flanked by large glands. Stigma inconspicuous. Fruit cup large, double-rimmed. TYPE SPECIES. •— U. verrucosum (Nees) Mez. This monotypic genus from Brazil is related to Licaria, but differs by the number of fertile stamens, which puts it near Aniba. The double- rimmed cup is similar to that of Licaria. *9 Local name Licari in French Guinea. 20 Named in honour of Ignatz Urban, a well-known German botanist, Director of the Botanic Garden and Museum at Berlin-Dahlem. •• 2 1 . S Y S T E M O N O D A P H N E21 Mez, 1889. Systemonodaphne Mez in Jahrb. bot. Gart. Berlin 5: 78. 1889. PRINCIPAL LITERATURE. — Kostsrmans in Rec. Trav. bot. neerl. 35: 104—108. 1938; in 3. sci. Res. Indon. 1: 145, 1952; in Bolet. teen. Inst. Ag-ron. Norte Brasil 28: 73—75. 1955. Flowers bisexual; tube short. Fertile stamens 9; anthers 2-celled. Fourth whorl of staminodes lacking. Outer two whorls with introrse cells and distinct, free filaments; third whorl connate, with small glands. Stigma small, discoid. Fruit cup disc-shaped-, double-rimmed with persis- tent perianth. TYPE SPECIES. — S. mezii Kosterm. Number of binomials 3; number of species 2. A small genus from Brazil; related to Licaria, but different by the number of fertile stamens and the fruit cup. 22. P H Y L L O S T E MONO D A P H N E 2 2 Kosterm., 1936. Phyllostemonodaphne Kostsrmans in Rec. Trav. bot. n,eerl. 33: 754. 1938. PRINCIPAL LITERATURE. — Kostermans in Eec. Trav. bot. neerl. 33: 754. 1936; in J. sci. Res. Indon. 1: 149. 1952. Flower characters as in Licaria, but the outer whorl of stamens is completely tepaloid; the second and third whorl are fertile and provided with glands. Anthers of the second whorl introrse, of the third whorl extrorse, 2-celled. Staminodes of the fourth whorl small of none. Stigma inconspicuous. Fruit cup flat, double-rimmed. TYPE SPECIES. — Ph. geminiflora (Meissn.)' Kosterm, This monotypic genus from Brazil is closely related to Licaria, but differs by the number of fertile stamens and tepals and by the shape of the fruit cup. It is also close to Dicypellium, which, however, has 4-celled anthers. III. Tribus L i t s e e a e Mez, 1889. Tribus Litseeae Msz in Jahrb. bot. Gart. Berl. 5: 6. 1889. Flowers in pseudo-umbels, surrounded by persistent, decussate, large bracts, forming an involucre. As a rule all stamens introrse. Tepals often " lore or less reduced. Tubs persistent and enlarged as a more or less devel- oped fruit cup on a usually not enlarged pedicel. a. Subtribus L i t s e i n e a e Kosterm., 1957. Anthers with 4 cells. 21 F r o m Greek aw, sun = t o g e t h e r ; orrj/xcov, stemon = stamen, and dawvri, daphne = l a u r e l ; the stamens a r e grown together. 22 F r o m Greek yvllov, phullon = leaf; ort]/Ltcov, stemon = stamen, and dawvn, daphne = l a u r e l ; the outer stamens a r e leaf-like. 240 R E I N W A R D T I A [VOL. 4 23. L I T S E A 2 3 Lamk. 1791 (nom. cons.). Litsea Lamarck, Encycl. meth. bot. 3: 574. 1791. SYNOX. — Malapoenna Adanson 1763. — Tomex Thunbsrg 1783. — Hexanthus Loureiro 1790. — Quinquedula Noronha 1790. — Fiwa Gmelin 1791. — Tetranthera Jacquin 1797. — Berrija (non Roxb.) Klein ex Willdenow 1800. — Litsaea Jussieu 1805. — Pipalia Stokes 1812. — Darwinia Dennstsdt 1818. — Dodecadenia Nees 1831. — Cylicodaphns Nees 1831. — Iozoste Nees 1831, p.p. ^- Jozosmene Nees ex Lind- ley 1836. — Lepidadenia Walker Arnott ex Nees 1833. — Cubeba Rafinesqus 1838. — Dccapenta Rafinesqus 1838. — Evslyna Rafinesque 1838, p.p. — Heckcria Rafinesque 1838. — Evosmus Rafinesque 1838, p.p. — Fiva Stsudel 1840. — Sebifera Blanco 1845. — Darwiniana "Dsnnstadt" ex Lindlay 1846. — Glabraria (non L.) Blume 1851. — Adenodaphne S. Is Moore 1921. — Pssudolitsea Yang 1945. PRINCIPAL LITERATURE. — Baillon, Hist. PI. 2: 440, 480. 1870; Bentham in B. & H., Gen. PI. 3: 181. 1880; Eooker f., Fl. Brit. Ind. 5: 154—182. 1883; Mez in Jahrb. bot. Gart. Bsrlln 5: 474—482. 1889; Koorders & Val. in Medsd. Lands Pl.tuin Buitan- sorg 68: 123—192. 1904; Bartlett in Proc. Am:r. Ac. 44: 597—001. 1909; Lecomt3, Fl. gin. Indochine 5: 130—142. 1914; Ridlsy, Fl. Mai. Pen. 3: 112—131. 1924; Liou- Ho, Laur. Chine et Indochine 1—207. 1932; Allen in Ann. Missouri Bot. Gard. 25: 331—400. 1938; in J. Arnold Arb. 26: 106. 1945; Nakai, Fl. sylv. Koreana 22: 49—61. 1939; Kostsrmans in J. sci. Res. Indon. 1: 93. 1952. Flowers dioecious in pseudo-umbels, surrounded by an involucre of persistent or subpersistent, large, decussate bracts. Tepals 6 or 0. Fertile stamens In male flower 9 or 12, sometimes more than 12 (subgenus Dode- cadenia/. Outer 2 whorls usually glandless; third and inner whorls flank- ed by glands. Filaments usually slender; anthers 4-celled, all introrse, or the basal pair of the third staminal whorl lateral. Ovary in the male flower stipitiform or 0. In the female flower an equal number of stami- nodes as stamens in the male flower; stigma peltate, conspicuous. Fruit seated in a more or less developed cup or disc; ths perianth usually cadu- cous. TYPE SPECIES. — L. chinensis Lamk. Number of binomials 474; number of species approximately 400. This large and widely distributed genus, only lacking in Africa and Europe, extends north to Japan, Korea and North America; south to New Zealand and subtropical South America. The diversity in such a large genus is appreciable, but does not warrant splitting it up into smaller entities, as has often been advocated in local floras. The flowers and umbels are remarkably uniform. The fruit cup is either shallow and disc-like or semi- globose, or may even completely ennvelop the fruit (in that case it differs from the fruit of Cryptocarya by the large apical aperture) ; sometimes it is very swollen and fleshy, sometimes thinly walled. The number of 1957] A. J. G. H. KOSTERMANS: Lauraceae 241 22 From the Chinese litse. stamens is the largest in sub-genus Dodecadenia, a small group, which is here, more or less arbitrarily, included in Litsea; the sub-genus Dode- cadenia has bisexual flowers. S u b g e n e r a three. 