f KEY TO THE GENERA OF A Henk van der Werff^ LAURACEAE THE IN NEW WORLDS Abstract The A key to the twenty-nine genera of Lauraceae naturally occurring in the western hemisphere is presented. A key based on and vegetative characters only. brief statement on distribution and size of the genera is floral is A key provided, together with a note on cupule shape. Spanish translation of the is included. Lauraceae have, general, the reputation of both floral and fruit characters (Koslermans, 1957; in being After having studied the Hutchinson, 1964), or share a combination of these difficult to identify. neotropical members of this family for the last six problems. Keys that are based on floral and fruit years, agree that the reputation well characters create special problems. As a rule, both I fully is deserved. There are various reasons for this. In floral and fruit characters are necessary for generic the place, many collections are sterile or fruit- identification, but a single specimen rarely has both first need ing and lack the floral characters needed for iden- flowers and fruits. Typically, fruits several To date, there are no keys for the generic months for development, by which time the flowers tification. from an identification of nonflowering specimens, and have long fallen Rarely, fruits earlier it is off". enough unlikely such a key will become available in the flowering season persist long to be collected may an such specimens be recognized to with fresh flowers, but that exception. In future. Sterile is same and do occur on species by a botanist familiar with the family, but cases, flowers fruits not the such instantaneous recognition does not occur, inflorescence. Occasionally, one finds flowers and if my specimen remain even on the same inflorescence. In expe- the likely to unidentified ''fruits'' is and genus and therefore inaccessible for a long time. rience, such "fruits'* are usually diseased de- to Fruiting specimens stand a better chance of being formed. Thus, because flowering specimens almost never have key requires both flowers because the cupule (or absence) con- fruits, a that identified its when and not workable and frustrate user. useful information, especially old sta- fruits will its tains is am mens remain on the cupule. However, not However, possible to construct a generic key I it is able to construct a workable key based on fruit for neotropical Lauraceae based solely on floral and vegetative characters. From a taxonomic point and vegetative characters. I present such a key American specimens even though below, which includes genera. of view, fruiting are valuable, all whereas specimens are The most important floral characters used in difficult to identify, sterile number of or no use. For botanists making forest this key, as well as in earlier keys, are of little stamens, number of anther per stamen, sexual inventories or studying species diversity, sterile cells voucher specimens are useful, hence collected, and condition of the flowers, and tepals equal or un- end up on the desk of a specialist, where they often equal. These characters, although readily visible, and remain are often variable within a genus, this variation unidentified. Even with good flowering material at hand, iden- has rarely been incorporated in earlier keys. For my tification to genus often difficult. In opinion, example, Persea is usually presented as having is main reason problem no workable nine 4-celled stamens, whereas the neotropical spe- the for this that is keys The published keys are partly obsolete cies can have nine or six stamens, these being all exist. some and some (such as Mez, 1889), are of regional scope (Mac- 4-celled, all 2-celled, or 4- 2-celled. 1938; Kostermans, 1936; 1945), use Moreover, the tepals can be equal or unequal and Bride, Allen, W. A. Gentry, M. Grayum, Rohwer, and D. Stevens critically read the manuscript and suggested improvement ' J. W. Myers made D. Stevens kindly translated the key into Spanish. the illustrations. J. Missouri Botanical Garden, P.O. Box 299, Louis, Missouri 63166-0299, U.S.A. ' St. Ann. Missouri Box. Gard. 78: 377-387. 1991 378 Annals the of Garden Missouri Botanical deciduous or persistent in fruit. In order to accom- and II 4-celled, III 2-celled or sterile; whorls and I niodate these permutations of characters, at II 2-celled, whorl III sterile), but Endiicheria it is limes necessary to have a genus appear several anomnla has whorls T and II 2-celled and whorl A times in the key III 4-ceIIed. further variation that any of is While constructing key, have completely whorls and be (In some species this I I, II, III sterile. among ignored any phylogenetic relationships the only whorl fertile, or only and II are fertile, I is I genera, mainly because phylogenetic schemes of or and are or only II III fertile, III fertile; is I Lauraceae are partially based on fruit characters have not yet seen a species with only whorl II (Kostermans, 1957) or are based on wood and Table presents the various androecial fertile.) 