Trichanthera gigantea (Bonpl.) Nees

Daniel, Thomas F., 2015, Synopsis of Trichanthera (Acanthaceae: Ruellieae: Trichantherinae), Proceedings of the California Academy of Sciences 62 (1), pp. 1-23 : 9-16

2. Trichanthera gigantea (Bonpl.) Nees in A. de Candolle, Prodr. 11:218. 1847.

Ruellia gigantea Bonpl. in Humboldt and Bonpland, Pl. Aequinoct. 2:75, t. 102. 1810–1811 (“1809”). Trichanthera gigantea Bonpl. ex Steud, Nomencl. ed. 1, p. 708. 1821, nom. illegit. (in syn.). Trixanthera angularis Raf., Sylva Tellur. 146. 1838, nom. illegit.

TYPE.— COLOMBIA. “Habitat frequentissime in sylvis fluvii Magdalenae , prope Badillas et juxta Ybague ,” (fide protologue), without locality or date (specimen), A. Humboldt & A. Bonpland s.n. (lectotype, designated here; see discussion below: P-00719181 !) . Figure 4 .

Clerodendrum verrucosum Splitg. ex de Vriese, Ned. Kruidk. Arch. 1:351. 1848 (as “ Clerodendron verrucosum ”). Besleria verrucosa (Splitg. ex de Vriese) Pulle, Recueil Trav. Bot. Néerl. 9:163. 1912.

TYPE.— SURINAME. “Crescit ad margines fluminum Parae cet. satis frequens,” January 1838, F. Splitgerber 523 (fide Wasshausen, 2006: holotype: L; isotype: P) .

Besleria surinamensis Miq., Linnaea 22:471. 1849.

TYPE.— SURINAME: without locality, Hostmann 764 (lectotype, designated by Wasshausen in 2006: U-image!; possible isolectotypes, see discussion below: MO!, P!, S!) .

Trichanthera gigantea var. guianensis Gleason, Bull. Torrey Bot. Club 54:617. 1927.

TYPE.— GUYANA. East Berbice-Corentyne: Greale, Corentyne River , Oct 1879, G. Jenman 371 (lectotype, designated here; see discussion below: K-image!) .

Shrubs to trees to 15 (–25) m tall and to 30 (–140) cm in diameter (DBH), often suckering from and/or with prop roots at base, sometimes with multiple trunks, branches sometimes long and pendant; young stems densely pubescent with flexuose to antrorse eglandular trichomes 0.05– 0.3 mm long, trichomes sometimes deciduous in patches, mature stems glabrate. Leaves petiolate, petioles to 75 mm long, blades ovate to elliptic 60–310 mm long, 34–150 mm wide, 1.4–2.6 × longer than wide, rounded to acute to subattenuate at base, acute-apiculate to acuminate at apex, surfaces covered with sessile and lenticular glands to 0.05 mm in diameter (sometimes inconspicuous; punctate-glandular), otherwise nearly glabrous and with any trichomes mostly restricted to major veins, margin entire to sinuate. Inflorescence an elongate terminal thyrse or panicle of thyrses, 50–200 mm long, rachis hidden by dense trichomes like those of young stems, dichasia expanded to a greater or lesser degree, pedunculate, peduncles 6–17 mm long, pubescent like rachis, secondary peduncles similar to peduncles. Bracts caducous or persistent, ovate to triangular, 2–5 mm long, 2–3 mm wide, proximal pair(s) sometimes somewhat leaflike and larger, pubescent like rachis. Bracteoles and secondary bracteoles triangular, 2–4 mm long, 1.5– 2.4 mm wide. Flowers pedicellate, pedicels 2–8 mm long. Calyx green or purplish (at least distally), 6–13 mm long during anthesis, tube 1–3 mm long, lobes homomorphic, imbricate, ovate-elliptic to elliptic to oblong to obovate-elliptic, 5–11 mm long, 3.5–7. 3 mm wide, rounded at apex, abaxial surface punctate-glandular and pubescent with antrorse eglandular trichomes, margin ciliate with erect to flexuose eglandular trichomes to 0.7 mm long. Corolla appearing ± glossy where not covered with trichomes, salmon-colored to dark reddish to brownish red to maroon in throat with lobes similarly colored or often tinged with yellow (along margins or distally) or entirely yellowish internally, 27–45 mm long, externally punctate-glandular and distally densely pubescent with appressed eglandular trichomes (such that the obscured surface appears pale) while lacking eglandular trichomes proximally, narrow proximal portion of tube 7–20 mm long, throat (6.5–) 9–20 mm long, 8–16 mm in diameter near midpoint, limb (15–) 20–30 mm in diameter, lobes often reflexed to recoiled, ovate, 6–11 mm long. Stamens 25–35 mm long, filaments salmon-colored to reddish or maroon proximally and often yellowish distally, pubescent (the longer stamen of each pair more densely so) with glandular (sometimes sparse or inconspicuous) and eglandular trichomes, thecae yellowish, 4. 5–7 mm long, pubescent with flexuose eglandular trichomes to 3 mm long and dorsally puberulent with subsessile glands to 0.1 mm long (also on connective), pollen (Daniel et al. 5474) 67–79 µm (polar axis) × 63–70 µm (equatorial axis, apertural face) × 40 µm (equatorial axis, interapertural face). Style reddish, 25–47 mm long, pubescent near base, glabrous distally, stigma with 1 lobe 2–4. 5 mm long, straight to coiled, other lobe vestigial, 0.3– 0.5 mm long. Capsule 16–20 mm long, densely pubescent with antrorsely appressed eglandular trichomes. Seeds 4.5– 4.7 mm long, 4– 4.3 mm wide.

PHENOLOGY.— Flowering throughout the year; fruiting: January–August (and probably other months as well).

