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BLUMEA 46 125-140 (2001) An annotated key in to the genera of Henk Missouri Lauraceae the flora Malesiana region Botanical Garden, van der P.O. Box Werff 299, St. MO Louis, USA 63166, Summary important component of wet tropical Lauraceae are an Malesiana region. poorly defined and together on a problems fication Key in are words. Their identification has been existing keys require Here specimen. generic a key delimitation are both based forests and flowers and almost discussed and fruits, entirely well are in represented two factors: several of the hampered by on vegetative which are, are however, rarely present flowering specimens characters the Flora genera in helpful is presented, generic identi- listed. Lauraceae, genera, characters. key, Malesia, flowers, vegetative Introduction Lauraceae are often Flora Malesiana wood or are Lauraceae to important component of wet, tropical an form region exception. Many species no commercially exploited. Still, of variety a remains very difficult. species genus or for forests and forests in the locally are used for their identification of reasons, Existing keys frequently require both flowers and fruits and because fruits of Lauraceae need several months before reaching maturity, both flowers and fruits are Therefore, keys requiring flowers and fruits rarely present are essentially on herbarium specimens. worthless for identification purposes. Recently, the FM area some keys to genera found in the FM were genera written. 1 and area boiling up few flowers when I also list several fruit and generic have identification. not surveyed the range of variation in in the list of fruit and The brief the FM area. published for small suggest; for example key based for which areas on of flowering they material the number of characters which seen many Cryptocarya stamens seem species in and anther locules. helpful indicators for the smaller genera, I R.Br, and Litsea Lam. and the full probably not expressed characters. for each genus larger species a characters in these genera is vegetative descriptions given work outside the present parts include all necessary tool for identification. I recommend I have genera vegetative the a vegetative large Especially is counting Although not to not Because of the small size of the flowers of Laura- area. good dissecting microscope a might it would be useful thought for all genera found in the FM ceae, a of Lauraceae have been (Kochummen, 1989; Middleton, 2000). These keys do apply only genera may be of Persea Mill, and more to the species occurring variable than the Caryodaphnopsis in descriptions Airy Shaw may 126 BLUMEA have 2-locular stamens No. 46, 6 instead of 9 fertile or 2001 1, and stamens few Chinese a species in Litsea have clustered leaves. placed Sterile, fruiting below. In pistillate specimens or vegetative some cases while in other specimens, the —Vol. characters generic has cases one to help can the recognize key presented in identification of such specimen species, to because visible. not are be identifiedwith the cannot fruit characters or CHARACTERS Flowers For better a of the understanding key and discussions I will All the other a small in with unisexual flowers species sex are stigma, without stipitiform pistillode flowers the staminodia may have pistillate broadened a absent. The of reduction of degree and is worth not all, genera flowers are floral parts: whorls, which Tepals: than the in arranged sometimes although they two whorl; the reverse, the outer are again opposite whorl the their base 2 at may form a cushion Endiandra R.Br.) Pleurothyrium their base whorl III glands at form enclosing the all due all enclosing to the few stamens (if more and glands that are either In 1 locule. The anther locules most genera the (introrse) stamens extrorse; in others all or are are said exserted. Because in twisted, it is difficult to of the on or and of species neotropical three) having glands or all glands stamens see, not at of have surprising small lauraceous flower. 2 locules; flaps rarely the side of the anther so char- do anthers facing the towards the outside of the flower (extrorse). of whorl I and II have introrse locules, while whorl III is stamens genera with umbellate inflorescences, the anthers are facing 4 of whorl enlarged some in small chambers, which open with have of the flower in (as stamens Stamens of stamens become most a and rule whorl III tepals, greatly enlarged fitted in a (I, II, III, IV, tepals. small in size and difficultto are typically only than of the as tepals. Thus, species most In Cinnadenia Kosterm. glands pollen glands of whorl III (as in stamens the genera the the whorls of than the inner whorl, is the inner most acteristic for Lauraceae. The anthers have center a may be shape inner whorl is the inner opposite cases in in four whorls opposite In misinterpretation stamens. stamens produce a a stamens Nees). Reports of all mostly in the larger opposite whorl of six. globose glands; their base, but these stamens of shape they following usually equal case whorl whorl II and whorl IV one surrounding or the number of given The are to are arranged and inside) tepals, outer tepals outer I and II appear III bear the opposite outer Stamens: counted from the outside towards the of whorl I or well documented not the encounters In that unequal. are (Triadodaphne Kosterm.). rare anther trimerous, with the whorls of floral parts in center one very an and staminodes