BONPLANDIA 13(1-4): 21-29. 2004

SYSTEMATICS OF (BURRET) D. LEGRAND & KAUSEL ()

LESLIE R. LANDRUM1 & ANDREW SALYWON2

Summary: Landrum, L. R. & Salywon, A. 2004. Systematics of Amomyrtus (Burret) D. Legrand & Kausel (Myrtaceae). Bonplandia 13(1-4): 21-29. ISSN: 0524-0476.

The systematics of Amomyrtus, an endemic of southwestern temperate South America, is reviewed. In molecular phylogenetic studies the genus is supported as monophyletic, with its closest relatives being other American genera such as Legrandia and Pimenta. A lectotype for Myrcia lechleriana Miquel is chosen.

Key words: , Amomyrtus , Amomytus meli, South America.

Resumen: Landrum, L. R. & Salywon, A. 2004. Sistemática de Amomyrtus (Burret) D. Legrand & Kausel (Myrtaceae). Bonplandia 13(1-4): 21-29. ISSN: 0524-0476.

Se revisa la sistemática de Amomyrtus, un género endémico del sudoeste de América del Sur. En los estudios moleculares el género aparece como monofilético, siendo sus parientes más cercanos los géneros americanos Legrandia y Pimenta. Se elige un lectotipo para Myrcia lechleriana Miquel.

Palabras clave: Taxonomía, , Amomytus meli, Sudamérica.

Amomyrtus (Burret) D. Legrand & Kausel relatively large (4-6 mm long), hard seeds, is a genus of two species endemic to the wet moderately to strongly aromatic , and temperate forests of southern Chile and pentamerous flowers (Landrum 1988). adjacent . Both species are known (Phil.) D. Legrand & Kausel for their hard wood and aromatic leaves. No and A. luma (Molina) D. Legrand & Kausel unique morphological synapomorphies distin• are both that reach a size of up to 20 m. guish the genus, but it can be distinguished Amomyrtus meli grows only in Chile from from other genera of Myrtaceae through a Arauco to Chiloe at elevations from sea level combination of characters of the embryo, seed to ca. 600 m; andA luma grows from near sea coat, flowers and inflorescence (McVaugh level to ca. 800 m from Maule to Aisen in 1968; Landrum 1988; Landrum and Sharp Chile and in the Andes of Argentina from 1989) and specifically from other genera of Neuquen to Chubut. The species commonly southern South America by a combination of grow near each other in their region of

1 School of Life Sciences, Arizona State University, P. O. Box 874501, Tempe, Arizona 85287-4501, USA 2 U. S. Department of Agriculture, Agricultural Research Service, U.S. Water Conservation Laboratory, Phoenix, Arizona, 85040-8807, USA