DISTRIBUTION AND HABITATS.— Southern Central America ( Panama), northeastern South America ( Brazil, French Guiana , Guyana, Suriname, and Venezuela), and northwestern South America ( Colombia, Ecuador, and Peru)—occurring from the Province of Veraguas in centralwestern Panama (ca. 08°7ʹ49.33ʺN, 080°56ʹ52.89ʺW) southward to the Amazonian lowlands near Belém in northeastern Brazil (ca. 01°26ʹ55.04ʺS, 048°22ʹ45.41ʺW) in the east and to the eastern slope of the Andes in the region of San Martín in north-central Peru (ca. 06°34ʹ7.19ʺS, 076°18ʹ28.88ʺW) in the west ( Fig. 5 ); plants occur in swampy ground and agricultural lands (pastures, cafetales), and especially along streams, fencerows, and roadsides in regions of tropical (dry to) moist to wet, lowland to montane, primary and secondary forests at elevations from sea level to 2250 m (to 3500 m fide Wasshausen 2013). Because Trichanthera gigantea is often cultivated (e.g., as a living fence, or for forage), the anthropogenic distribution of this species is undoubtedly more extensive than that suggested by the localities noted in the specimens cited; collections explicitly indicating that plants were cultivated are not included in that list. The species is especially common (or at least commonly collected) in Colombia. In Venezuela, plants are restricted to two regions on opposite sides of the country—both branches of the Cordillera Oriental in the west and lowlands of the Orinoco delta in the northeast.

The species has been noted to occur as far north as lowlands in Costa Rica ( Leonard 1938; McDade 1983; Durkee 1986). The only two specimens from Costa Rica attributed to this species ( Leonard 1930), both from Guanacaste in the northwestern part of the country (Tilarán, P. Standley & Valerio 46569 at US; Nicoya, Tonduz s.n. in 1900 at US), are sterile, although Standley & Valerio 46569 has immature inflorescences. Both appear superficially similar to Trichanthera gigantea . However, comparison of these plants to those of T. gigantea reveals that they lack the very conspicuous foliar cystoliths, triangular bracts, and feltlike pubescence of the young stems of the latter species. Thus, I remain unconvinced that either of these specimens represents T. gigantea , and Costa Rica is excluded from the known geographic range of this species.

LOCAL NAMES.— Wasshausen (1992), Record and Hess (1943), and Cook et al. (2005) listed the following local names for the species (including those from countries where it is only cultivated): aro blanco, cajeto, cenicero, fune, madre de agua, nacedero, quiebrabarriga/quiebrabarrigo, suiban ( Colombia); tuno ( Guatemala); naranjillo ( Venezuela); palo de agua ( Colombia and Panama); and beque, canella de Garca, pau santo ( Brazil). “Nacedero” appears to have the widest usage in both Colombia, where the plant is abundant, and in agricultural literature concerning Trichanthera gigantea . Indeed, this name and “cajeto” were listed by Bonpland (1810 –1811). Additional local names and their sources from Colombia include: rompebarriga ( Archer 523); arbol de agua, aro, cafetero, cafeto, chumbaguás, cuchiyuyo/cuchuyuyo, güibán, naceró, paloesal, sanantigua, sietenudos, tumbaguás, yátago, and zanca de araña ( Bernal et al. 2013); zanco (Forero & Jaramillo 461); and cajón de fraile (von Sneidern 5698). Other names encountered on herbarium specimens include: canela de velho ( Brazil, Rabelo et al. 2009), curuta ( Venezuela, L. Ruiz Terán 458), janau ( Brazil, Pires 51848), montonero ( Ecuador; Pennington & de la Cruz 10516), naiang ( Venezuela, M. Lizarralde 306), sapote yacu ( Peru; Williams 4894), and watra-oedoe ( Suriname; Werkhoven & v. Troon UVS 16450).

USES.— Leaves and green stems of Trichanthera gigantea have been shown to be a useful forage crop in its native geographic range and elsewhere. Numerous studies have documented its ecological parameters for cultivation, nutritive value as fodder, relative usefulness for various domesticated mammals, and harvest times and yields (e.g., Rosales 1997; Cook et al. 2005). Due to the broad ecological tolerance of this species (e.g., see above for ranges in elevation and biotic communities) and ease of propagation from cuttings, it can be grown in diverse habitats ( Rosales 1997). Outside of its native range, the species has been grown for its agricultural use in other parts of tropical America, southeastern Asia, and Malesia.

Other uses attributed to this species include: windbreak ( Leonard 1930), living fence (e.g., D’Arcy & Sytsma 14473; Wasshausen 1992), and cultivated ornamental ( Wasshausen 2013). Rosales (1997) indicated that “sprouts” of the species are used in maize porridge for human consumption. The wood is considered to be of relatively low quality (i.e., only suitable for unfinished wood products), but it is used for fuel and to make charcoal ( Fern 2014). Wood anatomy of Trichanthera gigantea was studied and described by Williams (1928; based on Pittier 12,056), Carlquist and Zona (1988; based on Forest Products Laboratory, Madison, Wisconsin sample 1117; likely from Williams 4894), and Mennega (2006; Utrecht Wood coll. 175a and 2001). Corothie (1961; without citation of voucher) described wood anatomy of “ Trichanthera sp. ” Wasshausen (1992) noted that woodcutters usually leave plants of T. gigantea standing to protect springs and streams.

Numerous medicinal uses for Trichanthera gigantea have been recorded for humans and domesticated animals. The following uses have been ascribed to humans: infusions of leaves used to treat flu and plants used for treatment of white vaginal discharges (de la Torre et al. 2008; Ecuador); a decoction of leaves used by women in labor to speed delivery (Lescure 2236; Ecuador); “en infusión como colagogo y diurético y en cocimiento como antiflogístico” (Puentes s.n.; Colombia); used as a remedy for fevers ( Wasshausen 1992; Colombia); plants used as a blood tonic, to treat nephritis, and as a lactogenic drink for nursing mothers ( Cook et al. 2005); and green stems used to cure nephritis and roots used as a blood tonic ( Rosales 1997; Colombia). Medicinal uses for domesticated animals include: leaves used to treat hernias (de la Torre et al. 2008; Ecuador); hot poultices of leaves used to cure abdominal hernias of horses ( Wasshausen 1992; Colombia); and plants used to treat colic and hernia in horses, retained placenta in cows, and intestinal obstructions in domestic animals ( Rosales 1997; Colombia).

CONSERVATION.— Trichanthera gigantea is known from more than 200 collections from a broad geographic area (extent of occurrence = ca. 4,591,000 km 2; north-south linear distance = ca. 1,980 km; east-west linear distance = ca. 3,625 km). Even if the EOO is reduced by one-half to two-thirds to account for open water (marine and terrestrial) and large regions from which the species has not been recorded, the geographic distribution of T. gigantea remains quite large, and there would appear to be significant amounts of suitable habitat for this species in much of the region (i.e., the northern and central portions of the Amazon basin) in which it could potentially occur. Based on its wide distribution (both natural and anthropogenic), local abundance (plants are sometimes noted to be common where found, e.g., Piedad R. et al. 40), occurrence in several protected areas in portions of its geographic distribution, and broad ecological amplitude, T. gigantea appears to be a taxon of least concern (LC) based on IUCN criteria ( IUCN 2014).

