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Two new species and a new combination in Protium (Burseraceae) from Costa Rica

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Two new species of Protium (Burseraceae) are described and illustrated: Protiumaguilariisp. nov., from the Pacific slope of the Osa Peninsula, Puntarenas Province, Costa Rica; and Protiumhammeliisp. nov., from wet forests on the Caribbean slopes of Nicaragua and Costa Rica. In addition, Protiumbrenesiicomb. nov., is proposed as a new combination based on Trichiliabrenesii, 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 Protiumcostaricense, 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.
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Two new species and a new combination in Protium (Burseraceae) from Costa Rica 89
Two new species and a new combination in Protium
(Burseraceae) from Costa Rica
Daniel Santamaría-Aguilar1, Laura P. Lagomarsino2
1 Current address: Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, USA 2 Missouri
Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, USA, and University of Missouri–St. Louis,
Biology Department, One University Blvd., Research Building, St. Louis, MO 63121, USA
Corresponding author: Daniel Santamaría-Aguilar (daniel.santamaria366@gmail.com; Daniel.Santamaria@mobot.org)
Academic editor: Pavel Stoev|Received 25 August 2016|Accepted 9 December 2016|Published 18 January2017
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
Abstract
Two new species of Protium (Burseraceae) are described and illustrated: Protium aguilarii sp. nov., from
the Pacic 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 owers 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.
Resumen
Se describen e ilustran dos nuevas especies de Protium (Burseraceae): Protium aguilarii sp. nov., de la
vertiente del Pacíco 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 ores 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.
Keywords
Burseraceae, Costa Rica, Nicaragua, Protium, Sapindales, taxonomy
PhytoKeys 76: 89–113 (2017)
doi: 10.3897/phytokeys.76.10298
http://phytokeys.pensoft.net
Copyright D. Santamaría-Aguilar, L.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.
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Daniel Santamaría-Aguilar & Laura P. Lagomarsino / PhytoKeys 76: 89–113 (2017)
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Introduction
e genus Protium (Burseraceae), with approximately 160 species, is nearly pantropi-
cal 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 ve 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 Engl. subsp.
sessiliorum (Rose) D. M. Porter on the Pacic slope]. e 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 owers 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.
e two new species described below and the need for a new combination in Pro-
tium were discovered during preparation of the Burseraceae treatment for the Manual
de Plantas de Costa Rica. is study was based on an examination of Protium specimens
deposited at A, CR, F, GH, MO and USJ (acronyms according iers 2016, continu-
ally updated), along with consultation of digital images in national and international
virtual herbaria and relevant literature on Protium (e.g., Rose 1911; Standley 1937;
Swart 1942; Standley and Steyermark 1946; Cuatrecasas 1957; Porter 1970; Daly
1987, 1989, 1997, 1999, 2002, 2014, 2016; Porter and Pool 2001). Additional eld
collections were conducted in Costa Rica in February through March 2016. e distri-
bution map was generated using the program SimpleMappr (Shorthouse 2010) from
coordinates reported on specimen labels; specimens whose label data did not indicate
coordinates are shown in brackets.
Taxonomy
Protium aguilarii D.Santam., sp. nov.
urn:lsid:ipni.org:names:77159804-1
Figs 1, 2, 3
Diagnosis. Protium aguilarii most closely resembles P. costaricense (Rose) Engl. and
P.pilosissimum Engl., for their leaves with petiole, rachis, and leaets that are hispidu-
lous on the abaxial side and leaets of comparable in size and coloration in herbarium
material, but diers from both for its glabrous pistil and fruits (vs. pubescent).
Type. 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: MO-6664125!; isotypes: CR-
51404! [ex-INB], NY-01189275, digital image!).
Two new species and a new combination in Protium (Burseraceae) from Costa Rica 91
Figure 1. Protium aguilarii. A Branch with inorescences B Venation C Pubescence on the leaf rachis
and petiolules D Inorescences E Female ower, with perianth partially removed F Fruits. A–D from
R. Aguilar 4593, CR E from R. Aguilar & X. Cornejo 11115, CR F from K. omsen 226, CR. Drawing
by Jessica Jiménez.
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 tri-
Daniel Santamaría-Aguilar & Laura P. Lagomarsino / PhytoKeys 76: 89–113 (2017)
92
chomes 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 leaets 5.2–10 × 2–4.3 cm, elliptic to ovate, obtuse at the base; other lateral
leaets 6.7–13.2 × 2.6–5 cm, elliptic to ovate, obtuse to subcuneate at the base (with
one side sometimes asymmetric); the terminal leaet 7.3–14.2 × 2.7–5.8 cm, ovate, el-
liptic, base obtuse to subcuneate and symmetrical; apex acuminate, the acumen 0.7–1
cm long; margin entire; leaets 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 percur-
rent 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 at, hispidulous, secondary veins at, scattered hispidulous to
almost glabrous, the higher-order veins at almost glabrous, the rest of surface nearly
glabrous. Inorescences fasciculate, axillary (sometimes at leaess nodes), staminate in-
orescences unknown, pistillate inorescences ca. 0.6 cm long (0.9–1.2 cm in fruits),
much shorter than the petiole, branching at the base, not exuous, all axes densely
pubescent with simple, yellowish brown or whitish trichomes; bracts subtending the
inorescences 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 owers 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 owers unknown. Pistillate owers 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,
inexed-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 equal-
ing than antesepalous, the laments ca. 0.9 mm long, at, 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
Two new species and a new combination in Protium (Burseraceae) from Costa Rica 93
Figure 2. Protium aguilarii. A Trunk and bark B Twigs C Branch with leaves D Leaves showing abaxial side
of leaets E Leaf showing adaxial side F Leaet 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.
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.
Habitat and distribution. Protium aguilarii is endemic to Costa Rica, where it is
restricted to the Osa Peninsula, on the southern Pacic 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. omsen
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 (Ru-
biaceae), Hirtella papillata Prance and Licania corniculata Prance (Chrysobalanaceae),
Oecopetalum greenmanii Standl. & Steyerm. (Metteniusaceae), and Ruptiliocarpon car-
acolito Hammel & N. Zamora (Lepidobotryaceae).
