Protium hammelii D.Santam, D. Santam, 2017

Santamaria-Aguilar, Daniel & Lagomarsino, Laura P., 2017, Two new species and a new combination in Protium (Burseraceae) from Costa Rica, PhytoKeys 76, pp. 89-113: 93-99

publication ID

http://dx.doi.org/10.3897/phytokeys.76.10298

persistent identifier

http://treatment.plazi.org/id/E5110F08-D390-56B6-B95A-1F151D716B6C

treatment provided by

PhytoKeys by Pensoft

scientific name

Protium hammelii D.Santam
status

sp. nov.

Protium hammelii D.Santam   sp. nov. Figs 3 View Figure 3 , 4 View Figure 4 , 5 View Figure 5 , 7A View Figure 7

Diagnosis.

Protium hammelii   is similar to Protium multiramiflorum   Lundell and Protium panamense   for their nearly glabrous leaves, petals, and usually pistil or pistillode (always glabrous in Protium panamense   ; sometimes glabrous in Protium multiramiflorum   ). However, the new species it is distinguished from Protium multiramiflorum   by the short calyx in both sexes (0.7-1.3 vs. 1.4-2 mm long), pyrenes with thick walls (0.6-1.1 vs. 0.3-0.5 [-0.6] mm thick), and a short and shallow scar (0.3-0.45 [-0.5] vs. 0.4-0.7 cm long). It is distinguished from Protium panamense   by its smaller lateral (11-22.2 × 3.3-6.4 vs. 16-32.5 × 6.7-10.3 cm) and terminal (11.2-17.7 × 3.9-8.5 vs. 15-28.3 × 7-13.6 cm) leaflets and shorter petiolules.

Type.

COSTA RICA. Limón: Parque Nacional Tortuguero, 5 km N de La Aurora, Guápiles, límite sur del Parque , junto rio Sierpe , 10°22'00"N, 083°31'00"W, 30 m, 11 Apr 1990 (♂ fl), J. Solano 77 (holotype: CR-152860!; isotypes: CR-51496! [ex- INB], F-2 sheets 2081330!, 2127441!, MO-6664125!, NY-01189417, digital image!) GoogleMaps   .