1. Litsea: flowers monoecious. 2. Dodecadenia (Nees) Kosterm.: flowers bisexual. 3. Octolitsea Liou-Ho: flowers dioecious, with 8 tepals. 24. N E O L I T S B A - ' 4 Merr., 1906. Neolitsea Merrill in Philip. J. Sci., Bot., Suppl. 1 (1) : 56. 1906. SYNON. — Tetradenia (non Benth.) Nees 1831. — Balanopsis Rafinesque 1838, p.p. — Bryantea Rafinesque 1838. — Litsea sect. Neolitsea Bentham 1880. PRINCIPAL LITERATURE. — Lecomte, Fl. gen. Indoch. 5: 142:—144. 1914; Ridley, Fl. Mai. Pen. 3: 131—133. 1924; Liou-Ho, Laur. Chine & Indoch. 139—155. 1932; Allen in Ann. Missouri Bot. Gard. 25: 415—431. 1938; Kostermans in J. sci. Res. Indon. 1: 147. 1952. Flowers dioecious, involucrate, involucral bracts large, persistent, decussate. Flowers dimerous; fertile stamens in male flowers and stami- nodes in female flowers 6 in 3 whorls; the inner whorl with glands. Anthers 4-celled, introrse. Stigma in female flower conspicuous, peltate. Fruit on a disc-like cup; pedicel often slightly thickened. TYPE SPECIES. — N. cassia (L.) Kosterm. Number of binomials 93; the number of species will be about 80. The genus is restricted to the Asiatic mainland and Malaysia. It is related to Litsea. b . Subtribus L a u r i i n e a e Kosterm., 1957. Anthers with two cells. 25. L I N D E E A 2 5 (non Adans.) Thunb., 1783 (non. cons.). Lindera T h u n b e r g , N o v . Gen. PL 3: 64. 1783. SYNON. — Benzoin B o e h m e r 1760. — Bistania N o r o n h a 1790 ( ? ) . — Euosmus N u t t a l l 1818, p.p. — Calosmon B e r c h t . & P r e s l . 1823. — Sassafras B e r c h t . & P r e s l . 1823. — Daphnidium N e e s 1 8 3 1 . — Polyadenia N e e s 1831. — Evelyna R a f i n e s q u e 1838, p.p. — Ozanthes R a f i n e s q u e 1838. — Evosmus R a f i n e s q u e 1838, p.p. — Aperula B l u m e 1 8 5 1 . — Iteadaphne B l u m e 1 8 5 1 . —• Parabenzoin N a k a i 1924. P R I N C I P A L L I T E R A T U R E . — H o o k e r f., F l . B r i t . I n d . 5 : 182—189. 1 8 8 6 ; Mez in J a h r b . bot. G a r t . B e r l i n 5 : 486—489. 1889; K o o r d e r s & V a l e t , i n Meded. L a n d s P l . t u i n 24 p r o m G r e e k ve.og, neos = n e w , a n d L i t s e a . 25 N a m e d a f t e r J o h a n n L i n d e r (1678, K a r l s t a d , S w e d e n ; f 172,3, S t o c k h o l m ) who w a s e l e v a t e d t o n o b i l i t y u n d e r t h e n a m e o f L i n d e s t o l p e ; a u t h o r o f a b o t a n i c a l p a m p h l e t . 242 R E I N W A R D T I A [VOL. 4 Buitenzorg 58: 229—246. 1904; Lecomte, Fl. gen. Indochine 5: 152—158. 1914; Ridley, Fl. Mai. Pen. 3: 133—137. 1924; Liou-Ho, Laur. Chine & Indoch. 117—139. 1932; Kostermans in J. sci. Res. Indon. 1: 90. 1952. Flowers and fruit characters as in Litsea, but anthers 2-celled (very rarely partly 4-celled) ; usually the fruit cup very shallow; perianth some- times persistent. TYPE SPECIES. — L. umbellata Thunb. Number of binomials 135; the number of species will be about 100. Like Litsea, to which this genus is closely related, it is widely distributed, also outside the tropical area; like Litsea it is not represented in Africa. S u b g e n e r a two. 1. Lindera: umbels many-flowered. 2. Iteadaphne (Bl.) Kosterm.: involucre with one flower. 26. L A U R U S 2 6 L., 1753. Laurus Linnaeus, Spec. pi. 369. 1753. SYNON. — Apella Adanson 1763. — Adaphus Necker 1780 (?). PEINCIPAL LITERATURE, — Webb & Berth., Phytogr. Canar. 3 (3) : 229. 1836—40; Gandoger, Fl. Europae 20: 44. 1890; Thome, Fl. Deutschl. 1: 157. 1&03; Coste, Fl. France 3: 215. 1906; Bonnier, Fl. France, etc. 9: t. 536. 1927; Hegi, Fl. Europa ed. 2, 4 (1) : 11. 1935; Kostermans in Rev. Univ. Chilena 24: 204—206. 1939. Flowers dioecious or bisexual, involucrate. Flowers dimerous. Male flowers with 12 stamens in 3 whorls; those of the outer whorl glandless; anthers 2-celled, introrse; ovary abortive. Female flowers with 4 large staminodes, alternating with the tepals, all with glands. Stigma discoid. Fruit on a fleshy, thickened pedicel, disc-like at the apex. TYPE SPECIES. — L. nobilis L. Number of binomials 334; number of species 2. The genus is restricted to the Mediterranean region and the Canary islands. Because of its involucre of decussate bracts it belongs near Litsea. The reduced number of staminodes in the female flower is an uncommon feature in Lauraceae. IV. Tribus C r y p t o c a r y e a e Meissn. 1864, p.p. Tribus Cryptocaryeae Meissner in DC, Prodr. 15 (1) : 5. 1864, p.p. F r u i t completely included in the accrescent flower tube. a . Subtribus E u s i d e r o x y l i n e a e Kosterm., 1957. Anthers with 4 cells. 26 Old latin plant name. 1957] A. J. G. H. KOSTERMANS: Laui-aceae 243 27. E u s i D E R O X Y L O N - 7 Teijsm. & Binn., 1863. Eusideroxylon Teijsmann & Binnendijk in Natuurk. Tijdschr. Ned. Ind. 25: 292. 1863. SYNON. — Bihania Meissner 1864. PRINCIPAL LITERATURE. — Teijsm. & Binn. in Natuurk. Tijdschr. Nederl. Ind. 25: 292. 