Daniel Santamaría-Aguilar & Laura P. Lagomarsino / PhytoKeys 76: 89–113 (2017)
94
Phenology. Protium aguilarii is known from only six fertile collections (one of
these with owers in bud). Pistillate owers have been collected in April, and fruits in
February, May, June, and December.
Common name. Copalillo (Spanish; Costa Rica, K. omsen 683).
Etymology. e epithet of this new species honors Reinaldo Aguilar Fernández
for his important contributions to botany and his dedicated study and devoted col-
lection of the plants of the Osa Peninsula for more than 25 years. He has become the
world’s expert in the ora of this remarkably species-rich and beautiful corner of the
world. is species is further dedicated to him in appreciation of his support and intel-
lectual stimulation.
Discussion. Protium aguilarii can be recognized by the combination of leaves with
5–9 leaets with hispidulous pubescence on the petiole, rachis, petiolules, abaxial side of
the leaets, and inorescence axes; leaets with a distinct marginal vein that is visible on
the abaxial side; short inorescences and infructescences; owers that are 4-merous, with
the petals appressed-pubescent on the abaxial side and glabrous on the adaxial side; and
glabrous pistils. e 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 leaets and inores-
cences, but P. aguilarii has a glabrous pistil (vs. densely pubescent in P. costaricense), short
inorescence (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. eucamptodro-
mous). Protium aguilarii also has usually shorter (non-basal) leaets 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 diers 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 inorescences and pubescent leaets but it dif-
fers by its pubescent pistil or pistillode and fruit (vs. glabrous in P. aguilarii).
Additional material examined. COSTA RICA. Puntarenas: Osa, Bahía Chal,
La Parcela, 08°43'50"N, 083°27'17"W, 150 m, 25 Jul 1996 ( bud), R. Aguilar 4593
(CR-2 sheets, F); Bahía Chal, La Parcela, 08°43'50"N, 083°27'17"W, 150 m, 12 Dec
1996 (fr), R. Aguilar 4746 (CR); Rancho Quemado, camino a Drake, parte mas el-
evada del camino, 200 m al Este de la torre del ICE, en una trocha que lleva al Tierra
de Conservación de Rancho Quemado, 08°41'33"N, 083°35'35"W, 350 m, 04 Apr
2008 ( ), R. Aguilar & X. Cornejo 11115 (MO, NY-digital image, USJ); Rancho
Quemado, siguiendo a la Ganado, 08°43'30"N, 083°35'30"W, 200–450 m, 26 Nov
1991 (st), J. Marín & G. Herrera 306 (CR-2 sheets); Sierpe, Península de Osa, subien-
do hacia el Cerro Chocuaco, desde el Bajo de San Juan, 400 m, 12 Jan 1991 (st), C.O.
Morales 262 (USJ); Distrito de Sierpe, Península de Osa, entre Rancho Quemado y
Drake, trocha al sur, sobre la la, antes de llegar a la torre, 08°41'30"N, 083°35'28"W,
Two new species and a new combination in Protium (Burseraceae) from Costa Rica 95
Figure 3. Distribution of Protium aguilarii and P. hammelii.
375 m, 21 Mar 2016 (st), D. Santamaría et al. 9851 (CR); Península de Osa, Agu-
abuena, 3.5 km W of Rincón, 1 km N of BOSCOSA station, 08°43'N, 083°31'W,
350 m, 09 Jan 1993 (fr), K. omsen 226 (CR, NY-digital image, USJ); Península
de Osa, Aguabuena, 3.5 km W of Rincón, 1 km N of BOSCOSA station, 08°43'N,
083°31'W, 350 m, 13 Nov 1992 (st), K. omsen 683 (CR); Península de Osa, Agu-
abuena, 3.5 km W of Rincón, 1 km N of BOSCOSA station, 08°43'N, 083°31'W,
350 m, 30 May 1993 (st), K. omsen 707 (CR); Península de Osa, Aguabuena, 3.5
km W of Rincón, 1 km N of BOSCOSA station, 08°43'N, 083°31'W, 350 m, 18 Jun
1993 (fr), K. omsen 807 (CR, NY-digital image).
Protium hammelii D.Santam., sp. nov.
urn:lsid:ipni.org:names:77159805-1
Figs 3, 4, 5, 7A
Diagnosis. Protium hammelii is similar to P. multiramiorum Lundell and P. pana-
mense for their nearly glabrous leaves, petals, and usually pistil or pistillode (always
glabrous in P. panamense; sometimes glabrous in P. multiramiorum). However, the
new species it is distinguished from P. multiramiorum 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) leaets
and shorter petiolules.
Type. COSTA RICA. Limón: Parque Nacional Tortuguero, 5 km N de La Auro-
ra, Guápiles, límite sur del Parque, junto río Sierpe, 10°22'00"N, 083°31'00"W, 30m,
Daniel Santamaría-Aguilar & Laura P. Lagomarsino / PhytoKeys 76: 89–113 (2017)
96
Figure 4. Protium hammelii. A Branch with inorescences B Venation on the abaxial side and marginal
teeth of leaets C Fruits D Pyrene E Fruit valves F Flower G Staminate ower with two petals removed,
showing the pistillode and stamens. A and B from J. Solano 77, CR C–E from J. Gómez-Laurito et al.
10998, CR F, G from W.D. Stevens 23769, CR. Drawing by Jessica Jiménez.
Two new species and a new combination in Protium (Burseraceae) from Costa Rica 97
11 Apr 1990 ( ), J. Solano 77 (holotype: CR-152860!; isotypes: CR-51496! [ex-
INB], F-2 sheets 2081330!, 2127441!, MO-6664125!, NY-01189417, digital image!).