1 among bark anatomy (Richter, 1981). These supposedly combinations and their distributions neo- phylogenetic classifications (which are quite differ- tropical Lauraceae, ''^Gamanthera'' excluded. If a known ent from each other) cannot readily be followed in key were to include combinations of all number number the construction of a key based on characters. of stamens, their position and of floral my Instead, aim to provide a workable key for anther cells in each whorl, one would end up with is How- the identification of flowering Lauraceae. a very long and tedious (but workable) key. Before presenting the key, I will briefly describe ever, I found that a shorter and more practical made the flower structure of Lauraceae. Examples are key can be by excluding the condition of the Mez based on American genera, and unusual conditions stamens of whorl as has been done by III, first of palaeotropical genera (such as dimerous flowers (1889). Exceptional species or groups of species, Potameia) are not discussed. Typically, each such as the 2-celled species of Per sea and Carj- in flower has two whorls of three tepals each. The odaphnopsis and the Aiouea species with three whorls are usually equal in size and shape, but stamens are keyed out separately. there are exceptions. Tepals are unequal Following key, the distribution, approximate in all tlie Caryodaphnopsis^ many number and species of in species of of species fruit type are briefly de- When Persea (Fig. Ic), in Anaueria, and in a few species scribed. an unusual leaf position occurs in When of Anlba and Licaria. tepals are unequal, a genus, this also mentioned. is A the outer three are smaller, except in fAcaria, good magnifying glass, or preferably a dis- where the outer three tepals can be larger (Fig. secting microscope, a necessity for identifying is unusual having Lauraceae. Id). *'*'Chlorocardiu!n'' in the is tepals in whorls of four or irregular. The stamens Included in the key are several taxa which are are arranged in four whorls of three each (with the not yet published {''''Chlorocardium,^^ ^^Gaman- exception of ''''Chlororardium,'''' which has stamens thera,^^ and ^^Paraia''), Their inclusion in the key in whorls of four) and are opposite the tepals. The absolutely not intended as their publication, but is whorls are counted from the outside to the center done to prevent the key from being obsolete at is Whorl moment of the flower. opposite the outer tepals, the of publication. I is whorl II is facing the inner tepals, whorl III is opposite whorl and whorl IV opposite whorl I, II. W and the exception of Litsea ith ''''Cklorocar- Key New World to Genera of Lauraceae diuni^^^ whorl IV always staminodial or lacking. is Cassylha Parasitic vine leafless 1 The stamens of whorl nearly always have two III Shrubs or trees with green leaves 2 1 globose glands at their base, the exceptions being 2(1) '^Stamen" by complete fusion of 3 sta- 1, Mezilanrus and Williamodendron which mens these in ''''Gamanthera'''' , 2 Stamens more than 3 glands are absent. In Phyllostemonodaphne and 1 Flowers 4 3(2) bisexual or staniinate Urbanodendron stamens have glands at their all 42 3 Flowers pistillate base. In Pleiirothyriiim the glands are strongly may enlarged and become completely fused, sur- 4 Stamens more than 3 8 rounding stamens. In general, the stamens of 5(4) Stamens with 4 anther cells W^illiamode ndron all 5 Stamens with 2 anther 6 cells whorls and are (nearly) identical and seemingly II I 6(5) Leaves clustered near tips of branches; an- The form one whorl of stamens. exceptions are six ther cells extrorse; inflorescence a panicle, Dicypellium and Phyllostemonodaphne, where MezUaurus racemose ultimate divisions the stamens of whorl have become tepaloid and 6 Leaves not clustered^ but alternate or op- I with only whorl and Stamens posite; position of anther cells variable (in sterile, II III fertile. L. cannella leaves somewhat clustered, but of whorl occasionally from those of whorls III differ anther cells introrse); inflorescence panicu- and number of anther II in cells. If diff'erent, I cymose liform, ultimate division 7 whorl III has usually fewer anther cells (whorls I 7(6) Only stamens of Whorl fertile, the cells I 2 1 379 Volume Number 2 van der Werff 78, New Key World Lauraceae 1991 to N', >.'..' — — & Mez Figure Flowers A. Nectandra grandiflora Nees {Gentry et ai 5976.3); B. Ocotea atirrensis 1. of: — Donn. Smith {Grayum 6868);— C. Persea Uebmannii Mez (Lshiki 1614); D. Licaria bracteata van der Werff {Kunkel 56). lateral and large, or lateral basal and small; 9(8) Flowers unisexual 1 inflorescences and flowers glabrous Aioiiea 9 Flowers bisexual 1 7 Only stamens of Whorl III fertile, the cells 10(9) Leaves deciduous in winter; flowers appear- ± Lindera extrorse, introrse or apical, not lateral; ing before leaves ..