21 BONPLANDIA 13(1-4). 2004 overlap, but have slightly different habitats especially subtribe Myrtinae, are being as discussed below. No hybrids are known investigated using molecular phylogenetic even though the flowering periods overlap, analyses of sequence data from the nuclear suggesting some type reproductive isolating ribosomal DNA Internal Transcribed Spacer mechanism. Region (ITS) of both Old and New World The phylogenetic affinities of Amomyrtus fleshy fruited and capsular fruited genera are not clear but various kinds of evidence (Salywon, 2003; Salywon & al, 2004). In have become available in recent years. these studies, maximum parsimony, Bayesian native to temperate South America analyses and neighbor-joining methods all often have relatives in Australasia (e.g., strongly support (100 % bootstrap support Nothofagus, Eucryphia, Laurelia, Gevuina, and 100% Bayesian posterior probability) the Tepualia) or temperate eastern South Ame• monophyly of Amomyrtus, which is not rica or the central or northern Andes (e.g., surprising for a group of two similar species, Myrceugenia, Blepharocalyx, Myrcianthes, but it is the first time that the monophyly of Azara, Escallonia) (Landrum 1981). One the genus has been tested using molecular might expect that relatives of Amomyrtus methods. In addition, the neighbor-joining would be found in one of those areas. trees, several of the most parsimonious trees, Secondary chemical data indicate that and highest likelihood Bayesian trees suggest Amomyrtus may have relatives in either that Amomyrtus has a sister group relationship area. Weyerstahl & al. (1992) found that with Legrandia, a morphologically distinct, Amomyrtus has certain 2-alkylchromones monotypic genus endemic to temperate Chile. otherwise unknown in plants and 1-pheny lalkan- In some analyses (i.e., neighbor-joining and 3-ones, only one of which (l-phenylhexan-3- Baysian majority rule) the Amomyrtus- one) had previously been found in Stellera Legrandia clade is sister to a clade of two chamaejasme L. (Gentianaceae). The latter Caribbean species of Pimenta, with the New compound was also found by Weyerstahl Zealand endemic Lophomyrtus and other & al. in Amomyrtella guili (Speg.) Kausel Australasian genera more distantly related. In (Myrtaceae), of northwestern Argentina. all of these analyses Lophomyrtus is resolved A similar 2-alkylchromone (2-isopropyl- as sister to Lenwebbia of Australia. The sister chromone) has been found in Lophomyrtus group relationship of these two Australasian bullata (Soland. Ex A. Cunn.) Burret genera is also supported by their quite similar (Myrtaceae) of New Zealand (Briggs and morphology (Snow & al. 2003). The higher White, 1971). The distribution of these level generic relationships of the fleshy- compounds is still of course poorly known, fruited genera in these phylogenetic analyses but if they prove to be unusual and restricted lack significant bootstrap support or clade to a few genera, then phylogenetic affinities credibility values and are thus largely between Amomyrtus, Amomyrtella and unresolved, due to short internodes between Lophomyrtus may be indicated. most genera. This condition is most likely due to an early rapid radiation of Myrtoideae. In Burret (1941) considered the species of any case molecular studies conducted so far Amomyrtus and Amomyrtella to be part of a seem to indicate an American origin for mainly Brazilian genus Pseudocaryophy- Amomyrtus and no detectable close relation• llus. All share a similar subapical protruding ship with any Australasian genera. More data placenta, which was probably the basis of from other segments of DNA and/or from Burret's placement. Recently Landrum more genera are needed for more robust (1986) submerged the type species of phylogenetic hypotheses. In particular, se• Pseudocaryophyllus in Pimenta, a mainly quence data from Amomyrtella would be Caribbean genus, but excluded Amomyrtus interesting as it is another American genus that and Amomyrtella. His reasons for excluding earlier studies (Burret 1941; Weyerstahl & al. the latter genera were differences in seed 1992) have indicated might be closely related coat, embryo and inflorescence structure. to Amomyrtus. Generic relationships of the Myrtaceae,