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 gla-
brescent, 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 pu-
bescent 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 tri-
chomes; terminal petiolule 1.8–4.3 (–5.5) cm long, pulvinulus conspicuous; basal pair
of leaets 10.1–19 × 3.7–6.7 cm, ovate or lanceolate, obtuse to subcuneate at the
base (sometimes asymmetric); other lateral leaets 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; leaets 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 irregu-
lar, 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 at with trichomes similar to the midrib or glabrous, the higher-
order veins at to lightly impressed, with scattered trichomes, the rest of surface with
scattered trichomes to glabrous. Inorescences axillary (sometimes at leaess nodes),
generally branching at the base, not exuous, the staminate inorescences 3.4–8.5
cm long, shorter or exceeding the petiole, minutely pubescent with simple and mal-
pighiaceous, whitish trichomes on all axes, the pistillate inorescences 1–1.8 cm long
[(1.4)– 2.5–12.5 cm in fruit], much shorter than the petiole; bracts subtending the
inorescences 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 owers 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 owers with calyx 0.7–1.3 × 1.3–2.4 mm, much
Daniel Santamaría-Aguilar & Laura P. Lagomarsino / PhytoKeys 76: 89–113 (2017)
98
Figure 5. Protium hammelii. A Stilt roots B Branch with inorescences C Adaxial side of the leaets
DInorescences E Flower F Fruits. Photo credits: Orlando Vargas (A–E); and N. Zamora (F).
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, inexed-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 laments more or less at, 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, gla-
brous, the style 0–1.5 mm long, stigma lobes 4, subglobose to weakly angulate, densely
papillose. Pistillate owers with calyx 1–1.2 × 2–2.5 mm, the lobes ca. 0.8 mm long,
all parts similar to that of the staminate owers; petals green, greenish yellow or white,
distinct, 3–3.5 × 1.16–1.5 mm, suberect to reexed at anthesis similar to those of the
staminate owers; 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 laments at, 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 papil-
lose, 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
Two new species and a new combination in Protium (Burseraceae) from Costa Rica 99
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. is 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 at or relatively at 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 Pentacle-
thra 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 ower. In Costa
Rica, it has been collected with staminate owers in January, February, April, and De-
cember; pistillate owers 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. e specic epithet honors Barry E. Hammel, curator at the Missouri
Botanical Garden and co-editor of the Manual de Plantas de Costa Rica, in recogni-
tion of his extensive work on the Costa Rican ora, as well as his personal support and
motivation.
Discussion. Protium hammelii is recognizable by its almost glabrous vegetative
parts, leaves with 5–7 leaets, commonly with a sparsely denticulate margin, promi-
nent tertiary veins on the abaxial side, 4(5)-merous owers with glabrous petals, pistils,
and pistillodes, the pistillate owers 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 identied as P. glabrum
(Rose) Engl., a species that is widespread from Belize to Panama, or P. panamense,
from Costa Rica, Panama, and Colombia. e rst of these is common in Costa Rica,
while the second is quite rare (Fig. 6A–F); both have leaets that are always entire.
Protium hammelii diers from P. 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 P. hammelii (vs. obtuse or rounded), and the glabrous (vs. with very
small trichomes). Protium panamense shares occasional stilt-roots, glabrous owers and
fruits that are apiculate at the apex with P. hammelii, but P. hammelii usually has
smaller lateral (11–22.2 × 3.3–6.4 vs. 16–32.5 × 6.7–10.3 cm), and terminal leaets
(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); P. 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 multiramiorum from Mexico to
Daniel Santamaría-Aguilar & Laura P. Lagomarsino / PhytoKeys 76: 89–113 (2017)
100
Honduras is similar to P. hammelii in its nearly glabrous leaves, petals, and sometimes
pistil or pistillode. (Although the pistil of P. multiramiorum was originally described
as glabrous [Lundell 1937], almost all collections studied, including one of the isotypes
Figure 6. Protium panamense. A Stilt roots B Branch with inorescences C Leaets D Inorescences
EStaminate owers F Fruits. Protium aracouchini G Trunk base H Branch with inorescences; inset
showing dry resin on cut twig I Inorescences; also see the suberose petiole base J Abaxial side of the
leaets. Protium ravenii K Trunk L Adaxial side of the leaets; inset dry resin on cut twig M Abaxial side
of the leaets 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 9836 K–N photos by Reinaldo Agui-
lar, from R. Aguilar 12067, except L, inset by Orlando Vargas.
Two new species and a new combination in Protium (Burseraceae) from Costa Rica 101
[Lundell 6212, GH-2 sheets!] and the paratype [Schipp 1021; A!, F!, GH!, MO!], have
the pistil or pistillode with tiny, scattered trichomes). e new species diers 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). Others species in Costa
Rica with glabrous pistils or pistillodes are P. aracouchini (Figs 6G–J), P. ravenii (Figs
6K–N), and P. aguilarii. e rst two species can be distinguished from P. hammelii
by their exuous inorescences and twigs and abundant exuding resin that becomes
whitish and chalky (Figs 6H and L, inset), while P. 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 P. panamense (Vargas 2000), but the observation likely corresponds to P.
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 Blueelds, río Escon-
dido, 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 Cam-
pamento Germán Pomares, 11°36'N, 083°52'W, 60–90 m, 08 Feb 1982 ( 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. Balles-
tero 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 ( ), 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 ( , 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
Daniel Santamaría-Aguilar & Laura P. Lagomarsino / PhytoKeys 76: 89–113 (2017)
102
Figure 7. Comparison between the pyrenes of Protium hammelii (A) and P. multiramiorum (B). Afrom
G. Davidse & G. Herrera 30879; and B from G.M. Aguilar et al. 4754. Drawing by Alex M. Campos.
Sarapiquí, Southwest trail, 1600 m line, 100 m, 16 Feb 1981 ( ), 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,
Two new species and a new combination in Protium (Burseraceae) from Costa Rica 103
El Swampo Trail, 100 m, 27 Apr 1981 (fr), J.P. Folsom 9882 (F, MO, NY-digital im-
age); 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, la Car-
rillo, 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, nca 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 (  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, vicin-
ity 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 ( bud),
A. Jiménez 3602 (F-2 sheets, CR); Finca Hermanos Vargas, 1 km al Suroeste de Puerto
Viejo, 125 m, 04 Feb 1966 ( ), 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 (  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-dig-
ital image, MO). Limón: Bosque Lluvioso [nca 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.