is pistillode may be absent; investigating. In most, but larger of of with pistillode a pistillode unisexual. remnants of an anther in the trace a find may or a a remnant threes. From the outside towards the size, one stigma a may be threadlike without tip, they or dioecious. In unisexual flowers are In staminate flowers usually present. describe the briefly flower structure of Lauraceae. Flowers of Lauraceae are either bisexual to have introrse locules. The latter is the in which the stamens have these genera the filaments ascertain if the locules are all introrse or long are not case in filaments and long and often and I have not H. der van Werff: An used this character in the are the lateral. In case, stamens ments, all the anthers whorls of stamens 6 3 fertile or same are is stamens. to some (Endiandra, Eusideroxylon 3 only IV all fertile) not are shorter and and stamens one or disappear to genera it is staminodial, in others it has stamens disappear fila- long have usually stamens staminodia. The first whorl is com- and then flowers have present, these represent whorl III are & Binn. and Teijsm. the locules instance) number of locules; if this is space for 12 fertile enough reduced If 127 have fewer locules than those of whorl I and II. to not Lauraceae for (in Endiandra, have the Sometimes additional whorls pletely disappeared. of genera exserted from the flower. When the whorl IV, the innermost whorl; in only the to (whorls I, II, III, stamens are included in the flower, there more key some cases stamens of whorl III tend Flowers with 12 fertile so In key. species most annotated An Triadodaphne). of this example trend towards reduction of number of stamens in smaller flowers is found in the genera Kosterm. Potoxylon and Eusideroxylon. all By these accounts genera closely are related; Potoxylon has larger flowers and 9 fertile stamens, while Eusideroxylon much smaller flowers with 3 fertile and with one unprotected a occupies an plate-like completely Inflorescence its base, or is seated in Eusideroxylon and Poto- receptacle. Cryptocarya in flowers of this genus the ovary is enclosed in the fruits technically superior; receptacular tube and thus inferior. are types The inflorescences of the Lauraceae (excluding three main types (Van der Werff& Richter, to one In at and thus hemi-inferior; fruits receptacle fused with it and thus not and the fruit is seated superior cup-shaped. or enclosed in and fused with the are is enclosed in and fused with the position; tube, but ovary has unilocular unicarpellate, persistent tepals embedded in the partially are receptacular however be shallow, intermediate the exceptions sometimes with pedicel, can the ovary is in these genera the the which cupule xylon ovule. With few on The ovary is stamens. of these groups, because in reduced while a few species or some cases genera Cassytha L.) be divided into can be assigned inflorescences have become strongly 1996). Not all inflorescences have inflorescences which can simply do not match of the three types. any In the first type the flowers are arranged in when young, umbels, which, in decussate bracts. All flowers with this type of inflorescence bels can be single in the axils of leaves shortshoots. Umbels can be sessile or or bracts or grow into can are be in In bracts a the shortshoot has normal easily pairs, vegetative a raceme whereas the tepals are not of single Inflorescences of the second type by are a dichasium racemose are are reduced to a um- leafless arranged along a contrast flowering, can the decussate bracts, single flower; case the umbels the presence of the inflorescence consists of flowers. are paniculate-cymose. These inflorescences in dichasia. The always strictly opposite (Fig. 1). through are surrounding branched, with the flowers ultimately arranged flowers of such rescences arranged along of umbels, but in still covered enclosed in whorls of three. Blume the umbels 1-flowered umbels and are unisexual. The terminal bud, which, after leafy twig. Young umbels, the base of the flower indicates that in this repeatedly be mistaken for flower buds. However, the bracts Iteadaphne at a can stalked. When the umbels leafless shortshoot, the whole structure looks like to a raceme, they are reduction of the lateral axes. are lateral Sometimes the inflo- BLUMEA 128 Vol. 46, Fig. No. 2001 1, 1. Detail of cencewith a paniculate-racemose the lateral flowers of the inflores- cymes strictly opposite (type 2). Inflorescences of the third type the ultimate cymes alternate short always not also not paniculate. However, especially to see, easy not of Beilschmiedia Nees and Endiandra tend makes it easier racemose to recognize this inflorescence inflorescence with rarely three or Two genera ( area Actinodaphne Nees and in not to have has together by in somewhat with very Cryptocarya). a on predominantly the basal lateral reduction of the lateral type. Cinnadenia) occur in the Flora Malesiana which have inflorescences unlike the three types described above. The type ofActinodaphne, as well as several other which longer pedicels type. Potoxylon four flowers are species uncommon branches of the inflorescence. This inflorescence is derived branches from the inflorescence of the third the lateral flowers of and instead strictly opposite, and subsessile flowers (a condition pedicels Species This is (Fig. 2). are frequently are species placed in that genus, has a species paniculate inflorescence which is enclosed in bracts when very young. These