A reappraisal of Barnebydendron (Leguminosae: Caesalpinioideae: Detarieae)
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- Warwick, M.C., Lewis, G.P. & Lima, H.C.d. Kew Bull (2008) 63: 143. doi:10.1007/s12225-007-9001-y
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Barnebydendron J. H. Kirkbr. (Leguminosae – Caesalpinioideae – Detarieae), when first described under the name Phyllocarpus Riedel ex Tul. (1843), was considered to be a monospecific genus restricted to the Atlantic forest in the vicinity of Rio de Janeiro, Brazil. A later collection from Guatemala in 1908 expanded the genus to two species. Many collections have been made since then in other parts of Central and South America and the distribution of the genus merits close examination. Recent publications have not provided full descriptions or illustrations, or cited specimens seen. A reappraisal of the genus is presented here and the monospecific status of the genus is confirmed.
KeywordsAtlantic forestBarnebydendronColoralliloGuacamayoLeguminosaemonospecificneotropicsseasonally dry forest
Kirkbride (1999), responding to a note by Barneby (1996), published the new generic name Barnebydendron for the caesalpinioid legume previously known as Phyllocarpus. The nomenclatural history linked to this decision is adequately laid out in Kirkbride’s paper (1999). Two species have been described in the genus Phyllocarpus: P. riedelii Tul. and P. septentrionalis Donn. Sm., but Barneby (1996) and Kirkbride (1999) recognised Barnebydendron as monospecific; neither of the authors, however, included a detailed description of the species or provided an illustration or numbered exsiccatae.
Examination of herbarium collections of Barnebydendron (many still housed under Phyllocarpus) has permitted us to test the monospecific status of the genus. Recent collections made available to us have clearly shown that the genus is more geographically widespread than previously thought. The main aim of this paper is thus to test whether Barnebydendron contains one or more species and to fill gaps in our knowledge of its distribution.
Barnebydendron is a genus of tropical trees found in Central America (from where most specimens in herbaria have been collected) and South America. There are small populations in Venezuela, in the Parque Nacional Manú in Peru, and in Brazil in the state of Acre and in Atlantic forest from southern Bahia to Rio de Janeiro, this forest being the location of the type specimen of B. riedelii (Tul.) J. H. Kirkbr.
Barnebydendron is a tree with abundant, striking scarlet flowers that open when the branches are leafless; it is used as an ornamental in urban areas, particularly in Rio de Janeiro, and notably in the Plant Introduction Garden at Summit in Panama and in the Escuela Agrícola Panamericana in Honduras. Barnebydendron was first collected in 1827 by L. Riedel and named Phyllocarpus riedelii by Tulasne in 1843. When W. A. Kellerman made collections in Guatemala in 1908, 6,000 km from the type locality, these were thought to represent a second species. J. Donnell Smith published Phyllocarpus septentrionalis in 1913 (Donnell Smith 1913), distinguishing it by the larger and fewer pairs of leaflets and citing the Wilmatte P. Cockerell collection of 1912 from Zacapa as the type. Even though there have been several relatively recent collections of Barnebydendron from across its geographical range, its distribution remains widely disjunct. The trees also grow in a wide variety of habitats, including humid coastal forest in Brazil, Purús basin tropical forest in Peru and Brazil, and sub-tropical forest in Guatemala. By contrast, it also grows in seasonally dry forest in Guatemala and other parts of Central America and in some areas of coastal southeastern Brazil. Microhabitats include dry gullies and sandy and rocky beaches in Central America, and riversides, floodplains and stream pastureland across the range of the genus. Cultivated trees are found in all areas except Peru. Collections made in Rio de Janeiro State were presumed by Barneby (1996) to be naturalised from material introduced into the Rio de Janeiro Botanical Garden, but recent collections from several sites in this state by one of us (H.C.L.) suggest that these populations are of wild origin because the trees are growing in remnants of undisturbed forest. In fact, along the Brazilian Atlantic coast from Rio de Janeiro northwards to southern Bahia, wild Barnebydendron trees grow in patches of lowland humid forest and in seasonally dry forest. Populations of Barnebydendron in Guatemala are largely native, although cultivated specimens have also been collected. Hughes & Lewis, on specimen (1224, K) field notes, described the genus as locally common in highly degraded dry forest with Acacia picachensis Brandegee, Caesalpinia pulcherrima (L.) Sw. and Thouinidium decandrum Radlk. Likewise, Hughes (1104, K) described Barnebydendron as frequent in small valleys with Caesalpinia velutina (Britton & Rose) Standl. and Cordia dentata Poir. In Panama, Allen, quoted in Woodson, Schery and collaborators (1951), described a specimen collected in Tumba Vieja in 1934 (Steyermark & Allen 16742) as coming from ‘wild trees which are quite common there in areas of climax rain forest’ and definitely indigenous to the Madden Lake forests.