1863; de Wit in Bull. Bot. Gard. Buitenzorg 3, 18: 200—207. 1949; Kostermans in J. sci. Res Indon. 1: 141. 1952; in Penggemar Alam 35: 57. 1955. - Flowers bisexual. Tepals 6, equal; flowers tube shallow. Fertile stamens 3 (third whorl), 4-celled, extrorse; outer 2 whorls petaloid; all glandless; stigma discoid. Fruit completely enclosed in and adnate to the accrescent flower tube; seedcoat horny, furrowed. TYPE SPECIES. — E. zwageri T. & B. Number of species and binomials 2. E. zwageri T. & B. occurs in East Sumatra, Bangka, Biliton, and Borneo. E. melagangai Sym. is re- stricted to North, West, and Central Borneo. In its fruit characters the genus resembles Cryptocarya, but the num- ber and shape of the stamens is different. The timber (ironwood) is one of the heaviest and most durable in Malaysia and resembles in this quality Ocotea rodioei Mez from Guiana, b. Subtribus C r y p t o c a r y i n e a e Kosterm., 1957. Anthers with two cells. 28. C R Y P T O C A E Y A 2 S R. Brown, 1810. Cryptocarya R. Brown, Prodr. FL Nov. Holland. 402. 1810. SYNON. — Peumus Molina 1782, p.p. — Cryptocaria Gay 1849. — Caryodaphne Blume ex Nees 1836. — Salgada Blanco 1845. — Icosandra R. A. Philippi 1857. — Pseudocryptocarya Teschner 1923. PRINCIPAL LITERATURE. — Baillon, Hist. PL 2: 472. 1870; Bentham in B. & H., Gen. 3: 150. 1880; Hooker f., FL Brit. Ind. 5: 117—121. 1890; Bailey, Queensl. Fl. 1297. 1901; Lecomte, Fl. gen. Indoch. 5: 144—148. 1914; Liou-Ho, Laur. Chine et Indochine 95—102, 1932; Kostermans in Rec. Trav. bot. neerl. 34: 557—575. 1937; in Bull. Jard. bot. Bruxelles 15: 91—108. 1938; in Humbert, Fl. Madagascar, 81e famille: 74—84. 1950; in 3. sci. Res. Indon. 1: 94. 1952. Leaves alternate or opposite. Flowers bisexual; tube slender, con- spicuous; tepals 6, equal. Fertile stamens 9, 6, or 3, anthers 2-celled; anthers of two outer whorls introrse, of third whirl extrorse and filaments flanked by glands. Fourth whorl consisting of conspicuous, stipitate sta- minodes. Stigma small or inconspicuous, rarely peltate. Fruit entirely 2T From Greek sv, eu = good, ocdtjQog, sideros = iron, and £vlov, xulon = 28 From Greek y.Qvnzoi, kruptos = hidden, and y.ngvov, karyon = nut; the art is covered by the accrescent flower tube. 244 R E I N W A R D T I A [VOL. 4 1957] A. J. G. H. KOSTERMANS: Lauraceae 245 included in the enlarged flower tube, leaving only a minute orifice at apex. Endocarp and exocarp often bony and ribbed. TYPE SPECIES. — C. glaucescens R. Brown. Number of binomials 303; number of species perhaps between 200 and 250. This pantropic genus, (only lacking in Central Africa) has its centre in Malaysia. Species occur as far as Chile and Australia. C. mosch- ata Mez yields saleable fruit (nox moschado) with nutmeg smell and taste. S u b g e n e r a three. 1. Cryptocarya, syn. Enneanthera, Kosterm., 1957: fertile anthers 9. 2. Hexanthera Kosterm.: fertile anthers 6. 3. Tnandra Kosterm.: fertile anthers 3. 29. R A V E N SARA29 Sonn., 1782. Bavensara Sonnerat, Voy. Indes et Chine 3: 248. 1782. SYNON. — Euodia Gaertner 1791. — Agathophyllum (non Blume) Jussieu 1789. PRINCIPAL LITERATURE. — Kostermans in Humbert, Notul. system. Paris 8: 96—112. 1939; in Humbert, PI. Madag., 81e Fam.: 46—74. 1950. Floral characters as in Cryptocarya. Fruit included in the accrescent flower tube; the basal part of the fruit divided into 6 (rarely 12) com- partments by 6 (rarely 12) false dissepiments, growing out from the inner wall of the flower tube; the dissepiments ruminating the seed. TYPE SPECIES. — R. aromatica Sonn. Number of binomials 27; number of species 18. The genus is endemic in Madagascar. It differs from Cryptocarya only by its ruminate seed and basal fruit septa. V. Tribus H y p o d a p h n e a e Kosterm., 1957. Ovary inferior. 30. H Y P O D A P H N I S 3 0 Stapf, 1909. Hypodaphnis Stapf in Dyer, PI. trop. Africa 6 (1) : 185. 1909. PRINCIPAL LITERATURE. — Kostermans in Bull. Jard. bot. Bruxelles 15: 88—91. 1938; in J. sci. Res. Indon. 1: 1952. Leaves alternate. Flowers bisexual. Fertile stamens 9 in three whorls; anthers 4-celled, those of the two outer whorls introrse, of the third whorl extrorse and extrorse — lateral, third whorl with glands. The fourth whorl lacking. Ovary inferior; stigma small, discoid. TYPE SPECIES. — H. zenkeri (Engl.) Stapf. This monotypic genus is endemic in Cameroon, Gaboon, and Nigeria. 29 From the local Madagascar name: ravin-tsara. so P r o m the Greek vno, hypo = under, and dayns, daphms = laurel. The name alludes to the inferior ovary. B . Subfamily C a s s y t h o i d e a e K o s t e r m . , 1957. SYNON. — Cassytheae Nees 1831; Cassythaceae Dumortier ex Lindley (Nat. Syst., ed. 2: 202. 1836). Parasitical or partly autotrophic monotypical twiners with small haustoria. Stems filiform, containing chlorophyll. Leaves reduced to minute scales, arranged spirally (1/3). Inflorescence indefinite, spicate or racemose, or reduced to heads. Flowers sessile or pedicellate within a minute bract with 2 similar bracteoles close under the perianth, bisexual or semi-dioecious (?). Tepals 6 (the outer 3 smaller and resembling the bracts), persistent; tube shallow, enlarged, and enveloping the fruit. Fer- tile stamens 9, two-celled; the outer two whorls without glands and with introrse anthers (rarely the second whorl staminodial) ; the third whorl flanked by glands and with extrorse anthers; the fourth whorl of distinct, sessile or stipitate staminodes; stigma small, obtuse or capitellate. Fruit completely included in the enlarged and succulent flower tube, with small orifice at apex, usually surrounded by the persistent, erect perianth. Testa membranous or coriaceous; cotyledons thick, flehy, often unequal, distinct at an early stage, but later more or less consolidated. TYPE GENUS. — Cassytha L. 31. C A S S Y T H A 3 1 L., 1753. Cassytha [Osbeck] Linnaeus, Sp. PI. 1: 35. 