bracts do not appear to be decussate, but inflorescence, rescences 1-2 cm in A. long are but leave alternate. a pilosa (Lour.) and finally pseudo-umbellate They cluster of there Merr. are fall off very scars are several to at 10 cm long, species placed inflorescence, this still with bract ference between the early Actinodaphne species in the development of the the very base of the inflorescence. Inflo- with a in A. malabarica Balakr. in Actinodaphne scars at the very pseudo-umbellate with a only sessile base. The dif- inflorescence and H. van der Werff: An annotated key to Fig. the genera 2. Detail of a of Lauraceae paniculate 129 inflorescence the ultimate flowers not arranged in strict with cymes (type 3). Litsea species with sessile umbels is difficult clustered leaves. area Fortunately, have alternate or the great to see, majority (if subopposite leaves, while are with species of Litsea in the FM all) species Actinodaphne species generally whorled leaves. The inflorescences of Cinnadenia are niculate, the flowers (unisexual) in Litsea especially not paniculate racemosely arranged along or racemose. the secondary have If pa- axes of the inflorescences. Fruit and fruit and mind some floral characters vegetative though that there characters vegetative Although of prime may be many I have used the usually term exceptions vegetative perulate some internodes. importance for the determinationof genera, times also be very only give to an helpful. indication of One should keep generic assignment these indications. Still, it is worthwhile in and to list characters. buds for fall off after the buds twigs. Only rarely (in the base of the at can that these characters of these fruit and bracts are characters vegetative elongate Neolitsea buds covered by bracts. These and leave distinct clusters of scars species) do the bracts persist as a on the short sheath at BLUMEA 130 Vol. In triveined leaves the main lateral veins the lamina, while in No. 46, depart 2001 1, from the midrib at the very base of leaves these veins leave the midrib tripliveined a little above the base of the lamina. - Fruits enclosed in the floral tube and with the fruit: large, Fruits less generally seated in a than 5 cupshaped of the fallen tepals scars Potoxylon. Eusideroxylon rather slender fruits (more than 5 has fruits - and Cryptocarya, Eusideroxylon cm long, 2-3 cm on top of the Potoxylon have wide) whereas Cryptocarya long. cm Neolitsea Merr. and bowlshaped cupule: Litsea, or and Actinodaphne. - - - Fruits with persistent, spreading Fruits with persistent, appressed tepals Fruits seated unprotected to the on reflexed tepals at the base: Persea. the base: Phoebe Nees. at pedicel, without persistent tepals or a cupule: Alseodaphne Nees, Beilschmiedia, Brassiodendron C.K. Allen, Dehaasia Blume, Endiandra. - with Twigs pale, whitish bark, this with the dark contrasting petioles: Alseodaphne, Dehaasia. - Leaves clustered, without Alseodaphne, Dehaasia, - Leaves clustered, with Leaves clusters or of the branches: along scars buds and clusters of perulate scars along thebranches: Actino- Persea. daphne, Neolitsea, - buds perulate Phoebe. opposite: Caryodaphnopsis, CinnamomumSchaeff. p.p., Iteadaphne Blume, Litsea p.p. - - - - - Leaves triveined tripliveined or opposite: Cinnamomum. Caryodaphnopsis, triveined, alternate: Lindera Thunb. p.p., Neocinnamomum H. Liu. Leaves tripliveined, Terminalbuds alternate: working comment p.p., Dehaasia A with Lauraceae, briefly scarcity on of good, Lauraceae collections good ments (for IN two flowering are problems are trees soon Caryodaph- become apparent and I and have small, often among the ten (Hyland, 1989) 1986, in which these collections in which It took nearly more a team together. to want to the fact that many flowers. Even inconspicuous most common few. This is reflected in great majority of species. bring p.p., STUDYING LAURACEAE from fewer than 10 collections). A welcome to Alseodaphne collections. No doubt this is due instance Kostermans, in Australia p.p., Neolitsea p.p., Litsea p.p., p.p. those: Lauraceae grow into rather tall though Lindera p.p., Litsea p.p., Neolitsea, Persea. glaucous: Actinodaphne PROBLEMS When p.p., Lindera p.p. Cryptocarya perulate: Actinodaphne, Lower leaf surface nopsis, Cryptocarya 1. and Leaves families in forest all revisions most exception species or floristic treated are plots, treat- known is the Revision of Lauraceae than 20 collections were studied of the of dedicated field collectors several years For the FM area, good flowering collections are very much needed. 2. The poor delimitation of genera. In Werff& Richter, tem: earlier classifications (reviewed 1996) generic delimitationwas largely based on a in Van der numerical sys- number of fertile stamens, number of anther locules, combined with inflores- H. these characters results in with its mostly 6 2-locular in use forming delimitation will groups of it I would mately the relationships not yet at acters. One I was premature taking into unable to recognize species not key. In the other case, our understanding More and better collections THE GENERA la. Herbaceous b. Erect a a a parasitic woody plants sometimes two 1989) allows or Alseodaphne needed to was pointed solve these IN THE vine with leaves reduced regular based Nothaphoebe OF LAURACEAE with Nothaphoebe, to floral char- of the two, I include and have therefore included two of this group, as are on character used in earlier separation and Alseodaphne genera in the