Thorough examination was made of the leaves, and when available, also of the flowers and fruits of each specimen to ascertain if there were correlations amongst morphological characters, geographical areas, and different types of habitat. Cultivated collections were compared against wild collections. After analysis of over 90 specimens, we find no significant differences in leaves, flowers or fruits and certainly no consistent correlation of character differences with geography. There is continuous morphological variation between all of the specimens and we consider that there is no justification for recognition of more than one species. Our study found a consistently higher number of ovules, (3 – )5 – 10(– 12), per ovary than previous descriptions, 1 – 2( – 4), ‘few-ovuled’. We agree with Kirkbride (1999) and Barneby (1996) that the genus is monospecific.
Barnebydendron J. H. Kirkbr. (1999: 817).
Phyllocarpus Riedel ex Tul. (1843:142), non Phyllocarpus Riedel ex Endl. (1842). Type: Phyllocarpus riedelii Tul. (= Barnebydendron riedelii (Tul.) J. H. Kirkbr.).
Deciduous trees. Leaves paripinnate. Inflorescences racemose on older branches. Flowers conspicuous, pedicellate; calyx four-lobed. Corolla with three free petals and occasionally two vestigial ones. Androecium of nine fused stamens and one free; anthers dorsifixed. Gynoecium slightly shorter than androecium; ovary short-stipitate, with (3 –)5 –10(– 12) ovules (in contrast to earlier reports of 1– 2(– 4) ovules per ovary). Fruit flattened, winged along upper suture, indehiscent, 1 – 2 (– 3)-seeded.
Barnebydendron riedelii (Tul.) J. H. Kirkbr. (1999: 817).
Type: Brazil, Rio de Janeiro, L. Riedel s.n. (Catal. Herb. Bras. Guillemin no. 1022 [Tulasne 1844]) (holotype P!; isotypes NY (2 sheets)!, RB!).
Syn.: Phyllocarpus riedelii Tul. (1843: 142). Phyllocarpus septentrionalis Donn. Sm. (1913: 433). Type: Guatemala, Dept. Zacapa, prope Gualán, Feb. 1912 (fl.) W. P. Cockerell s.n. (lectotype US , selected by Standley & Steyermark (1946) in the Flora of Guatemala, labelled by Standley as ‘type’).
Barnebydendron was dedicated to Rupert C. Barneby (1911 – 2000) by Joseph H. Kirkbride (1999); phyllocarpus, Greek, phyllo = leaf, carpus = fruit, apparently referred to the laterally compressed, somewhat leaf-like, winged fruit.
Distribution (including cultivated material).
BRAZIL. Acre, Púrus basin, Bahia, Espírito Santo, Rio de Janeiro. COSTA RICA. Península de Osa, Puntarenas. CUBA. Havana Botanic Gardens (cult., pers. obs. G. P. L.). EL SALVADOR. San Salvador (cult.). GUATEMALA. Jalapa, Zacapa. GUYANA. Demerara (cult.). HONDURAS. Cortés, Morazán. Canal Zone (cult.). PERU. Madre de Dios, Manu Province. VENEZUELA Amazonas.