1753. SYNON. — Cussuta Rumphius 1747. — Rombut [Rumph.] Adanson 1763. — Aeat- sjavalli [Rheede] Adanson 1763. — Cassyta L. 1764. — Cassita Hill 1765. — Volu- tella Forskal 1775. — Calodium Loureiro 1790. — Rumputris Rafinesque 1836. PRINCIPAL LITERATURE. — Meissner in DC, Prodr. 15 (1) : 252. 1864; Bentham, PL Austral. 5: 308. 1870; Baillon, Hist. PI. 2: 444, 483. 1870; Bentham in B. & H., Gen. PL 3: 164. 1880; Hackeberg- in Verh. Natur. Ver. Rheinl. und Westfalen 46: 98—138. 1889; Pax in Engl. & Prantl, Pfl.Pam. 3 (2) : 124. 1889; Boewig in Bot. Confer. Univ. Pennsylv. 2: 399—416. 1904; Black, Fl. S. Austr. 338. 1922—29; Kienholz in Proc. Ann. Phil. Soc. 65, Suppl. 58—100. 1926; Metcalfe and Chalk, Anat. Dicot. 2: 1152. 1950. TYPE SPECIES. •— Cassytha filiformis L. Number of binomials 50, of which perhaps not more than one third will stand after critical revision. The genus is mainly Australian; one species (C. filiformis L.) is pantropical; Africa has a few species. By its aberrant habit and ecology, it has often been treated as a separate family, but in floral characters it is not different from Lauraceae and approaches Cryptocarya. The plants contain superabundant (even in the cotyledons) mucilagi- nous material. 31 Greek name (xadviag, kasytas or Hadvxas kadytas) for Cuscuta. (dodder). 246 R E I N W A R D T I A [VOL. 4 1957] A. J. G. H, KOSTBRMANS : Lauraceae 247 The epidermis is composed of heavily cutinized, square cells. Stomata in rows; pores at right angles to the axis, rubiaceous, deeply sunken, with heavy cuticular ridge, forming an outer vestibule to the narrow stomatal pore. Outer part of cortex of 1—6 layers of small rounded cells. Endo- dermis not differentiated. Phloem in the form of strands situated in furrows in the outer periphery of the xylem. Xylem in the form of a continuous cylinder, bounded internally by protoxylem groups extending into the pith. Vessels in the inner part of the secondary wood 120 »J, or more in diameter, and much larger than those in the denser peripherpal part; lateral pits large, circular, bordered; perforations simple. A ring of separate vascular bundles, sometimes consisting wholly of phloem, recorded by Solereder in the hypocotyl and young axis. Acicular crystals in the cortex. Fruit enclosed by a hard, horny layer of the endocarp and/or exocarp. The cotyledons remain quite distinct morphologically, but they seem to be adherent by means of some cementing substance. The seeds germinate best in almost pure sand. After germination the food substance is rapidly passed from the cotyledons into the hypo- cotyl, causing it to become very turgid; the food remains dissolved in the cell sap as sugar. Side roots develop, which soon outstrip the main root. Above, the hypocotyl attenuates rapidly into a thin, almost filamentous, bright green stem; it carries up with it the empty seed, which, because of its firm, elastic shell, is difficult to strip off. The seedling show's active circumnutating movements in clockwise direction; the sweep is fairly rapid. The roots have no root cap. ADDITIONS TO: A HISTORICAL SURVEY OF LAURACEAE p. 85. Laurus cassia L. is based on no. 146 in his Flora zeylanica, which in turn is based on Hermann's specimens. In Hermann's Herba- rium (British Museum) these represent a mixture of Litsea zeyl- anica Nees and the wild C i n n a m o m u m zeylanicum (Trimen in J. Linn. Soc, Bot. 24: 140. 1887). Nyctandra Prior (in Proc. Linn. Soc. (1883—86) : 5) is a synonym of Nectandra Rol. ex Rottb. 91. Almost simultaneously with my recombination Dehaasia incrassata (Jack) Kosterm. (April 1952), Merrill published the same new combination in J. Arnold Arb. 33: 230. June 1952. p. 89. p. 94. Through the courtesy of Dra. Ida de Vattimo of the Botanical Garden in Rio de Janeiro, I obtained exhaustive information on the genus Linharea Arruda de Camara. Camara's paper: "Ensaio sobre a utilidade de estabelecer jardins nas principals provincias do Brasil" was never published. This paper was translated in part in Henri Koster's: "Travels in Brazil" (1810). Koster's book was later translated into French (1818) and into Portuguese. On page 584 of the Portuguese edition, corres- ponding to page 493 of the English and page 491 of the French edition, we find the following (translation by de Vatimo) : Plants of Pernambuco * In vol. 1 of J. sci. Res. Indon. (1952). Canela do mato, Linharea aromatica, Arrud. Cent. Plant. Pern. Catinga Branca, Linharea tinctoria, Arrud. Cent. Plant. Pernam. In the first plant the leaves and cortex have a pleasant perfume, resembling the scent of pinkies (clove smell). The plant is not employed by man. I obtained a tasteful extract from the leaves and cortex by distillation. I learned by experience that the liquid extracted from the leaves is not only agreeable to smell and taste but also is very good (therapeutic) for the stomach. It is very abundant in the "taboleiros" of the States of Paraiba and Ceara, in the margins of the Pinhanco; I saw them also in Piani (17). The