Daniel Santamaría-Aguilar & Laura P. Lagomarsino / PhytoKeys 76: 89–113 (2017)
104
(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 Par-
ismina, 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
( 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 ( ), R. Soto 2758 (CR); Cerro Coronel, E of Laguna Danto, 10°41'N,
083°38'W, 20–170 m, 16–13 Jan 1986 ( ), W.D. Stevens 23769, 23770 (CR, MO);
Cerro Coronel, along río Colorado at and below outow of Laguna Danto, 10º42'N,
83º39'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. omsen 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).
Protium brenesii (Standl.) D.Santam., comb. nov.
urn:lsid:ipni.org:names:77159806-1
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 [ ], 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 Pacic 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 owers have been made from March to May,
and December; female owers in February; and fruits in March and April, and from
June to December.
Two new species and a new combination in Protium (Burseraceae) from Costa Rica 105
Figure 8. Types of Protium brenesii (A) and P. costaricense (B). Images courtesy of Museo Nacional de
Costa Rica.
Common name. Copal (Spanish; Costa Rica, E. Bello 473).
Discussion. In the course of examining material identied 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 identied that diered
from the rest. is material has twigs and leaets with inconspicuous pubescence on
the abaxial side; leaves with more numerous and usually narrower leaets; and longer
inorescences 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). erefore, a new combination is proposed here,
transferring T. brenesii to Protium.
e rst 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). is species is similar in some aspects to P. costaricense, with
which it has been confused for more than 90 years. ese two taxa share the following
morphological characteristics: twigs and leaves with dense pubescence; entire leaets;
4(5)-merous owers with pubescent petals, pistil, and pistillode; and rugose pyrene.
e leaets and fruits of the two species are also more or less similar in shape and size,
Daniel Santamaría-Aguilar & Laura P. Lagomarsino / PhytoKeys 76: 89–113 (2017)
106
but tend to be narrower in P. brenesii. However, P. brenesii can be distinguished from
P. costaricense by its longer inorescences [(5–) 7–11.5 vs. 1.6–6.5 cm], with mal-
pighiaceous and simple trichomes (vs. only simple) on the axes, and male and female
owers with the disk equal or taller than the calyx (vs. shorter), a feature that is also
evident on collections with fruits. Additionally, the terminal leaets 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. con-
fusum (Rose) Pittier (or its synonym, P. schippii Lundell), the latter distinguished by its
distally undulate or serrulate leaets (vs. entire), inorescences usually with a mixture
of malpighiaceous, reddish trichomes and apparently glandular, whitish trichomes (vs.
yellowish to pale brown malpighiaceous and simple trichomes), and owers 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 P. costaricense (E. Bello 473) correspond to P.
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 Ángeles, camino de Los Ángeles a la Laguna de Hule,
10°17'55"N, 084°12'20"W, 740–900 m, 28 Oct 1995 ( bud), J. González & G.
Perera 995 (CR-2 sheets, MO, NY-digital image); Cantón de San Ramón, Reserva
Biológica Monteverde, río Peñas 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 Biológica Monteverde, río Peñas Blancas, 10°19'N, 084°43'W,
850 m, 06 Sep 1988 (fr), E. Bello 353 (CR, MO, NY-digital image); Reserva Biológica
Monteverde, río Peñas Blancas, 10°19'N, 084°43'W, 820 m, 10 Oct 1988 (fr), E.
Bello & E. Cruz 458 (CR, MO, NY-digital image); Reserva Biológica Monteverde, río
Peñas 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 Biológica Monteverde,
río Peñas Blancas, 10°18'36"N, 084°43'12"W, 900 m, 21 Apr 1993 ( ), E. Bello
5014 (CR-2 sheets); San Pedro de San Ramón, 1000 m, 06 May 1923 ( ), A.M.
Brenes 19953 [Hb. Brenes 3883], (CR, F, NY-digital image); Colinas de San Pedro de
San Ramón, 1050–1075 m, 27 May 1925 ( bud), A.M. Brenes 19955 [Hb. Brenes
4222], (CR, F, NY-digital image); Colinas de San Pedro de San Ramón, 04 Jul 1925,
1075 m, A.M. Brenes 4827 [612], (F); Colinas de San Pedro de San Ramón, 19 May
1927 ( ), A.M. Brenes 19954 [Hb. Brenes 5445], (CR, NY-digital image); Bajos del
Jamaical, Reserva de San Ramón, 700–1000 m, 10 May 1985 ( ), I. Chacón 1800
(CR-4 sheets); Reserva Forestal de San Ramón, Colonia Palmareña, 800–950 m, 19–
22 Sep 1985 (fr), J. Gómez-Laurito 10528 (CR, USJ); Reserva Forestal de San Ramón,
sendero a la la 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,
Two new species and a new combination in Protium (Burseraceae) from Costa Rica 107
Figure 9. Distribution of Protium brenesii and P. costaricense.
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, Peñas 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, río Peñas Blancas, 10°20'N, 084°43'W,
850 m, 13 Mar 1987 ( ), W. Haber & E. Bello 6801 (CR, MO, NY-digital image);
Reserva Biológica Monteverde, río Peñas Blancas, 10°20'N, 084°43'W, 800 m, 14 Apr
1987 ( ), W. Haber & E. Cruz 6979 (CR, MO, NY-digital image); Reserva Bi-
ológica Monteverde, río Peñas Blancas, 10°18'N, 084°45'W, 900 m, 21 May 1987 (
with galls), W. Haber & E. Bello 7169 (MO, NY-digital image); Reserva Biológica
Monteverde, río Peñas Blancas, 10°20'N, 084°43'W, 820 m, 10 Jun 1997 (fr), W.