„ inflorescences and/or flowers usually with 10 Plants evergreen, flowers and leaves present same Endlicheria amounts pubescence Licaria the time varying of at Outer 6 stamens each with 2 anther 9 11(9) Leaves opposite 8(4) cells „ 1 8 Outer 6 stamens each with 4 anther cells 23 11 Leaves alternate or clustered „ 15 m 49 380 Annals of the Garden Missouri Botanical Table Androecial configurations of American 1 Lau raceae. & Whorls Caryodaphnopsis pro I, II, III 4-celled: parte, C.innamomum Chlororardiuniy pro Nec- parte, Litscdy tandrriy Oroten, Pnraia, Persea pro parte, Pleuroth- Povcdodaph Rhodostemonodaph yriu ne, nc^ Sas- , safras, Umhcllularia, Urhanodendron pro parte & Whorls Aiouea pro Aniba I, II, III 2-celled: parte, pro parte, Bcilsihmiedia, Caryodaphnopsis pro parte, Cassythay Cryplocarya, Endlicheria pro parte, Ku- blizkia pro parte, Lindera, Persea pro parte, IJrban- odendron pro parte & Cinnamomum Whorls pro I II 4-ceIled, III 2-celled: parte, Kubitzkia pro parte, Persea pro parte f. & Whorls 11 4-celled, III 0-celled: Persea pro parte I & Whorls Endlicheria anomala I II 2-celled, III 4-celled; & Whorls Aiouea pro II 2-celled, III 0-celled: parte, I AnaueriGj Aniba pro parte, Caryodaphnopsis pro parte & Whorl Aiouea 2-celled, II III 0-celled: pro parte I & Whorl I 0-celled, II III 4-celled; Dicypelliuni & P Whorl hy mono- I 0-celled, II III 2.celled: Host e daphne & Wliorl Williamodendron II 0-ceIled, III 4-celled: I & Wliorl II 0-celled, III 2-celled: Licariay Mezilaurus I 12(11) Stamens 6, triangular, dark red, the fila- ments wider than the anthers, united at the hase; outer tepals about V^ as long as the Anaueria ones inner 12 Stamens 6 or 9, filaments free, narrower stamens never dark than the anthers; red; tepals equal or strongly unequal, outer ones >- "'^ <14 ones 13 the size of inner <V4 13(12) Outer tepals the size of inner tepals .... Caryodaphnopsis E .__ 3 Tepals equal 1 1 14(13) Floral tvihe deep, about as long as the tepals Figure 2. Stamens (whorl or II) of:— A. Ncctandra I Cryptocarya — ..„ & Mez (Kayap reticuhita (R. P.) 161); B. Nectandra much 14 Floral tube shallow, shorter than the — niembranacea (Sw.) Griseb. {Cuadros 2839); C. Ocotea Beilschmiedia — tepals Mez veraguensis (Meissner) (Stevens 6476); D. Ocotea Whorl stamens — 15(11) Filaments of of 111 united; pulchella Mart. (Goetzke 210); E. Rhodostenwno- Kubitzkia flowers red daphne kunthiana Rohwer (Nees) {Perry s.n.). 15 Filaments flower color variable, free; in- some 16 cluding red species in Stamens Whorl 16(15) of sterile, tepaloid; fertile I stamens 6, representijig Whorls II and 111 short, much shorter than tepals; Andean m Phyllostemonodaphne 1,000-2,500 Persea species, elev. 16 Stamens of Whorl fertile; fertile sta- 19 Staminodia (whorl IV) lacking; floral tube I mens 9 Whorl and Amazonian or 6; 6, representing well developed; lowland species if I 17 Aniba canelilla or A. parviflora II 17(16) Fertile stamens 9, each with 2 glands at the 20(18) Floral tube very shallow, scarcely visible at Urhanodendron Beilschmiedia base anthesis 17 stamens 6 or only those of Whorl 20 Floral tube well developed, about as long as Fertile 9, III with glands at their base 18 the tepals 21 18(17) Outer tepals ca. V2 as long as inner ones 21(20) Leaves with a fine, raised reticulation; floral ± tube deep and narrow, the tepals spread- (Fig. 1 c) 1 18 Tepals equal or nearly so (Fig. la and b) ing and flowers clearly wider than floral tube; .. 20 much filaments narrower than anthers; _ Cryptocarya 19(18) Staminodia (whorl IV) well developed, with staminodia well developed sagittate or cordate apices; floral tube very 21 Leaves with lax reticulation or this not ap- Volume Number 2 van der Werff 381 78, New 1991 Key World Lauraceae to parent; floral tube shallow or deep; if deep, 34 Stamens of Whorl I fertile, not tepaloid 35 „.„. tepals erect and flowers about as wide as 35(34) Outer tepals about Vi the size of the inner 36 tube; filaments frequently poorly ones floral dif- ferentiated from anthers; staniinodia present 35 Tepals equal or nearly so 37 or absent 22 36(35) Stamens of Whorls and II with a sterile, I _ 22(21) Flowers (minutely) tomentellous, tomentose triangular leaf apex rounded or obtuse; tip; or with appressed pubescence, but never leaves whorled; Guyanas and adjacent Brazil stamens and Venezuela Ocotea rubra pruinose; filaments of usually densely pubescent, poorly differentiated from an- 36 Stamens of Whorls and without a I II sterile, thers; leaves alternate or clustered Aniba triangular tip, the anther cells occupying the 22 Flowers usually glabrous, infrequently with entire anther; leaf apex various; leaves al- SE scattered sometimes pruinose; ternate or whorled; widespread, from hairs, fila- ments of stamens usually glabrous and U.S.A. to Brazil and Chile Per sea dif- ferentiated from anthers; leaves alternate 37(35) Staminal glands greatly enlarged, protruding .. Aiouea between the outer 6 stamens and sometimes 23(8) Flowers unisexual 24 fused into a large, pillowlike mass; stamens 23 Flowers bisexual 27 of Whorls and with at least 2 lateral I II 24(23) Young inflorescences enclosed decussate anther Pleurothyrium in cells bracts; flowers arranged in pseudo-umbels 37 Staminal glands not enlarged, not protruding Lit sea between outer stamens; free; outer stamens 24 Young 38 inflorescences not enclosed in de- with introrse cells cussate bracts; flowers arranged in panicles 38(37) Staminodia representing Whorl IV well de- or racemes 25 veloped, with a cordate or sagittate apex; 25(24) Deciduous trees; flowers appearing before filaments of stamens as long as anthers or or with young leaves, leaves often lobed longer 39 „„. Sassafras 38 Staminodia representing Whorl IV small or 25 Evergreen plants with mature leaves when lacking; present, mostly without a cordate if flowering; leaves never lobed 26 or sagittate filaments of stamens shorter tip; 26(25) Anther cells arranged in a low arch (Fig. than or as long as anthers (a few species of anthers poorly from Ocotea have large staminodia with a cordate 2e); diff*erentiated fil- Rhodostemonodaphnc aments but these have stamens with a very short tip, 26 Anther arranged 2 rows 2c and filament) 40 cefls in (Fig. d); anthers clearly diff"erentiated from 39(38) Leaves usually tripliveined, alternate, fre- much the narrower filament (rarely quently tufts of hairs present in the axils of fila- Cinnamomum ments very short) Ocotca the lowermost veins 27(23) Flowers arranged in pseudo-umbel, this, when 39 Leaves pinnately veined, alternate or clus- young, covered by bracts Umbellularia tered; axillary tufts of hairs lacking Persea 27 Flowers arranged in paniculiform inflores- 40(38) Leaves clustered; twigs with clusters of scars cences, these never enclosed by bracts 28 from fallen bracts; tepals roundish, hyaline, 29 28(27) Leaves opposite with a pubescent base, otherwise glabrous "Pa 28 Leaves alternate or whorled 31 ra a •ii f < 40 29(28) Outer tepals Vi the size of the inner ones; Leaves alternate; twigs without clusters of from leaves often strongly triveined scars faflen bracts; tepals not roundish, Caryodaphnopsis not hyaline, glabrous or with difi'erent dis- 29 Tepals equal; leaves pinnately veined 30 tribution of pubescence 41 30(29) Stamens 12 or more; leaves glabrous or 4 (40) Anther cells arranged in 2 vertical rows (Fig. 1 nearly so on lower surface ''''Chlorocardiuin^ 2c and stamens and inner face of tepals d); 30 Stamens leaves ferruginous tomentose or glabrous or variously pubescent, rarely pa- 9; tomentellous on lower surface piflose papillose, the anther cells clearly (if _ Nectandra oppositifoUa in 2 vertical rows); te[)als free at base, falling 31(28) Anther cells on the flat tip of columnar sta- individually in old flowers; a few old stamens mens Povedadaphne sometimes present on cupule of young fruits; 31 Anther cells not apical; stamens not colum- tepals at anthesis erect (Fig. lb) or spreading 32 Ocotea nar „ 32(31) All stamens with 2 glands at their base, the 41 Anther cells arranged in an arc (Fig. 2a and glands free Urbanodendron b); stamens and inner face of tepals papil- 32 Only stamens of Whorl III with glands (in lose; tepals united at the base, usually falling Pleurothyrium glands greatly enlarged and as a unit (together with stamens) in old flow- sometimes fused) 33 ers; stamens rarely present on cupule of Stamens Whorl young 33(32) of III with fused filaments; fruits; tepals at anthesis spreading flowers red Kubilzkla (Fig. la) Nectandra 33 Stamens of Whorl III with free filaments; if 42(3) Plants cold-season deciduous and/or inflo- filaments seemingly fused some Ocotea rescences subumbellate and young flowers (in 43 species), then flowers white to yellow, never enclosed in involucrate bracts 42 red 34 Plants evergreen and inflorescences panic- Stamens Whorl 34(33) of devoid of anther ulate, the flowers not enclosed involucrate I cells, in m tepaloid DicypeUiu bracts 45 2 46 1 382 Annals of the Garden Missouri Botanical 43(42) Inflorescences racemose or paniculate; leaves 12 Estambres 6 o 9, nunca rojo-obscuros, con mas delgados que anteras often trilobed Sassafras filamentos las y muy con 43 Inflorescences subunibeflate; leaves never libres; tepalos iguales o desiguales menos trilobed 44 los exteriores !4 o del largo de los 13 44(43) Plants deciduous; pseudo-umbels sessile or interiores menos nearly so Lindera 13(12) Tepalos exteriores 14 o del largo de Caryodaphnopsis 44 pseudo-um- Plants deciduous or evergreen; los interiores pedunculate Litsea 3 Tepalos iguales bels 1 _ 1 Tubo como 45(42) Staminodes straplike, the filaments about as 14(13) floral profundo, tan largo los Cryptocarya wide as the anther; traces of four on tepalos locelli mucho mas Rhoilostemonodaphne 4 Tubo poco profundo, corto each anther floral 1 Beilschmiedia 45 Staminodes club-shaped, the filaments nar- que los tepalos rower than the anthers 46 15(11) Estambres del Verticilo III con filamentos Kuhitzkia 46(45) Anthers with 4 remnants of leaves unidos; flores rojas locelli; Ocolea 5 Estambres con filamentos libres; flores rara- alternate _ 1 46 Anthers with 2 remnants of leaves mente rojas locelli; 1 whorled Endlichcria 16(15) Estambres del Verticilo esteriles, tepa- alternate or I estambres representando loides; fertiles 6, Phyllostenwnodaphne los Verticilos II y III 16 Estambres estambres Clave Para los Generos de del Verticilo I fertiles; 9 6 6 entonces representando Mundo fertiles 6, si Nuevo Lauraceae del 17 los Verticilos I y II Trepadoras parasitas y afilas CassyUia 17(16) Estambres fertiles 9, cada uno con 2 glan- 1 Urbanodendron Arbustos o arboles con hojas verdes 2 dulas en base 1 la 2(1) Estambre aparentemente 1 por fusion com- 7 Estambres fertiles 6 6 9, solo los del Verticilo 1 pleta de 3 estambres ^^Gamanthera'*^ con glandulas en base 18 III la Estambres claramente mas de 3 18(17) Tepalos exteriores cerca de del largo de 1 ^/z 3(2) Flores bisexuales o estaminadas 4 los interiores (Fig. Ic) 19 _ 42 3 Flores pistiladas 18 Tepalos iguales o casi iguales (Fig. la y b) 20 Estambres 3 5 4(3) _ mas 4 Estambres de 3 8 19(18) Estaminodios bien desarrollados, con apices muy 5(4) Anteras con 4 tecas Williamodendron sagitados o cordados; tubo