Research Article |
Corresponding author: Daniel Santamaría-Aguilar ( daniel.santamaria366@gmail.com ) Academic editor: Pavel Stoev
© 2017 Daniel Santamaría-Aguilar, Laura P. Lagomarsino.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Santamaría-Aguilar D, Lagomarsino LP (2017) Two new species and a new combination in Protium (Burseraceae) from Costa Rica. PhytoKeys 76: 89-113. https://doi.org/10.3897/phytokeys.76.10298
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Two new species of Protium (Burseraceae) are described and illustrated: Protium aguilarii sp. nov., from the Pacific slope of the Osa Peninsula, Puntarenas Province, Costa Rica; and P. hammelii sp. nov., from wet forests on the Caribbean slopes of Nicaragua and Costa Rica. In addition, Protium brenesii comb. nov., is proposed as a new combination based on Trichilia brenesii, a name that was based on a specimen collected with flowers in the mountains near San Ramón, Alajuela Province, Costa Rica. It is compared with P. costaricense, a similar species with which it has been confused for more than 90 years. Finally, illustrations and specimen citations are provided for all the aforementioned taxa, and some others with which they have been confused.
Se describen e ilustran dos nuevas especies de Protium (Burseraceae): Protium aguilarii sp. nov., de la vertiente del Pacífico en la Península de Osa, provincia de Puntarenas, Costa Rica; y P. hammelii sp. nov., de los bosques húmedos de la vertiente del Caribe en Nicaragua y Costa Rica. Además, se propone la combinación Protium brenesii comb. nov., basada en Trichilia brenesii, un nombre que fue descrito en base en un ejemplar con flores recolectado en las montañas de San Ramón, provincia de Alajuela, Costa Rica. Se compara con P. costaricense, especie similar, con la cual se confundió por más de 90 años. Finalmente, se proveen ilustraciones y listas de los ejemplares examinados para todos los taxones antes mencionados, y además algunos otros similares.
Burseraceae , Costa Rica, Nicaragua, Protium , Sapindales , taxonomy
The genus Protium (Burseraceae), with approximately 160 species, is nearly pantropical in distribution, though absent from continental Africa. Twelve species (including those described in this paper) have been recorded from Costa Rica, making this the largest of the five native genera in the country. Protium is distributed in Costa Rica from sea level to 1500 m, mainly in humid lowland forests, though some species occur in montane forest or (more rarely) relatively dry areas [e.g., P. tenuifolium subsp. sessiliflorum (Rose) D. M. Porter on the Pacific slope]. The genus is characterized in general by its arborescent or less often shrubby habit; resin that is usually aromatic; imparipinnately compound, trifoliolate, or rarely unifoliolate leaves, with petiolules that are commonly pulvinulate at both ends, (3) 4–5-merous flowers with distinct or weakly connate petals and 8–10 stamens inserted outside the base of the nectary disk, and dehiscent fruits with 1–5 pyrenes.
The two new species described below and the need for a new combination in Protium were discovered during preparation of the Burseraceae treatment for the Manual de Plantas de Costa Rica. This study was based on an examination of Protium specimens deposited at A,
Protium aguilarii most closely resembles P. costaricense (Rose) Engl. and P. pilosissimum Engl., for their leaves with petiole, rachis, and leaflets that are hispidulous on the abaxial side and leaflets of comparable in size and coloration in herbarium material, but differs from both for its glabrous pistil and fruits (vs. pubescent).
Protium aguilarii. A Branch with inflorescences B Venation C Pubescence on the leaf rachis and petiolules D Inflorescences E Female flower, with perianth partially removed F Fruits. A–D from R. Aguilar 4593,
Protium aguilarii. A Trunk and bark B Twigs C Branch with leaves D Leaves showing abaxial side of leaflets E Leaf showing adaxial side F Leaflet bases G Venation H Flower. Photo credits: Reinaldo Aguilar (A–G) from D. Santamaría et al. 9851; and Xavier Cornejo (H) from R. Aguilar & X. Cornejo 11115.