Flowers of Trichanthera gigantea are reputed to be the major food (nectar) source for the endangered (IUCN category EN) chestnut-bellied hummingbird, Amazilia castaneiventris , the abundance of which appears to be affected by the availability of these trees and which exhibits territorial behavior where flowers are common ( Cavanzo 2011; BirdLife International 2012). Cavanzo (2011) also noted that reproduction of A. castaneiventris was observed in periods of increased floral abundance of T. gigantea .

NOMENCLATURE.— According to the protologue of Ruellia gigantea , the type locality is in the basin of the Río Magdalena in Colombia. Plants were either collected or noted to occur near Badillas (sometimes cited as “Badillo” or “Badilla,” ca. 07°58′20.00″N, 073°51′11.60″W) and in the valley of the Río Combeima near Ibagué at 1300 m (04°27′14.65″N, 075°15′20.15″W). Both Pérez A. (1956) and Wasshausen (1992) noted that Mutis had previously described and illustrated this plant, and that Bonpland had likely made use of these materials. At least five specimens of Trichanthera gigantea resulted from Humboldt and Bonpland’s trip up the Río Magdalena, and it is likely that there were at least two collections (based on numbers noted in handwritten descriptions of the specimens at P). The known extant specimens are discussed below.

Leonard (1930) indicated that the type locality of Trichanthera gigantea was “in sylvis fluvii magdalenae prope Badillas.” He subsequently noted that type material of T. gigantea was at B and K-Hooker ( Leonard 1951), from where specimens had been seen and noted by Nees (1847). The holotype is sometimes cited as having been a specimen destroyed at B ( Wasshausen 1992; Wasshausen 2006). Field Museum photo 5887 from the Berlin Negatives database (emuweb.fieldmuseum.org/botany/berDisplay.php?irn=240003&QueryPage=%2Fbotany%2Fsearch_berlin.php) of the destroyed specimen at B reveals that this specimen lacked collection data (at least on the face of the specimen bearing the plant). This specimen was undoubtedly part of the set of collections given to Kunth by Humboldt prior to his return to Berlin in 1829 ( Hiepko 2006), and included in the general collection at B. It would have been a duplicate of one of the collections at P, and thus an isosyntype. There are no specimens of T. gigantea (under that name or R. gigantea ) in the Willdenow herbarium at B ( Hiepko 1972). Among the major sets of collections of Humboldt and Bonpland ( Hiepko 2006), there are three specimens of T. gigantea at P (where Bonpland began working up the primary set of their collections prior to Willdenow and subsequently Kunth taking on that task). From the specimens and extensive notes of Bonpland (on at least two of three sheets of paper attached to one of the specimens; these two bear extensive descriptions, which generally correspond to information in the protologue, and are apparently based on collections numbered 1545 and 1828), it seems reasonable to assume that the original material for this species is at P. At least one of the specimens at P likely was in Bonpland’s set that was incorporated into the general herbarium at P in 1832 ( Hiepko 2006). Because there are at least two discernible localities in the protologue and undoubtedly at least two different collections, there appear to be syntypes. The specimen from the general herbarium to which all of the descriptive materials are attached, and which is the most complete of those at P, is designated as the lectotype. The other specimens at P, at least one of which could be an isolectoype, are P- Bonpl.-00670081 and P-00719182. A duplicate of one of the Humboldt and Bonpland collections (an isosyntype or an isolectotype) is extant at K.

Neither specimens nor images of types of Clerodendrum verrucosum have been seen. The herbaria of deposit noted above is derived from Wasshausen (2006).

From among the syntypes of Besleria surinamensis (i.e., Kappler 1639 and Hostmann 764), Wasshasuen (2006) designated Hostmann 764 at U as the lectotype. The presumed isolectotypes at MO, P, and S indicate Hostmann’s number as “764a.” Neither the protologue nor the presumed isolectotypes provide place or date of collection. Locality data for the other syntype, Kappler 1639, which was collected in 1844, is noted in the specimens cited below. The lectotype is mounted on the same sheet as a specimen of Kappler 1639 at U.

Gleason (1927) did not designate a type for Trichanthera gigantea var. guianensis . Among the five collections of T. gigantea he cited in the protologue, at least two of them appear to pertain to his new taxon. Leonard (1930) did not indicate a type for var. guianensis . Although Wasshausen (2006) cited Jenman 371 at K as the lectotype of this variety, he did not indicate that it was being designated as such by him and it is not listed among the new lectotypifications made in his treatment ( Wasshausen 2006:163); thus, his indication does not constitute a lectotypification ( McNeill et al. 2012). Although I searched the literature on Acanthaceae in the region in which this taxon was collected, I did not find a lectotypification for this species conforming to the rules of nomenclature. Thus, I have designated Jenman 371 at K as the lectotype of T. gigantea var. guianensis .

MORPHOLOGICAL VARIATION.— Gleason (1927:617) noted that Trichanthera gigantea var. guianensis consisted of lowland plants that differed by their “larger flowers, more hirsute filaments, smaller and more loosely branched inflorescences, and somewhat glaucous leaves, with the veins not elevated on the upper side.” Leonard (1930) provided more precise distinctions (inflorescences 3–8 vs. 5–15 cm long and 2–3 vs. 4–5 cm wide, and filaments pilose throughout vs. filaments pilose proximally and glabrous distally), but noted corollas of the same size as those of the nominate variety. He indicated that var. guianensis occurs in Brazil, Guyana, and Suriname whereas var. gigantea occurs in Colombia, Costa Rica, Ecuador, Panama, Peru, and Venezuela. Bremekamp (1938a, 1938b) did not make reference to this variety in his detailed account of the genus for the Flora of Suriname, and his description noted that filaments of plants from Suriname are densely pubescent proximally but glabrescent distally. Most recently, Wasshausen (2006) recognized var. guianensis for the plants occurring in Guyana, Suriname, and French Guiana. Based on my studies, similar variation in corolla length, filament pubescence, and inflorescence length is evident among plants in the eastern portion of the species’ range ( Brazil, Guyana, Suriname), its western range in South America ( Venezuela, Colombia, Ecuador), and in Panama. There appears to be some geographic tendencies in pubescence of the filaments. For example, 1) all plants from Ecuador have few or no eglandular trichomes distally on the filaments (but this condition is also evident in other parts of the species’ geographic range), and 2) plants from Guyana, Suriname, and Brazil generally have trichomes on the filaments, but these are more abundant in some plants than others (and this variation is also seen elsewhere in the geographic range of the species). Thus, no infraspecific taxa are recognized in this account.