second plant is a shrub that grows very abundant in the slopes of mountains and water courses margins in the interior of Brazilian states of Pernambuco, Paraiba and Ceara. When boiled the plant yields a yellow colour, long lasting over skins. May be it to be possible to find a way of fixation of this colour in cotton cloth, the same way we do with the tatajuba (Motrus tinctorea). It is also used in the treatment of itch, an eruptive disease. As I could not include these plants in none of the known genera I put them in a new genus and gave it the name Linharea in honour to D. Rodrigo de Souza Coutinho, Count of Linhares, culta and protector of Arts". The footnote 17 refers to a note of Koster and of Camara, which runs as follows: "17. Labat speaks about a species of "canelle batarde" and adds: "On se sert beaucoup, en Italie, d'une canelle semblable a celle que je viens de decrire. Les Portugais l'apportent du Bresil dans des paniers de roseaux refendus et a jour; on l'apelle canelle giroffle 248 REINWARDTIA [VOL. 4 (canella garofanata). On la met en poudre avec un peu de girofle, de veritable canelle, de Poivre, et de graines tout-a-fait resemblables a celles de nos bois d'Inde des Isles, et on en fait un debit assez considerable" — Nouveau Voyage, Tome III, p. 92 (K). Pinhanco is the same as Pianco (C). From the description and from the fact that Linharea aromatica is a common plant, it is possible that it represents Ocotea pretiosa Benth. Nothing can be said of Linharea tinctoria and it may be even not Lauraceous. In Roster's book and the French translation the generic name is wrongly spelt Linharia. p. 95. 1812 Stokes, Jonathan. Through the courtesy of Dr C. V. Morton (Smithsonian Institution, Washington), I obtained a transcript of the original description of Pipalia Stokes in "A botanical materia medica consisting of the generic and specific characters of the plants used in medicine and diet" (vol. 4, page 456. 1812). "884. Pipalia. Calyx of the male flowers none, of the female polyphyllous, corol- la none. Style curved at the base. Stigma capitate. Berry monosper- mous. From Jones. 1. Pipalia solitaria. Gajapippali. Jones in as. res. IV. 313" The genus and the single species are validly published (combined generic and specific description of a monotypic genus. Stokes prob- ably did not know this plant at all; he copied the data from a paper on Indian plants by Sir William Jones, which was published in the Transactions of the Royal Asiatic Society of Bengal (vol. 4: 227. 1801). Of Jones' paper I also obtained a transcript from Dr. Morton. The paper does not bear Jones' name, but merely states that it is by "the late President" (this was definitely Sir William Jones). Jones' paper bears as title: A catalogue of Indian plants, comprehending their Sanscrit and as many of their Linnaean gene- ric names as could with any degree of precision be ascertained" (p. 225). On page 227 we find 169 "Gajapippali, a new genus" On page 295 and 296 1957] " A. J. G. H. KOSTERMANS : Lauraceae "ON SELECT INDIAN PLANTS. 295 249 66. Gajapippali: Syn. Carippali, Capiballi, Colaballi, Srevasi, Vasira. Some add, Chavica, or Chavya; but that is named in the Amara- cosh as a distinct plant, vulgarly Chava, or Chavi. Vulg. Pippal-fhanca, Maidah. MALE FLOWERS. Cal. Common Perianth four-leaved; leaflets roundish, concave; the two exterior, opposite, smaller, containing from eight to four- teen florets. Partial calyx none. Cor. None. Nectary, many yellow glands on the pedicel of the fila- ments. Stam. Filaments from eight to eighteen in each floret, connected by a short villous pedicel, thread-form, very hairy. Anthers large, netted, irregular, inflated, containing the pollen. Pist. Rudiments of a germ and style withering. FEMALE FLOWERS. Cal. Common Perianth as in the male, but smaller; containing from ten to twelve florets. Partial calyx none, unless you assume the corol. Cor. many-petaled, umbelled, Petals erect, lance-linear, fleshy, covered within and externally with white hairs. Nectary, yellow glands sprinkling the receptacle. BOTANICAL OBSERVATIONS Pist, Germ oval. Style cylindric, curved at the base, Stigma headed, Per. Berry globular, one-seeded. Seed spherical, smooth. Flowers umbelled, yellow from their anthers. Leaves mostly oblong-lanced, but remarkably varying in shape, alternate. Both flowers and fruit have an agreeable scent of lemonpeel; and the berries, as a native gardener informs me, are used as a spice or condiment. It was from him that I learned the Sanscrit name of the plant; but as balli means a creeper, and as the Pippal-jhanca is a tree perfectly able to stand without REINWARDT1A [VOL. 4 1957] A. J. G. H, KOSTERMANS : Lauracea? 251-: support, I suspect in some degree the accuracy of his informa- tion; though I cannot account for his using a Sanscrit word without being led to it, unless he had acquired at least traditional knowledge. It might be referred, from the imperfect mixed flow- ers, to the twenty-third class." From this description it is evident, that Pipalia represents either Litsea or Lindera (the number of anther cells is not indi- cated) . The shape of the leaf and their scent points to Litsea cubeba Pers. p. 113. 1823 C. G. Nees and Th. Fr. L. Nees von Esenbeck, de Cinnamo- momo disputatio (a supplement: "Berichtigung zur Disputatio de Cinnamomo" appeared in Allg. Bot. Zeit. 34: 1—30. 