22 L.R. Landrum & A. Salywon, Systematics of Amomyrtus (Myrtaceae)

Amomyrtw (Burret) D. Legrand & cm wide; calyx-lobes linear, linear oblong, or Kausel narrowly triangular; seed 4-4.5 mm long, grey-green; hypanthium 2-2.5 mm long, not Kausel, E, Lilloa 13: 145. ("1947") 1948. noticeably glandular; usually growing near Pseudocaryophyllus Berg, sect. Amomyrtus water. Burret, Notizb. Bot. Gart. Berlin-Dahlem 15: 1. A. luma 514. 1941. Type. Eugenia darwinii Hooker f, lectotype designated by McVaugh (Taxon 17: 403. T. Trunk more or less uniformly whitish; twigs 1968). obscurely pubescent to glabrous; leaves strongly aromatic, often over 3 cm long and 1.5 cm wide; Aromatic or trees up to ca. 20 m calyx-lobes approximately triangular, slightly high, with smooth or somewhat scaly bark; wider than long; seed ca. 5-6 mm long, light tan; hairs simple, unicellular; leaves opposite, hypanthium 2.5-3.5 mm long, strongly glandular; persistent, coriaceous, petiolate, usually lan• usually growing as an understory in upland ceolate or ovate; inflorescence a solitary habitats. flower or bracteate shoot of ca. 6 flowers, the 2. A. meli peduncles solitary (or rarely superimposed); flowers perfect, pentamerous; bracteoles caducous at or before anthesis; calyx lobes 1. Amomyrtus luma (Molina) D. Legrand & persistent; white; whitish, 30- Kausel 80; anthers with 0-1 gland in the connective; Figs. 1A, D, E & F and 2B ovary 2-3- locular, the placentae attached to the upper septum, more or less shield-shaped, Kausel, E, Lilloa 13: 146. ("1947") 1948. the ovules radiating from the placental Myrtus luma Molina, Saggio sulla storia natu• margin; fruit a , the seeds hard, 1-4, rale del Chili, ed. 1. 173. 1782. Type. Description shaped like a flattened snail's shell with a of Molina. single coil, 4-6 mm long; embryo tinged Myrtus multiflora Jussieu ex Jaume Saint-Hillaire, purple, C-shaped, the hypocotyls about as in Duhamel, Traite arbr. arbust, ed. 2, 1: wide or slightly wider than the cotyledons, the 208. 1800-1803. Type. Chile. "Rapporte par cotyledons less than 1/5 as long as hypocotyl, Dombey. Herb, de Jussieu," Dombey s.n. membranous, folded back against the hypoco- (holotype, P, ex herb. Jussieu, = MICH neg. 1967!; tyl. F neg. 7924 of apparent isotype at G!). Amomyrtus is distinguished from all other Eugenia darwinii Hook, f, Fl. Antarctica genera of southern South American Myrtaceae 277. 1846. Type. "South Chili, Cape Tres Mün• by its pentamerous flowers, hard seeds 4-6 tes," Darwin s.n. (lectoype CGE, designated by mm long, and moderately to strongly aromatic Porter [1986]; isolectotype, K!, = ASU photo!). leaves. Burret's name for this group appears to Myrtus darwinii (Hook, f.) Barneoud, in Gay, be a union of the names Amomis Berg (= Fl. chil.2: 383. 1847. Pimenta Lindley) and Myrtus L. Myrcia lechleriana Miquel, Linnaea 25: Both species of Amomyrtus have very hard 651. 1852. Type. Chile. "Prope coloniam Arique & wood that is utilized for tool handles and prope urbem Valdivia," Lechler 181 & 576, lumbering and farming equipment or wherever (Lechler 576, syntype, U, = ASU photo!; Lechler very strong wood is needed. 181, syntype at U!, hereby designated as lectotype, = ASU photo!; isolectotypes, SGO!, W!). Myrtus valdiviana Phil, Linnaea 28: 688. 1858. Key to the species of Amomyrtus Type. Chile. "Prope Corral," Krause s.n. (holotype, SGO!; isotype, W!; Fneg. 19707 of isotype at M!). 1. Trunk reddish-brown with lighter spots; twigs Pseudocaryophyllus darwinii (Hook, f.) Burret, densely to moderately pubescent; leaves mode• Notizbl. Bot. Gart. Berlin-Dahlem 15: 514. 1941. rately aromatic, seldom over 3 cm long or 1.5

23 BONPLANDIA 13(1-4). 2004

Shrub or tree up to ca. 20 m high, moderately moderately pubescent, the bark at first light aromatic, moderately to densely pubescent on reddish-brown to grey, with age splitting in a young growth, the trunk smooth, reddish-brown reticulate pattern, revealing reddish-brown with lighter large spots where the bark has inner bark. Leaves elliptic, elliptic-oblong, recently fallen; hairs whitish, most ca. 0.3-0.5 ovate, or lanceolate, 1.4-3(-5) cm long, mm long, erect to antrorse; twigs densely to 0.6-1.5(-2.5) cm wide, 1.5-3 times as long as

Fig. 1. Amomyrtus luma (A, D, E from Gunckel 1779 at UC; F from Landrum 3144 at NY): A, twig showing bracteate shoot (raceme) inflorescences. D, bud. E, section of ovary showing placenta and ovules. F, seed. Amomyrtus meli (B, C from Kausel 1434 at F; G, H from Landrum 3193 at NY): B, bud. C, section of ovary showing placenta and ovules. G, seed. H, embryo. Drawn by Terry Bell.