Tree, 4-30 tall × 9-40 cm DBH, sometimes with stilt roots; external bark grayish. Resin transparent when fresh, aromatic. Twigs 2-5 mm diam, appressed-pubescent with simple or malpighiaceous, whitish yellow trichomes 0.1-0.5 mm long, to glabrescent, sparsely lenticellate, solid, never white-stained with resin that crystallizes on the stem (except weakly in W.D. Stevens 31653; also on the fruits). Leaves (2-) 4-6 jugate, 21.5-43.5 cm long; petiole (2.7-) 4.7-7.4 (-8.2) cm long, 0.1-0.3 cm diam, semi-terete except weakly sulcate at the base, striate; rachis 4-8.2 (-9) cm long or absent, terete, striate on both sides; petiole and rachis nearly glabrous or sparsely pubescent with simple and malpighiaceous, usually whitish yellow trichomes 0.1-0.3 mm long; lateral petiolules 0.7-2.1 cm long, with pulvinuli evident on both ends, striate, canaliculate adaxially, glabrous or minutely pubescent with whitish yellow trichomes; terminal petiolule 1.8-4.3 (-5.5) cm long, pulvinulus conspicuous; basal pair of leaflets 10.1-19 × 3.7-6.7 cm, ovate or lanceolate, obtuse to subcuneate at the base (sometimes asymmetric); other lateral leaflets 11-22.2 × 3.3-6.4 cm, lanceolate, oblong, ovate or elliptic, obtuse to subcuneate at the base; the terminal 11.2-17.7 × 3.9-8.5 cm, broadly elliptic to obovate, lanceolate, obtuse or subcuneate at the base (usually with both sides equal); apex acuminate, the acumen 0.7-1.3 cm long; margin entire or much more commonly sparsely denticulate; leaflets drying more or less pale brown, olivaceous to amber on both sides; secondary venation eucamptodromous or slightly brochidodromous, secondaries in 12-17 pairs of secondary veins, ascending, weakly arcuate, sometimes discolored on abaxial side, ascending, the spacing irregular, perpendicular intersecondary veins often 1 per pair of successive secondary veins or absent, intercostal tertiaries alternate or mixed percurrent tertiary; on abaxial side the midrib prominent, glabrous or minutely pubescent, with trichomes ca. 0.03-0.6 mm long, whitish yellow or reddish, secondary veins prominent, with trichomes similar to the midrib or glabrous, the higher-order veins prominulous with scattered trichomes or glabrous, the rest of surface almost glabrous to glabrous, not papillate; on adaxial side, the midrib prominent, minutely and scattered pubescent, secondary veins impressed to flat with trichomes similar to the midrib or glabrous, the higher-order veins flat to lightly impressed, with scattered trichomes, the rest of surface with scattered trichomes to glabrous. Inflorescences axillary (sometimes at leafless nodes), generally branching at the base, not flexuous, the staminate inflorescences 3.4-8.5 cm long, shorter or exceeding the petiole, minutely pubescent with simple and malpighiaceous, whitish trichomes on all axes, the pistillate inflorescences 1-1.8 cm long [(1.4)- 2.5-12.5 cm in fruit], much shorter than the petiole; bracts subtending the inflorescences 1-1.7 mm long, triangular, acuminate at the apex, densely pubescent abaxially; those on primary axes 0.6-1.3 mm long, triangular, acuminate or obtuse at the apex, densely pubescent abaxially; bracteoles subtending flowers 0.3-0.8 mm long, triangular or broadly ovate, obtuse to acuminate at the apex, pubescent or nearly glabrous abaxially. Flowers 4(5)-merous, the pedicel 1.5-2.8 mm long (2-6 mm in fruit), generally glabrous. Staminate flowers with calyx 0.7-1.3 × 1.3-2.4 mm, much taller than the disk, the lobes 0.4-1 mm long, rounded to depressed-deltoid, glabrous on abaxial side, papillate marginally, often persistent in fruit; petals variously reported as green, greenish yellow or white, 3-4 × 1.4-1.8 mm, distinct, erect to suberect at anthesis, lanceolate or ± triangular, glabrous on the abaxial side, glabrous but papillose on the adaxial side, papillose and weakly involute marginally, inflexed-apiculate at the apex (the apiculum 0.15-0.25 mm long); disk 0.23-0.46 tall × 0.26-0.5 mm thick, glabrous; stamens 8, (sub) equal, the antesepalous 1.8-2.4 mm long, the antepetalous 1.5-2 mm long, exceeding the pistillode, the filaments more or less flat, papillate, the anthers 0.7-0.9 mm long, lanceolate, obtuse to subcordate at the base, apiculate at the apex; pistillode 0.5-0.83 × 0.5-0.8 mm at the base, ovoid, globose or conical, glabrous, the style 0-1.5 mm long, stigma lobes 4, subglobose to weakly angulate, densely papillose. Pistillate flowers with calyx 1-1.2 × 2-2.5 mm, the lobes ca. 0.8 mm long, all parts similar to that of the staminate flowers; petals green, greenish yellow or white, distinct, 3-3.5 × 1.16-1.5 mm, suberect to reflexed at anthesis similar to those of the staminate flowers; disk 0.4-0.5 tall × ca. 0.3 mm thick, glabrous; staminodes 8, (sub) equal, the antesepalous 1.5-1.7 mm long, the antepetalous 1.3-1.5 mm long, shorter or longer than the pistil, the filaments flat, not papillate, the anthers 0.6-0.8 mm long, lanceolate, cordate at the base; pistil 1.16-1.3 × 1-1.23 mm (at the base), ovoid or conical, glabrous, the style 0.3-0.8 mm long, stigma lobes-4, globose, densely papillose, each lobe sulcate on the middle ( ± as an inverted “C”). Fruits 1.6-2.4 × 1.1-2.1 cm, globose to ovoid, reddish or green (M. Ballestero 71) when ripe, obtuse at the base, the apex generally conspicuously apiculate at the apex, smooth or (more commonly) lenticellate, glabrous, stipitate [the stipe (0.1-) 0.2-0.5 cm long]; pseudoaril white; pyrene 1(2), 1.3-1.6 × 0.9-1.2 cm, smooth, ovoid to very widely ovate, obtuse at the base, apiculate at the apex, the wall 0.6-1.1 mm thick, whitish or yellowish; funicular scar 0.3-0.45 (-0.5) cm long, usually not very deep, without a rib in the middle or the rib inconspicuous.

Habitat and distribution.