floral corto, 5 Anteras con 2 tecas 6 mucho mas corto que los tepalos; especies msnm 6(5) Hojas agrupadas en los extremus de las ra- andinas de 1,000-2,500 Persea „. mas; tecas extrorsas; inflorescencia una pa- 19 Estaminodios ausentes; tubo bien de- floral nicula con las divisiones ultimas racemosas sarrollados; especies amazonicas de eleva- Mezilaurus ciones hajas Aniha ranelilla o A. parvijlora 6 Hojas alternas u opuestas, no agrupadas; 20(18) Tubo poco profundo, escasamente floral vis- orientacion de las tecas variable (en L. can- en antesis Beilschmiedia ible la mas menos nella hojas algo agrupadas, pero entonces 20 Tubo bien desarrollado, o floral mismo que 21 las tecas introrsas); inflorescencia paniculi- del largo los tepalos forme, divisiones ultimas cimosas 7 21(20) Hojas con una reticulacion fina promi- y 7(6) Solamente los estambres del Verticilo I fer- nente; tubo floral profundo y angosto, te- ± Umbo mas tiles, tecas laterales y grandes o lateral-ba- palos patentes y el claramente mucho mas sales petjuenas; inflorescencias y flores ancho que tubo; filamentos y el glabras Aiouea angostos que anteras; estaminodios bien las Solamente estambres Cryptocarya 7 los del V^erticilo III desarrollados tecas extrorsas, introrsas o inapica 21 Hojas con una reticulacion laxa o inconspi- fertiles, y/o nunca laterales; inflorescencias cua; tubo poco profundo o profundo, les, flo- floral res usualmente pubescentes Licaria profundo entonces los tepalos erectos si y 8(4) Anteras de los 6 estambres exleriores con limbo tan ancho como el tubo; filamentos el 2 tecas 9 frecuentemente poco diferenciados de las 8 Anteras de los 6 estambres exteriores con anteras; estaminodios presentes o ausentes 23 22 4 tecas menudamente 9(8) Flores unisexuales 10 22(21) Flores tornentosas o aplicado- 9 Flores Disexuaies pubescentes, nunca prulnosas; filamentos con hi 1 1 10(9) Hojas deciduas en invierno, flores produci- frecuencia densamente [)ubescentes, poco el das antes que las hojas Lindera diferenciados de las anteras; hojas alternas Aniba 10 Hojas no deciduas, hojas presentes o agrupadas flores y a vez Endlicheria 22 Flores usualmente glabras, infrecuente- la 11(9) Hojas opuestas mente con tricomas dispersas, a veces prui- 1 11 Hojas alternas o agrupadas 15 nosas; filamentos usualmente glabros y di- 12(11) Estambres rojo-obscuros, con filamentos ferenciados de anteras; hojas alternas 6, las mas anchos que anteras unidos en Aiouea las y la 24 base; tepalos exteriores cerca de V2 del largo 23(8) Flores unisexuales de los interiores Anaueria 23 Flores bisexuales 2 7 Volume Number 2 van der Werff 383 78, New 1991 Key World Lauraceae to 24(23) Inflorescencias jovenes envueltas en brac- pujando entre los 6 estambres exteriores y teas decusadas; flores en pseudo-umbelas a veces fusionadas en una masa grande como ... Litsea almohada; estambres de los Verticilos y II I 24 Inflorescencias jovenes no envueltas en con al menos 2 de las tecas laterales m bracteas decusadas; flores en paniculas o Plea rot hyriu racimos 25 37 Glandulas estaminales no agrandadas ni fu- 25(24) Arboles deciduos, flores producidas antes o sionadas; estambres exteriores con todas las junto con las hojas; hojas frecuentemente tecas introrsas 38 lobadas Sassafras 38(37) Estaminodios representando Verticilo IV el 25 Arboles o arbustos perennifolios, flores prod- bien desarroUados, con apices cordados o ucidas con las hojas maduras; hojas nunca sagitados; filamentos iguales o mas largos lobadas 26 que anteras 39 las „ 26(25) Tecas dispuestas en un arco bajo; anteras 38 Estaminodios representando Verticilo IV au- poco diferenciadas de los filamentos (Fig. sentes o presentes pequenos y mayor- si Rhodostemonodaphne 2e) mente sin apices cordados o sagitados; fi- .. 26 Tecas dispuestas en 2 hileras; anteras clara- lamentos iguales o mas cortos que las anteras mas mente anchas que los filamentos (rara- (unas pocas especies de Ocotea tienen es- mente filamentos niuy cortos) (Fig. 2c y d) taminodios grandes con apices cordados, pero muy Ocotea 40 estas tienen filamentos cortos) 27(23) Flores en pseudo-umbelas, estas cubiertas 39(38) Hojas usualmente trinervias, alternas, fre- por bracteas cuando jovenes Umbellularia cuentemente con fasciculos de tricomas pre- 27 Flores en inflorescencias paniculiformes, es- sentes en las axilas de los nervios basales ., tas nunca envueltas por bracteas 28 Cinnamornum 28(27) Hojas opuestas 29 39 Hojas pinnatinervias, alternas o agrupadas, 28 Hojas alternas o verticiladas 31 sin fasciculos de tricomas axilares Persea 29(28) Tepalos exteriores del tamafio de los in- 40(38) Hojas agrupadas; ramitas con fasciculos de !