24 L.R. Landrum & A. Salywon, Systematics of Amomyrtus (Myrtaceae) wide, essentially glabrous except for the with the sides concave, 1-1.5 mm long, the midvein that is usually pubescent above; apex sinuses between the lobes broadly U-shaped; acute to abruptly acuminate or rounded, often petals suborbicular, ca. 3 mm long; hypan- mucronate; base acute or cuneate; thium obconic, 2-2.5 mm long, not noticeably unchanneled to shallowly channeled, 2-4 mm glandular, reddish-brown, glabrous or long, 0.5-1 mm wide, densely pubescent to nearly so; disk 2.5-4 mm across, glabrous; glabrous; midvein flat to slightly raised, style 4-8 mm long, glabrous; stamens prominent below; lateral veins indistinct or up ca. 30-45, ca. 4-5 mm long; anthers 0.5 mm to ca. 10 pairs faintly visible; marginal veins long, usually with 0-1 gland in the more or less equaling the laterals in promi• connective; ovary 2-3-locular; ovules nence, paralleling the margin; blades usually ca. 4-6, the placenta ca. 0.3 mm wide. Fruit stiffly coriaceous, drying grey-green to dark black, subglobose, ca. 5-10 mm diameter; reddish-brown, often with many recessed seed coat smooth, hard, grey-green, pores above. Flower buds 3-4 mm long; ca. 4-4.5 mm long, the seed-coat wall peduncles solitary, 0.5-2 cm long, 0.3-0.5 mm ca. 0.2 mm thick. wide, densely to sparsely puberulent, usually grouped together on bracteate shoots, the Distribution: A tree, usually of wet shoot-axes up to ca. 2.5 cm long, the bracts habitats from Maule to Aisen in Chile and scale-like to elliptic, ca. 1-2.5 mm long, from Neuquen to Chubut in the Andes of deciduous at or before anthesis; bracteoles Argentina. narrowly linear, ca. 1 mm long; calyx-lobes linear, linear-oblong, or narrowly triangular Phenology: Flowering from September to BONPLANDIA 13(1-4). 2004