number of anther cells, along on out did not s.l. and, with of new collections has Kochummen (1989). problems. FLORA MALESIANA AREA to . . minute bracts 6. Cassytha 2 or rarely (in some or bracts, Actinodaphne species 3 paniculate, rarely 8 racemose dimerous 4 b. Flowers trimerous 5 Inflorescences umbellate; terminal buds b. Inflorescences paniculate; terminal buds perulate; not stamens surrounded 6 18. Neolitsea .... by bracts; stamens 18- 7. Cinnadenia 32 5a. Leaves whorled b. Leaves alternate, hnd Nothaphoebe description fruiting by I leaves leafless shortshoots b. Flowers bisexual, inflorescences 4a. Ulti- classification reflect- a Cinnadenia) paniculate 3a. Flowers so on towards the recog- 2a. Flowers unisexual, arranged in umbels, these single in the axils of leaves and still monophyletic. are genera and groups are classification based practical key if the genera or likely way of practical of Lauraceae species Dehaasia is very similar Dehaasia Alseodaphne, KEY TO make want develop to will monophyletic and poly- forming monophyletic and Phoebe; because the fruit difference between the helped Species stamens genera of recognition generally accepted two instance Kochummen, Alseodaphne. Vegetatively, in to try account differences in flower or fruit between the only to rec- between these genera, but the information needed for this is is Persea case them both in my the Because taxa. be Van der Werff & Richter, the are species hand. publications (for in of can Caryodaphnopsis, monophyletic than those who prefer to recognize monophyletic ing cases between seems nition of genera without result in that genera require and I have therefore tried relationships is example within One of the main aims of species. relationships express ordening and distinct wood anatomy. examples (see likely generic circumscriptions incompletely known, two This and other genera. Those who have different 131 have 9 4-locular stamens, but sometimes 9 2-locular stamens. generic paraphyletic In genera and allows the recognizable indicate that androecial characters vary 1996) Lauraceae of genera bisexual), and fruit type. A classification based or opposite leaves, strongly unequal tepals or even to to the key other characters than those listed above. An of that genus their annotated such, it is practical. Some genera, however, groups. As recognizable ognized by is der Werff: An type, flowers (unisexual cence on in van 1. rarely subopposite or opposite Actinodaphne 6 132 BLUMEA Vol. No. 46, 1, 2001 6a. Anthers 4-locular 16. Litsea 7 b. Anthers 2-locular 7a. Umbels 1 -flowered; leaves subopposite or 14. alternate b. Umbels with several flowers; leaves alternate Iteadaphne 15. Lindera 8a. Anthers 2-locular 9 b. Anthers 4-locular 17 9a. Fertile stamens 3 b. Fertile stamens 6 10a. Tepals equal 10 9 or the or 11 three outer smaller than the inner three slightly ll.Endiandra b. Tepals the unequal, flowers apparently 1 la. Fertile stamens 6 b. Fertile stamens 9 12a. Tepals longer outer with three much only three larger and covering the inner three, thus 22. tepals 12 13 than stamens, included in the flower; anther cells lateral, stamens 4. Brassiodendron slit-like b. shorter than Tepals Triadodaphne stamens stamens, exserted; anther cells extrorse, 13. roundish 13a. Leaves opposite b. Leaves alternate 14a. Outer or subopposite or clustered tepals clearly smaller small, Hexapora 8. Cinnamomum 14 half the (about size) than the inner ones 10. Dehaasia b. Tepals equal or 15a. Flowers with a nearly 15 so tubular deep, receptacle, this enclosing the ovary and later the fruit 9. b. Flowers with quently a shallow with small, persistent tepals 16a. Lateral flowers of the ultimate light-colored and the fruit receptacle, contrasting at cymes same stamens 3 b. Fertile stamens 9 opposite, b. Leaves alternate, 19a. b. the with the much darker usually ... twigs and petioles 12. Eusideroxylon 18 tri- or 19 tripliveined pinnately or triveined 20 8. Cinnamomum Tepals unequal, the pinnately 21a. Inflorescences b. Inflorescences outer tepals much less than half the inner ones Caryodaphnopsis 17. Neocinnamomum veined racemose 21 or nearly 21. so the outer ones about half as long as the inner ones 2. or nearly so Potoxylon 22 paniculate Tepals unequal, Tepals equal twigs usually 3. Beilschmiedia 20a. Leaves triveined b. infre- 10. Dehaasia petioles subopposite; 5. 22a. or 16 Tepals equal b. Leaves Cryptocarya pedicel bark of the color 17a. Fertile 18a. Leaves on the base strictly opposite; b. Lateral flowers of the ultimate cymes about the unprotected Alseodaphne 23 H. 23a. van der deciduous Tepals Werff: An annotated persisting or as key the to small scales at Lauraceae of genera 133 the base of the fruit 2. b. 24a. b. 1. Tepals persisting Tepals clasping Inflorescences 19. Persea Nees paniculate umbellate with clusters of bract or stamens scars at the very base. 9, each with 4 locules, tepals equal. Fruit seated small, flat cupule. Leaves whorled, usually pinnately veined, sometimes glaucous below. Terminal buds immediately perulate terminal buds. whorled leaves, but differ from having clusters of usually perulate; above the whorls of leaves. Best whorled leaves, and in 20. Phoebe reflexed in fruit to Flowers triinerous, unisexual, on a 24 the base of the fruit Tepals spreading Actinodaphne Alseodaphne fruiting stage, conspicuous in the umbels generally the inflorescence. About 100 present scars its recognized by Some on twigs rather large, of Litsea may also have species with umbellate inflorescences Actinodaphne species stalked and in the absence of bract Asia with species, tropical the older usually few a scars at the base of in southern species China. 