This species is known so far only from wet forest on the Caribbean slope of Nicaragua and Costa Rica. It occurs mainly between 10 and 200 m in elevation, although some collections were made between 300 and 700 m. In Costa Rica, this species is common in the Sarapiquí region, where it seems to prefer alluvial soils on flat or relatively flat ground, sometimes on river banks. In this area, Protium hammelii   grows sympatrically with the following species: Carapa guianensis   Aubl. ( Meliaceae  ), Dipteryx panamensis   (Pittier) Record & Mell ( Fabaceae  ), Euterpe precatoria   Mart. ( Arecaceae  ), Minquartia guianensis   Aubl. ( Coulaceae  ), and Pentaclethra macroloba   Kuntze ( Fabaceae  ), among others species (N. Zamora, pers. comm.; May 2016).

Phenology.

In Nicaragua, Protium hammelii   has been collected with fruits in January, February, from May to July, and in October, but never in flower. In Costa Rica, it has been collected with staminate flowers in January, February, April, and December; pistillate flowers in January and February; and fruits in January, March, April, June, from August to October, and in December.

Common name.

Alcanfor (Spanish; Nicaragua, R. Rueda et al. 2642, 2701; J.C. Sandino 3424).

Etymology.

The specific epithet honors Barry E. Hammel, curator at the Missouri Botanical Garden and co-editor of the Manual de Plantas de Costa Rica, in recognition of his extensive work on the Costa Rican flora, as well as his personal support and motivation.

Discussion.

Protium hammelii   is recognizable by its almost glabrous vegetative parts, leaves with 5-7 leaflets, commonly with a sparsely denticulate margin, prominent tertiary veins on the abaxial side, 4(5)-merous flowers with glabrous petals, pistils, and pistillodes, the pistillate flowers with globose stigma lobes that are sulcate in the middle, and glabrous, usually lenticellate fruits that are stipitate and apiculate at the apex. Specimens of Protium hammelii   have frequently been identified as Protium glabrum   (Rose) Engl., a species that is widespread from Belize to Panama, or Protium panamense   , from Costa Rica, Panama, and Colombia. The first of these is common in Costa Rica, while the second is quite rare (Fig. 6A-F View Figure 6 ); both have leaflets that are always entire. Protium hammelii   differs from Protium glabrum   by its consistently glabrous petals on both sides, pistil, and pistillode (vs. petals on the adaxial side, pistil, and pistillode always pubescent). Fruiting material can usually be distinguished by the apiculate apex of the fruits of Protium hammelii   (vs. obtuse or rounded), and the glabrous (vs. with very small trichomes). Protium panamense   shares occasional stilt-roots, glabrous flowers and fruits that are apiculate at the apex with Protium hammelii   , but Protium hammelii   usually has smaller lateral (11-22.2 × 3.3-6.4 vs. 16-32.5 × 6.7-10.3 cm), and terminal leaflets (11.2-17.7 × 3.9-8.5 vs. 15-28.3 × 7-13.6 cm) that are also usually thinner, as well as thinner lateral and terminal petiolules that are also shorter (1.8-4.3 [-5.5] vs. 5.2-8.7 cm); Protium hammelii   further has smaller (1.6-2.4 × 1.1-2.1 cm), globose to ovoid fruits (vs. usually 2.2-3 × 0.9-1.7 cm lanceolate. Protium multiramiflorum   from Mexico to Honduras is similar to Protium hammelii   in its nearly glabrous leaves, petals, and sometimes pistil or pistillode. (Although the pistil of Protium multiramiflorum   was originally described as glabrous [ Lundell 1937], almost all collections studied, including one of the isotypes [Lundell 6212, GH-2 sheets!] and the paratype [Schipp 1021; A!, F!, GH!, MO!], have the pistil or pistillode with tiny, scattered trichomes). The new species differs in its short calyx in both sexes (0.7-1.3 vs. 1.4-2 mm long), stigma lobes that are sulcate (vs. not sulcate), and pyrenes with thick walls (0.6-1.1 vs. 0.3-0.5 [-0.6] mm thick) and short scar (0.3-0.45 [-0.5] vs. 0.4-0.7 cm long) (Fig. 7B View Figure 7 ). Others species in Costa Rica with glabrous pistils or pistillodes are Protium aracouchini   (Figs 6G-J View Figure 6 ), Protium ravenii   (Figs 6K-N View Figure 6 ), and Protium aguilarii   . The first two species can be distinguished from Protium hammelii   by their flexuous inflorescences and twigs and abundant exuding resin that becomes whitish and chalky (Figs 6H and L View Figure 6 , inset), while Protium aguilarii   can be distinguished by its petals that are pubescent abaxially.