Haber & E. Cruz 7248 (CR, MO); Reserva Biológica Monteverde, río Peñas 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, río
Peñas 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,
río Peñas 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 Mon-
teverde, río Peñas Blancas, 10°18'N, 084°44'W, 900, 15 Dec 1987 (fr), W. Haber &
E. Bello 7914 (CR, MO); Reserva Biológica Monteverde, río Peñas Blancas, 10°19'N,
084°43'W, 800 m, 14 Dec 1987 ( ), W. Haber & E. Bello 7899 (CR, MO, NY-
digital image); San Ramón, Bosque Eterno De Los Niños, 4 km SW of La Tigra de
San Carlos, valley of río La Esperanza, nca 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 Ramón, 10°12'53"N, 084°36'28"W, 03 May 1987 ( ), G. Herrera
617 (CR, F, MO); San Ramón, Los Ángeles, Reserva de San Ramón, 2 km al Norte de
la Estación, 10°12'40"N, 084°36'20"W, 1000 m, 18 Oct 1993 (fr), G. Herrera 6604
Daniel Santamaría-Aguilar & Laura P. Lagomarsino / PhytoKeys 76: 89–113 (2017)
108
(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, río Peñas 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 Ramón, 20 Oct 1969 (fr), S. Salas
et al. 1378 (USJ); Reserva de San Ramón, 13 May 1985 ( ), L. Umaña s.n. (USJ-
026360); Cantón de Upala, Bijagua, El Pilón, Cabeceras del río 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 ( bud), P. Sánchez & L.J. Poveda 1282 (CR, F);
Parque Nacional Guanacaste, Sector San Ramón, Dos Ríos, sendero a Níspero y Ar-
gentina, 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, río Colón, 10°52'50"N, 085°24'05"W,
550 m, 23 Mar 1994 ( ), D. García 196 (CR-2 sheets, MO, NY-digital image);
Cantón 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, Sen-
dero La Lava, río 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 im-
age). Guanacaste: Cantón de La Cruz, Parque Nacional Guanacaste, Estación 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, Es-
tación 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, Es-
tación Pitilla, Sendero El Mismo, Finca La Pasmompa, 11°02'00"N, 085°24'30"W,
700 m, 09 Dec 1990 ( ), 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 Na-
cional Guanacaste, Estación Pitilla, Sendero El Mismo, 10°59'26"N, 085°25'40"W,
700 m, 15 Jun 1991 ( ), P. Ríos 364 (CR-2 sheets, MO); Cantón 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 Blan-
ca, 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 Plant-
ación, 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 ( ), 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 ( bud), D.
Two new species and a new combination in Protium (Burseraceae) from Costa Rica 109
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 ( bud),
D. Santamaría & J.F. Morales 900 (CR); Cantón de Pérez Zeledón, vicinity of El Gen-
eral, [09°23'42"N, 083°38'26"W], 1040 m, Feb 1936 ( ), 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 ( ), A.F. Skutch 4244 (A, MO, NY-digital im-
age); Pérez Zeledón, vicinity of El General, [09°22'20"N, 083°39'12"W], 675–900 m,
Mar 1940 ( ), A.F. Skutch 4849 (A, CR, F-2 sheets, MO); basin of General, 675–
900 m, 10 Feb 1942 (), A.F. Skutch 5024 (F).
In view of the long history of confusion involving Protium brenesii and P. costari-
cense, the following information is provided to clarify some important parameters of
the latter species, as it is here interpreted:
Protium costaricense (Rose) Engl., Nat. Panzenfam., ed. 2 [Engler & Prantl] 19a:
414. 1931.
Figs 8B, 9
Icica costaricensis Rose, N. Amer. Fl. 25(3): 259. 1911.
Type. COSTA RICA. [Alajuela:] Santa Clara: Las Delicias, [500 m], Jan 1897 [
], P. Biolley 10665 [T. Biolley (sic), in the protologue] (holotype: US-digital image!
[herbarium of Capt. John Donnell Smith, in the protologue]; isotypes: CR!, F!).
Habitat and distribution. Protium costaricense it is known from wet forests on the
Caribbean slopes of Nicaragua (Atlántico Sur and Río San Juan Departments), Pana-
ma (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).
Phenology. Collected with staminate owers in February and June; pistillate ow-
ers in February, July, and August; and fruits in January and from June to December.
Common name. 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 (Condit et al. 2011).
Additional material examined. 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 (MO). Río San Juan: Sábalo, 1 km al norte
de río San Juan, 11°02'N, 084°27'W, [100 m], 09–10 Jul 1985 (fr), P.P. Moreno
26057 (MO); sobre el río Indio, entre San Juan del Norte Nuevo y La Casa de Narciso
Orozco, incluyendo Caño Negro, 10°58'N, 083°44'W, 0–100 m, 01 Jul 1994 ( ),
R. Rueda et al. 1612 (MO); Reserva Indio-Maíz, Mpio. de San Juan del Norte, Laguna
de Silico, 10°51'N, 083°46'W, 0–10 m, 02 Aug 1996 (fr), R. Rueda et al. 4850 (MO);
Reserva Indio-Maíz, Mpio. de El Castillo, a lo largo del río Bartola entre el caño la Lar-
Daniel Santamaría-Aguilar & Laura P. Lagomarsino / PhytoKeys 76: 89–113 (2017)
110
garta y la cabecera del río Bartola, 11°16'N, 084°16'W, [50–100 m], 29 Dec 1996 (fr),
R. Rueda et al. 5127 (MO); Reserva Indio-Maíz, Mpio. de El Castillo, a 8 km de la ca-
becera del río Bartola, en dirección al Cerro el Diablo, 11°01'N, 084°14'W, 120 m, 04
Jan 1997 (fr), R. Rueda et al. 5338 (MO); Reserva Indio-Maíz, Mpio. de El Castillo, 3
km al norte de la desembocadura del Caño Chontaleño, 11°05'N, 084°15'W, 24 Feb
1997 ( ), R. Rueda et al. 6285 (MO); Mpio. El Castillo, Reserva Indio-Maíz, río San
Juan, entre la desembocadura del río Bartola y el Caño Sarnoso, 10°56'N, 084°20'W,
100 m, 04 Dec 1998 (fr), R. Rueda et al. 9462 (MO). COSTA RICA. Alajuela: along
road between Cañas (Guanacaste) and Upala, near río Zapote, 1.8–2.7 km south of
río Canalete, ca. 100 m, 25 Jun 1976 ( ), T.B. Croat 36355 (MO, NY-digital
image); San Carlos, San Luis de Cutris, 23 Sep 1983 (fr), L.J. Poveda 3663 (USJ).
