Protium brenesii (Standl.) 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 : 102-106

Protium brenesii (Standl.) D.Santam comb. nov. Figs 8A , 9

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: CR-2 sheets! [Hb. Brenes 20009, both with the same herbarium number], NY-00054791, digital image!).

Habitat and distribution.

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.

Phenology.

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.

Common name.

Copal (Spanish; Costa Rica, E. Bello 473).

Discussion.

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 Trichilia 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 Protium 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 Protium brenesii . However, Protium brenesii can be distinguished from Protium 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 Protium 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, Protium brenesii is a species of montane forests at elevations of 640-1500 m, while Protium costaricense is most frequent in lowland wet forests from 0-200 m. Some collections of Protium brenesii have been confused with Protium confusum (Rose) Pittier (or its synonym, Protium 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 ( Haber 2014), Protium sp. A. (7508 [W. Haber & E. Cruz]) and Protium costaricense (E. Bello 473) correspond to Protium brenesii ; the same applies to the collection cited by Gómez-Laurito and Ortiz (2004) (J. Gómez-Laurito et al. 12278).

Additional material examined.

COSTA Rica. Alajuela: Cantón de Grecia, Cordillera Central, Los Angeles , 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 (CR-2 sheets, MO, NY-digital image); Canton de San Ramon , Reserva Biologica Monteverde , rio Penas Blancas , parcela de los Enanos , 10°18'00"N, 084°43'48"W, 850 m, 02 Sep 1988 (fr), E. Bello 332 (CR-2 sheets, F, MO, NY-digital image, USJ); Reserva Biologica Monteverde , rio Penas Blancas , 10°19'N, 084°43'W, 850 m, 06 Sep 1988 (fr), E. Bello 353 (CR, MO, NY-digital image); Reserva Biologica Monteverde , rio Penas Blancas , 10°19'N, 084°43'W, 820 m, 10 Oct 1988 (fr), E. Bello & E. Cruz 458 (CR, MO, NY-digital image); Reserva Biologica Monteverde , rio Penas Blancas , parcela de Badilla, 10°19'N, 084°43'W, 850 m, 22 Oct 1988 (fr), E. Bello 473 (F, MO, NY-digital image, USJ-2 sheets); Reserva Biologica Monteverde , rio Penas Blancas , 10°18'36"N, 084°43'12"W, 900 m, 21 Apr 1993 (♂ fl), E. Bello 5014 (CR-2 sheets); San Pedro de San Ramon , 1000 m, 06 May 1923 (♂ fl), A.M. Brenes 19953 [Hb. Brenes 3883], (CR, F, NY-digital image); Colinas de San Pedro de San Ramon , 1050-1075 m, 27 May 1925 (fl bud), A.M. Brenes 19955 [Hb. Brenes 4222], (CR, F, NY-digital image); Colinas de San Pedro de San Ramon , 04 Jul 1925, 1075 m, A.M. Brenes 4827 [612], (F); Colinas de San Pedro de San Ramon , 19 May 1927 (♂ fl), A.M. Brenes 19954 [Hb. Brenes 5445], (CR, NY-digital image); Bajos del Jamaical , Reserva de San Ramon , 700-1000 m, 10 May 1985 (♂ fl), I. Chacón 1800 (CR-4 sheets); Reserva Forestal de San Ramon , Colonia Palmareña, 800-950 m, 19-22 Sep 1985 (fr), J. Gómez-Laurito 10528 (CR, USJ); Reserva Forestal de San Ramon , sendero a la fila al S. O. de la Estación, 10°13'N, 084°37'W, 05 Sep 1992 (fr), J. Gómez-Laurito 12278 (CR, F, USJ); Barranquilla, Falda Noroeste del Cerro