Fruit dispersal by birds and mammals has been reported at the La Selva Biological Station for Protium panamense   ( Vargas 2000), but the observation likely corresponds to Protium hammelii   .

Additional material examined.

NICARAGUA. Atlántico Norte [Zelaya]: Reserva Bosawas, Mpio. de Bonanza, Cerro Cola Blanca, entre el cacerío de Vitinia y empalme de la Comarca de Panamá, 14°04'N, 084°34'W, 200 m, 02 Jun 1997 (fr), R. Rueda & I. Coronado 6595 ( MO). Atlántico Sur [Zelaya]: área de la Bahía de Bluefields, río Escondido, camino entre El Pool y Abardeen Hills, 0-30 m, 22 Mar 1949 (st), A. Molina 1899 (F); Monkey Point, Caño El Pato, 1.5 km sobre la ribera del Caño, 11°35'N, 083°42'W, 10 m, 25 Oct 1981 (fr), P.P. Moreno 12411 ( MO); Caño Montecristo, al este del Campamento Germán Pomares, 11°36'N, 083°52'W, 60-90 m, 08 Feb 1982 (fl bud), P.P. Moreno 15132 ( MO); Mpio. de Rama, Loma Buena Vista, 12°08'N, 084°12'W, 100-150 m, 23 May 1984 (st), W. Robleto 613 ( MO); a lo largo del río Maíz, 11°16'N, 084°07'W, [25-50 m], 08 Jan 1995 (fr), R. Rueda et al. 2642 ( MO); Mpio. Laguna de Perla, río Wawanshang, 12°40'N, 083°42'W, 50 m, 15 Feb 2002 (fr), R. Rueda & R. Dolmus 16851 ( MO); 1 km de Colonia Serrano, río Serrano, 11°34'N, 084°22'W, 70-80 m, 31 Jul 1982 (fr), J.C. Sandino 3424 ( MO). Chontales: Along road from Ciudad Sandino toward El Guabo, 0.7 km SW of El Porvenir, 12°09'02"N, 084°52'43"W, 345 m, 17 May 2011 (fr), W.D. Stevens 31653 ( MO). Matagalpa: Falda norte del Cerro Musún, frente a trocha a Wanawás, [13°02'N, 085°15'W], 200-500 m, 16 May 1980 (fr), M. Araquistain & P.P. Moreno 2789 ( MO). Río San Juan: Mpio. de San Juan del Norte, del Delta 1 km al este y después 2 km al norte, 10°46'N, 083°46'W, [40-80 m], 08 Jun 1995 (st), R. Rueda et al. 2701 ( MO). Costa Rica. Alajuela: Los Chiles, Finca La Urraca, Los Lirios, camino a los Chiles, ca. 100 m, 11 Dec 1985 (fr), J. Gómez-Laurito et al. 10998 ( CR, F, USJ). Heredia: Sarapiquí, OET, La Selva, 14 Jun 2004 (fr), R. Aguilar et al. 8337 ( LSCR-digital image); Parque Nacional Braulio Carrillo, frente al Puesto La Ceiba, 10°19'47"N, 084°04'48"W, 400-700 m, 23 Dec 1988 (fr), M. Ballestero 71 ( CR, MO); about 5 km north of Puerto Viejo along the road to El Muelle, 10°28'N, 083°58"W [10°30'36"N, 084°00'36"W], 100 m, 08 Jan 1967 (♂ fl), W.C. Burger & G. Mata 4307 (F-2 sheets, MO); about 5 km north of Puerto Viejo along the road to El Muelle, 10°28'N, 083°58"W [10°30'36"N, 084°00'36"W], 100 m, 08 Jan 1967 (♀ fl, fr), W.C. Burger & G. Mata 4315 (F, MO, NY-digital image); Finca La Selva, the OTS Field Station on the río Puerto Viejo just E of its junction with the río Sarapiquí, Southwest trail, 1600 m line, 100 m, 16 Feb 1981 (♂ fl), J.P. Folsom 8965 (F, MO); Finca La Selva, the OTS Field Station on the río Puerto Viejo just E of its junction with the río Sarapiquí, central trail