Limón: cantón de Pococí, Refugio Nacional de Fauna Silvestre Barra del Colorado,
Sardinas, 10°38'24"N, 083°43'48"W, 15 m, 25 Nov 1992 (fr), F. Araya 61 (CR, MO,
NY-digital image); Refugio Nacional de Fauna Silvestre Barra del Colorado, Puerto
Lindo, 10°41'24"N, 083°39'00"W, 200 m, 24 Jul 1995 (fr), F. Araya & J. Corrales
803 (MO, NY-digital image); Southwestern-most ridge of Cerro Coronel, NW-facing
slope, just S of the río Colorado, 10°40'30"N, 083°39'30"W, 10–80 m, 17–18 Sep
1986 (fr), G. Davidse & G. Herrera 31469 (MO); Hacienda Tapezco-Hds, La Suerte,
29 air km W of Tortuguero, 10°30'N, 083°47'W, 40 m, 20 Aug 1979 ( ), C. Da-
vidson & J. Donahue 8514 (MO); Parque Nacional Tortuguero, Estación Agua Fría, 3
km al Sur, Sendero Real, 10°27'N, 083°34'W, 40 m, 18 Jan 1988 (fr), R. Robles 1624
(MO, NY-digital image); Refugio Nacional de Fauna Silvestre Barra del Colorado,
sector Cocorí, 30 km N de Cariari, 10°35'40"N, 083°48'00"W, 100 m, 15 Jun 1991
(fr), E. Rojas 227 (CR, MO, NY-digital image); río Santa Clara, 1 ½ mi NW. of Los
Diamantes, ne. of Guápiles, ca. 980 ft [298 m], 18 Aug 1961 (fr), G.B. Rossbach 3823
(GH); Cerro Coronel, E of río Zapote, 1 km of río Colorado, 10°40'N, 083°40'W,
10–40 m, 13–14 Sep 1986 (fr), W.D. Stevens & O.M. Montiel 24333 (MO); Pueblo
Nuevo, 17 km NE of Guácimo, 10°20'N, 083°36'W, 100 m, 07 Sep 1994 (fr), K.
omsen 1112 (CR, NY-digital image). PANAMA. Colón: Donoso, río Hoja, UTM:
E544656; N985023 [08°54'39"N, 080°35'38"W], 23 Aug 2009 (fr), B. Araúz & J. De
Gracia 2118 (MO). Panamá: Zona del Canal: Barro Colorado Island, Frank Drayton
Trail, [09°09'15"N, 079°51'05"W], [10–150 m], 22 May 1968 ( bud), T.B. Croat
5769 (MO); Barro Colorado Island, William Morton Wheeler Trail, [09°09'20"N,
079°51'10"W], [10–170 m], 22 Sep 1968 (fr), T.B. Croat 6295 (MO); Barro Col-
orado Island, Drayton House, [09°08'29"N, 079°50'35"W], [10 m], 28 Feb 1969
(st), T.B. Croat 8262 (MO); Barro Colorado Island, Drayton House, [09°08'29"N,
079°50'35"W], [10 m], 16 Jul 1970 (fr), T.B. Croat 11337 (F, MO); Drayton House,
[09°08'29"N, 079°50'35"W], [0–5 m], 28 Aug 1970 (fr), T.B. Croat 11939 (MO);
Barro Colorado Island, James Zetek Trail, [09°09'31"N, 079°52'05"W], [10–100 m],
07 Jun 1971 ( ), T.B. Croat 14926 (F, GH, MO-2 sheets); Barro Colorado Island,
Drayton House, 06 Jul 1969 (fr), R. Foster 1093 (F, GH); Pipeline Road, 4 mi. N of
Gamboa, [09°10'N, 079°46'W], [50–100 m], 21 Dec 1971 (st), A.H. Gentry 3230
(MO); Pipeline Road, [09°10'N, 079°46'W], 100 m, 10 Aug 1971 (fr), E. Lao et al.
Two new species and a new combination in Protium (Burseraceae) from Costa Rica 111
16 (GH, MO-2 sheets); Pipeline Road, 16.6 km from beginning of road, N along río
Agua Salud, [09°14'38"N, 079°48'57"W], 0–100 m, 22 Sep 1974 (fr), S.A. Mori &
J.A. Kallunki 2039 (GH, MO); along río Mendosa near Pipeline Road, 8 km NW of
Gamboa, [09°09'36"N, 079°44'44"W], 95 m, 01 Nov 1973 (fr), M. Nee 7743 (MO);
Isla Barro Colorado, 120 m, 12 Oct 2006 (fr), R. Pérez & S. Aguilar 1632 (MO); Barro
Colorado Island, W of Drayton House, [09°08'29"N, 079°50'35"W], [0–10 m], 16
Feb 1932 ( ), R.H. Woodworth & P.A. Vestal 605 (A, MO).
e pseudarils of Protium costaricense are reported to have a pleasant avor (L.J.
Poveda 3663, USJ).
Acknowledgments
We are grateful for the facilities and hospitality extended by the following herbaria:
A, CR, F, GH, MO, and USJ. We appreciate the photographers and institutions that
made their photographs available: Reinaldo Aguilar, Xavier Cornejo, Carmen Gal-
dames, Steven Paton, Rolando Pérez, Orlando Vargas and Nelson Zamora; CR, NY,
OTS, and STRI. Photographs from the eld are reproduced with permission from
the following sources: Flickr by Reinaldo Aguilar (https://www.ickr.com/photos/),
Flórula Digital de la Estación Biológica La Selva (http://sura.ots.ac.cr/orula4/), Mu-
seo Nacional de Costa Rica, (http://ecobiosis.museocostarica.go.cr/), and Smithsonian
Tropical Research Institute Herbarium (http://biogeodb.stri.si.edu/herbarium/). Jes-
sica Jiménez (Figs 1, 4) and Alex Mauricio Campos (Fig. 7) prepared the illustrations.