COSTA RICA. Puntarenas: Reserva Forestal Golfo Dulce, Osa Peninsula, Rancho Quemado, ca. 15 km W of Rincón, on forested slopes at NW end of valley, near Fila Ganado, 08°33'N, 083°34'W, 300–400 m, 29 May 1988 (fr), B. Hammel, G. Herrera, M. M. Chavarría & Á. Solís 16885 (holotype:
Tree, 8–15 m tall × 6–18 cm DBH, lacking stilt roots; external bark white (D. Santamaría et al. 9851). Resin transparent when fresh, a little sticky, aromatic. Twigs 2–4 mm diam, appressed-pubescent with simple or malpighiaceous, pale brown trichomes 0.05–0.6 mm long, sparsely lenticellate, solid, never stained white. Leaves 2–4-jugate, (12.5–) 19.5–33.8 cm long; petiole (2.3–) 3.3–6.3 cm long, 0.2 cm diam, semi-terete, smooth or slightly striate; rachis (2.6–) 3.5–11.8 cm long, terete, smooth or slightly striate; petiole and rachis hispidulous with simple, pale brown trichomes 0.1–0.5 mm long; lateral petiolules 0.3–1.7 cm long, with pulvinuli evident on both ends, smooth or slightly striate on both sides, rounded, hispidulous with yellowish brown trichomes; terminal petiolule 1.9–4.1 cm long, pulvinulus conspicuous; basal pair of leaflets 5.2–10 × 2–4.3 cm, elliptic to ovate, obtuse at the base; other lateral leaflets 6.7–13.2 × 2.6–5 cm, elliptic to ovate, obtuse to subcuneate at the base (with one side sometimes asymmetric); the terminal leaflet 7.3–14.2 × 2.7–5.8 cm, ovate, elliptic, base obtuse to subcuneate and symmetrical; apex acuminate, the acumen 0.7–1 cm long; margin entire; leaflets drying dark brown or olivaceous above and pale brown or olivaceous below; secondary venation brochidodromous, secondaries in 7–10 pairs of secondary veins, ascending, weakly arcuate, the spacing irregular, perpendicular intersecondaries sometimes present, intercostal tertiaries alternate or mixed percurrent tertiary; on abaxial side the midrib prominent, dense hispidulous, with trichomes 0.06–0.55 mm long, yellowish brown, the secondary veins prominent, with trichomes similar to the midrib, the higher-order veins prominulous, with scattered trichomes, the rest of the surface with scattered trichomes, not papillae; on adaxial side the midrib lightly prominent to flat, hispidulous, secondary veins flat, scattered hispidulous to almost glabrous, the higher-order veins flat almost glabrous, the rest of surface nearly glabrous. Inflorescences fasciculate, axillary (sometimes at leafless nodes), staminate inflorescences unknown, pistillate inflorescences ca. 0.6 cm long (0.9–1.2 cm in fruits), much shorter than the petiole, branching at the base, not flexuous, all axes densely pubescent with simple, yellowish brown or whitish trichomes; bracts subtending the inflorescences ca. 1.3 mm long, lanceolate, acuminate to obtuse at the apex, densely pubescent abaxially; those on primary axes ca. 0.6 mm long, broadly ovate, obtuse at the apex, densely pubescent abaxially; bracteoles subtending flowers ca. 0.8 mm long, triangular, acuminate at the apex, pubescent abaxially. Flowers 4-merous, the pedicel ca. 2.2 mm long (ca. 3 mm in fruit), sparsely pubescent with trichomes ca. 0.2 mm long. Staminate flowers unknown. Pistillate flowers with calyx 1–1.16 × 2.3 mm, sparsely pubescent on abaxial side with trichomes ca. 0.1 mm long, the lobes ca. 0.4 mm long, rounded to depressed-deltate, much taller than the disk, often persistent in fruit; petals white, ca. 2.6 × 0.9 mm, distinct, suberect at anthesis, lanceolate, sparsely appressed-pubescent on abaxial side with yellowish brown to whitish trichomes ca. 0.1 mm long, glabrous but papillose on the adaxial side, papillose and involute marginally, inflexed-apiculate at the apex (the apiculum ca. 0.2 mm long); disk ca. 0.4 tall × 0.2 mm thick, glabrous; staminodes 8, 1.66–1.8 mm long, the antepetalous almost equaling than antesepalous, the filaments ca. 0.9 mm long, flat, the anthers ca. 0.6 mm long, lanceolate, subcordate at the base; pistil ca. 1.5 × 1 mm (at the base), ovoid, glabrous, the style ca. 0.8 mm long, stigma lobes 4, globose, densely papillose. Fruits 1.6–1.8 × 1.2–1.4 cm, subglobose to slightly obliquely ovoid, green (possibly immature), obtuse at the base, acuminate at the apex, sometimes weakly curved, smooth and glabrous, stipitate (the stipe ca. 0.2 cm long); pseudoaril present, color unknown; pyrenes, 1 or 2 usually developing, 1.1–1.4 × ca. 1.2 cm, smooth, broadly ovate, obtuse at the base, acuminate at the apex, bony, the wall ca. 0.75 mm thick, yellowish; funicular scar ca. 0.7 cm long, usually not very deep, without a rib in the middle.