/4 mas teriores o pequenos; hojas con frecuen- cicatrices de bracteas caidas; tepalos algo cia fuertemente trinervias Caryodaphnopsis redondos, hialinos, pubescentes solamente 29 Tepalos iguales; hojas pinnatinervias 30 en base "Pa raia i» la 30(29) Estambres 12 o mas; hojas glabras o casi 40 Hojas alternas; ramitas sin fasciculos de ci- glabras en el enves ''''Chlorocardium'''* catrices; tepalos ni redondos ni hialinos, gla- 30 Estambres 9; hojas ferrugineo-tomentosas bros o con la pubescencia distribuida de otra en enves Nectandra oppositifolia forma 41 el 31(28) Tecas en el apice piano de los estambres 41(40) Tecas dispuestas en 2 hileras (Fig. 2c y d); columnares Povedadaphne estambres caras adaxiales de tepalos y los 31 Tecas no apicales, estambres no columnares glabros o variadamente pubescentes, rara- 32 mente papilosos papilosos, las tecas clara- (si ._ Todo 32(31) los estambres con 2 glandulas en la mente en 2 hileras); tepalos libres en la base base, las glandulas libres Urbanodendron y cayendose individualmente; unos pocos 32 Solamente los estambres del Verticilo III con estambres viejos frecuentemente presentes glandulas (en Pleurothyrium las glandulas en la cupula de los frutos jovenes; tepalos muy agrandadas y a veces fusionadas) 33 erectos o patentes en la antesis (Fig. 2b) ... 33(32) Estambres del Verticilo III con filamentos Ocotea fusionados; flores rojas Kubitzkia 41 Tecas dispuestas en un arco (Fig. 2a y b); 33 Estambres del Verticilo III con filamentos estambres y caras adaxiales de los tepalos o aparentemente fusionados (en papilosas; tepalos unidos apenas en base, libres, si la algunas especies de Ocotea)^ entonces usualmente cayendose juntos juntos con las (y flores blancas a amarillas, nunca rojas 34 los estambres); estambres raramente pre- 34(33) Estambres del Verticilo I esteriles y tepa- sentes en la cupula de los frutos jovenes; Dicypellium en loides tepalos patentes antesis (Fig. la) la 34 Estambres no Nectandra del Verticilo I fertiles y te- paloides 35 42(3) Plantas deciduas en invierno y/o el inflo- 35(34) Tepalos exteriores cerca de ^A del tamano rescencias subumbeladas y flores jovenes en- de los interiores 36 vueltas por bracteas involucrales 43 35 Tepalos iguales o casi iguales 37 42 Plantas perennifolias con inflorescencias 36(35) Estambres de los Verticilos y II con apices paniculadas, flores no envueltas por bracteas 1 triangulares y esteriles; hojas con apices re- involucrales 45 dondeados u obtusos; Guayanas y areas ad- 43(42) Inflorescencias racemosas o paniculadas; ho- yacentes de Brasil y Venezuela Ocotea rubra jas frecuentemente trilobadas Sassafras __. 36 43 Estambres de Verticilos I y II sin apices Inflorescencias subumbeladas; hojas nunca ocupando 44 triangulares y esteriles, las tecas trilobadas toda la antera; hojas raramente con apices 44(43) Plantas deciduas; pseudo-umbelas sesiles o redondeados u obtusos; ampliamente Lindera distri- casi sesiles 44 buida desde el sur de los Estados Unidos Plantas deciduas o perennifolias; pseudo- hasta Brasil Chile Persea umbelas pedunculadas Litsea y muy 37(35) Glandulas estaminales agrandadas, em- 45(42) Estaminodios ligulados, filamentos igual de 384 Annals of the Garden Missouri Botanical ancho que las anteras, cada antera con ves- ed on a naked pedicel. It occurs in lowland forests Rhodostcmonodaphne tigios de 4 tecas from Costa Rica to Peru and Brazil. The seven mas 45 Estaminodios claviformes, filamentos occur Indochina, with one palaeotropical species in 46 delgados que anteras las New Guinea (Kostermans, extending species to 46(45) Anteras de estaminodios con vesligios de los 1974). Ocotea 4 tecas; hojas alternas 46 Anteras de estaminodios con vestigios de los Cassytha L. 2 tecas; hojas alternas o verticiladas Endlicheria One from Mexico species the Neotropics, to in Aiouea often found near the coast, but also locally Aublet Brazil, common More Mauritia swamps. species occur in Probably a polyph,yl1etic genus in the currently c 4r u u .l t-u . . ,. andi South • Ihe genus has been Australia Africa. •^ in ,1 accepted sense, as has been noted by several1 au- • u ^ •. own r r ' : ipli aiceidi in its ta.mij-li y-be^cauuste ot its parasitic 1988: Rohwer van thors (Burner, et in press; al., ^ ^ habit, but hi ere retained in the Lauraceae. iS r^, der Werff, 1987b, 1988).V The genus ,has ,been ty monographed by Renner (1982) and consists of '^Chlorocardium about 20 species, raneine from southern Mexico n r j A . genus wi.t,h ^two species, one trom Lruyana and ^ to southern rB^razil1 andI Paraguay. MostIspIeciI.es have a a^^ ^^a S^^uri• nam, the ot.hu er ftrom Aa mazonian th?cuador and „ , , n the seated on a shallow cupule with a thickened fruit ^ ,. -- adjacent, Coli ombiia• , previ• ousli y i• nciludi edi i• n iJcot# ea, pedicel. wood, and but differing in characters of flowers^ The Ecuadorian/Colombian species Anaiieria Kostermans leaf position. incompletely known. Fruits are described as large is A known Amazonian monotyyp^ic^^genus from for- ^ /, -77 X 5r cm)\, wi-t^hu a normal1 or very 1lar„ge ^cup^ul1e^. (to and The near the border of Brazil Peru. seeds ests Cuinamomum A Schaeffer cupule are said to be edible after roasting. is lacking Flowers have green tcpals and dark in fruit. A poorly understood genus occurring in large, stamens, very unusual Lauraceae. red in Asia, Australia, and the Americas. Possibly with known Aniba Aublet more than 50 species in the Neotropics, .11 West from Mexico and the Indies to southern Brazil ™ Aniba Recently revised by Kubitzki (1982). in- ^^^, tP>araguay. tT.h! e r -. • suub»tendj ed1 iby a smal1l1 fruit •^ is •' 40-50 many eludes species, with pleasantly aro- . . i^^^^io ^ ^ cupule, t.hi. s of^^ten crowned, wi• t,h persi•st, e.nt tepals. ' 1. r matic yellow wood (rose.wo.od. 1 • disti1l1led1 from oil is ,,,. ^ c^ ^..