Jabonal , 10 °09'40"N, 084°39'30"W, 1500 m, 04 Nov 1997 (fr), J. González et al. 2081 (CR-2 sheets, MO); Monteverde Reserve , Penas Blancas river valley, Eladio Cruz farm, 10°20'N, 084°43'W, 800 m, 01 Nov 1986 (fr), W. Haber & E. Bello 6176 (CR, NY-digital image); Reserva Biológica Monteverde, rio Penas Blancas , 10°20'N, 084°43'W, 850 m, 13 Mar 1987 (♂ fl), W. Haber & E. Bello 6801 (CR, MO, NY-digital image); Reserva Biológica Monteverde, rio Penas Blancas , 10°20'N, 084°43'W, 800 m, 14 Apr 1987 (♂ fl), W. Haber & E. Cruz 6979 (CR, MO, NY-digital image); Reserva Biológica Monteverde, rio Penas Blancas , 10°18'N, 084°45'W, 900 m, 21 May 1987 (fl with galls), W. Haber & E. Bello 7169 (MO, NY-digital image); Reserva Biológica Monteverde, rio Penas Blancas , 10°20'N, 084°43'W, 820 m, 10 Jun 1997 (fr), W. Haber & E. Cruz 7248 (CR, MO); Reserva Biológica Monteverde, rio Penas Blancas , Finca Wilson Salazar, 10°18'N, 084°43'W, 800-900 m, 20 Aug 1987 (fr), W. Haber & E. Cruz 7391 (CR, MO, NY-digital image); Reserva Biológica Monteverde, rio Penas Blancas , Finca Wilson Salazar, 10°18'N, 084°43'W, 860 m, 20 Oct 1987 (fr), W. Haber & E. Cruz 7508 (CR-2 sheets), 7509 (MO); Reserva Biológica Monteverde, rio Penas Blancas , Finca Wilson Salazar, 10°18'N, 084°43'W, 800 m, 06 Nov 1987 (fr), W. Haber & E. Cruz 7691 (CR, MO, NY-digital image); Reserva Biológica Monteverde, rio Penas Blancas , 10°18'N, 084°44'W, 900, 15 Dec 1987 (fr), W. Haber & E. Bello 7914 (CR, MO); Reserva Biológica Monteverde, rio Penas Blancas , 10°19'N, 084°43'W, 800 m, 14 Dec 1987 (♂ fl), W. Haber & E. Bello 7899 (CR, MO, NY-digital image); San Ramón, Bosque Eterno De Los Ninos , 4 km SW of La Tigra de San Carlos, valley of rio La Esperanza , finca Araya Ledezma, 10°18'N, 084°37'W, 600-800 m, 01 Jul 1992 (fr), W. Haber et al. 11232 (CR-2 sheets, MO); Reserva Forestal de San Ramon , 10°12'53"N, 084°36'28"W, 03 May 1987 (♂ fl), G. Herrera 617 (CR, F, MO); San Ramón, Los Ángeles, Reserva de San Ramon , 2 km al Norte de la Estación, 10°12'40"N, 084°36'20"W, 1000 m, 18 Oct 1993 (fr), G. Herrera 6604 (MO); area of the Reserva Biológica Alberto M. Brenes, 10°13'N, 084°36'W, 1010 m, 29 Apr 2001 (st), J. Homeier & A. Wolter 723 (USJ); area of the Reserva Biológica Alberto M. Brenes, 10°13'N, 084°36'W, 1010 m, 29 Apr 2001 (st), J. Homeier & A. Wolter 1010 (USJ); Reserva Forestal Arenal, rio Penas Blancas , Quebrada Agua Gata, Finca Francisco, 10°20'N, 084°42'W, 1200 m, 19 Sep 1990 (fr), N. Obando 122 (CR-2 sheets, MO, NY-digital image); Reserva Biológica Monteverde, Estación Eladio’s, 10°18'30"N, 084°43'10"W, 820 m, 02 Oct 1990 (fr), N. Obando et al. 187 (CR-2 sheets, MO, NY-digital image); San Rafael de San Ramon , 20 Oct 1969 (fr), S. Salas et al. 1378 (USJ); Reserva de San Ramon , 13 May 1985 (♂ fl), L. Umaña s.n. (USJ-026360); Canton de Upala , Bijagua, El Pilón, Cabeceras del rio Celeste , 10°49'N, 084°57'W, 700 m, 21 Apr 1988 (fr), G. Herrera 1852 (CR, MO, NY-digital image); Volcán Tenorio, Pilón, 19 Nov 1987 (fl bud), P. Sánchez & L.J. Poveda 1282 (CR, F); Parque Nacional Guanacaste, Sector San Ramón, Dos Ríos, sendero a Níspero y Argentina, 10°52'50"N, 085°24'30"W, 550 m, 04 Mar 1995 (fr), R. Espinoza et al. 1298 (CR-2 sheets, MO, NY-digital image); Parque Nacional Guanacaste, Nueva Zelandia, Estación San Ramón, La