1831). In this paper the authors described new Cinnamomum species under the generic name Laurus and an exhaustive historical survey is given of Cinnamomum zeylwnicwm Bl. I N D E X T O L A U R A C E A E Names published as new in the original publication, Comm. For. Res. Inst. Indon. no. 57, March 22, 1957, of which this article is a reprint, are in this index still given in bold face type. The page number in the original publication is found by subtracting 192 from the. present number. Synonyms are given in italics. The names on pages 218 and 219 are not listed in this index. F I G . 1. Relationships within the Lauraceae. Acatsjavalli (Rheede) Adans. 245. Sub gen. Acrodiclidium (Nees) Kosterm. 238. Acrodiclidium Nees 238. Actinodaphne Nees 194, 195, 199, 200, 203, 205, 210, 214, 217, 234, 235; prui- nosa Nees 234. Adapting Neck, 242. Adenodaphne S. Moore 240. Ade?iotrachelima Baill. 232. Adenotrachelium Nees ex Meissn. 231. Afrodaphne Stapf 229. Agathophyllum (non Bl.) J u s s . 244. Agathophyllum (non Willd., nee Juss.) Bl. 231. Agriodaphne Nees ex Meissn. 231. Sub gen. Aiouea (Aubl.) Kosterm. 236. Aiouea Aubl. 1951 197, 198, 203, 207, 208, 210, 217, 220, 233, 236, 237; guia- nensis Aubl. 236. Subgen. Alseodaphne (Nees) .Benth. 195, 196, 197, 199, 200, 201, 203, 206, 210, 211, 212, 227. Alseodaphne Nees 225, 226, 227, 228; coriacea Kosterm. 208; paludosa King 210; semecarpifolia Nees 226. Ampelodaphne Meissn. 237. Anaueria Kosterm. 229. Aniba Aubl. 196, 198, 200, 201, 202, 203, 207, 209, 210, 214, 217, 236, 237, 238; canelilla Mez 208, 216; coto Kosterm. 216; duckei Kosterm. 216, 237; guia- nensis Aubl. 237; kappleri Mez 206, 208; ridleyana Mez 214; rosaeodora Ducke 199, 216, 237. Subtrib. Anibineae Kosterm. 236. Annonales 215. Apella Adans. 242. Aperula Bl. 241. Aperiphracta Nees ex Meissn. 231. 252 R E I N W A R D T I A [VOL. 4 1057] A. J. G. H. KOSTEKMANS: Lauraceae 253 Apivea Steud. 236. Apollonias Nees 196, 209, 217, 228; bar- busana (Cav.) A. Braun 228. Aydendron Nees & Mart. 236, 238. Balanopsis Raf. 231, 241. Beilschmidtia Rchb. 229. Siibgen. Beilschmiedia 229. Beilschmiedia Nees 193, 195, 196, 198, 199, 200, 201, 202, 203, 205, 207, 208, 209, 211, 214, 217, 228, 229, 230; bul- lata Kosterm. 199; endiandraefolia Kosterm. 229; mannii Benth. 202; myrmecophila Kosterm. 213; pendula Benth. 202; roxburghiana Nees 200, : 207, 208, 209, 229; tarairi Benth. 217; tawa Benth. 217; variabilis Rob. & Wilcz. 196, 211. Subtrib. Beilschmiediineae Kosterm. 228. Bellota Gay 229. Benzoin B.oehm. 214, 241. Berberidaceae 215. Berberis 205. Berniera Baill. 229. Berrija (non Roxb.) Klein ex Willd. 240. Bihania Meissn. 243. Bistania Nor. 241. Boldu (non Feuillee) Nees 229. Boldus (non Adans.) O. Ktze 229. Borbonia [Plum.] Boehm. 231. Brassiodendron Allen 229. Bryantea Raf. 241. Calodium Lour. 245. Calosmon Bercht. & Presl. 241. Calycanthaceae 215. Calycodaphne Boj. 231. Camphora [Bauh.] Boehm. 233. Camphorina Nor. 233. Camphoromoea Nees ex Meissn. 231. Cannella Schott ex Meissn. 231. Cansiera (non Juss.) Spr. 231. Caryodaphne Bl. ex Nees 243. Caryodaphnopsis Airy-Shaw 225. Cassita Hill 245. Cassyta L. 245. Cassytha L. 195, 196, 197, 203, 205, 206, 211, 212, 217, 220, 221, 245; filiformis L. 245. Cassythaceae Dum. ex Lindl. 245. Subordo Cassytheae Meissn. 220, 221. Trib. Cassytheae Nees 220, 245. Subfam. Cassythoideae Kosterm. 245. Cecidodaphne Nees 233. Ceramocarpium Nees ex Meisnn. 231. Ceramophora Nees ex Meissn. 232. Cedrota Schreb. 236. Chanekia Lund. 238. Chlorantaceae 215. Subtrib. Cinnamomineae Kosterm. 231. Cinnamomoides 214. Trib. Cinnamomeae Baill. emend. Kos- term. 220, 231. Cinnamomum Boehm. 194, 195, 196, 198, 199, 200, 201, 202, 203, 208, 210, 213, 214, 217, 226, 227, 232, 233, 234; bur- mannii Bl. 203, 213; camphora Nees & Eberm. 198, 202, 213, 216; cassia Bl. 216; iners Bl. 197, 203; japonicum 213; javanicum Bl. 203; linearifolium 213; obtusifolium 213; pedunculatum 213; porrectum Kosterm. 216; sericeum Sieb. 208; sieboldii 213; zeylanicum Bl. 200, 213, 216, 233, 250. Sub gen. Clavipersea Kosterm. 225, 226, 227, 228. Clinostemon Kuhl & Swamp. 238. Colomandra Neck. 236. Cricula trifenestrata 213. Crowella 22. Cryptocaria Gay 243. Subgen. Cryptocarya 244. Cryptocarya R. Br. 195, 196, 198, 199, 200, 201, 202, 203, 206, 208, 209, 210, 211, 212, 217, 221, 240, 243, 244, 245; caloneura Kosterm. 213; corrugata C. T. White 202; glaucescens R. Br. 202, 244; konishii Hay. 203; massoy Kos- term. 216; moschata Nees & Mart. 216, 244; perrieri Danguy 208. Trib. Cryptocaryeae Meissn. 220, 242. Subtrib. Cryptocaryneae Kosterm. 243. Cubeba Raf. 240. Cussuta Rumph. 245. Cyanodaphne Bl. 228. Cylicodaphne Nees 214, 240. Cyna/momum Den. 233. Damburneya Raf. 231. Subtrib. Daphnidieae Meissn. 220. Daphnidium Nees 241. Daphnogene 214. Daphnophyllum 214. Darwinia Dennst. 240. Darwiniana "Dennst." ex Lindl. 240. Decapenta Raf. 240. Dehaasia -Bl. 196, 200, 203, 210, 228; caesia Bl. 195, 216; incrassata (Jack) Kosterm. 228, 246. Dendrodaphne Beurl. 231. Dictyodaphne Bl. 229. Dicypellium Nees & Mart. 195, 201, 203, 205, 208, 217, 235, 239; caryophyllatum Nees 216, 236. Dipliathus Raf. 238. Subgen. Dodecadenia (Nees) Kosterm. 241. Dodecadenia Nees 240. Drimophyllum Nutt. 235. Douglassia Schreb. 236. Ehrhardia Scop. 236. Endiandra R. Br. 195, 198, 199, 201, 202, 203, 207, 208, 209, 214, 217, 229, 230; fragrans Kosterm. 229, 230; glauca Kosterm. 