January; fruiting from December to March. 10648 (F, NY, UC, US); Chiloé, Alto Palena, Mallín Local names: Luma, palo madroño, cau- de Amaro, 21 Jan 1976 (fr), Gajardo s.n. (EIF); chao, caochao, caucharme (C. Muñoz, 1966). Valdivia, Llancacura, cancha 4, 500 m, 10 Jan 1967 (fr), Gajardo 164 (EIF); Valdivia, 20 km W of La Argentina. Chubut: W end of Brazo Norte Union, M. N. Alerce Costero (40° 14'S, 73° Lago Menendez (42° 37'S, 71° 53'W), 550 m, 8 22'W), 600 m, 6 Feb 1988 (fr), Gardner & Knees Mar 1975 (fr), Hjerting & al. 75- 175 (MICH); 4087 (ASU); Valdivia, 38 km SW of San José de Lago Kruger, 510 m, 10 Dec 1945 (fl), Meyer la Mariquina (39° 30'S, 73° 10'W), 30 m, 15 Mar 9544 (H, MICH); Parque Nacional Los Alerces, 1986 (st), Gentry & al. 53450 (ASU); Chiloé, Lago Menendez, El Cafe, 10 Feb 1949 (fr), Soria- Chepu, 8 Oct 1958 (fl), Godley 151 (SGO); Chiloé, no 3448 (CTES). Neuquén: P. N. Nahuel Huapi, Cordillera San Pedro, 2000 ft, 13 Nov 1958 (st), picada Los Cántaros, 6 Jan 1952 (fl), Boelcke & Godley 461 (SGO); Llanquihue, La Ensenada, 18 Correa 5286 (CONC). Río Negro: Laguna Frias, 5 Sep 1930 (fl), Gunckell779 (MO, UC); Valdivia, Mar 1946 (fr), de Barba 1026 (F, W); Nahuel Corral, La Aguada, 15 Nov 1930 (fl), Gunckel Huapi, arroyo Panqueco, 16 Mar 1948 (fr), De 1841 (CONC, GH); Chiloé, Piriquina, 18 Nov 1931 Barba 2290 (H); Lago Nahuel Huapi, Puerto Blest (fl), Junge 91 (MO); Riñihue, ca. 600 m, 17 Feb (ca. 41° 2'S, 71° 50'W), ca. 800 m, 9 Mar 1982 1941 (st), Kausel 941 (H); Llanquihue, Maullín, (fr), Landrum 4374 (NY). Chile. Region VII: Parga, Río Peuchen, 22-23 Nov 1944 (fl), Linares, ca. 5 miles N of Parral, 18 Feb 1958 (fr), Kausel 1430 (F, H, SGO); Parga, Fundo Porve• Eyerdam 10732 (UC, US); Maule, Constitución, nir, 22 Nov 1944 (st), Kausel 1454 (H); Chiloé, Quebrada Honda, 250 m, 7 Feb 1949 (st), Kausel Yaldad, ca. 10 km W of Quellón (ca. 43° 15'S, 2725 (H, SGO); Maule, Chanco, Cordillera Costa 73° 45'W), near sea level, 7-8 Feb 1978 (fr), de Cauquenes, 450 m, 8 Nov 1957 (fl), Schlegel Landrum 3129 (MICH, NY); Frutillar, forest re• 1399 (CONC); Maule, Cordillera Costa de Cau• serve of U. de Chile (ca. 41° 10'S, 73° 5'W), 50 quenes, 450 m, 15 Nov 1958 (fl), Schlegel 1780 m, 7 Mar 1982 (st), Landrum 4357 (NY); Valdi• (CONC). Region VIII: Curanilahue, Nov 1927 via, ca. 18 km E of Hueicolla along rd to Cordi• (fl), Claude-Joseph 5397 (US); Concepción, llera Pelada (ca. 40° 10'S, 73° 35'W), ca. 600 environs de Concepción, 1855 (fl), Germain s.n. m, 17 Apr 1982 (st), Landrum 4487 (NY); (GH); Laraquete, Río Cruces, 24-25 Mar 1940 Valdivia, Niebla, along coastal rd. to N, ca. 0.5 (st), Kausel 750 (H); Nuble, San Carlos, Las Mer• km N of La Misión, ca. 15 m, 17 Jan 1988 (st), cedes, camino a la Laguna El Valiente, 700 m, 6 Landrum 5903 (ASU); Llanquihue, Los Muermos, Mar 1944 (fr), Kausel 1328 (F, H); Arauco, ca. 0.3-1 km along road to Cumbre Alta off road Nahuelbuta, faldeos del Cerro Pino Huacho (37° to Quenuil, ca. 50 m, 18 Jan 1988 (fr), Landrum 41'S, 73° 14'W), 500 m, 24 Nov 1978 (fl), 5920 (ASU); ca. 10 km S of Valdivia on rd. to Marticorena & al. 1578 (CONC). Region IX: Paillaco, parcela de C. Donoso, 18 Oct 1991 Cautín, Huerquehue, near volcán Llaima, Mar (fl), Landrum 7591 (ASU); P. N. Puyehue, 1970 (fr), Donoso & Landrum 934 (EIF, MICH); Aguas Calientes (ca. 40° 45'S, 72° 20'W), ca. Malleco, Angol, Purén, Nov 1938 (fl), Kausel s.n. 300 m, 9 Nov 1993 (fl), Landrum 8060 (ASU); (F); Malleco, Coyancahuin, 14-16 Feb 1939 (fr), Chiloé, Isla Talcán, 23 Feb 1961 (st), Marticorena Kausel 536 (SGO); Toltén, Río Puculon, 21 Feb 1745 (CONC); Chiloé, Quellón, Punta Lapa (43° 1942 (st), Kausel 1071 (SGO); Temuco, Cerro 8'S, 73° 38'W), 12 Jan 1975 (fr), Marticorena & al. Ñielol, ca. 140 m, 25 Sep 1939 (fl), Montero 3761 100 (CONC, CTES); Valdivia, Panguipulli, cami• (GH); Cautín, just E of Pucón, from Río Turbio no de Neltume a Puerto Fui (39° 51'S, 71° along road to Termas de Huife (39° 20'S, 71° 55'W), 490 m, (fr), Marticorena & al. 613 45'W), 29 Nov 1991 (fl), Taylor & Taylor 10869 (CONC, CTES); Valle de Peulla, 8 Feb 1945 (ASU). Region X: Valdivia, San José de la (fr), Rudolph 4314 (VALD); Valle de Cayutue, Mariquina, 24 Oct 1939 (fl), Aravena 18025 (UC); 10 Jan 1945 (fl), Rudolph 4315 (VALD); Lago Chiloé, Río Yelcho, Nov 1939 (fl), Bernath s.n. (F, Rupanco, Encanto, 9 Nov 1947 (fl), Rudolph H); Osorno, Pucatrihue, 29 Jan 1958 (fr), Eyerdam 4316 (VALD); Lago Riñihue, Riñihue, 18 Sep 10526 (F, MICH, UC, US); Valdivia, Cordillera de 1940 (fl), Santesson 997 (NY); ca. 10 km NE of Alerce, 500-900 m, 10 Feb 1958 (fr), Eyerdam Pto. Octay, 26 Dec 1980 (fr), Schmid 1980- 66