BRAZIL. Acre, Mouth of Río Macauhán, 9°20′S 69°W, 25 Aug. 1933, Krukoff 5651 (F, K, MO, RB); Mun. Rio Branco, Seringal Corredeira, Colocação Nova Morada, Lago da Corredeira, 8 Jan. 1984, de Lima et al. 2087 (RB); Mun. Rio Branco, Seringal Corredeira, Colocação Paraíso, 9 Jan. 1984, de Lima et al. 2110 (RB); FUNTAC, Area de Estudos, 9 May 1990, Saraiva et al. 319 (MO); Seringal, Auristella 10°56′S 69°34′W, 11 July 1911, Ule 9448 (F, K, RB); Ule 9447 (F, RB). Bahia. Prado, Faz. Estrela, 24 May 1972, Monteiro 23697 (RB). Espirito Santo. Proximo Linhares, 6 May 1966, Duarte 9743 (RB); Caxixe Quente, 18 May 1999, Hatschbach et al. 69141 (MO, RB); Marx s.n. (K, RB); Mun. Aracruz, Rod. Coqueiral-Santa Cruz, 19 May 1993, Oliveira 513 (RB). Rio de Janeiro. Horto Floresta, perto do Rumo, 2 May 1941, de Almeida et al. s.n. (RB); Estrada do Rumo, 13 May 1941, de Almeida 348 (RB); Corcovado, 2 May 1931, Campos Porto s.n. (P, RB ); Serra do Tingua, 22 Sept. 1837, Glaziou 6839a (P, RB); 1839, Guillemin cat. No.1002, Type (P); mata em frente de Jardim Botânico, 27 April 1987, Lewis et al. 1615 (K, MO, RB); Horto Florestal, 18 May 1984, de Lima et al. 2147a (RB); São Pedro da Aldeia, São Mateus, Faz. da Caveira, 6 May 1987, de Lima et al. 2873 (RB); Corcovado, 15 July 1931, Rosa & Lourenço s.n. (RB); Horto Florestal, 11 June 1932, Victorio & Lourenço s.n. (RB); 1827, Riedel (NY). COSTA RICA. Puntarenas. Península de Osa, Playa Carbonera, Puerto Jiménez, 8°24′N 83°17′W, 4 Jan. 1994, Aguilar 2952 (K, MO); Península de Osa, between Rio Esuinas and Palmar Sur de Osa, 12 Dec. 1950, Allen 5698 (F); Fila Retinto, Quebrada Benjamín, 8°59′N 83°28′W, 3 Jan. 1990, Hammel et al. 17710 (F, MO, P); Fila Retinto, Quebrada Benjamín, 8°59′N 83°28′W, 3 Jan. 1990, Hammel et al. 17711 (P); Cantón de Golfito, Valle de Coto Colorado, 8°38′N 83°10′W, 20 Oct. 1993, Morales et al. 1898 (MO); Carbonera, 8°24′N 83°17′W, 12 April 1992, Zamora & R. Aguilar 1828 (F, K, MO); Costado Noreste, 7°39′N 83°10′W, 2 Feb. 1992, Zamora et al. 1772 (F). EL SALVADOR. Depto. San Salvador. 13°40′N 89°11′W, 22 Jan. 1990, Villacorta 515 (F, K, MO). GUATEMALA. Jalapa. Guastatoya, 20 Feb. 1908, Kellerman 8056 (F); Vivero del El Progreso, 13 May 1983, Kennedy 1–7 (MO); 1 mile above El Progreso, 3 Feb. 1917, Popenae 759 (F); Barranquillo, 1932, Salas 1467 (F); Barranquillo, 1942, Steyermark 43325 & 46453 (F). Zacapa. Juan Ponce, Gualán, 11 Feb. 1994, Castillo 2187 (MO); 7 km W of El Loba, 9 Feb. 1970, Harmon & A. Fuentes 1842 (MO); 3 km NE of Gualán, 6 Feb. 1984, Hughes 415 (K); S of Zacapa on road to Chiquimula, 8 Feb. 1983, Hughes 292 (K); 8 March 1983, Hughes 317 (K); 9 May 1985, Hughes 708 (K); Agua Blanca, 14°54′N 89°32′W, 1 March 1988, Hughes 1104 (K); Road to Chiquimula, 14°53′N 89°32′W, 13 Feb. 1989, Hughes & Lewis 1224 (K). Road from El Progreso to Guacomayo, 15°04′N 89°37′W, 21 Feb. 1991, Macqueen & Pennington 69 (K). GUYANA. Demerara. Georgetown, 6°46′N 58°10′W, 2 April 1940, Martyn s.n. (K, RB). HONDURAS. Cortés. Vicinity of La Lima, 11 April 1947, Standley & Chacón P. 7251 (F); La Lima, 14 April 1947, Zamora & Aguilar 1828 (F, K, MO). Morazán. Dept. Yoro, 15°26′N 86°46′W, 25 March 1991, Hughes 1449 (K); El Zamorano, 30 Dec. 1961, Molina 10209 (F); Valle del Río Yeguare, 13 April 1967, Molina 20622 (F, K); Campus of Escuela Agrícola Panamericana, 17 Jan. 1986, Molina 33901 (MO); 35 km SE of Tegucigalpa, 6 Feb. 1986, Nelson 9502 (MO). PANAMA. Canal Zone. Summit, Feb. 1940, Allen 2053 (F, K, MO); Plant Introduction Garden, Summit, 6 Jan. 1935, Steyermark s.n. (MO); between Tumba Vieja and Salamanca, 3 Dec. 1934, Steyermark & Allen 16742 (MO, P). PERU. Madre de Dios. Río Cumerjali, Parque Nacional Manú, 71°32′W 11°49′S, 21 Oct. 1986, Foster & d’Achille 11963-B (F, K); Puerto Maldonado, Los Amigos Biological Station, c. 7 km upriver from mouth of Río Los Amigos, 19 May 2004, Maceda 1395 (K); Cocha Cashu Station, Río Manú, 5 Nov. 1986, Foster 12120-B (F, K); Rodal Semillero, km 104, 20 May 1980, Begarzo 150 (US); Cocha Cashu, Río Manú, 11°55′S 77°18′W, 16 Oct. 1991, Núñez et al. 14419 (M); Pumahuasi, 17 June 1964, Vasquez A. 32EVA (MO). VENEZUELA. T. F. Amazonas. Tayari, 24 Nov. 1975, Lissots.n. (US).