^^a ;^ y Ihe neotropi.cal, speci. es ,have otten \been ,treated in ^ Aniba from Costa rosaeodora). distributed It is Cinnamomum. Phoebe, seem placed but better in Rica southern Brazil and Bolivia. Most species ,, r to Many A. si. an speci.es, i• ncl\ uad*ing„ .v,e^r.u,,n«i^, .w.n.\i.c\^nU ^ (.. occur the lowlands, but has also been collected in it cinnamon, have opposite leaves, but neo- yields all Andes up 1,800 m. The ,1s1eated in the to fruit is r ^-^^a r ' ^ . , speci.es .have al1.ternat.e, most*li y *tnpliveined tropical a rather deep cupule, which often lenticellate ^. in is u ^ ^ l, eaves. Cinnamomum, as accept* edj here, • possibuily is .' have or warty. Several species clustered leaves. polyphyletic. Beilschmiedia Nees Brown Cryptocarya R. In the Neotropics a poorly understood genus A poorly understood pantropical genus, large, 10-20 Kostermans by with species, last revised ^^^ ^^ fewer neotropical species, mostly in ^j^j^ (1938), who accepted 15 species. It has been col- southern Brazil and Chile, but also known from lected from Mexico to southern Brazil and in the French Guyana and adjacent Andean Ven- Brazil, West Ecuador, and P The almost ezuela, fruit is Heilschmiedia equally poorly un- laeotropics, is completely enclosed the cupule, which has only in many by more derstood, but represented species. ^ is small, apical pore. & Caryodaphnopsis Shaw DirypeUium Nees Martius Airy A Only recently reported from the Neotropics (van small genus of two rarely collected species. & der Werff Richter, 1985), this genus includes restricted lo Amazonian Brazil, reviewed by Roh- published neotropical species, with several more wer (1988). The cupule cupshaped, and has a five is The awaiting publication. The strongly unequal tepals double margin. tepals are persistent in fruit, One Dicypellium caryophyllaceum, has and opposite leaves are diagnostic. Fruits (in neo- species, tropical species) are round or pearshaped, and seat- been heavily exploited because of its aromatic bark. Volume Number 2 van der Werff 385 78. New 1991 Key World Lauraceae to Endlicheria Nees shaped cupule. Leaves of Litsea glaucescens are bay used as a spice, similar to leaves. A 40 neotropical genus of about species, re- ported from Costa Rica, the Lesser Antilles south Mezilaurus Taubert Paraguay, and southern The genus to Brazil. is A neotropical genus of 16 species occurring in (Rohwer The cupule polyphyletic et in press). al., Amazon the basin (including Bolivia, Peru, Colom- most have and variable; species a thick, fleshy is Venezuela, and the three Guyanas), revised bia, smooth cupule. Several species have whorled leaves. by van der Werff (1987a). The seated on fruit is Gamanthera^ The '''' a small, platelike cupule. flowers are very two genera similar to those of Licaria, but the A monotypic genus recently discovered Costa in differ in cupule shape, leaf position, inflorescence Rica, characterized by the presence of only one and wood anatomy. have type All species leaves "stamen," synandrium formed by com- actually a Two clustered at tips of branches. species with 4- stamens comm.). plete fusion of three (Endress, pers. were celled anthers recently transferred to Wil- The number synandrium can be of of the locelli 3, & liamodcadron (Kubitzki Richter, 1987). The or cupule rather deeply cupshaped. 2, 1. is margin and Nectaadra with a double persistent, reflexed te- Rol. ex Rottb. The monoecious. only species pals. is A large, neotropical genus of about 120 species, currently under revision by Rohwer. occurs J. It Kubitzkia van der Werff* from southern Florida and Mexico to Argentina. A genus neotropical of one, possibly two, species. Fruits are seated in a cupshaped (sometimes small) known from Guyana, Surinam, Venezuela, and cupule. Tepals are spreading at anthesis. Rohwer who Brazil, reviewed by (1988), accepts Systemonodaphne name Ocotea as the valid for this genus. Aublet The cupule cupshaped, with a double margin is The largest genus of Lauraceae in the Neotrop- and Arguments persistent, reflexed tepals. for re- 300 known from with at least species. also ics, It is name Systemonodaphne jecting the are discussed Madagascar and The genus very tropical Africa. is by van der Werff' (1986). and dumping ground variable serves as a for species accommodated that cannot be readily other gen- in Licaria Aublet era. Its distribution in the Neotropics from Mex- is A 40 neotropical genus of about species, revised ico and southern Florida to Argentina. Shape and by Kurz (1983). occurs from southern Florida of cupule variable and ranges from small It size is and Mexico to southern Brazil and BoUvia. The and platelike to cupshaped, sometimes with a dou- cupule cupshaped and has a double margin; ble margin or with persistent tepals. Tepals are is however, in some species the double margin is erect or spreading at anthesis. A scarcely noticeable. few species have opposite leaves. ''^Paraia'' A monotypic genus known from Amazonian Bra- Lindern Thunberg The cupule has a double margin with persistent zil. Lindera is represented by three species in east- tepals. Tepals are erect at anthesis. ern North America and an hundred additional in Persea The Miller on Asia. fruit seated a small, platelike cupule. is Two of the American species are rare; one was An incompletely understood genus with at least only recently described (Wofford, 