Protium aguilarii is endemic to Costa Rica, where it is restricted to the Osa Peninsula, on the southern Pacific coast in Puntarenas Province. It occurs in primary forest, at 150–400 m elevation. In Aguabuena, Rincón de Osa, this species occurs in well-drained forest on undulating terrain, with many palms and large lianas; here, it co-occurs with Brosimum utile (Kunth) Oken (Moraceae), Carapa Aubl. (Meliaceae), and Symphonia L. f. (Clusiaceae) (see, for example, K. Thomsen 226). In Rancho Quemado, P. aguilarii is a small, infrequent tree on mountain ridges, where it is sympatric with tree species that are not very common in the area, or even the country as a whole, including Faramea permagnifolia Dwyer ex C. M. Taylor (Rubiaceae), Hirtella papillata Prance and Licania corniculata Prance (Chrysobalanaceae), Oecopetalum greenmanii Standl. & Steyerm. (Metteniusaceae), and Ruptiliocarpon caracolito Hammel & N. Zamora (Lepidobotryaceae).
Protium aguilarii is known from only six fertile collections (one of these with flowers in bud). Pistillate flowers have been collected in April, and fruits in February, May, June, and December.
Copalillo (Spanish; Costa Rica, K. Thomsen 683).
The epithet of this new species honors Reinaldo Aguilar Fernández for his important contributions to botany and his dedicated study and devoted collection of the plants of the Osa Peninsula for more than 25 years. He has become the world’s expert in the flora of this remarkably species-rich and beautiful corner of the world. This species is further dedicated to him in appreciation of his support and intellectual stimulation.
Protium aguilarii can be recognized by the combination of leaves with 5–9 leaflets with hispidulous pubescence on the petiole, rachis, petiolules, abaxial side of the leaflets, and inflorescence axes; leaflets with a distinct marginal vein that is visible on the abaxial side; short inflorescences and infructescences; flowers that are 4-merous, with the petals appressed-pubescent on the abaxial side and glabrous on the adaxial side; and glabrous pistils. The new species resembles, and has been confused with, P. costaricense, which, as treated here, is known only from the Caribbean slope of Nicaragua, Costa Rica, and Panama. Both species share hispidulous pubescence on the leaflets and inflorescences, but P. aguilarii has a glabrous pistil (vs. densely pubescent in P. costaricense), short inflorescence (ca. 0.6 vs. 1.6–6.5 cm long), petals that are glabrous on the abaxial side (vs. sparsely pubescent), and secondary venation brochidodromous (vs. eucamptodromous). Protium aguilarii also has usually shorter (non-basal) leaflets than P. costaricense (6.7–13.2 vs. 10.5–17.5 cm), and glabrous fruits (vs. minutely pubescent with scattered trichomes), with a stipitate base, the stipe ca. 0.2 cm long (vs. sessile or with stipitate ca. 0.1 cm long). Additionally, P. aguilarii differs from P. costaricense by its smooth (vs. rugose) pyrene. In Costa Rica, others species with glabrous pistils or pistillodes are P. aracouchini Marchand, P. hammelii (described here), P. panamense (Rose) I. M. Johnst., and P. ravenii D. M. Porter. Unlike P. aguilarii, these species have nearly glabrous leaves and other vegetative parts. Protium aguilarii also shares some similarities with the South American P. pilosissimum including short inflorescences and pubescent leaflets but it differs by its pubescent pistil or pistillode and fruit (vs. glabrous in P. aguilarii).