26 L.R. Landrum & A. Salywon, Systematics of Amomyrtus (Myrtaceae)

(NY); Valdiva, Chaiguín, Los Alerzales, 800 m, 17 2. Amomyrtus meli (Phil.) D. Legrand & Dec 1953 (fl), Smith & Sparre 264 (CONC); Kausel Chiloé, ca. 20 km S of Ancud, route 5, 80-100 Figs. IB, C, G&Hand2A m, 15 Jan 1981 (fr), Sobe! & Strudwick 2742 (ASU); Osorno, Pucatrihue, 11-12 Jan 1948 (fr), Sparre 3902 (SGO); Llanquihue, Maullin, Kausel, E., Lilloa 13: 146. ("1947") 1948. 19-21 Jan 1948 (fr), Sparre 4070 (SGO); Myrtus meli Phil., Linnaea28: 638. 1857. Type. Llanquihue, Llaguepe, Monte Yate, ca. 600 m, Chile. "In sylvis densis provinciae Valdiviae," 29-31 Jan 1948 (fr), Sparre 4200 (SGO); Salto Philippi s.n, (holotype, SGO!, =A.SU photo!; del Pilmaiquén, 13 Feb 1948 (st), Sparre 4456 possible isotypes G!, = ASU photo!, MICH!, W!; F (SGO); Osorno, Cordillera de La Reina, ca. 750 neg. 19704 of possible isotype at M!). m, 20 Feb 1948 (st), Sparre 4567 (SGO); Pseudocaryophyllus meli (Phil.) Burret, Notizbl. Bot. Valdivia, Curiñanco, 7-9 Mar 1948 (st), Sparre Gait Berlin-Dahlem 15:515. 1941. 4708 (SGO); Osorno, Fundo Mulpulmo (40° 33'S, 72° 57'W), 90 m, Mar 1967 (fr), Zollitsch Tree up ca. 20 m high, strongly aromatic, 224 (CONC). Region XI: Puerto Gato, 15 Oct usually only obscurely pubescent on young 1947 (fl), Behn 7 (CONC, H); Puyuhuapi, 2 Nov growth, the trunk with smooth whitish bark, 1947 (fl), Behn 93 (CONC); Quitralco, 10 Dec not strongly mottled; hairs whitish, up to 1947 (fl), Behn 118 (CONC); Lago General Ca• ca. 0.3 mm long, usually curved antrorsely; rrera (Buenos Aires), Bahia Murta, Río Tranqui• twigs only obscurely pubescent to glabrous, lo, 3 Feb 1943 (fr), Grosse H-l 110766 s.n. (H); the bark greyish-white or light reddish-brown, road from Coihaique to Puerto Aisén, Puente smooth at first, becoming longitudinally Cascadal, ca. 30 km W of Coihaique (45° 25'S, broken, with longitudinal strips persisting 72° 20'W,), 15 Nov 1993 (fl), Landrum 8099 beneath the scars. Leaves ovate, lanceo• (ASU); Coihaique, at bridge ca. 12 km W of late, or elliptic, 2-5(-9) cm long, 0.7-2.5(-3) school of Lago Atravesado (ca. 45° 40'S, 72° cm wide, 1.4-3.5 times as long as wide; apex 20'W), 19 Nov 1993 (fl), Landrum 8175 (ASU); acute, often mucronate; base acute; petiole Pen insula Tres Montes, Pto. Barraso, 2 Apr unchanneled to shallowly channeled, 2-4 mm 1929 (st), Roivainen H-l386491 s.n. (H, SGO); long, 1-2 mm wide, yellowish-brown; midvein entre estancia Teresa y Bahia Exploradores (46° about flat or slightly raised above, moderately 25'S, 73° 10'W), 40 m, 3 Mar 1967 (fr), Seki prominent below; lateral veins indistinct or up 325 (CONC); San Rafael, Rada Quesahuen (46° to ca. 7 pairs faintly visible, the lower pairs 22'S, 73° 44'W), 50 m, 4 Feb 1959 (fl), Schlegel ascending, the upper pairs leaving the midvein 2057 (CONC). at an angle of 45 degrees or more; marginal veins about equaling the laterals in prominence; Amomyrtus luma can be distinguished blades stiffly coriaceous, drying grey-green to from other species of Chilean Myrtaceae by yellow-green, dull to slightly lustrous above. its reddish brown bark with lighter spots. Flower buds 4-5 mm long; peduncles solitary The bark is not so orange as it is in Luma (rarely superimposed), 0.4-2 cm long, ca. 0.5 apiculata (DC.) Burret and Blepharocalyx mm wide, glabrous to minutely puberulent, cruckshanksii (Hooker & Arnott) Niedenzu. moderately glandular, usually grouped together Amomyrtus luma has moderately aromatic in bracteate shoots, the bracts deciduous leaves, somewhat less pungent than those of before anthesis, elliptic, 1.5-2 mm long; A. meli, but with a distinctive fragrance. bracteoles linear, ca. 1-1.5 mm long; calyx-lobes According to Murillo (1889) and M. Muñoz approximately triangular, ca. 1-1.5 mm long, & al. (1981) Amomyrtus luma has been used slightly wider than long, the sides nearly medicinally in Chile as an "aromatic, stimu• straight or slightly concave, glabrous; petals lant, and astringent." Gay (1847) mentions suborbicular, ca. 3-4 mm long, glabrous, or that the fruits of A. luma (=Myrtus luma) the margin ciliate; hypanthium obconic, ca. were used by Indians to make fermented 2.5-3.5 mm long, ca. 3 times as long as the drinks. calyx-lobes, more or less smooth when dry,