1983). Twigs, 80 neotropical species, but better represented in common and American leaves, fruits of the species and one Canary Asia with species in the Islands. are used prepare a fragrant to tea. The fruit can be seated on a naked pedicel (as in Persea aniericana, the type species) or sub- is Lamarck Litsea tended by persistent, indurate tepals (as in the great New World A A In the Litsea represented by about majority of species). cupule never present. is is known five species, from the U.S.A., the mountains worldwide survey needed to determine Machi- is if of Mexico, and Costa Rica. Most species (several /a5 belongs in Persea (as accepted here) or should is The hundred) occur in Asia south to Australia and the be recognized as a distinct genus. neotropical Pacific Islands. The fruit subtended by a cup- species have been revised by Kopp(1966). Persea is 386 Annals of the Garden Missouri Botanical Mez americana Urbanodendron is widely cultivated for its edible fruit. A few neotropical species have clustered leaves. A neotropical genus of three species, restricted The known two Phoebe Nees southern cupules are to Brazil. in more species: double-rimmed, with or less persis- Phoebe here considered a palaeotropical ge- is The genus has been revised by Rohwer tent tepals. Phoebe nus. Neotropical species formerly placed in (1988). Cinnamomurn; are, for the larger part, included in & a few belong to Ocotea. Williamodendron Kubitzki Richter A Ph yUostenwnodaphne Kostermans genus two neotropical with species, infre- known quently collected, but from Costa Rica, An monotypic genus from infrequently collected, northern Colombia, Amazonia, and southern Brazil. The southern the Atlantic rainforests in Brazil. fruit The number more of species increase as will likpely double-rimmed The genus seated a cupule. in is become The cupule small collections available. is Rohwer was revised by (1988). and platelike, resembling that of Mezilaurus. Leaves Pleurothyrium Nees are clustered at the tips of branches. A 40 neotropical genus with close species, to Literature Cited known from Guatemala south to Peru, Bolivia, and A Allen, C. K. 1945. Studies in Lauraceae VI pre- The seated a rather deep, cup- Brazil. fruit in is Mexican and Amer- liminary overview of the (.entral n f s,haped, cupul1e, wh1 i- ch1 • often warty or l1 enti• cellate. us 280-434. .^^^ ^^^^^^ ^^^^^ 26: j ^^^^^,^j have pronoimced marginal Several species a vein, Burger, W. 1988. A new genus Lauraceae from C. of an unusual feature in Lauraceae, or clustered leaves. Costa Rica, with comments on problems of generic The genus heing revised by van der WerfT. and specific delimitation within the family. Brittonia is 275-282. 40: Povedadaphne Burger Hutchinson, 1964. The Genera of Flowering Plants, J. Volume Clarendon Oxford. Press, I. A monotypic genus apparently Cos- A restricted to Kopp, 1966. taxonomic revision of the genus L. The Persea Western Hemisphere. Mem. New York ta Rica. rather large fruit subtended by a in the is 11 Bot. Card. 14: 20. The small, platelike cupule. fruiting pedicel swol- is Kostermans, A. G. H. 1936. Lauraceae. Flora J. /;/. columnar stamens, len. Distinctive are the nine of Suriname. Meded. Kolon. Amsterdam, Afd. Inst. each with four apical locelli. Handelsmus. Volume 244-337. 11. 1938. Revision of the Lauraceae V. Rec. . & R hodostenionoddphne Rohwer Kubitzki 834-931. Trav. Bot. Neerl. 35: Conmiun. 1957. Lauraceae. Forest Res. Inst. . A known poorly geims of about a dozen species, 1-64. 57: common, A two of which are rather while several 1974. monograph of Caryodaphnopsis A. . 123- The Shaw. Reinwardtia 137. need genus has 9: species are of description. in *]f r-^D- JO *r> Kubitzki, K. 1982. Anlba. Flora Neotropica 31: In: Ibeen reported from Costa Kica -i and reru. ^ to Brazil ^^ 1-84. ^ Vhe cupule deeply cupshaped and rather large. & is Williamodendron H. G. RiCHTER. 1987. & Kubitzki Richter, a new genus of neotropical Lau- Sassafras Presl 49-58. raceae. Bot. Jahrb. Syst. 109: Kurz, H. 1983. Fortpflanzungsbiologie einiger Gattun- A genus one widespread small with three species, gen neotropischer Lauraceen und Revision der Gat- in the eastern U.S.A., the other two in China, tung Licaria (Lauraceae). Ph.D. Thesis, Univ. of The Leaves are frequently lobed. cupule cup- Hamburg, Hamburg. is MacBride, 1938. Lauraceae. Flora of Peru. Field shaped, often crowned with remnants of the tepals. F. In: 819-931. Mus. Nat. Hist. Bot. Ser. 13(2): Bark and root have been used to prepare tea or Mez, 1889. Lauraceae Americanae. Jahrb. Konigl. C. root beer. 1-556. Bot. Cart. Berlin 5: Renner, 1982. Aiouea. Flora Neotropica 31: S. In: System onodaphne 85 116. Richter, H. G. 1981. Anatomic des sekundaren Xy- See Kubitzkia. lems und der Rinde der Lauraceae. Sonderb. Natur- Hamburg 1-148. Vereins wiss. 5: VmbeUalaria Nut tall Rohwer, G. 1988. The genera Dicypellium, Phyl- J. lostenwnodaphney Systemonodaphne and Urbano- A monotypic genus occurring California and in 157- drndron 110: (Lauraceae). Bot. Jahrb. Syst. southern Oregon The subtended (U.S.A.). fruit is — 171. The by a small, platelike cupule. leaves are used & H. G. Richter H. van der Werff. 1991. , as a spice; the wood valued tor cabinetmaking. Two new genera of neotropical Lauraceae and critical is