to Holdridge Trail 3000 m line, 100 m, 08 Mar 1981 (immat fr), J.P. Folsom 9279 ( MO); Finca La Selva, the OTS Field Station on the río Puerto Viejo just E of its junction with the Río Sarapiquí, junction South Boundary and Western Boundary, 21 Mar 1981 (immat fr), J.P. Folsom 9435 ( CR); Finca La Selva, the OTS Field Station on the río Puerto Viejo just E of its junction with the Río Sarapiquí, El Swampo Trail, 100 m, 27 Apr 1981 (fr), J.P. Folsom 9882 (F, MO, NY-digital image); Magsasay, near La Selva, 10°24'N, 084°03'W, 150 m, 16 Jul 1990 (st), A.H. Gentry 71766A ( MO); La Selva, río Sarapiquí near Puerto Viejo, junction SSO and LOC Trails, 10°26'N, 084°01'W, 100 m, 05 Jan 1993 (all st), A.H. Gentry et al. 78485 ( CR, MO), 78507 ( CR, MO), 78522 ( CR, MO), 78543 ( CR, MO); La Selva, río Sarapiquí near Puerto Viejo, junction SSO and LOC Trails, 10°26'N, 084°01'W, 08 Jan 1993 (st), A.H. Gentry & R. Ortiz 78630 ( CR, MO); Parque Nacional Braulio Carrillo, fila Carrillo, 700 m, 30 Mar 1984 (fr), L.D. Gómez et al. 21135 ( CR); Parque Nacional Braulio Carrillo, estación El Ceibo, 10°20'00"N, 084°04'00"W, 450-500 m, 13 Mar 2003 (fr), J. González 3147 ( CR); Parque Nacional Braulio Carrillo, estación El Ceibo, 10°20'00"N, 084°04'00"W, 450-500 m, 14 Mar 2003 (immat fr), J. González 3176 ( CR, MO, USJ); Chilamate, Cerros de Sardinal, finca propiedad de Isaias Alvarado, 100 m, 26 Aug 2007, J. González et al. 9326 ( LSCR-digital image); Finca La Selva, Arboretum tag #522, 10°26'N, 084°01'W [10°25'53"N, 084°00'13"W], [40 m], 30 Dec 1970 (♂ fl bud), G.S. Hartshorn 968 (F, MO-2 sheets); Finca La Selva, 10°26'N, 084°01'W, [100 m], 25 Jan 1973 (fr), G.S. Hartshorn 1108 ( CR); Finca La Selva, Holdridge Arboretum tag #119, 10°26'N, 084°01'W, [100 m], 22 Aug 1975 (fr), G.S. Hartshorn 1476 ( CR, F, LSCR-digital image, MO); Finca La Selva, the OTS Field Station on the río Puerto Viejo just E of its junction with the río Sarapiquí, quebrada El Sura, 100 m, 06 Jun 1984 (fr), B. Jacobs 2148 ( CR); Finca La Selva, the OTS Field Station on the río Puerto Viejo just E of its junction with the río Sarapiquí, camino circular Lagano, at bridge across Q. [Quebrada] Salto, 100 m, 15 Jun 1928 (fr), B. Jacobs 2362 (F); Finca La Selva, the OTS Field Station on the río Puerto Viejo just E of its junction with the río Sarapiquí, vicinity Sendero Jagarunda and Lindero Sur intersection, [10°25'53"N, 084°00'13"W], 100 m, 28 Jun 1984 (fr), B. Jacobs 2644 ( MO); río Sarapiquí, 125 m, 18 Jan 1966 (fl bud), A. Jiménez 3602 (F-2 sheets, CR); Finca Hermanos Vargas, 1 km al Suroeste de Puerto Viejo, 125 m, 04 Feb 1966 (♂ fl), A. Jiménez 3602 ( CR); Estación Biológica La Selva, LOC 600, 10°25'47"N, 084°01'00"W, 55 m, 29 Jul 2004 (st), S. Letcher 77 ( USJ); Finca La Selva, the OTS Field Station on the río Puerto Viejo just E of its junction with the río Sarapiquí, Far