Costa Rica. Puntarenas: Osa, Bahía Chal, La Parcela, 08°43'50"N, 083°27'17"W, 150 m, 25 Jul 1996 (fl bud), R. Aguilar 4593 (
Protium hammelii is similar to P. multiramiflorum Lundell and P. panamense for their nearly glabrous leaves, petals, and usually pistil or pistillode (always glabrous in P. panamense; sometimes glabrous in P. multiramiflorum). However, the new species it is distinguished from P. 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 P. 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.
COSTA RICA. Limón: Parque Nacional Tortuguero, 5 km N de La Aurora, Guápiles, límite sur del Parque, junto río Sierpe, 10°22'00"N, 083°31'00"W, 30 m, 11 Apr 1990 (♂ fl), J. Solano 77 (holotype:
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.
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).
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.
Alcanfor (Spanish; Nicaragua, R. Rueda et al. 2642, 2701; J.C. Sandino 3424).
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.
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 P. glabrum (Rose) Engl., a species that is widespread from Belize to Panama, or P. panamense, from Costa Rica, Panama, and Colombia. The first of these is common in Costa Rica, while the second is quite rare (Fig.
Protium hammelii. A Branch with inflorescences B Venation on the abaxial side and marginal teeth of leaflets C Fruits D Pyrene E Fruit valves F Flower G Staminate flower with two petals removed, showing the pistillode and stamens. A and B from J. Solano 77,
Protium panamense. A Stilt roots B Branch with inflorescences C Leaflets D Inflorescences E Staminate flowers F Fruits. Protium aracouchini G Trunk base H Branch with inflorescences; inset showing dry resin on cut twig I Inflorescences; also see the suberose petiole base J Abaxial side of the leaflets. Protium ravenii K Trunk L Adaxial side of the leaflets; inset dry resin on cut twig M Abaxial side of the leaflets N Fruits. Photo credits: Rolando Pérez (A); Carmen Galdames (B); Steven Paton (C–F). G–J photos by Reinaldo Aguilar, from D. Santamaría & R. Aguilar 9836K–N photos by Reinaldo Aguilar, from R. Aguilar 12067, except L, inset by Orlando Vargas.
Fruit dispersal by birds and mammals has been reported at the La Selva Biological Station for P. panamense (
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 (
Basionym: Trichilia brenesii Standl. Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 583. 1937.
Type. COSTA RICA. [Alajuela:] colinas del Tremendal (San Pedro) de San Ramón, [09] Apr 1935 [♂ fl], A.M. Brenes 20510 (holotype: F-866066!; isotypes:
Protium brenesii is only known from Costa Rica, where it grows mainly in the Cordilleras de Guanacaste, de Tilarán and Central on both the Caribbean and Pacific slopes, though it also has been collected in the Cordillera de Talamanca (Dota region) and the Valle del General. It is found in primary forest and along roads and rivers between 640 and 1500 m elevation. Protium brenesii is found at the highest elevations of any species of its genus in Costa Rica.
Collections with male flowers have been made from March to May, and December; female flowers in February; and fruits in March and April, and from June to December.
Copal (Spanish; Costa Rica, E. Bello 473).
In the course of examining material identified as Protium costaricense, a notable number of collections from Costa Rica, mainly from 640–1500 m elevation in the Cordilleras de Guanacaste, de Tilarán and Central, were identified that differed from the rest. This material has twigs and leaflets with inconspicuous pubescence on the abaxial side; leaves with more numerous and usually narrower leaflets; and longer inflorescences and infructescences. Protium costaricense, as interpreted here, is a species of the lowlands of the Caribbean slope in Nicaragua, Costa Rica, and Panama, while the other collections represent a distinct montane taxon. An appropriate name already exists, and had been applied to some material collected in the Costa Rican cordilleras: Trichilia brenesii Standl. (1937: 583). Therefore, a new combination is proposed here, transferring T. brenesii to Protium.