27 BONPLANDIA 13(1-4). 2004 densely glandular, the glands noticeable as white rd. to Paillaco, parcela de C. Donoso, 18 Oct 1991 spots when dry; disk ca. 4 mm across, glabrous; (st), Landrum 7589 (ASU); Valdivia, ca. 20 km S style 7-8 mm long, glabrous; stamens ca. 40-80, of Valdivia on road to Osorno (39° 55'S, 73° ca. 5-7 mm long; anthers ca. 0.5-0.8 mm long, 10'W,), 11 Nov 1993 (fl), Landrum 8098 (ASU); usually without glands in connective; ovary camino de Panguipulli a Los Lagos, 275 m, 13 Jan 2-locular; ovules 5-8 per locule, the placenta 1976 (fr), Marticorena & al. 304 (CONC); ca. 0.5-0.7 mm wide. Fruit black, subglobose, Pichilafquen, Lago Rupanco, 16 Nov 1956 (st), ca. 1-1.5 cm in diameter; seed coat hard, light Pfister s.n. (CONC); Valdivia, Chunimpa, Feb colored, smooth, somewhat shiny, ca. 5-6 mm 1875 (st), Philippi 661b (SGO); Osorno, Nadi de long, the seed-coat wall ca. 0.25 mm thick. Octay, 21 Nov 1945 (fl), Rudolph 4327 (VALD); Osorno, Contaco, 25 Dec 1950 (fl), Rudolph 7123 Distribution: A tree, usually of upland habitats (VALD); La Union, Llancacura, 200 m, 20 Jan in wet forests of Chile from Arauco to Chiloé. 1957 (st), Schlegel 1269 (CONC); Valdivia, Curinaco, 200 m, 7-9 Mar 1948, Sparre 4760 Phenology: Flowering from late October to (SGO). December; fruiting from January to May. Amomyrtus meli is usually distinguishable Local name: Meli, luma, luma blanca (C. Do• from other Chilean Myrtaceae by its light noso, pers. comm.). smooth bark. The strong aroma of the crushed leaves is distinctive, being pleasantly fruity, and Chile. Region VIII: Lago Lleu-Lleu, 24 Feb once experienced, is easily recognized. The 1939 (st), Kausel 601 (H); Nahuelbuta, orillas del leaves lose most of their aroma upon drying. The Río Caramávida (37° 40'S, 73° 16'W), 150 m, 24 fruits are pleasant tasting when ripe. In the area Nov 1978 (fl), Marticorena & al. 1566 (CONC); of Valdivia, Chile, tea of the leaves of A. meli Toltén, Río Boldo, 13 Feb 1942 (st), Kausel 1027 has the reputation of being effective against high (CONC, H). Region IX: Cautín, Imperial, Toltén, cholesterol (C. Donoso, pers. comm.). Puculon, 21 Feb 1942 (fr), Kausel 1074 (CONC, H, SGO). Region X: 38 km SW of San José de la Mariquina (39° 30'S, 73° 10'W), 30 m, 15 Mar Acknowledgments 1986 (st), Gentry & al. 53451 (ASU); Osorno, Salto Pilmaiquén, 19 Feb 1942 (st), Kausel 1435 (H); Llanquihue, Parga, 22-23 Nov 1944 (st), We thank two anonymous reviewers and the Kausel 1435 (H); ca. 20 km S of Valdivia on road editors for many helpful suggestions. to Osorno (39° 55'S, 73° 10'W,), 11 Nov 1993 (fl), Landrum (ASU); Chiloé, road to Chepu S of Ancud, 8 May 1972 (fr), Landrum 932 (MICH); Literature Cited Chiloé, Ancud, near airport, 7 May 1972 (fr), Landrum 933 (MICH); Chiloé, Quemchi, ca 1 km W of town (ca. 42° 10'S, 73° 30'W), near sea BRIGGS, L. H. & G. W. WHITE. 4-methyl-l- level, 10 Feb 1978 (fr), Landrum 3193 (MICH, phenylpentane-l,3-dione and 2-isopropylchromone NY); Frutillar, forest reserva of the U. de Chile, ca. from the essential oil of Lophomyrtus bullata. J. 50 m, 7 Mar 1982 (fr), Landrum 4356 (NY); Chem. Soc. C, 3077-3079. Valdivia, ca. 18 km E of Hueicolla (ca. 40° 10'S, BURRET, M. 1941. Myrtaceen-Studien. Notizbl. Bot. 73° 35'W), ca. 600 m, 17 Apr 1982 (st), Landrum Gart. Berlin-Dahlem 15: 479-500. GAY, C. 1847. Florachilena(Myrtaceae), vol. 2: 376-399. 4488 (NY); Valdivia in park, 22 Apr 1982 (fr), LANDRUM, L. R. 1981. The phylogeny and geography of Landrum 4561 (MICH); Valdivia, Niebla, along Myrceugenia (Myrtaceae). Brittonia33: 105-129. coastal rd. to N, ca. 0.5 km N of La Misión, ca. 15 . 1986. Campomanesia, Pimenta, Blepharocalyx, m, 17 Jan 1988 (st), Landrum 5902 (ASU); Los Legrandia, Acca, Myrrhinium, and Luma (Myrtaceae). Muermos, ca. 0.3-1 km along road to Cumbre Alta Flora Neotropica Monographs 45: 1 -179. off road to Quenuil, ca. 50 m, 18 Jan 1988 (st), . 1988. The myrtle family (Myrtaceae) in Chile. Landrum 5913 (ASU); ca. 10 km S of Valdivia on Proc. Calif. Acad. Sci. 45(12): 277-317.