2. Alseodaphne (including Nothaphoebe) Nees Inflorescences of type 2. Flowers trimerous, bisexual, tepals unequal and to Fruit seated equal. brightly colored, or small unprotected tepals persisting in several this light-colored bark, About 90 with the darker Various authors have tried to separate at pedicels thick, fleshy vs. in fruit tepals vs. equal vs. unequal most a club-shaped; Bentham and or (1880) qualitative, and Kostermans subgenera opinion and a tepals Tropical one was was as a distinct American vs. fleshy and only have deciduous tepals tepals persistent vs. quantitative (tepals subequal tepals and and the intensively as Nothaphoebe Alseodaphne in his Kostermans Nothaphoebe more and Alseodaphne key Nothaphoebe valid genera. Even than Nothaphoebe. has been separating Alseodaphne as the type published, and as in Persea (1973a) changed anybody else, I found in BO annotated in 1960 as the type of Nothaphoebe names stami- vs. tepals equal; fruiting ones are genus (however, genera). annotated in 1992 Kosterm. Neither of these the difficulties of not species treated clear concept of Alseodaphne and Kosterm., the other perulate. vague). (1957) recognized Alseodaphne of Clemens 29777; unequal the other but several Kostermans studied Asian Lauraceae developed twigs, Characters Alseodaphne. of Persea; Hutchinson (1964) included recognized Nothaphoebe included among the not conspicuous, cordate-sagittate in fruit. Of these characters, distinction becomes therefore rather sections plate-like a of the conspicuous, Terminalbuds from Nothaphoebe qualitative difference; looks tips often with Twigs petioles. fruiting pedicels cylindrical, persistent deciduous in fruit is this sometimes swollen Asia. used for this include staminodia of whorl IV nodia much smaller, pedicel, the base of the fruit; at below. species glaucous contrasting species, tropical 9, each with 4 locules, stamens the Leaves clustered near the cupule rarely present (A. insignis Gamble). pinnately veined, on of Alseodaphne though he two is his never sheets clemensiae kinabaluensis but these annotations underscore Nothaphoebe. It seems therefore best BLUMEA 134 to include needed to in Nothaphoebe Vol. and would placed Asia. Good is needed for their with in fruit of all study very similar had but towards s.l. tropical three undescribed species vegetative characters clustered leaves which clearly point pointing Alseodaphne to been collected in they species (apparently description, involved is species several genera. It is worth are Alseodaphne pale bark, dark, long petioles, large, and thickened pedicels a one or Neotropics material of these flowering involved) are be described in certainly in 2001 1, Alseodaphne although determine if all shouldbe that several collections from the out No. 46, are (thick twigs glaucous below) Alseodaphne. 3. Beilschmiedia Nees Inflorescence tepals equal, seated I have paniculate (type 3). erect unprotected not seen About 250 This Flowers trimerous, bisexual, stamens 9, 2-locular; anthesis, deciduous in older flowers. Fruit without at the on pedicel. with species Leaves alternate glaucous or cupule, a veined. subopposite, pinnately lower leaf surfaces. Terminal buds perulate. not species, pantropical. confused with has been genus with 9 2-locular stamens. The best Dehaasia, which has also bisexual flowers difference between the is the inflorescence two type; type 3 in Beilschmiedia, type 2 in Dehaasia. Dehaasia species have also frequent- ly clustered leaves and the note with twigs under Endiandra. A persistent (but small) tepals placed in distinct genus, a contrasting species placed the base of the fruit; such at in New Guinea. Whether these be bark and pale few Australian species requires dark species can be expected best included in Beilschmiedia are further See also petioles. in Beilschmiedia have also should or study. 4. Brassiodendron C.K. Allen Inflorescence stamens thesis. Fruit without a 6, 2-locular, tepals equal, half-erect seated cupule, lower leaf surface pinnately veined, all from Australia with These reduction sometimes appearing paniculate (type 3), by trimerous, bisexual, do species not one not species form unprotected a Brassiodendron a closely 5. related as to key which is difficult Airy cupule, unprotected group and to use sole on the one pedicel. Caryodaphnopsis species is strongly unequal tepals. having (type 2). outer below. Terminal buds but differ in species, Hyland (1989) placed them and for practical species reasons occurring I recognize in New Guinea is Shaw paniculate-cymose tropical America, 6 Endiandra. 4-locular; tepals unequal, the glaucous perulate. solutionmay well express theirrelationships distinct genus. The Caryodaphnopsis Inflorescence not at an- Leaves alternate, pedicel. Terminalbuds glaucous. in Endiandraand Beilschmiedia. Hyland's better, but results in the Flowers spreading also in New Guinea. homogeneous a on racemose. or easily not Flowers trimerous, bisexual, 3 about 1/5 Leaves area recognized long opposite, perulate. in the FM as by as tri- About 15 flowers with equal tepals. or tripliveined, species in (Philippines, Borneo, its opposite Cinnamomumspecies often have stamens 9, the inner 3. Fruit without more or tropical New a less Asia and Guinea). leaves and flowers with opposite, tripliveined leaves, H. 6. Inflorescence the der Werff: An annotated key the to genera Flowers bisexual, trimerous, a raceme. outer in Australia, This herbaceous, perulate. not enlarged About 20 un- floral species, species pantropical. one parasitic vine be confused with any other genus of Laura- cannot sometimes it is confused with Cuscuta ceae; 135 9, 2-locular; tepals stamens 3 much smaller than the inner 3. Fruit enclosed in the tube. Leaves alternate, minute, scale-like. Terminal buds mostly of Lauraceae L. Cassytha equal, van (Convolvulaceae). 