Plants that are somewhat morphologically intermediate with T. corymbosa , at least in some characters, are discussed under that species.

FLORAL BIOLOGY AND POLLINATION.— The following information pertaining to the floral ecology of Trichanthera gigantea is based on personal observations, information noted on herbarium specimens, and published studies. The relatively large, somewhat fleshy corollas ( Fig. 2 ) open in the afternoon (Perez A. 1956; pers. obs., Schmalzel 372, Judziewicz 4493) when nectar is present (Perez A. 1956; Schmalzel 372) and detectable odors absent (pers. obs., Judziewicz 4493, Pennington & de la Cruz 10516, Piedad R. et al. 40). Floral rewards include pollen and nectar. Corollas dehisce and fall from the inflorescence before morning of the next day. The style remains attached to the ovary for at least several days following anthesis; ovaries that do not set fruit fall from the plant within three days ( McDade 1983). Working at a site in Panama, McDade (1983) demonstrated that flowers are neither autogamous nor apomictic (at least not showing autonomous agamospermy), that at least eight pollen grains are necessary for fruit and seed set, and that mean seed set per fruit was very low at this site, probably because of pollen limitation (low vigor of pollen, low numbers of grains deposited on stigmas, or both).

Pollination of Trichanthera gigantea by bats ( Glossophaga soricina ) was documented by Steiner (1981) and photographed by Merlin Tuttle ( Anonymous 1984; image can be viewed at: <http://www.scientificamerican.com/slideshow/bats-in-history-and-world/> and <http://www. wbur.org/npr/181634051/this-bat-knows-how-to-drink?ft=3&f=181634051>). These bats hover in front of flowers, gather nectar with their highly specialized tongues ( Harper et al. 2013), and contact stigmas and/or pollen with the top of their heads. Visitation to flowers by bats was also noted on labels of several herbarium specimens (e.g., Monslave B. 807 from Colombia). Elsewhere among Acanthaceae , bat pollination has been noted for Harpochilus neesianus Mart. ex Nees (Acanthoideae: Justicieae) by Vogel et al. (2004). Flowers that share the characteristic syndrome of floral adaptations associated with bat pollination have been noted for several other Neotropical species of Louteridium (e.g., Vogel et al. 2004) and Ruellia (e.g., Vogel et al. 2004; Ramamoorthy 1991; Tripp 2010).

Other floral visitors to Trichanthera gigantea include red wooly opossum ( Caluromys derbianus ; Steiner 1981), hummingbirds (Pérez A. 1956; McDade 1983; Cavanzo 2011, which show s a photograph of Amazilia castaneiventris visiting and possibly pollinating a flower of T. gigantea ; Henry Stockwell in Panama, pers. comm.; Nee 10446; Tripp & Lujan 520, which indicates hummingbirds as pollinators), large bees (Perez A. 1956; McDade 1983), and ants (Perez A. 1956; Rosales 1997). Some of these floral visitors likely effect pollination as well. Collectors have been particularly attentive to the presence of aggressive ants on inflorescences or flowers of plants (e.g., Almeda & McPherson 6022, Daniel et al. 5475, Haught 4549, Judziewicz 4493, Luteyn & Pipoly 9378, Phillippe et al. 21186). The ants have been described as “swarming,” “stinging,” and “guarding flowers.” Some collections note that the ants make “mudlike” nests in the inflorescences.