Loop Trail at about 350 m, [10°25'53"N, 084°00'13"W], 100 m, 11 Feb 1996 (♂ fl bud), R.L. Wilbur 65080 (F, MO); Parque Nacional Braulio Carrillo, El Ceibo, 10°22'29"N, 084°02'10"W, 200-300 m, 24 Aug 2004 (fr), R. Kriebel et al. 4857 ( CR); Estación Biológica La Selva, 10°26'00"N, 084°00'30"W, 0-100 m, 05 Feb 2004 (st), A. Rodríguez 8395 ( CR, USJ); OET La Selva, a orillas del río Sarapiquí, [10°25'53"N, 084°00'13"W], [100 m], 30 Oct 2005 (fr), N. Zamora 3871 ( LSCR-digital image, MO). Limón: Bosque Lluvioso [finca propiedad de INBio], 10°11'28"N, 083°51'28"W, 350 m, 16 Aug 2005 (st), L. Acosta et al. 3550 ( CR); Pococí, 300 m al sur del Hotel Vista Al Mar, Tortuguero, 10°35'51"N, 083°31'40"W, 10 m, 22 Oct 2011 (fr), M. Argueta 107, 109 ( USJ); Guápiles, La Leona, 10°09'45"N, 083°49'37"W, 478 m, 31 May 2005 (fr), C. Benavides & A. Chacón 160 ( USJ); North shore of the mouth of the río Colorado at Barra del Colorado, 10°47'40"N, 083°35'30"W, 1-5 m, 12 Sep 1986 (fr), G. Davidse & G. Herrera 30879 ( CR, F, MO); Guápiles, Cariari, El Zota, Finca El Progreso, 10°30'35"N, 083°44'39"W, 40 m, 11 Jun 2011 (fr), M. Díaz s.n. (USJ-99607); Parque Nacional Tortuguero, Cerro Tortuguero, 10°35'37"N, 083°31'31"W, 5-120 m, 22 Oct 2011 (fr), J. Gómez-Laurito 15689 ( USJ); near río Parismina, 8 km W of Dos Bocas, [10°14'46"N, 083°27'21"W], 8 m, 31 Mar 1972 (fr), R.W. Lent 2457 ( CR, F, MO); Matina, Colonia Puriscaleña, Sendero Cerro Azul, 09°59'44"N, 083°23'08"W, 400-500 m, 15 Mar 2000 (fr), E. Mora 984 ( CR); Parque Nacional Tortuguero, Estación Agua Fría, rumbo Noreste, a orillas del río Agua Fría, 10°27'N, 083°33'W, 40 m, 02 Feb 1988, R. Robles 1586 ( CR, MO, NY-digital image); Llanura de Santa Clara, Chiporrisito, 10°36'10"N, 083°47'20"W, 400 m, 30 Jan 1995 (fl bud), A. Rodríguez 507 ( INB); Parque Nacional Tortuguero, Estación Agua Fría, Sendero El Aguacate, a 500 m de la entrada, 10°26'40"N, 083°34'40"W, 20 m, 11 Jan 1990 (fr), J. Solano 62 ( CR, MO, NY-digital image); Caño Chiquero, Tortuguero, 31 Jan 1986 (♂ fl), R. Soto 2758 ( CR); Cerro Coronel, E of Laguna Danto, 10°41'N, 083°38'W, 20-170 m, 16-13 Jan 1986 (♂ fl), W.D. Stevens 23769, 23770 ( CR, MO); Cerro Coronel, along río Colorado at and below outflow of Laguna Danto, 10°42'N, 83°39'W, 5-10 m, 25 Jan 2016 (fr), W.D. Stevens 24002 ( CR, MO); Cerro Coronel, E of Laguna Danto, 10°41'N, 083°38'W, 16 Mar 1987 (st), W.D. Stevens et al. 24912 ( MO); Monte Verde, 300 [ft?], 25 Apr 1928 (fr), H.E Stork 1682 (F); Zapota Dos, ca. 20 NW of Tortuguero village, on farm of Ronulfo Vargas, 10°38'N, 083°41'W, 90-110 m, 16 Mar 1995 (st), K. Thomsen 1377 ( CR); Parque Nacional Tortuguero, Agua Fría, 10°26'35"N, 083°34'38"W, 32 m, 14 Jun 2007 (fr), L.D. Vargas et al. 2379 ( CR); Parque Nacional Tortuguero, Agua Fría, 10°26'20"N, 083°34'47"W, 30 m, 18 Oct 2007 (fr), L.D. Vargas et al. 2813 ( CR).