We would also like the thank the following people for their generous assistance with
various aspects of this study: Reinaldo Aguilar, Isidro Chacón, Miguel Chaves, Arman-
do Estrada, Michael H. Grayum, Barry E. Hammel, José Esteban Jiménez, M. Lúcia
Kawasaki, Ronald Liesner, Isabel Pérez, Amy Pool, and Peter F. Stevens. We thank
Michael H. Grayum for kindly reading earlier versions of the manuscript. Finally, a
special thanks to John D. Mitchell and, especially, Douglas C. Daly and their careful
revision, edits and helpful suggestions that considerably improved this contribution;
DCD also provided additional features to dierentiate P. aguilarii from its congeners.
LPL was funded by an NSF Postdoctoral Research Fellowship in Biology under Grant
No. 1523880.
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... Etymology:-The specific epithet honors Daniel Santamaría-Aguilar, a very talented Costa Rican botanist with an extensive knowledge of the Neotropical flora, having described several new species of different angiosperm families (e.g., Santamaría-Aguilar 2015, Santamaría-Aguilar & Ortiz 2016, Santamaría- Aguilar & Lagomarsino 2017. He is also working on the taxonomic treatment of Burseraceae for the Flora of Costa PERDIZ eT AL. ...
... bicolored, green below and red above) when mature (see Table 1, Fig. 2), and broadly and slightly obliquely ovoid or broadly ellipsoid when more than one pyrene develops (vs. narrowly oblique-ovoid), and inflorescence usually ramified to the third (vs. the second) order (see pictures of P. santamariae in Santamaría- Aguilar & Lagomarsino (2017), Fig. 6G-J, cited as P. aracouchini). ...
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... However, curiously, in very wet lowland forests at the Península de Osa (a region adjacent to Golfito), no Lepanthes species has previously been recorded. Nonetheless, this region potentially habored Lepanthes species because of its very humid forests (Luer, 2003 (Bogarín et al., 2008;Hammel, 2015;Santamaría-Aguilar et al., 2016, 2019Santamaría-Aguilar and Lagomarsino, 2017;Santamaría-Aguilar and Aguilar Fernández, 2017;Callejas Posada, 2020;Rodríguez A NEW LEPANTHES (ORCHIDACEAE: PLEUROTHALLIDINAE) FROM PENÍNSULA DE OSA, PUNTARENAS, COSTA RICA and Santamaría-Aguilar, 2020). In addition, some of these plant species could inhabit areas highly vulnerable to habitat fragmentation and loss (Karremans and Lehmann, 2018;Bogarín, et al., 2020). ...
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Lepanthes is one of the most species-rich genera of orchids in the Neotropics, with most of the species found in medium to high elevation forests and few species in lowlands. We describe and illustrate Lepanthes osaensis, a new species from the very wet lowland forest of Península de Osa, Costa Rica. It is similar to Lepanthes cuspidata but differs mostly in the vinous leaves; smaller sepals; the narrower, bilobed petals; and the smaller lip with triangular blades. Notes on its distribution, habitat, flowering, and conservation status, as well as discussion of a taxon with similar morphology, are provided.
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This review of Neotropical Burseraceae emphasizes developments since the last major review of the family in 2011. The Burseraceae comprise a Laurasian group (represented by Eocene fossils in the Northern Hemisphere) that originally dispersed through Central America into Amazonia. During cooling and drying events in the Oligocene, the frost-intolerant northern American progenitors were likely driven extinct; subsequently, the family experienced several vicariant events and later several long-distance dispersals across the Southern Hemisphere. From Amazonia, the family re-colonized Central America and the Caribbean. The most rapid diversifications in the Americas for the Burseraceae occurred during the Miocene in Protium and Bursera, much of it through geological events, dispersal, and habitat specialization. A number of taxonomic advances were made in Neotropical Burseraceae since 2011; these included 59 published new species overall, re-drawn generic limits in tribe Protieae, new genus records for Burseraceae in Central America and the Cerrado of Brazil, new taxa that more than doubled the number of Neotropical Dacryodes, and a recently recognized center of diversity for Protium in the Andes. Revised generic descriptions and a new key to the New World genera of Burseraceae are provided. Special attention is given to the implications of leaf architecture for characterization of clades. Monoecy (rare) and parthenocarpy (possibly frequent) are discussed, and the close relationship of dioecious trees and small bee pollination is highlighted. Most Burseraceae are dispersed by birds or arboreal mammals that carry pyrenes relatively short distances away from the mother tree; however, other modes are found in the family, including wind dispersal (rare in New World Burseraceae), clumped dispersal of pyrenes by ants and lizards, and oilbirds that can disperse Dacryodes fruits more than 30 km; some dispersers that ingest pyrenes also aid in germination. The diversity and abundance of Burseraceae in a number of regions and habitats (but especially in moist forests of Amazonia and dry forests of Mexico) are striking. This, plus the fact that the taxonomy and phylogeny of New World Burseraceae are relatively well-resolved, spotlights the Burseraceae as an important model organism for researching mechanisms of diversification, species limits, cryptic species, and “hyperdominance” in tropical forests. High chemical diversity and differences in biological activity make sense in the context of diversification and coexistence. Studies of chemical defenses support the idea of a “growth defense trade-off” and suggest that selection by different natural enemies could be implicated in the speciation process; they also show that closely related species often display high chemical divergence, and plants with the most chemical defenses have a lower number and diversity of insect herbivores. The range of physical and chemical properties of Burseraceae resin is reflected in their cultural uses, which are diverse while showing strong ethnobotanical convergence.