7. Cinnadenia Kosterm. Inflorescence a plate-like cupule. rarely collected The with 9 Leaves alternate, genus of two which differs in (18-32 stamens type species having as having in C. C. as a a seen an synonym flowers. malayana stamens isotype are malayana and found it seen were smaller towards the have 4 center Kochummen (1989) dimerous, but Kostermans suggesting genus; species (and to A Kostermans (1973b) described the that the flowers which Kostermans (Hook.f.) Kosterm., known poorly a small, inflorescence is Actino- Long (1984). liyuyingii H. Liu, of Cinnadenia paniculata Cinnadenia is on a perulate. not terminal buds, and flowers 1 have not are (1973b) and this has dimaterial of seen thus the concept of Cinnadenia in Kochummen's (1989) observations. Dr. J. Rohwer on flower of C. progressively did he had paniculate a in whorls of 3, of Litsea (see discussion), Malaysia. perulate malayana Kosterm.). and base my view of this malayana this article) in Peninsular as dimerous veined. Terminal buds whorled leaves, trimerous flowers, (1977) mentioned that the trimerous. I have merous one or 4-locular. Fruit seated (9-32), pinnately species, mentioned that the flowers of C. treated variable stamens other genus with unisexual flowers and only daphne, Flowers unisexual, trimerous panicle. number of tepals equal, tepals and 28 (pers. comm.) stamens checked which became of the flower. 8. Cinnamomum Schaeff. Inflorescences paniculate-cymose (type 2). 4-locular; tepals equal, half-erect or without persistent tepals tripliveined About 350 or Species nately Jaya alternate and species, mostly Oceania and or tropical with in tepals partly persistent. pinnately veined; rarely and tropical tripliveined leaves perulate usually lack buds. A few most pinnately veined leaves and tepals equal, small but also stamens cupule, Leaves either perulate buds, species 9, this with opposite and tripliveined. in Australia, present species Philippines with pin- and Irian 1986) instead of 4-locular anthers. The Cinnamomum species in the FM perulate while from the leaves combined with flowers with fication of Cryptocarya a leaves alternate and subtropical Asia, have 9 2-locular anthers (Kostermans, Inflorescences Flowers trimerous, bisexual, anthesis. Fruit seated in America. veined leaves have opposite, tripliveined 9. the at equal tepals area; one or two allow species easy identi- have alternate, terminal buds. R.Br. paniculate (type 3). half-erect to Flowers trimerous, spreading at anthesis. Fruit bisexual, entirely stamens 9, 2-locular; enclosed in the enlarged 136 BLUMEA floral tube, at the with small tip ribs. Leaves alternate, perulate. Several species have among Lauraceae, but this are not rare. species in A monotypic in few a somewhat also leaf uncommon not shape Glaucous lower leaf surfaces with the species, pantropical, their 9 genera, 2-locular Eusideroxylon and stamens and floral tube deep of majority larger (more than 5 distinctive. are have also fruits Potoxylon, floral tubes, but both have much enlarged an longitudinal Terminal buds species tripliveined. in Litsea. occur with frequently oblong leaves, than 300 more 2001 1, Asia. tropical The flowers with Two in or might genus of large No. 46, from the fallen tepals, scars pinnately Vol. fruits. long) cm enclosed fully 10. Dehaasia Blume Inflorescences deciduous or but persistent, sometimes with very small, species not trimerous, bisexual, 3 much smaller than the inner outer in fruit. Fruit enlarged, persistent tepals sometimes whitish bark, which species). Terminal buds Dehaasia is closely related to and without assigning stamens a the it is specimen pedicel, in some a pale to in Alseodaphne Asia. in only indistinguishable either of these to have twigs case from which it differs Vegetatively, the equal, of the branches, tips is also the species, tropical Alseodaphne, 2-locular instead of 4-locular anthers. on 9, stamens almost to fruiting pedicel species some petioles (this About 35 perulate. not below. In glaucous with the dark contrasts 3) unprotected the base, the at swollen and colored. Leaves alternate, clustered near the pinnately veined, daphne Flowers paniculate-cymose (type 2). 