28 L.R. Landrum & A. Salywon, Systematics of Amomyrtus (Myrtaceae)

& W. P. SHARP. 1989. Seed coat characters of Biology, Arizona State University, Tempe, some American Myrtinae (Myrtaceae): Psidium and Arizona, United States. related genera. Syst. Bot. 14(3): 370-376. SALYWON, A., N. SNOW, M. F. WOJCIECHOWSKI, MCVAUGH, R. 1968. The genera of American J. CSIZMADI & L. R. LANDRUM. 2004. Phyloge- Myrtaceae-An interim report. Taxon 17: 354-418. netic relationships of Myrtaceae as inferred from MUÑOZ P., C. 1966. Sinopsis de la flora chilena, second nrDNA ITS sequence data. Botany 2004, Snowbird, edition. Universidad de Chile, Santiago. Utah, http://www.botanyconference.org/engine/ MUÑOZ S., M., E. BARRERA M. & I. MESA P. 1981. search/index.php? func=detail&aid=:795. El uso medicinal y alimenticio de plantas nativas y SNOW, N.. G. P. GUYMER & G. SAWVEL. 2003. naturalizadas en Chile. Mus. Nac. Hist. Nat. Publica• Systematics of Austromyrtus, Lenwebbia, and the ción Ocasional 33:3-89. Australian species of Gossia (Myrtaceae). Syst. Bot. MURILLO, A. 1889. Plantas Medicinales du Chili. Sec. Monographs 65: 1-95. Chilienne, Exposition Universelle de Paris. WEYERSTAHL, P., H. MARSCHALL & L. R. PORTER, D. M. 1986. Charles Darwin's LANDRUM. 1992. Constituents of the Leaf Extract specimens from the voyage of HMS Beagle. J. Linn. of Amomyrtus meli (R. A. Philippi) Legrand & Soc, Bot. 93: 1-172. Kausel, A. luma (Molina) Legrand & Kausel, and of SALYWON, A. 2003. A monograph of Mosiera Amomyrtella guili (Speg.) Kausel (Myrtaceae). (Myrtaceae). Ph.D. dissertation, Department of Flavour and Fragrance Journal 7: 247-251.

Original recibido el 7 de octubre de 2004; aceptado el 10 de noviembre de 2004.

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