Panamá. Prov. Colón: forested slopes of Cerro El Jefe, near large coffee finca, 9°14'02"N, 79°22'30"W, 700–1000 m elev., 24 Jan 1970, R. L. Wilbur et al. 11327 (holotype: MO [acc. 2027894; barcode MO-554811]!; isotypes: F [cat. 1694605]!, GH!, NY [barcode 04112199]!, US [cat. 2640360; barcode 03009254]!).

Rinorea callejasii is most similar to R. pubiflora var. pubiflora s.s. by the lamina with symmetrical bases and bearing domatia, filaments free and fruit symmetrical, but it differs by the branchlets densely pubescent with ferruginous trichomes (vs. branchlets glabrescent in R. pubiflora var. pubiflora s.s.), primary and secondary veins on the abaxial lamina surface densely villous with ferruginous trichomes (vs. primary and secondary veins on the abaxial lamina surface glabrescent), fruit 1.5–2 cm long with 2 ovules per carpel (vs. fruit 2–3.5 cm long with 3 ovules per carpel) and the seeds glabrous (vs. seeds pubescent).

Shrubs or trees 2–12 m tall, terminal branchlets pubescent with erect ferruginous trichomes 0.4–0.5 mm long. Leaves opposite, petiolate; petioles 2–10 mm long, pubescent with erect ferruginous trichomes 0.2–0.3 mm long; stipules deciduous, free, elliptic, 1.6–4.8 × 0.8–1.3 mm, pubescent with appressed ferruginous trichomes 0.5 mm long; lamina elliptic, 4.6–15 × 2–6.5 cm, adaxial surface pilose on the mid-vein and secondary veins with erect ferruginous trichomes 0.4–0.5 mm long, abaxially pubescent on the mid-vein, trichomes on the secondary and tertiary veins, trichomes 0.3–0.5 mm long, erect, ferruginous, base cuneate, symmetrical, margin subserrate, apex acuminate, acumen 0.7–1.2 cm long, mucronate; with 8–10 major secondary vein pairs, major secondary veins eucamptodromous becoming brochidodromous distally, with regular spacing and uniform angle of the major secondary vein; major secondary attachment to mid-vein decurrent, leaf domatia present. Inflorescence axillary, terminal or subterminal, racemose, 3–5 cm long, 0.7–1.5 cm diam., axis pubescent with erect ferruginous trichomes 0.3–0.5 mm long; pedicels 1.8–3.6 mm long, pubescent with erect golden trichomes 0.3–0.5 mm long, articulated near the base; peduncle bracts deciduous, narrowly triangular, 1.5–1.8 × 1.5–1.8 mm, herbaceous, pubescent along the costa with appressed golden trichomes 0.3–0.5 mm long, margin ciliolate; bractlets persistent below articulation, subopposite, 1 × 1 mm. Flowers 3–3.6 × 3–3.4 mm; sepals subequal, triangular, 1.3–2 × 1.4–1.6 mm, 5- to 9-veined, pubescent along the costa with appressed golden trichomes 0.3 mm long, margin ciliolate, apex apiculate; petals lanceolate, 3.4–3.8 × 0.7–1.4 mm, pubescent along the costa with appressed golden trichomes 0.3–0.4 mm long, margin entire, apex acute, white; stamens 2.9–3.3 mm long, all filaments free, dorsal gland pilose with spreading golden trichomes 0.2–0.3 mm long covering lower half of the filament, anthers elliptic, 1.1–1.4 × 0.3–0.5 mm, apex rounded, connective 0.7–0.8 mm long, dorsal connective scale conspicuous, lateral as well as apical, lanceolate, 2.2–2.5 × 0.8–1 mm, margin entire, orange-brown; ovary globose, 0.8–1 × 0.9–1 mm, pubescent with erect, golden trichomes 0.4–0.5 mm long; style erect, filiform, 2–2.3 × ca. 0.1 mm, pubescent proximally with appressed golden trichomes 0.2–0.3 mm long, stigma undifferentiated. Fruit a symmetrical, subligneous capsule dehiscent along three sutures, ellipsoid, 1.5–2 × 0.5–0.6 cm, apex acuminate, pubescent with erect golden trichomes 0.2–0.3 mm long, green in vivo, brown when dry. Seeds two per valve, globose, 4–5 mm diam., glabrous, with maculae, brown when dry. (Fig. 1)

Figure 1. 

Rinorea callejasii Hoyos-Gómez A habit B detail of young twig with pubescence C detail leaf showing domatia, abaxial surface D leaf architecture E inflorescence F flower G sepals, abaxial surface H petals, abaxial surface I stamens, adaxial surface J gynoecium K infructescence L fruit, showing two seeds per valve M seed. (A–J: M. D. Correa 765 [MO]; K–M: G. D. McPherson 6851 [MO]).