2-locular; tepals unequal (the having from Alseois genera guess. a 11. Endiandra R.Br. Inflorescences paniculate (type 3). Flowers trimerous, bisexual, tepals equal, rarely subequal (with ones), half-erect to spreading at the tepals slightly outer anthesis. Fruit unprotected deciduous in fruit. Leaves alternate, pinnately veined, buds the not greatly enlarged About 100 area can 12. only two be not disc not the of the surrounding the to the the pedicel, glaucous the base 3, 2-locular; below. Terminal stamens stamens erect at with small the tepals outer either genus anthesis. Fruit Vegetatively cences entirely from fallen below. Terminal buds not tepals. perulate. very similar and flowers with pistil. to with much large larger stamens: than the inner Triado- ones, and staminodia (representing whorl Beilschmiedia, and fruiting specimens certainty. & Binn. laxly paniculate (type 3). scars become and Flowers trimerous, bisexual, stamens locular; staminodes representing whorl I and II well developed; tepals equal, less inner tepals Australia and the Pacific. Endiandra is similar Teijsm. at on other genera have flowers with 3 fertile having assigned Eusideroxylon Inflorescence a which has 4-locular stamens and II). Vegetatively often glands united, forming are which differs in Eusideroxylon, I and and species some species, tropical Asia, In the FM daphne In perulate. stamens smaller than to only enclosed in the accrescent Leaves alternate, Wood very hard. 1 Potoxylon, three fertile species, but differs in its stamens. floral tube, pinnately veined, not 3, 4- more or at the tip glaucous Borneo and Sumatra. laxly branched inflores- H. 13. der Werff: An van annotated to the key of Lauraceae genera 137 Hexapora Hook.f. Inflorescence paniculate (type 3) 6,2-locular; tepals equal, tepals and stamens at erect at the base clustered, pinnately veined, typic genus, rarely (Kochummen, recognized whorl I and II and by Flowers trimerous, bisexual; known from only its 6 2-locular by the 1989). stamens. Fruit with Leaves alternate below. Terminal buds glaucous not collected and is best Hexapora senting or racemose. anthesis, shorter than the somewhat or with round locules repre- locules, which is very unusual for extrorse 14. long, slit-like anther cells. Neither tepals Inflorescence umbellate, the umbels 1-flowered, shortshoot. Flowers trimerous, unisexual; Fruit not seen, said genus be seated to opposite, pinnately veined, typic not glaucous from Java and Sumatra, S Chinaand NE India has also been This species has also on a few related related to I. caudata and that the buds of L. I expect that umbels I. to confusa, lated to a more thorough study opposite or A perulate. subsub- mono- differing level. A generic on in its species stamens along leafless short- leaves. It is likely closely young, bracteate inflorescences. are find that from [I. caudata (Nees) H.W. Li]. tripliveined or the reduction of multi-flowered 1-flowered umbels in Lindera has taken with its or or Kosterm. (Kostermans, 1992) is spicata will not Lindera, but to in Iteadaphne spicata opposite 1981). recognition placed slender, leafless a Leaves cupule. 1-floweredumbels with 2-celled described Lindera recently together along below. Terminal buds shoots, but differs in having alternate, strongly trithat the persist lateral are in fruit. 6-9, 2-locular, tepals equal small (Richter, certain Iteadaphne deserves am not a stamens closely 1-flowered umbels and wood anatomy I nor stamens stamens Blume Iteadaphne equal. mono- Malaysia. of whorl I and II. Brassiodendron has also 6 2-locular stamens, but these and have A perulate. Peninsular Penang, stamens not stamens persistent place subopposite, pinnately more than veined leaves is I. caudata and L. spicata,, which have alternate, tri- or once and that closely not re- leaves. tripliveined 15. Lindera Thunb. Inflorescence umbellate, umbels with several flowers, of leaves. Flowers nearly so. trimerous, unisexual; Fruit with or without a small stamens 9 along or more, a shortshoot Leaves alternate, cupule. pinnately veined, the lower surface sometimes glaucous. Terminal buds perulate of about 100 in species, mostly tropical and temperate N America; relatively few species Lindera differs from Litsea specimens or specimens with only subtropical in the FM in axils or 2-locular, tepals equal Asia with a or not. few or A genus species flowers are in area. in its 2-locular (not 4-locular) anthers and pistillate or tripli- difficult to determine to fruiting genus. 16. Litsea Lam. Inflorescence umbellate, umbels with several flowers, leaves. Flowers lacking. trimerous, Fruit with a small unisexual; to rather stamens along shortshoots or in axils of usually 9, 4-locular, tepals equal large cupule. Leaves alternate or or rarely opposite, 138 BLUMEA pinnately veined, more genus, sometimes than 300 Vol. glaucous in species, mostly No. 46, 1, 2001 below. Terminal buds perulate A not. or large tropical Asia, also present in Oceania, Australia and in N and C America. Litsea is characterized 4-locular anthers. It is area, the inflorescences bearing shortshoot genus. leaves in the FM tripliveined shoots genus in the FM variation in the morphological or its umbellate inflorescences, trimerous flowers by large a illogical) in some can species of umbels; however, there is raceme 'raceme' is twig bearing a Although I would be and area I have be not not always may expect if they 10 cm, and to and of great deal with clustered exist. The short- making terminal bud vegetative a the term shoot looks much like umbel-bearing a species seen surprised quite long (up and the one indicating a the umbels. 