ADDITIONAL SPECIMENS EXAMINED (only images were seen for specimens cited from CDMB and COL).— BRAZIL: Amapa: Macapá, Rio Vila Nova , B. Rabelo et al. 2009 ( NY, US). Pará: Belém , igapó do I. A. N., G. Black 826 ( NY, US) ; IPEAN grounds, Belém , A. Gentry & A. Pinheiro 13102 ( MO, US) ; Amazon estuary, Breves , E. Killip & A. Smith 30230 ( F, NY, US) ; Crauateua, Rio Guamá , R. de Lemos Fróes 20392 ( NY, US) ; beira do Guamá, J. Pires 3454 ( CAS, US) ; vic. of Belem, J. Pires 51848 ( NY, US) ; Belém, R. Guamá , N. Silva 57808 ( NY, US). Roraima: Mun. Alto Alegre, Reserva Ecológica de Maracá , N tip of island at Três Igaripés, Corredeira de Rapariga, ca. 1 km upriver from jct. Furo de Santa Rosa with Rio Trairão , P. Edwards 2649 ( C, MO, NY, UPS) ; Canto Galo, Rio Mucajaí between Pratinha and Rio Apaiú, G. Prance et al. 3980 ( CAS, NY, S, US) ; 10 km SE of Serra de Lua, 02°25–29′ N, 060°11–14′W, G. Prance et al. 9336 ( NY, S, US).— COLOMBIA: Antioquia: 0–5 km S of Q. La Tirana along the river, vic. Planta Providencia, 28 km S of Zaragoza , W. Alverson et al. 328 ( MO, NY, WIS) ; 4 km N of Fredonia, W. Archer 523 ( NY) ; Mpio. Frontino, San Andrés, vía Dabeiba – Fuemia , 18–33 kms., 06°40′N, 076°23′W, R. Callejas et al. 5850 ( MO, NY, US) ; Mpio. Chigorodó, 2 km E of Chigorodó, Had. Pasatiempo , 07°40′N, 076°42′W, R. Callejas et al. 9721 ( NY) ; Mpio. Cocorná, La Piñuela, carretera a San Francisco, 06°02′N, 075°08′W, D. Cañas 810 ( MO) ; Mpio. San Luis, camino de la vereda Las “Confusas” a la autopista Medellín–Bogotá, 06°00′N, 074°45′W, D. Cárdenas L. & J. Ramírez 2570 ( COL, MO) ; Parque Nacional Natural “Las Orquideas,” Sector Venados, margen derecha del Río Venados , 06°33′N, 076°19′W, A. Cogollo et al. 3010 ( COL) ; near Río León , ca. 20–30 km upstream and S of river mouth and ca. 15 km W of Chigorodó (ca. 07°45′N, 076°50′W), C. Feddema 1917 ( MICH, NY, US) ; Pavarandó Grande, Río Pavarando , R. Fonnegra et al. 1716 ( MEXU, MO, NY, US) ; Mpio. Río Negro, 7 km from Turbo on road to Necocli, 08°9′0″N, 076°41′48″W, A. Gentry 9223 ( COL, MO, NY, US) ; Turbo , 10 km E of Turbo, O. Haught 4549 ( P, US) ; cerca de Villa Arteaga, F. López & M. Sánchez M. 61 ( NY) ; de Puerto Bélgica por la carretera hasta el Río Man , R. Romero-Castañeda 2333 ( COL) ; Andes, carretera Andes – Jardín, 05°40′N, 075°55′W, D. Sánchez et al. 1155 ( MO) ; vic. of Medellín, R. Toro 78 ( NY) ; Mpio. Carepa , 2 km N of Carepa, 07°52′N, 076°42′W, J. Zarucchi et al. 5005 ( MO, NY) ; Mpio Andes, km 13 of road Jardín – Andes (3 km before Andes), 05°39′N, 075°52′W, J. Zarucchi et al. 7023 ( COL, MO, US). Bolívar: 1 km de Arenal , E. Forero & R. Jaramillo 461 ( NY) ; Sahagun, F. Pennell 4101 ( NY) ; Mpio. Barranco de Loba , corr. El Pueblito, sector Las Payayas, 08°40′N, 074°10′W, F. Roldán et al. 1837 ( NY) ; Cordillera Occidental, Guimarí , K. von Sneidern 5698 ( PH). Boyacá: Mpio. Puerto Boyacá, Inspección de Puerto Boyacá, Quebrada La Cristalina , 05°50′60″N, 074°19′60″W, R. Bernal et al. 2247 ( COL) ; Mpio. Santa María, Arrayanes, Puerto de Agua Caliente, en la ruta a San Luis de Gaceno , cercanias del Río Lengupá , 04°50′57.8″N, 073°13′45.3″W, J. Betancur et al. 11530 ( COL) ; Mpio. Zataquirá, Hormigas , A. Cadena G. 237 ( COL) ; Valle de Soatá , J. Cuatrecasas & H. García B. 1085 (COL). Caquetá: Mpio. San Vicente del Caguán, Trazado de la carretera entre Neiva y San Vicente, Las Perlas, bajo Río Pato , Finca Galicia , J. Betancur et al. 2276 ( COL, MO, US). Casanare: Mpio. El Yopal , J. Campo K. & L. Pinzón P. 208 (COL). Cauca: Mpio. Guapi, Parque Nal. Natural Isla de Gorgona, camino a Playa Blanca , G. Lozano et al. 5641 ( COL). Cesar: Poponte, Magdalena Valley , C. Allen 876 ( MO). Chocó: Mpio. Acandí, Corregimiento San Francisco, Golfo de Urabá , 08°23′N, 077°07′W, J. Betancur et al. 1216 ( MO) ; Mpio. Acandí , corr. Triganá, Reserva Zazardí, 08°20′N, 077°10′W, F. Cardona N. et al. 1641 ( NY) ; Mpio. Quibdó, Corregimiento Bebará, sector La Calle en el Río Bebará , W. Córdoba 411 ( MO) ; Río Chintado, above La Nueva , J. Duke 9850 ( MO) ; Río Yuto between Lloró and La Vuelta, A. Gentry & E. Rentería A. 24340 ( MO) ; Río Mecana , 06°16′N, 077°21′W, A. Juncosa 1745 ( MO) ; Mpio. Quibdó, barrio Bahia Solano , R. Moreno et al. 5 ( MO) ; Río Tolo , región de Guayabal, al SE de Acandí, L. Quiñones et al. 4 ( COL, MO, US) ; Parque Nacional Natural Los Katíos, sector Bijao, S. Zuluaga R. 785 (COL). Córdoba: Mpio. San Antero, Cerro de Buenos Aires , 09º17’48.6ʺN, 075°50’02.9″W, J. Aguirre S. et al. 181 ( COL) ; Mpio. Lorica, Corregimiento Nariño, Quebrada Cardozo , O. Rivera D. et al. 1729 ( COL) ; Mpio. Chima, Corregimiento Sitio Viejo, El Cerro, Cerro Tofeme, 09°04′58″N, 075°35′13″W, O. Rivera D. et al. 2534 ( COL). Cundinamarca: Caqueza in descendu And. bogotens. orienteus versus, E. André 1875 ( P) ; Sierra de Subia, 