Rinorea callejasii is restricted to the Caribbean coast of Colón and San Blas Provinces, Panama and the Pacific coast of Chocó Department, Colombia, a region belonging to the biogeographic Province of Chocó-Darien in the Pacific Dominion (sensu Morrone (2014)). It occurs in lowland humid rainforests at elevations of 30–500 m. (Fig. 2)

Figure 2. 

Distribution of R. pubiflora f. grandifolia (triangles), R. pubiflora var. pubiflora (pentagons), R. pubiflora f. andersonii (stars), Rinorea callejasii (asterisks).

Rinorea callejasii honours Dr. Ricardo Callejas Posada, Professor of Biology at Universidad de Antioquia, Medellín, Colombia. Prof. Callejas has made many contributions to the taxonomic knowledge of the Piperaceae and expanded the understanding of the diverse flora of Colombia, especially in the Department of Antioquia. He was the major advisor of SEHG.

The species flowers January through March; specimens in fruit were collected in March and May.

Rinorea callejasii has a geographic range in the form of an EOO of 40,729 km² and AOO of 52 km². In Panama, it is represented by two locations within Chagres National Park and one location in Soberanía National Park. Outside of the protected areas, where the species exists at nine locations, it is threatened by the exploitation of natural resources, mineral extraction and energy development. In Colombia, it is represented by two locations, both outside protected areas. The species does not meet the criteria necessary to assign a threatened status. However, given ongoing habitat disturbance, it is likely that the species will qualify in the near future for a threatened status and, thus, it is preliminarily assessed with a status of “Near Threatened” (NT).

Rinorea pubiflora, as circumscribed by Hekking (1988), was morphologically variable across its large distribution in the Neotropics. He recognised a variety of R. pubiflora: var. grandifolia (Eichler) Hekking, which was further divided into two forms: f. grandifolia and f. andersonii (Sandwith ex Hekking) Hekking. Rinorea callejasii is a segregate of R. pubiflora var. pubiflora s.s. In attempting to explain the morphologically divergent specimens of var. pubiflora from Panama (e.g. G. C. de Nevers et al. 5075), Hekking (1988) invoked a scenario of hybridisation between R. squamata S.F.Blake and var. pubiflora. However, our analysis of morphological divergence and biogeographic distribution suggest that R. callejasii is a separately evolving lineage from the nominate species (sensu De Queiroz (2008)) and deserving of recognition as a new taxon at the rank of species.

Rinorea callejasii is sympatric with four other taxa of Rinorea. Rinorea squamata and R. hirsuta Hekking can be differentiated by their single glabrous seed per valve (vs. two glabrous seeds per valve in R. callejasii). Rinorea dasyadena is distinguished by its two pubescent seeds per valve and asymmetrical lamina base (vs. two glabrous seeds and symmetrical lamina base). Rinorea lindeniana var. fernandeziana can be separated by its one pubescent seed per valve (vs. two glabrous seeds per valve).