17. Neocinnamomum H. Liu Inflorescence or paniculate-cymose, inflorescence reduced to a 4-locular, tepals equal, half-erect species not perulate. anthesis. Fruit strongly A genus of 6 tri- species or in on an stamens 9, obconical, swollen pedicel, not tripliveined, tropical Asia, below. glaucous with 1 poorly known and the swollen pedicel from Sumatra. The fascicled flowers, triwith fascicle. Flowers trimerous, bisexual; at Leaves alternate, tepals persistent. Terminal buds the cymes often sessile and the flowers in fascicles single persistent tepals or tripliveined, in fruit make alternate leaves, rather easy. recognition 18. Neolitsea Merr. Inflorescence umbellate, umbels with several flowers, along shortshoot. Flowers dimerous, unisexual; a 4, spreading at anthesis. Fruit seated initially persisting, and sometimes tropical Asia, but has to extending leaves cupules. a off. Leaves generally clustered, tripliveined perulate. About 100 species, its dimerous flowers characters vegetative although help separate the along In sheath a at species of Neolitsea the bracts species in genera: Neolitsea usually veined leaves and stalked umbels, these shortshoots. Some Litsea few mostly in other characters it two leaves and sessile umbels, these the leafless parts of the innovations, while Litsea has or together Australia. recognized by Litsea. Some opposite), pinnately as to several close or 6, 4-locular; tepals equal, small, plate-like cupule, the tepals sometimes below. Terminal buds usually clustered, tripliveined along on a ultimately falling glaucous Neolitsea is best is close but single stamens borne alternate (rarely usually also have rather surrounding frequently in the axils of large, cup-shaped the terminalbuds persist the base of the leafless internodes. 19. Persea Mill Inflorescence paniculate-cymose 4-locular; tepals equal of the fruit, spreading or to (type 2). subequal, reflexed, a cupule pinnately veined, sometimes glaucous genus with regions c. 200 species of the Northern in tropical Flowers trimerous, half-erect at anthesis. Tepals persistent absent. Leaves alternate or below. Terminal buds Asia and America with Hemisphere. bisexual; a at 9, the base slightly clustered, perulate. few stamens A poorly defined species in subtropical H. The van der Werff: An above description given annotated applies only Asian Persea and Phoebe and these species characters. In Asian Persea species in fruit, in Phoebe persistent are the type P. americana species, tepals neither tepals (the and partly spreading Mill.), der Werff from van in fruit. A equal tepals others have Surinam) comprehensive study tablishment of clear spreading to on fruit reflexed the fruit. Difficulties with pressed against has deciduous in fruit in as Phoebe), strongly unequal tepals unequal or in fruit while 3) persistent (including in fruit (but equal tepals persistent the fruit pressed against nor and persistent due to the great variation found in American 3 much smaller than the inner outer (P. julianae are sometimes are usually separated are groups are which species, 139 found floral differences between not two tepals in Persea. Some of these have species placed uous they delimitationin this group generic the of Lauraceae genera the Asian to in the segregate genus Machilus. I have placed the to the key species one which decid- are of Persea s.l. and Phoebe is needed for the es- boundaries. generic 20. Phoebe Nees Inflorescence paniculate-cymose 4-locular; tepals equal and Tropical Asia, discussed clasping better quite 21. c. sometimes common Phoebe in the in the FM racemose or pinnately veined, in can be 9, slightly clustered, usually not from Persea separated It fruiting stage. than Persea, area with not a that is my impression perulate. only by lack the that Phoebe is with obovate leaves species species generally few flower clusters stamens glaucous vegetatively perulate are buds. near the base of the inflorescence. 9, 4-locular; tepals equal, accrescent more or less erect at floral tube. Leaves alternate below. Terminal buds not A perulate. monotypic very stamens similar to but the flowers Eusideroxylon, and the inflorescence is basically are racemose much (laxly Eusideroxylon). Triadodaphne Inflorescence Kosterm. paniculate (type 3). tepals strongly unequal, inner 3, more or veined, not only or stamens Tepals persistent from Borneo. and with 9 fertile paniculate 22. only is Potoxylon larger anthesis. absent. Leaves alternate anthesis. Fruit enclosed in and fused with the genus known erect at Kosterm. Flowers trimerous, bisexual; and less below. Terminal buds in Phoebe and that Phoebe Potoxylon Inflorescence cupule glaucous under Persea, represented a Flowers trimerous, bisexual; more or species. 100 spreading tepals vs. (type 2). subequal, thebase of the fruit, clasping pinnately veined, As or less the erect at glaucous outer Flowers 3 much anthesis. Fruit trimerous, bisexual; larger not below. Terminal buds and almost stamens 3, 2-locular; completely covering known. Leaves alternate and not perulate. A monotypic the pinnately genus known from Borneo. Triadodaphne nize it based on 3 and the much was its included by Rohwer strongly unequal tepals deeper receptacle. (1993 ) in Endiandra, but I with the Kostermans outer (1993) 3 much prefer larger to recog- than the inner included 2 additional species 140 in BLUMEA had the that No. 46, tepals outer species 1, of T. saw greatly so) initially placed. was I specimen shorter than the inner ones (but not in Endiandra, where it 2001 Kosterm. from New Guinea and T. inaequitepala (Kosterm.) Kosterm. from the Solomon Islands. The tepala T. T. Triadodaphne, —Vol. I have pachy- inaequitepala and I would include not seen material of pachytepala. ACKNOWLEDGEMENTS This article is based pleasant 1 and 2 largely observations made on thanks to the Director express my and one. I also thank Peter productive in previously published were during and Staff members Stevens for the Annals 4-week a who at BO and stay did their I would like best to make very to visit my a critically reading the manuscript. 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