10 km N of Cumaca on road to Viotá , A. Barclay et al. 3527 ( COL, US) ; Mpio. La Mesa, carretera de La Mesa a Anapoima, El Placer , A. Fernández & L. Mora 1373 ( COL, NY) ; Mpio. Arbeláez, 7 km antes de Pandi, J. Fernández A. et al. 7007 ( COL, NY) ; Mpio. Viotá, La Victoria, Finca El Retazo, J. Jácome 437 ( COL) ; Mpio. Sasaima, Río Agua Dulce , Quebrada Doroga , G. Lozano C. 654 ( COL) ; Mpio. Viotá, Las Palmas, Finca Pensilvania , A. Rodríguez A. 3 ( COL) ; La Mesa, vía La Mesa – San Javier , A. Salama et al. 251 ( COL) ; Mpio. El Colegio, Inspección El Triunfo, La Soledad , desvío por El Quiosco , A. Sanabria G. et al. 388 ( COL) ; Mpio. Nilo, Inspección de Pueblo Nuevo, camino al Cerro del Cualamaná , J. Torres R. et al. 774 ( COL) ; entre La Mesa et le Magdalena, J. Triana s.n. ( NY, P) ; Santandercito, a orillas del Río Bogotá , L. Uribe U. 334 ( COL) ; Pradilla near San Antonio de Tena , J. Wood 4137 ( COL, MEXU, US). Huila: along river, E of San Antonio Fortalecillas , E. Little 7930 ( UC) ; Mpio. La Argentina, El Progresso, 02°12′40.1″N, 075°56′40.1″W, G. Silva et al. GAS0358 A ( COL). Meta: floodplain of Río Metica just E of Puerto López, 3°55’43ʺN, 73°2′44ʺW, G. Davidse & F. Llanos 5470 ( COL, MO, US) ; caños cercanos a Villavicencio, J. Fernández A. et al. 5573 (MO) ; P.N.N. Tinigua, Serranía Chamusa, Centro de Investigaciones Primatológicas La Macarena , P. Stevenson 922 ( MO). Norte de Santander: región de Sarare, hoya del Río Chitagá, en La Cabuya , J. Cuatrecasas 13437 ( COL, US) ; Culagá Valley, near Tapatá ( N of Toledo ), E. Killip & A. Smith 20504 ( NY, US) ; W side of Culagá Valley, N of Labateca , E. Killip & A. Smith 20534 ( NY, US) ; between Chinácota and La Esmeralda, E. Killip & A. Smith 20891 ( US) ; km 20 carretera via Pamplona, D. Villamizar V. 72 (MO). Putumayo: Río Putumayo, Puerto Porvenir, arriba de Puerto Ospina , hacia La Loma , J. Cuatrecasas 10733 ( COL). Quindío: Mpio. Calarcá, La Bella, km 4 vía Calarcá–Barcelona , 04°30′03″N, 075°41′02″W, M. González 141 ( COL) ; Mpio. Filandia, El Placer vía a Filandia , L. Piedad R. et al. 23 ( COL) ; Mpio. Génova, Finca El Janeiro, L. Piedad R. et al. 40 (COL). Santander: 10 km N de Bacaramanga , J. Araque M. & F. Barkley 18 S. 212 ( NY, US) ; Mpio. Suaita, Corregimiento San José de Suaita , zona cercana a la quebrada La Vega, J. Fernández-Alonso et al. 20862 ( COL) ; Mpio. Floridablanca, predios del campus del la Universidad Pontificia Bolivariana , García & J. Benavides 3835 ( CDMB) ; Oiba, F. González 3591 ( NY) ; Bucaramana and vicinity, E. Killip & A. Smith 15452 ( NY, US) ; Río Suratá valley , between El Jaboncillo and Suratá, E. Killip & A. Smith 16426 ( US) ; La Corcova ( Tona ), E. Rentería et al. 692 ( NY) ; alrededores de Oiba, R. Puentes s.n. ( COL) ; Río Servita, vic. Málaga , J. Wood 4338 ( COL). Tolima: El Fresno , J. Cuatrecasas 9374 ( COL) ; Ibagué, I. Holton s.n. in 1853 ( NY) ; Mpio. Ibagué, Tres Esquinas, 04°29′6″N, 075°15′51″W, A. López 4 ( COL). Valle de Cauca: Río San Juan , E. Core 1501 ( US) ; Cordillera Occidental, vertiente occidental, hoya de Albán, entre Quebradita del Retiro y el Río Albán , J. Cuatrecasas 22631 ( F) ; El Frejito, E. Dryander 47 ( US) ; Cali, H. Garcia B. 4314 ( US) ; Mpio. Ginebra, Inspección de Policia, Costa Rica, Mina la Emilia , L. Jiménez et al. 06 ( COL) ; Bajo Calima, Concesión Pulpapel / Buenaventura, 03°55′N, 077°W, M. Monslave B. 807 ( MO, US) ; “La Manuelita,” Palmira , F. Pennell & E. Killip 6193 ( NY) ; cerca de Cali, E. Pérez A. & J. Cuatrecasas 6332 ( COL, US) ; Timba, K. von Sneidern 1240 ( NY).— ECUADOR: Carchi: between Chical and Peña Blancas, valley of Río San Juan on Colombian border, A. Gentry & G. Shupp 26496 ( MO) ; environs of Chical , 12 km below Maldonado on Río San Juan, 01°04′N, 078°17′W, M. Madison et al. 4474 ( F, MO) ; below Maldonado , ca. 80 km W of Tulcan, T. Pennington & R. de la Cruz 10516 ( US) ; Maldonado , L. Werling & S. Leth-Nissen 420 ( NY). Esmeraldas: Limones–Borbón, 5 km before Borbón , 01°07′N, 079°00′W, L. HolmNielsen et al. 26040 ( MO, NY) ; Macedonia, Esmeraldas–Atacames , C. Játiva & C. Epling 466 ( NY, UC, US) ; Atacames, 25 km SW de Esmeraldas, E. Little & R. Dixon 21002 ( NY, US) ; San Lorenzo Cantón, Reserva Etnica Awá, Centro Ricaurte, 01°10′N, 078°32′W, G. Tipaz et al. 2223 ( MEXU). Los Ríos: Río Palenque Biological Station, km 56 Quevedo–Santo Domingo , C. Dodson 5846 ( MO, US). Manabí: El Recreo , H. Eggers 14823 ( MA, US) ; 5 km E de Chone, carretera hacia Santo Domingo de los Colorados, 00°40’S, 080°05′W, D. Neill & M. Asanza 7979 ( MO). Santo Domingo de los Tsáchilas: near Santo Domingo , C. Jativa & C. Epling 530 ( NY, UC, US) ; carretera Quito – Chiriboga – Empalme, entre kms. 75 and 85, 00°15′S, 078°50′W, V. Zak & J. Jaramillo 2326 ( RSA, MEXU, MO, NY, US). Without locality: A. Gilmartin 221 ( MO). — FRENCH GUIANA: Cayenne: Rivière Camopi, en amont du Saut Yaniwé [ca. 03°5′20ʺN, 052°45′11.69ʺW], J. de Granville 2071 ( U). St. Laurent du Maroni: Saut Pierkourou sur le Tampok [ca. 02°49’59.88ʺN, 053°33′0.03ʺW], Moretti 1268 ( P). — GUYANA: Barima-Waini: Anabisi River , J. de la Cruz 1348 ( NY, US) ; Hossororo, near Port Kumaka via Aruka River , 08°10′07″N, 059°48′17″W, T. Hollowell et al. 453 ( MO) ; Barima River , G. Jenman 7037 (K-image, NY). Cuyuni-Mazaruni: Mazaruni Station, Forest Dept. of British Guiana F624 (3360) ( NY) ; Roraima, Schomburgk 998 ( P). Demerana-Mahaica: E bank of Demerara River at Atkinson Field , H. Irwin 167 ( US) ; Canaan, Demerara River, Jenman 5356 ( K, NY). Essequibo Islands-West Demerara: Naamryck Canal , ca. 3.5 km SW of Parika, 06°50′N, 058°27′W, L. Gillespie & D. Gopaul 1042 ( US) ; Naamrye Canal just W of Lookout, 06°50′N, 058°25′W, J. Pipoly & G. Samuels 11752 ( NY, P, US). Potaro-Siparuni: ca. 0.5 km from Paramakatoi, trail to Youwang and Monkey Mt. , 04°41′N, 059°42′W, H. Clarke & S. Grose 1261 ( NY, US). Upper Takutu-Upper Essequibo: Rupununi area, Surama , 04°05′N, 059°04′W, P. Acevedo et al. 3389 ( MO, NY, US) ; Essequibo, South Rupununi savanna, SE of Aishalton, 02°25′N, 059°10′W, T. Henkel & R. James 3723 ( MO, NY, US) ; NW Kanuku Mts., 2–4 km N of Nappi Mt., 03°19′N, 059°33′W, B. Hoffman & R. Foster 3597b ( MO) ; S Pakaraima Mts. , 3 km E of Tipuru village, 04°12′N, 059°32′W, B. Hoffman & R. Jacobs 1147 ( NY, US) ; Rupununi Distr., Kanuku Mts. , Crabwood Cr., Camp 23, 03°07′N, 059°06′W, M. Jansen-Jacobs et al. 3300 ( MO, NY, P, US) ; NW slopes of Kanuku Mts., drainage of Moku-Moku Creek ( Takutu tributary), A. Smith 3405 ( MO, NY, P, US). — PANAMA: “ Canal Zone:” without locality, P. Allen 1725 ( MO, NY) ; without locality, M. Correa A. et al. 1719 ( MO) ; Madden Forest Road 2, T. Croat 8957 ( MICH, MO). Coclé: lower portion of valley along R. Antón, El Valle de Antón , A. Hunter & P. Allen 385 ( MO). Colón: Juan Mina Plantation, Río Chagres, region above Gamboa , P. Allen 4106 ( MO) ; Boyd-Roosevelt Hwy., 5 mi W of Sabanitas, T. Croat 14073 ( MO, NY, RSA, UC, US) ; vic. of Río Indio on road from Portobelo to Nombre de Dios , T. Croat 33620 ( MO, NY) ; along Pipeline Road, 1–5 km NW of Gamboa, 09°08′N, 079°42′W, E. Judziewicz 4493 ( MO, WIS) ; Chagres River , ca. 3 mi above Gamboa Bridge, H. Kennedy et al. 2302 ( MO, RSA) ; Parque Nacional Soberania (Pipeline Road), 3–7 km NW of Gamboa, L. McDade 542 ( CAS) ; along Pipeline Road, 3.5 km NW of Gamboa, M. Nee 10446 ( MO, RSA, US). Darién: Serranía de Pirre, near Cana mining camp in region of Alturas de Nique on road to Boca de Cupe , F. Almeda & G. McPherson 6022 ( CAS, NY) ; vicinity of Canglon, 110 mi from Bayano Dam Bridge , T. Antonio 4578 ( MO) ; trail from Punta Guayabo Grande to Río Jaque , T. Antonio & W. Hahn 4432 ( MO) ; without locality, N. Bristan 124 ( MO, UC) ; Rancho Frio Station, Río Perrecénega , ca. 10 mi E of El Real, W. D’Arcy & G. McPherson 16165 ( MO) ; 10 km NE of Jaque, Río Tabuelitas above Birogueirá, village on Río Jaque below mouth of Río Pavarandó , W. D’Arcy & K. Sytsma 14473 ( MEXU, MO) ; 1–5 mi downstream from El Real, J. Duke 4924 ( MO, UC, US) ; Cerro Piriaque, J. Duke 8110 ( MO) ; 18 km SE of Jaqué, Ensenada del Guayabo , N. Garwood et al. 93 ( MEXU, MO) ; Ensenada del Guayabo , 16–19 km SE of Jaqué, N. Garwood 1201 ( MO) ; Río Tuira between Río Paya and Río Cube, A. Gentry 4354 ( MO) ; El Real, trail to Río Pirre , H. Kennedy 2828 ( F, MO, NY) ; trail from Canglón – Yaviza road to Río Chucanaque , 7.7 mi E of Canglón, 08°20′N, 077°50′W, S. Knapp & J. Mallet 3965 ( CAS, MEXU, MO) ; S of El Real along trail at base of Cerro Pirre, ca. 08°00′N, 077°45′W, G. McPherson 7081 ( CAS, MO) ; Sambú River above tide limit, H. Pittier 5541 ( NY) ; El Real airport, O. Sexton 260 ( MO), 261 ( MO) ; Cerro Piriaque, E. Tyson et al. 3814 ( MO) ; Rio Cocalito , C. Whitefoord & A. Eddy 118 ( MEXU, MO). Emberá: Marraganti and vicinity, R. Williams 1007 ( NY). Herrera: El Barrero de Pesé , M. Rodríguez 53 ( MO, NY). Los Santos: vicinity of Tonosí along Quebrada Ocho Paso tributary of Río Tonosí , W. Stern et al. 1834 ( MICH, MO, US) ; 10 mi N of Tonosí, E. Tyson et al. 2948 ( MO) ; road from Tonosí to Guánico , E. Tyson et al. 3118 ( MO). Panamá: drowned forest of upper Río Pequeni between Salamanca Hydrographic Station and Río Boquerón, P. Allen 17275 ( MICH, MO, P) ; Río Villalobos – Pedregal, D. Botello 26 ( MO) ; Barro Colorado Island , T. Croat 4632 ( NY), 8528 ( MO, NY, RSA, US) ; Río Maje, ca. 30 minutes by speedboat from confluence with Río Bayano , T. Croat 34378 ( MO) ; along Pipeline Road between entrance to Parque Nac. Soberania and fourth bridge, T. Daniel & H. Herrera 5490 ( CAS, MO) ; along road to Farfan Beach just W of Bridge of the Americas , T. Daniel et al. 5474 ( CAS, MICH) ; Barro Colorado Island , R. Dressler 3427 ( MO) ; between Río Pacora and Chepo, J. Dwyer et al. 5124 ( CAS, MO, US) ; Barro Colorado Island , R. Foster 1680 ( F, MO) ; Pipeline Road, 09°15′N, 079°45′W, C. Hamilton et al. 3253 ( MEXU, MO) ; San José Island, Camp Valley , I. Johnston 679 ( MO, P, US) ; along Gaillard Hwy., 1.5 km NW of Summit Garden, M. Nee 9392 ( MO, NY) ; Barro Colorado Island , R. Schmalzel 372 ( MEXU, MO) ; ca. 15 km SW of Cañaza near Río Torti, base of Serranía de Cañazas , 08°52′N, 078°22′W, B. Stein 1365 ( MEXU, MO) ; Barro Colorado Island , W. Stoutamire 2092 ( MICH). Guna ( San Blas ): W side of Loma Armila , W. D’Arcy & G. McPherson 16133 ( MO). Veraguas: Santiago, Loma de Regina , I. Gordon & F. Camarena 43 ( MO) ; Santiago, L. Urriola 30 ( MO). — PERU: Loreto: lower Río Huallaga, Santa Rosa [155–200 m], L. Williams 4894 ( US). San Martín: prope Tarapoto, Peruviae orientalis, R. Spruce 3951 ( K, NY, P, US). — SURINAME: Commewijne: Plant. Liberté , J. Florschütz & P. Florschütz 1007 ( NY). Nickerie: Wageningen I, km 172, M. Werkhoven & F. v. Troon USV no. 16450 ( US). Paramaribo: ad margines sylvarum pr. u. Paramaribo , A. Kappler 1639 ( MO, P, S). Wanica: Domburg, fluv. Suriname inferior, old plantation, K. Kramer & W. Hekking 2349 ( NY) ; bank of Para River , 2–4 km S of Houttuinen, K. Kramer & W. Hekking 2748 ( DAV, NY). — VENEZUELA: Barinas: las afueras de Puntax de Piedras, Bernardi 1099 ( NY) ; ca. 34 km NE of Altamira and 5 km NE of Caldas, ca. 08°55′N, 070°20′W, J. Luteyn & J. Pipoly 9378 ( CAS, MO, NY, US). Delta-Amacuro: medio Río Grande , 60 km NE de El Palmar, ca. 08°25′N, 061°45′W, G. Aymard C. 5378 ( MO) ; Dept. Tucupita, 13–14 km SE of Piacoa, along trail to Río San José , 8°32′N, 62°3′W, G. Davidse & A. González 16453 ( MEXU, MO) ; Dpto. Antonio Díaz, Cano Merejina, 08°46′N, 061°10′W, A. Fernández 3990 ( MO, NY). Falcón: Mpio. Jacura, Distr. Acosta, base del Cerro de La Mina , L. Ruiz T. 458 ( MO) ; Distr. Colina, Río Ricoa , S de Las Dos Bocas, 11°19′N, 069°24–25′W, J. Steyermark & A. Gonzáles 113647 ( MO). Lara: road from Guarico to Chabasaquén , 09°35′8.5″N, 069°50′54.3″W, E. Tripp & M. Lujan 520 ( CAS, RSA). Mérida: Zea, Bernardi 1051 ( MEXU, NY) ; San Cristoval, N. Funck & L. Schlim 1506 ( P) ; Río Caparo , ca. 1 km upstream from dam site, ca. 07°41′N, 071°28′W, R. Liesner & A. González 9393 ( MEXU, MO, US) ; 0.5– 2 km above dam site on Río Guaimaral , ca. 07°45′N, 071°29′W, R. Liesner & A. González 10578 ( MEXU, MO, NY, US) ; carretera Santa Cruz de Mora–El Portón, L. Valverde et al. 1119 ( MO). Portuguesa: Distr. Guanare , terrenos de la UNELLEZ, 09°04′N, 069°49′W, G. Aymard 1030 ( MO) ; 30 km (air) W of Guanare, along Río Tucupido , 09°2′N, 070°01′W, R. Liesner et al. 12460 ( MEXU, MO, NY, US) ; NW of Guanare on Río Guanare, 09°2.55′N, 069°48.38′W, L. Phillippe et al. 21186 ( MO) ; Paso del Guanare , H. Pittier 12056 ( NY) ; Mpio. Mesa de Cavacas, Distr. Guanare , el lecho del Río Guanare , tramo ca. 1 km desde el Puente de la carretera Guano–Barinas , B. Stergios et al. 7898 ( MO, NY) ; Río María, Boca del Monte , 23 km N del vado del Río Suruguapo , 38 km N por la autopista Guanare–Ospino, en el sitio Las Marías, N de Guanare , 09°18′N, 069°43′W, J. Steyermark et al. 127152 ( MO, US). Táchira: near Palo Grande , A. Alston 7097 ( NY). Trujillo: entre Campo Elías y Batatal , C. Benítez de Rojas 976 ( F). Zulia: límite Distr. Bolívar-Baralt, Río Misoa, 10 km SE del empalme de las carreteras Maracaibo– Carora–Valera , G. Bunting & L. Aristeguieta 5175 ( NY) ; Distr. Bolívar, Cuenca de la Represa Burro Negro (Pueblo Viejo), 12 km de la vía Campo Lara – Piedras Blancas–Río Chiquito , G. Bunting et al. 7204 ( NY) ; Sierra de Perijá , 3.8 km 50° of Saimadodyi, 09°36′N, 072°55′W, M. Lizarralde 306 ( MO, NY) ; near Riocito [10°34′59.72ʺN, 072°22′0.06ʺW], C. Mell s.n. in 1923 ( NY).

Figures

FIGURE 4. Trichanthera gigantea. A. Distal portion of branch with inflorescence (Luteyn & Pipoly 9378, CAS). B. Leaf from proximal portion of shoot (McPherson 7081, CAS). C. Dichasium (Daniel et al. 5474, CAS). D. Corolla opened to show stamens (Daniel et al. 5474, CAS). E. Stamen (Daniel et al. 5474, CAS). F. Style and stigma (Daniel et al. 5474, CAS). G. Calyx and capsule (Daniel & Herrera 5490, CAS). H. Capsule (Luteyn & Pipoly 9378, CAS). I. Seed (Daniel et al. 5474, CAS). Drawn by M. Logies.

FIGURE 5. Distribution of Trichanthera gigantea. A. Distribution in Central America (Panama). B. Distribution in South America.

FIGURE 2. Trichanthera corymbosa (A–C) and T. gigantea (D, E).A. Corymbose inflorescence (Romero C. 10753, MO). B. Flower showing calyx with heteromorphic lobes (Pittier 12828, NY). C. Seed (Trujillo & Fernández 16379, US). D. Inflorescence and flower (Daniel et al. 5474). E. Base of tree with suckering (Daniel et al. 5474).