habitat and ecology.
In Brazil, Barnebydendron riedelii occurs in Atlantic tropical forest, the Amazon basin and on dry, rocky hillsides, from 50 – 800 m; in Guatemala, in dry gullies and degraded seasonally dry tropical forest to floodplains and river banks, from 130 – 900 m. The Peruvian specimens occur in floodplains and ravines from 350 – 850 m. In Costa Rica, B.riedelii occurs on rocky, dry beaches, from 1 – 30 m, along waterways and in primary forest from 100 – 500 m; in Panama, one specimen was collected at 66 m in inundated forest. The specimens from Honduras and El Salvador were cultivated.
Usually flowering when branches are leafless; April – June in Brazil, Peru and El Salvador, December – February in Guatemala, Costa Rica and Honduras, December – March in Panama. Glaziou 6839a, apparently collected in September 1837 in the Serra de Tingua, Rio de Janeiro, is a specimen with immature flowers. There is, however, some doubt concerning the authenticity of the field label, as Glaziou collections are sometimes collections made by other field botanists with fictitious field labels added to the pilfered specimen (see, e.g., Smith 1966; Wurdack 1970).
Flowers are likely to be pollinated by humming birds (Arroyo 1981). Fruiting is from April – May in Guatemala, Costa Rica and Honduras, and is known to occur in August in Brazil and from October – November in Peru.
Brazil: Itapicuru, Guarabu, Guarabu Cebola, Favela and Guaribeiro. El Salvador: Flor de mico, Guacamayo and Papagayo. Guatemala: Guacamayo and Palo Negro. Honduras: Coloradillo and Coralillo. Peru: Pumaquiero Colorado.
Primarily as an ornamental tree, having spectacular displays of mostly scarlet flowers. Isley (1975) reported it as being an introduced cultivated ornamental in cities of subtropical Florida.
Least Concern (LC). The species occurs in three widely disjunct areas of Central and South America in both wet and dry vegetation types and is not currently threatened globally. The tree has been recently collected in areas of southern Bahia and Rio de Janeiro that are new locality records. Although much of the Atlantic forest of eastern Brazil was cut down in the last century, the species evidently survives in a number of remaining, although restricted, areas of this vegetation type. It seems probable that the species is more at threat in the eastern part of its wide Neotropical range than elsewhere.
Herendeen et al. (2003a), on the basis of combined molecular and morphological data, placed Barnebydendron in a moderately supported clade with Goniorrhachis (from Bahia, Brazil) in a basally branching position sister to the rest of tribe Detarieae except Schotia (from southern and South Africa). Fougère-Danezan et al. (2003) recovered a similar relationship based on molecular data, although without support, but concluded that they had no unequivocal evidence concerning the position of Schotia, Goniorrhachis and Barnebydendron. The overall similarity in flower morphology and colour (various shades of red) between Barnebydendron and Schotia is striking. The flowers of Goniorrhachis, on the other hand, are white and somewhat Rosaceous in appearance.
Although the geographical distribution of Barnebydendron has no parallel in the legumes, the combined distribution of five basally branching genera in the Detarieae (Mackinder 2005) displays an overall pattern similar to the genera within the Umtiza clade of the Caesalpinieae (Herendeen et al. 2003b; Lewis 2005; Schrire et al.2005). The detarioid genera Neoapaloxylon and Brandzeia are from Madagascar, Schotia occurs in southern Africa and Barnebydendron and Goniorrhachis are Neotropical.
Phyllocarpus pterocarpus Riedel ex Endl. (= Deguelia scandens Aubl.).
The authors thank the curators of the herbaria at F, MO, NY and P for kindly loaning material for study and two anonymous reviewers for their input. This work was supported in part by National Science Foundation grant DEB-0316375.