Colombia. Dept. Chocó. Mpio. de San José del Palmar: Hoya del río San Juan, 04°36'N, 76°54'W, 7 Apr 1979 (fr), E. Forero et al. 4780 (COL, MO, US); Mpio. Bahía Solano: hills behind Bahía Solano, 6°13'14"N, 77°24'27"W, 0–250 m elev., 5 Jan 1973 (fl), A. H. Gentry & E. Forero 7245 (COL, F, MO, U). Panamá. Prov. Colón: Santa Rita Ridge lumber road, 23 Feb 1968 (fl), M. D. Correa-A. & R. L. Dressler 765 (MO, NY, US); Santa Rita Ridge, 9°19'N, 79°39'W–9°24'N, 79°48'W, 1 Mar 1971 (fl), T. B. Croat 13831 (MO); El Llano-Cartí Road, 10 miles from Inter-American Highway near El Llano, 9°17'45"N, 78°56'15"W, 330 m elev., 28 Mar 1976 (fl, fr), T. B. Croat 33754 (COL, MO); El Llano-Cartí Road, 5–6 miles N of Inter-American Highway at El Llano, 09°15'30"N, 78°55'50"W, 350–375 m elev., 7 May 1976 (fl), T. B. Croat 34788 (COL, MO); El Llano-Cartí Road, km 26.5, trail NE from road, [coordinates on original label: 9°19'N, 78°55'W; corrected to 9°22'N, 78°58'W], 175 m elev., 9 Mar 1985 (fl), G. C. de Nevers et al. 5075 (MBM, MO); Santa Rita Ridge road, 4 miles from Transisthmian Highway to Agua Clara weather station, 9°21'N, 79°42'W, 11 Dec 1973 (fl), R. L. Dressler et al. 8846 (MO); Santa Rita Ridge road, 4 miles from Transisthmian Highway to Agua Clara weather station, 9°21'N, 79°42'W, 500 m elev., 11 Dec 1973 (fl), A. H. Gentry et al. 8846 (MO, COL); along road into Santa Rita, E of Agua Clara, 9°07'10"N, 79°42'02"W, 4 Mar 1973 (fl), H. Kennedy 2741 (MO); Santa Rita Ridge, 9°20'N, 79°45'W, 13 May 1986 (fl), G. D. McPherson 9168 (MG, MO); 12.7 km from Inter-American Highway, 350 m elev., 15 Feb 1975 (fr), S. A. Mori 4694 (COL); Santa Rita Ridge Road, 17 km from Boyd-Roosevelt Hwy, 9°22'N, 79°40'W, 450 m elev., 14 Mar 1975 (fl), S. A. Mori et al. 5041 (COL, MO); El Llano-Cartí Road, 20.7 km north from Inter-American Hwy, 9°17'58"N, 78°55'58"W, 20 Mar 1975, S. A. Mori & J. A. Kallunki 5115 (COL, MO); 9 km from Inter-American Highway, 9°15'50"N, 78°55'51"W, 350 m eelv, 22 Mar 1975 (fr), S. A. Mori 5154 (COL, MO); upper Río Piedras headwaters, along trail from end to Santa Rita Ridge road, ca. 11 km SW of Cerro Braja, 9°25'N, 79°35'W, 4 May 1981 (fr), K. J. Sytsma et al. 4311 (MO). Prov. San Blas: El Llano-Cartí Road, 7 miles on Interamerican Highway, 9°15'N, 79°00'W, 550 m elev., 14 Mar 1985 (fr), G. D. McPherson 6851 (MO).

Venezuela. Dept. Amazonas de Atabapo: “Caño Iguapo” Alto Orinoco, 15 km al SE de la Esmeralda, 3°8'N, 65°27'W, 100–180 m elev., 20 Feb 1990, G. Aymard et al. 8050 (holotype: MO [acc. 5983541; barcode MO-1980109]!; isotype: MA [cat. 543845]!).

Rinorea aymardii is similar to R. melanodonta S.F.Blake by the lamina with domatia and symmetrical bases, filaments free and style erect, but it differs by the smaller, herbaceous, lanceolate lamina, 4–11 × 1–2.7 cm (vs. larger, coriaceous, elliptic lamina, 6.5–17 × 1.7–5.2 cm in R. melanodonta), 7–8 major secondary vein pairs (vs. 9–12 major secondary vein pairs), abaxial tertiary venation percurrent, pubescent (vs. abaxial tertiary veins reticulate, glabrescent), petals 2.8–3 × 1–1.2 mm, with entire margin and pubescent at the apex (vs. petals 5–5.7 × 1.5–2.2 mm, with margin ciliolate and glabrous at the apex), shorter stamens, 2.8–3 mm long (vs. longer stamens, 4–4.5 mm long), dorsal gland glabrous (vs. dorsal gland pilosulous) and smaller anther connective scales, 2–2.1 × 0.7–0.8 mm (vs. larger anther connective scales, 3.2–3.5 × ca. 1.5 mm).

Treelets to 3 m tall, young branchlets strigose with erect golden trichomes 0.1–0.2 mm long, glabrescent. Leaves opposite, petiolate; petiole 1.8–3.8 mm long, pubescent with erect golden trichomes 0.1–0.2 mm long; stipules deciduous, free, lanceolate 2–2.4 × 0.9–1 mm, herbaceous, pubescent with appressed golden trichomes 0.1–0.2 mm long, apex mucronate; lamina elliptic-lanceolate, 4–11 × 1–2.7 cm long, adaxially glabrous, abaxially pubescent on the mid-vein and veins with appressed golden trichomes 0.2–0.4 mm long, with 7–8 major secondary vein pairs, secondary veins with irregular spacing, vein angles smoothly decreasing proximally, mixed epimedial tertiary veins, symmetrical and rounded base, margin crenate, apex acuminate, acumen 0.5–1 cm, mucronate, leaf domatia absent. Inflorescence axillary, terminal or subterminal, racemose, 1.7–4 cm long, 0.8–1 cm diam., axis pubescent with erect golden trichomes 0.2 mm long; pedicels 1.7–2 mm, pubescent with erect golden trichomes 0.2 mm long, articulated near the middle; bractlets persistent below articulation, opposite, ca. 1 × 1 mm, herbaceous, the costa pubescent with appressed golden trichomes 0.2 mm long, peduncle bracts triangular, ca. 1 × 1.2 mm, herbaceous, pubescent with appressed ferruginous trichomes 0.2 mm long, margin ciliolate. Flowers 3–3.5 × 3–3.5 mm, sepals subequal in size and shape, triangular, 1.4–2.4 × 1–1.3 mm, 3–5-veined, costa pubescent with appressed golden trichomes, 0.2 mm long, margin ciliolate, apex apiculate; petals lanceolate, 2.8–3 × 1–1.2 mm, pubescent with erect golden trichomes 0.1–0.2 mm long near the apex, margin entire, apex acute, cream-coloured in vivo, brown when dry; stamens 2.8–3 mm long, filaments free, glabrous, 0.4–0.6 mm long, dorsal glands glabrous, completely covering the filaments; anthers lanceolate, 1–1.2 × 0.4–0.5 mm, glabrous, apex rounded, connective 1–1.2 mm long, dorsal connective scale lanceolate, 2–2.1 × 0.7–0.8 mm, scarious, brown–orange, margin subentire; ovary globose, 0.8–1 × 0.8–0.9 mm, pubescent with erect golden trichomes 0.2–0.3 mm long; style erect, filiform, glabrous, 1.7–1.9 × ca. 0.1 mm, stigma acute. Fruit unknown. (Fig. 3)