Campana, Dos Ríos, rio Colon , 10°52'50"N, 085°24'05"W, 550 m, 23 Mar 1994 (♂ fl), D. García 196 (CR-2 sheets, MO, NY-digital image); Canton de San Carlos , hacia Quebrada “Corella” San Carlos, 650 m, 23 Jun 1966 (fr), A. Jiménez 4045 (CR, F, MO, NY-digital image); La Fortuna, Finca El Jilguero, Sendero La Lava , rio Aguas Calientes 0.5 km aguas arriba, 10°26'35"N, 084°42'20"W, 700 m, 23 Nov 1992 (fr), G. Herrera 5625 (CR-2 sheets); North side Arenal Volcano, 10°28'N, 084°42'W, 800 m, 11 Apr 1974 (fr), R. Lent 3862 (CR, F, NY-digital image) . Guanacaste: Cantón de La Cruz, Parque Nacional Guanacaste, Estacion Pitilla , al noroeste de la estación, 11°01'48"N, 085°25'12"W, 550 m, 16 Jun 1989 (fr), B. Hammel 17495 (CR, F, MO, NY-digital image); Parque Nacional Guanacaste, Estacion Pitilla , 9 km al S de Santa Cecilia, 10°59'25"N, 085°25'40"W, 700-1000 m, 06 Mar 1991 (fr), C.O. Moraga 315 (CR-2 sheets); Parque Nacional Guanacaste, Estacion Pitilla , Sendero El Mismo , Finca La Pasmompa , 11°02'00"N, 085°24'30"W, 700 m, 09 Dec 1990 (♂ fl), P. Ríos 216 (CR, MO); Parque Nacional Guanacaste, Estación Pitilla, Fila Orosilito y Sendero El Mismo, 10°59'26"N, 085°25'40"W, 700 m, 02 Mar 1991 (fr), P. Ríos 310 (CR-2 sheets, MO, NY-digital image); Parque Nacional Guanacaste, Estación Pitilla, Sendero El Mismo, 10°59'26"N, 085°25'40"W, 700 m, 15 Jun 1991 (♂ fl), P. Ríos 364 (CR-2 sheets, MO); Canton de Bagaces , Parque Nacional Rincón de la Vieja, Sendero de la toma de agua, a 3 km de la estación, 10°46'05"N, 085°17'40"W, 1000 m, 17 Sep 1990 (fr), G. Rivera 546 (CR-2 sheets, MO, NY-2 sheets, digital image); Parque Nacional Rincón de la Vieja, Colonia Blanca, 10°48'20"N, 085°17'50"W, 1300-1600 m, 08 Nov 1990 (fr), G. Rivera 847 (CR-2 sheets); Parque Nacional Rincón de la Vieja, Sector Santa María, Sendero La Plantacion , cabeceras Quebrada Zopilote, 10°46'48"N, 085°17'24"W, 950-1100 m, 14 Aug 1996 (fr), J. F. Morales 5667 (CR-2 sheets); Guatuso, Lago Coter, 5 km norte, Hotel Ecolodge, 10°35'20"N, 084°55'50"W, 700 m, 28 Apr 1997 (♂ fl), G. Rivera 3005 (CR) . San José: Reserva Forestal Los Santos, quebrada Bomba, cruce a Fila Mona y La Bomba , 09°30'00"N, 083°56'45"W, 500 m, 28 Feb 2005 (fl bud), D. Santamaría & J.F. Morales 751 (CR); Reserva Forestal Los Santos , Dota , Fila Vega , Sendero a Fila Seca , 09°29'40"N, 083°57'30"W, 800-950 m, 03 Mar 2005 (fl bud), D. Santamaría & J.F. Morales 900 (CR); Canton de Perez Zeledon , vicinity of El General , [09°23'42"N, 083°38'26"W], 1040 m, Feb 1936 (♀ fl), A.F. Skutch 2620 (A, GH, MO, NY-digital image); Pérez Zeledón, vicinity of El General, [09°20'48"N, 083°39'27"W], 640 m, Mar 1939 (♂ fl), A.F. Skutch 4244 (A, MO, NY-digital image); Pérez Zeledón, vicinity of El General, [09°22'20"N, 083°39'12"W], 675-900 m, Mar 1940 (♂ fl), A.F. Skutch 4849 (A, CR, F-2 sheets, MO); basin of General, 675-900 m, 10 Feb 1942 (fl), A.F. Skutch 5024 (F) .

In view of the long history of confusion involving Protium brenesii and Protium costaricense , the following information is provided to clarify some important parameters of the latter species, as it is here interpreted:

Figures

Figure 8. Types of Protium brenesii (A) and Protium costaricense (B). Images courtesy of Museo Nacional de Costa Rica.

Figure 9. Distribution of Protium brenesii and Protium costaricense.