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The Alberto Brenes Biological Preserve is located in northwest Costa Rica, situated on the Caribbean slope with elevations ranging from 800 to 1500 m. This area is subject to high rain fall throughout the year (4500 mm), and a somewhat lower rain fall from February to April. The moderate temperatures (due to elevation and cloud coverage) and the ample rainfall result in a Submontane Evergreen Tropical Rain Forest. This forest can also be considered as a transitional one between Montane Cloud Forest, found at higher elevations, and Lowland Rain Forest characteristic of lower elevations, as would be expected at this elevation. We present an annotated checklist for the Preserve accounting for 137 families of Angiosperms in 483 genera and 1294 species. The families with the most species are Araceae (50), Gesneriaceae (52), Melastomataceae (61), Orchidaceae (208), Rubiaceae (75), and Solanaceae (37). The most species-rich monocot genera are Pleurothallis (34), Anthurium (25) and Maxillaria (23); the most species-rich dicot genera are Peperomia (33), Piper (23), Miconia (16), Ocotea (21) and Psychotria s.s. (15).The following species were described as new from the preserve: Drymonia submarginalis (Gesneriaceae), Ocotea gomezii, Ocotea morae, Povedadaphne quadriporata (Lauraceae), Marlierea mesoamaricana (Myrtaceae), Prosthechea ortizii (Orchidaceae, as Encyclia ortizii), Coccoloba porphyrostachys, Coccoloba liportizii (Polygonaceae), Rudgea monofructus (Rubiaceae) and Ticodendron incognitum (Ticodendraceae); the latter described as a new family too.
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Questa è la seconda parte della flora sinottica Prima Flora Colombiana. Essa segue il medesimo modello della prima parte (Burseraceae in Webbia 12: 375–441, 1957) ma per le Malpighiaceae le chiavi delle specie sono di tipo dicotonico. Corne nelle Burseraceae vengono dati i sinonimi, la distribuzione basata sulle raccolte, osservazioni tassonomiche, nomi vernacolari ed usi; vengono anche aggiunte considerazioni circa le proprietà narcotiche di alcune specie di Malpighiaceae (per esempio yagé, caapi).
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div class="page" title="Page 1"> La Reserva Biológica Alberto Brenes está localizada al noroeste de Costa Rica, en la vertiente Caribe, con elevaciones que van de 800 a 1500 m. Esta área está expuesta a altas precipitaciones durante todo el año (4500 mm) con pocas lluvias de febrero a abril. Con temperaturas moderadas debido a la ele- vación, la cobertura de nubes y la alta precipitación dan como resultado un bosque tropical húmedo de pre- montano, que también se puede ubicar como un bosque nuboso de transición entre el bosque nuboso de montano, que se halla en elevaciones más altas, y el bosque tropical húmedo de elevaciones más bajas. Así, por su ámbito altitudinal intermedio, los bosques de la Reserva Biológica A. M. Brenes poseen una diversi- dad notable de especies que pertenecen a zonas bajas y altas de Costa Rica. Se presenta aquí una lista con anotaciones de las plantas de la Reserva distribuidas en 137 familias de angiospermas, representadas por 483 géneros y 1294 especies. Las familias con mayor número de especies son Araceae (50), Gesneriaceae (52), Melastomataceae (61), Orchidaceae (208), Rubiaceae (75) y Solanaceae (37). Los géneros de mono- cotiledóneas con mayor número de especies son Pleurothallis (34), Anthurium (25) y Maxillaria (23). Los géneros de dicotiledóneas con mayor número de especies son Peperomia (33), Piper (23), Miconia (16), Ocotea (21) y Psychotria s.s. (15). Las siguientes especies fueron descritas de la Reserva A.M. Brenes como nuevas para la ciencia: Drymonia submarginalis (Gesneriaceae), Ocotea gomezii , Ocotea morae , Povedadaphne quadriporata (Lauraceae), Marlierea mesoamericana (Myrtaceae), Prosthechea ortizii (Orchidaceae, como Encyclia ortizii ), Coccoloba porphyrostachys , Coccoloba liportizii (Polygonaceae), Rudgea monofructus (Rubiaceae) y Ticodendron incognitum (Ticodendraceae); ésta última, una familia nueva para la ciencia. </div
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Trattinnickia is closely related toDacryodes and is transferred to the tribe Canarieae. Additional characters are proposed to supplement the weak characters that traditionally have been used to separateCrepidospermum andTetragastris fromProtium. The assumed “intermediate” genusParaprotium is shown to be an unnatural assemblage and is here merged withProtium. This revised concept of generic limits in the Neotropical Protieae and Canarieae leads to the following new combinations:Tetragastris occhionii (Rizzini) Daly,Protium pilosum (Cuatrec.) Daly,Protium nitidifolium (Cuatrec.) Daly,Protium vestitum (Cuatrec.) Daly, andDacryodes cuspidata (Cuatrec.) Daly. A key to the Neotropical genera of Burseraceae is provided.
Article
Thesis (Ph. D.)--City University of New York, 1987. Includes bibliographical references (leaves 457-469). Photocopy.
Araya 61 (CR, MO, NY-digital image); Refugio Nacional de Fauna Silvestre Barra del Colorado, Puerto Lindo, 10°41'24-digital image); Southwestern-most ridge of Cerro Coronel, NW-facing slope, just S of the río Colorado, 10°40'30
Limón: cantón de Pococí, Refugio Nacional de Fauna Silvestre Barra del Colorado, Sardinas, 10°38'24"N, 083°43'48"W, 15 m, 25 Nov 1992 (fr), F. Araya 61 (CR, MO, NY-digital image); Refugio Nacional de Fauna Silvestre Barra del Colorado, Puerto Lindo, 10°41'24"N, 083°39'00"W, 200 m, 24 Jul 1995 (fr), F. Araya & J. Corrales 803 (MO, NY-digital image); Southwestern-most ridge of Cerro Coronel, NW-facing slope, just S of the río Colorado, 10°40'30"N, 083°39'30"W, 10–80 m, 17–18 Sep 1986 (fr), G. Davidse & G. Herrera 31469 (MO);
Estación Eladio's, 10°18'30"N, 084°43'10
  • Reserva Biológica Monteverde
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);