The first known collection of Protium brenesii was made by Alberto M. Brenes (1870–1948) in the mountains near San Ramón, Alajuela Province, Costa Rica, in May 1923 (Brenes 19953). This species is similar in some aspects to P. costaricense, with which it has been confused for more than 90 years. These two taxa share the following morphological characteristics: twigs and leaves with dense pubescence; entire leaflets; 4(5)-merous flowers with pubescent petals, pistil, and pistillode; and rugose pyrene. The leaflets and fruits of the two species are also more or less similar in shape and size, but tend to be narrower in P. brenesii. However, P. brenesii can be distinguished from P. costaricense by its longer inflorescences [(5–) 7–11.5 vs. 1.6–6.5 cm], with malpighiaceous and simple trichomes (vs. only simple) on the axes, and male and female flowers with the disk equal or taller than the calyx (vs. shorter), a feature that is also evident on collections with fruits. Additionally, the terminal leaflets of P. brenesii are smaller (6.8–10.7 × 2.5–3.7 vs. 10.5–17.5 × 4.5–7.7 cm) and have shorter petiolules (1.5–2.5 vs. 2.8–3.5 [–5] cm). Importantly, P. brenesii is a species of montane forests at elevations of 640–1500 m, while P. costaricense is most frequent in lowland wet forests from 0–200 m. Some collections of P. brenesii have been confused with P. confusum (Rose) Pittier (or its synonym, P. schippii Lundell), the latter distinguished by its distally undulate or serrulate leaflets (vs. entire), inflorescences usually with a mixture of malpighiaceous, reddish trichomes and apparently glandular, whitish trichomes (vs. yellowish to pale brown malpighiaceous and simple trichomes), and flowers with the petals, pistil, and pistillode densely covered by dark red trichomes (vs. with whitish or pale brown trichomes).
In the checklist of Plantas Vasculares de Monteverde (
COSTA Rica. Alajuela: Cantón de Grecia, Cordillera Central, Los Ángeles, camino de Los Ángeles a la Laguna de Hule, 10°17'55"N, 084°12'20"W, 740–900 m, 28 Oct 1995 (fl bud), J. González & G. Perera 995 (
In view of the long history of confusion involving Protium brenesii and P. costaricense, the following information is provided to clarify some important parameters of the latter species, as it is here interpreted:
Icica costaricensis Rose, N. Amer. Fl. 25(3): 259. 1911.
Type. COSTA RICA. [Alajuela:] Santa Clara: Las Delicias, [500 m], Jan 1897 [♀ fl], P. Biolley 10665 [T. Biolley (sic), in the protologue] (holotype:
Protium costaricense it is known from wet forests on the Caribbean slopes of Nicaragua (Atlántico Sur and Río San Juan Departments), Panama (Colón and Panamá Provinces), and Costa Rica. In Costa Rica, it is known from throughout the Caribbean coastal plain in Alajuela and Limón Provinces. It grows in primary forest and along river or forest edges, from 0 to 200 m in elevation (reportedly up to 500 m, according to the label of the type).
Collected with staminate flowers in February and June; pistillate flowers in February, July, and August; and fruits in January and from June to December.
In Nicaragua, this species is known as alcanfor (A. Laguna 73; R. Rueda et al. 5338). In Costa Rica and Panama, it goes by copal, chutra, kerosín and alcanfor (
NICARAGUA. Atlántico Sur [Zelaya]: Santa Fe, unión del Caño Agua Fría y Quebrada La Capilla, [11°41'N, 084°28'W], [100–200 m], 02 Oct 1982 (fr), A. Laguna 73 (
The pseudarils of Protium costaricense are reported to have a pleasant flavor (L.J. Poveda 3663,
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