229. Endlicheria Nees 196, 198, 201, 203, 204, 205, 207, 210, 236, 237; arunciflora Mez 204; bullata Ducke 199; glomerata Mez 204; longifolia Mez 216; paradoxa Mez 205, 208; sericea Nees 237. Subgen. Endocarpa (Raf.) Kosterm. 236. Endocarpa Raf. 236. Subgen. Enneanthera Kosterm. 244. Subgen. Ennearrhena Stapf 229. Euodia Gaertn. 244. Euosmus Nutt. 234, 241. Sect. Eupersea Pax 227. Euphoebe Bl. ex Meissn. 225. Subtrib. Eusideroxylineae Kosterm. 242. Eusideroxylon T. & B. 195, 196, 202, 208, 209, 210, 211, 212, 214, 217, 221, 242; melagangai Sym. 2,04, 243; zwageri T. & B. 196, 202, 216, 243. Evelyna Raf. 240, 241. Evonymodaphne Nees ex Lindl. 238. Evosmus Raf. 234, 240, 241. Farnesia Heist, ex Fiva Steud. 240. Fiwa Gmel. 240. Fabr. 225. Glabraria (non L.) Bl. 240. Sect. Gnesiopersea, Benth. 227. Goeppertia Nees 237. Gomortegaceae 215. Gymnobalanus Nees & Mart. 241. Subordo Gyrocarpeae Meissn. 220. Haasia Nees 228. Hamamelidaceae 215. Heckeria Raf. 240. Hernandia Plum. 220. Hernandiaceae 214, 220. Trib. Hernandiae Benth. 220. Subgen. Hexanthera Kosterm. 244. Hexanthus Lour. 240. Hexapora Hook. f. 217, 230; curtisii Hook f. 230. Subgen. Hexarrhena Stapf 229. Huberodaphne Ducke 237. Hufelandia Nees 193, 229. Trib. Hypodaphneae Kosterm. 224. Hypodaphnis Stapf 195, 196, 202, 203, 209, 211, 217, 231, 244; zenkeri Stapf 202, 244. Icosandra Phil. 243. lozoste Nees 234, 240. Subgen. Iteadaphne (Bl.) Kosterm. 242. Iteadaphne Bl. 241. , Jozosmene Nees ex Lindl. 240. Laurales 215. Laureaceae Lindl. 225. Laureae Rchb. 225. Subordo Laureae Mez 221. Lauri Juss. 220, 225. Subtrib. Lauriineae Kosterm. 241. Laurinae Vent. 225. Laurineae verae Zoll. 225. Laurineae St. Hil. 225. Subordo Laurineae Meissn. 220. Lauriniutn 214. Laurinoxylon 215. Lauriphyllum 215. K E I N W A R D T I A [VOL. 4 1957] A. J. G. H. KOSTEKMANS: Lauraceae 255 Laurocalyx 215. Laurocarpum 215. Subfam. Lauroideae Mez emend Kosterm. 225. Subfam. Lauroideae Pax 193, 221. Lauromerrillia Allen 207, 229. Laurus L. 195, 201, 202, 205, 213, 215, 220, 225, 242, 250; cassia 246; nobilis L. 213, 216, 242. Lepidadenia Walker Arn. ex Nees 240. Leptodaphne Nees & Mart. 231. Subgen. Licaria 238. Licaria Aubl. 195, 196, 199, 200, 201, 202, 203, 206, 208, 210, 212, 217, 220, 238, 239; camara Kosterm. 210; capitata Kosterm. 204; cinnamomoides Kosterm. 216; guianensis Aubl. 204, 238; puchury-major Kosterm. 216. Subgen. Lindera (Thunb.) Kosterm. 242 Lindera Thunb. 195, 198, 201, 202, 203, 204, 203, 213, 215, 216, 217, 220, 241, 250; aestivale 213; benzoin 213; bifaria Benth. 202; erythrocarpa Mak. 202; glauca 213; subumbelliflora Kosterm.; umbellata Thunb. 242, 203. Linharea A. de Cam. 231, 247; aromatica A. de Cam. 247, 248; finctoria A. de Cam. 246, 248. Linharia 248. Litsaea Juss. 240. Litsea sect. Neolitsea Benth. 241. Litsea Lamk 195, 196, 198, 200, 201, 202, 203, 204, 205, 206, 207, 209, 210, 215, 216, 217, 234, 235, 240, 241, 242, 250; ohinensis Lamk 240; cubeba Pers. 196, 250; odorifera Val. 216; zeylanica Nees 246. Subgen. Litsea (Lamk) Kosterm. 241. Litsaeeae Mez 221, 239. Trib. Litseaceae Meissn. 220. Subtrib. Litseineae Kosterm. 239. Subgen. Machilus (Nees) Kosterm. 227. Machilus Nees 194, 203, 225, 226; thun- bergii Sieb. & Zucc. 227. Magnoliaeeae 215. Magnoliales 215. Malapoenna Adans. 240. ; Menestrata Veil. 225. Mespilodaphne Nees & Mart. 215, 231. Mezia (nee Schw.) O. Ktze 230. Mezilaurus Taub. 196, 197, 199, 200, 202, 203, 206, 207, 208, 209, 217, 230; crassiramea Mez 199; ita-uba Mez 230; lindaviana Schw. & Mez 202, 203; na- valium (All.) Taub. 230. Micropora Hook. f. 230. Subgen. Misanteca (Cham. ' & Schl.J Kosterm. 238. Misanteca Cham. & Schl. 238. Monimiaceae 215. Morus tinctorea 247. Myristica 220. Myristicaceae 215. Subgen. Nectandra (Rol.) Kosterm. 199, 201, 205, 232. Nectandra. Rol. ex Rottb. 199, 214, 217, 231, 232, 246; leucantha Nees 204; sinuata Mez 199. Nemodaphne Meissn. 232. Neocinnamomum Liou-Ho 233. Neolitsea Merr. 195, 203, 205, 215, 217, 241; acuta-trinervia Kan. & Sas. 202; cassia (L.) Kosterm. 242. Neosilvia Pax 230. Nesodaphne Hook. f. 229. Nobeliodendron O. C. Schm. 238. Subgen. Nothaphoebe (Bl.) Benth. 226, 227, 228. Nothaphoebe Bl. 225, 226, 227; umbelli- flora Bl. 226. Nyctandra Prior 232, 246. Nyrophyllum Neck. 225. ' Subgen. Ocotea (Aubl.) Kosterm. 232. Ocotea Aubl. 194, 198, 199, 200, 201, 203, 205, 208, 209, 210, 214, 215, 217, 220, 231, 232, 233, 234, 236; aniboides Mez 204; bullata E. Mey. 216, 217, 232; clavigera Mez 210; funiculacea Mez 216; guianensis Aubl. 232; pretiosa Benth. 248; rodioei Mez 195, 202, 203, 216, 232, 243; veraguensis Mez 216, 232; verruculosa Mez 197. Trib. Ocoteae Baill. 220. Subgen. Octolitsea Liou-Ho 205, 241. Oreodaphne Nees & Mart. 215, 231. Trib. Oreodaphneae Meissn. 220, 281. Ozanthes Raf. 241. Parabenzoin Nakai 241. Parthenoxylon Bl. 243. Subgen. Persea (Boehm.) Kosterm. 227. Persea Boehm. 194, 197, 198, 200, 201, 202, 203, 205, 207, 208, 209, 213, 215, 217, 225, 227, 232, 233; americana Mill. 196, 199, 201, 204, 205, 210, 212, 213, 216, 226; carolinensis 213; lingue Nees 216, 227; pubescens 213; thunbergii (Sieb. & Zucc.) Kosterm. 227; tonki- nensis Kosterm. 212. Trib. Perseeae Mez 221, 225. Trib. Perseaceae Meissn. 220, 225. Subtrib. Perseineae Kosterm. 215. Perseoxylon 215. Subfam. Perseoideae Pax 193, 202, 221, 225. Petalanthera Nees & Mart 231. Peumus Mol. 243. Phoebe Nees 194, 196, 197, 198, 199, 201, 203, 209, 210, 217, 226, 227, 232, 233; amoena Mez 210; declinata Bl. 203; lanceolata Nees 227. Phyllostemonodaphne Kosterm. 195, 205, 208, 217, 236, 239; geminiflora (Me- issn.) Kosterm. 239. Pipalia Stokes 240, 248, 250; solitaria Stokes 248. Subgen. Pleurothyrium Nees Kosterm. 196, 203, 207, 232. Pleurothyrium Nees ex Lindl. 213, 217, 231, 232. Polydenia Nees 241. Polycarpicae 215. Pomatium Nees & Mart, ex Lindl. 231. Porostema Schreb. 231. Potameia Thou. 194, 195, 196, 205, 211, 217t 231; argentea Kosterm. 231; chi- nensis Kosterm. 206; thouarsii Roem. & Sch. 231; velutina Kosterm. 206. Potamica Poir. 231. Proberberideae 215. Protoravensara 215. pseudocryptocarya Teschn. 243. Pseudolitsea Yang 240. Pseudosassafras Lee. 234, 235. Purkayasthaea Purk. 229. Quinquedula Nor. 240. Ranales 215. Ravensara Sonn. 197, 198, 200, 201, 202, 209, 211, 212, 217, 220, 244; aromatica Sonn. 216, 244. Rombut Rumph. Adans. 245, Rumputris Raf. 245. Salguda Blanco 243. Sassafras Bercht. & Presl. 241. Sassafras [Kramer] Boehm. 194, 195, 199, 200, 201, 204, 213, 215, 216, 217, 234, 235; albidum (Nutt.) Nees 234; officinale 213. Sassa-fridiuni Meissn. 232. Schuera Nees 237. Schaueria Nees & Meissn. 237. Sciadiodaphne Rchb. 235. Sebifera Blanco 240. Senneberia Neck. 231. Sennebiera O. Ktze 232. Septina Nor. 233. Silvaea Meissn. 230. Silvia Allem. 230. Stremmatodaphne Gamble 225. Strychnodaphne Nees & Mart. 231. Symphysodaphne A. Rich. 238. Synandrodaphne Meissn. 232. Syndiclis Hook. f. 194, 231. Systemonodaphne Mez 200, 207, 210, 217, 239; mezii Kosterm. 239. Tamala Raf. 225. Teleindra Nees & Mart. 231. Tetradenia (non Benth.) Nees 241. Tetrwnthera Jacq. 215, 240. Subtrib. Tetranthereae Meissn. 220. Thouvenotia Dangny 229. Thymelaeaceae 215. Tomex Thunb. 220, 240. Subgen. Triandra Kosterm. 244. Subgen. Trianthera Mez 236. Trianthera Conwentz 214, 215. Triplomeia Raf. 238. Tylostemon Engl. 193, 229. 256 R E I N W A R D T I A [VOL. 4 Umbellularia Nutt. 195, 201, 202, 203, 216, 235; californica (Nees) Nutt. 202, 205, 235. Urbanodendron Mez 195, 200, 207, 210, 217, 238; verrucosum (Nees) Mez 238. Virola 220. Volutella Forsk. 245. Wimmeria Nees ex Meissn. 229. Yushuma Kam. 234. R E I N W A R D T I A Published by Herbarium Bogoriense, Kebun Raya Indonesia Volume 4, Part. 2, pp. 257-280 (1957) FLORAE MALESIANAE PRAECURSORES XVI ON THE TAXONOMIC SUBDIVISION OF THE GLEICHENIACEAE, WITH DESCRIPTIONS OF NEW MALAYSIAN SPECIES AND VARIETIES by R. E. HOLTTUM * SUMMARY A new subdivision is given of the fern family Gleicheniaceae. The genus Platyzoma R. Br is excluded from the family. The genus Stroma- topteris from New Caledonia is arranged in a distinct subfamily. In the remainder of the family, subfamily Gleichenioideae, two genera are recog- nized, Gleichenia (with subgenera Diplopterygium, Gleichenia, and Mertensia) and Dicranopteris (with subgenera Acropterygium and Dicra- nopteris) . The problem of subdividing the family is discussed with reference to former treatments and to new data, and a conspectus of the new system, with synonymy and key to the genera and subgenera, is given. A number of new species, new varieties, and new combinations is made both in Gleichenia and Dicranopteris. In preparing a taxonomic revision of the family Gleicheniaceae for Flora Malesiana, I have reviewed the status of the genera proposed within the family by previous authors, and as a result have been led to take a position midway between the arrangement of Christensen (Index Filicum 1905) and of Copeland (Genera Filieum 1947). The present paper gives a summary of the facts on which this decision was reached; a fuller comparative treatment of the subject, with a discussion on morphology and growth-habit in this and other families of primitive ferns, will be published elsewhere. The genus Platyzoma R. Br., which has usually be included in Gleich- eniaceae (even in the genus Gleichenia), appears to me so different that it should be excluded. A statement on this subject has been published separately (Kew Bulletin 1956; 551) ; the genus will not be further men- tioned in the present paper. The genus Stromatoptetis, confined to New Caledonia, is peculiar in various ways, but has superficial sori of sporangia which agree with 1 Formerly Professor of Botany in the University of Malaya. — 257 — Binder13 Rein.Vol 4,Part 2,pp 119-310_Page_01 Rein.Vol 4,Part 2,pp 119-310_Page_39 Rein.Vol 4,Part 2,pp 119-310_Page_40 Rein.Vol 4,Part 2,pp 119-310_Page_41 Rein.Vol 4,Part 2,pp 119-310_Page_42 Rein.Vol 4,Part 2,pp 119-310_Page_43 Rein.Vol 4,Part 2,pp 119-310_Page_44 Rein.Vol 4,Part 2,pp 119-310_Page_45 Rein.Vol 4,Part 2,pp 119-310_Page_46 Rein.Vol 4,Part 2,pp 119-310_Page_47 Rein.Vol 4,Part 2,pp 119-310_Page_48 Rein.Vol 4,Part 2,pp 119-310_Page_49 Rein.Vol 4,Part 2,pp 119-310_Page_50 Rein.Vol 4,Part 2,pp 119-310_Page_51 Rein.Vol 4,Part 2,pp 119-310_Page_52 Rein.Vol 4,Part 2,pp 119-310_Page_53 Rein.Vol 4,Part 2,pp 119-310_Page_54 Rein.Vol 4,Part 2,pp 119-310_Page_55 Rein.Vol 4,Part 2,pp 119-310_Page_56 Rein.Vol 4,Part 2,pp 119-310_Page_57 Rein.Vol 4,Part 2,pp 119-310_Page_58 Rein.Vol 4,Part 2,pp 119-310_Page_59 Rein.Vol 4,Part 2,pp 119-310_Page_60 Rein.Vol 4,Part 2,pp 119-310_Page_61 Rein.Vol 4,Part 2,pp 119-310_Page_62 Rein.Vol 4,Part 2,pp 119-310_Page_63 Rein.Vol 4,Part 2,pp 119-310_Page_64 Rein.Vol 4,Part 2,pp 119-310_Page_65 Rein.Vol 4,Part 2,pp 119-310_Page_66 Rein.Vol 4,Part 2,pp 119-310_Page_67 Rein.Vol 4,Part 2,pp 119-310_Page_68 Rein.Vol 4,Part 2,pp 119-310_Page_69 Rein.Vol 4,Part 2,pp 119-310_Page_70 Rein.Vol 4,Part 2,pp 119-310_Page_71