Figure 3. 

Rinorea aymardii Hoyos-Gómez A habit B leaf architecture C detail of leaf, abaxial surface D infructescence E flower F sepals, abaxial surface G petals, abaxial surface H stamens, showing both surfaces I gynoecium. (A–I: G. Aymard 8050 [MO]).

Rinorea aymardii, which is only known from the type specimen, occurs the Alto Orinoco–Casiquiare Biosphere Reserve in Amazonas State, Venezuela, a biogeographic area classified as the Province of Imerí in the Boreal Brazilian Dominion (sensu Morrone (2014)). It grows in terra firma humid forest at elevations of 100–180 m. The landscape of the type locality is characterised by gently sloping (10–20%) hills composed of granitic and quarzitic rocks (Aymard 2000). (Fig. 4)

Figure 4. 

Distribution of R. melanodonta (triangles), R. macrocarpa (asterisks), R. betancurii (pentagons), Rinorea aymardii (squares).

Rinorea aymardii honors the Venezuelan botanist Gerardo Aymard, who has made significant contributions to the taxonomic knowledge of the flora of Venezuela, especially the south-western part of the country in the Río Negro and Orinoco River Basins.

The species is only known from one flowering specimen collected in February.

Rinorea aymardii has a geographic range in the form of an EOO of 4 km² and AOO of 4 km². It is currently known only from the type collection, which represents one location within the terra firma forests of the Alto Orinoco–Casiquiare Biosphere Reserve in Venezuela where it is threatened by illegal logging and mining. Based on its limited AOO, the single location and the projected continuing decline in the quality of habitat due to uncontrolled deforestation and resource extraction, the species might be preliminarily assigned to the “Critically Endangered.” However, given the difficulty accessing this inadequately explored area, it is likely there are additional occurrences that would allow a more accurate assessment of the risk of extinction than is possible at this time. The species is therefore preliminarily assigned to the IUCN (2012) Category of “Data Deficient” [DD].

Rinorea aymardii is sympatric with R. macrocarpa from which it can be differentiated by the smaller lamina 4–11 × 1–2.7 cm with crenate margins (vs. larger lamina 5.5–21.5 × 2–9.2 cm with subentire to subcrenate margins in R. macrocarpa), filaments free (vs. filament fused into a tube) and style filiform (vs. style conical).

Colombia. Dept. Caquetá: alrededores del refugio la Selva, 1°4'0.23"N, 72°44'24.6"W, 630 m elev., 1 Dec 1992, M. Velayos et al. 6509 (holotype: MO [acc. 4644279; barcode MO-1590549]!; isotypes: COL!, MA [cat. 543845]!).

Rinorea chiribiquetensis is similar to R. ovalifolia (Britton) S.F.Blake by the lamina base symmetrical, margin subserrate, ovary globose and style filiform, but it differs by the smaller lamina, 3.2–7.2 × 1.6–3.6 cm, with appressed pubescence on the abaxial surface and domatia present (vs. the larger lamina, 3–16 × 2–7.5 cm, glabrescent and lacking domatia in R. ovalifolia), shorter inflorescence, 1.5–5 cm long (vs. longer inflorescence, 4–12 cm long), shorter pedicels, 1.7–3.3 mm long (vs. longer pedicels, 4–6 mm long) and stamens with filaments free (vs. stamens with filaments fused).

Shrubs to 1.5 m tall, terminal branchlets pubescent with appressed golden trichomes 0.3 mm long, lenticels 0.5–0.6 mm. Leaves opposite, petiolate; petiole 3–6 mm long, pubescent with erect golden trichomes 0.2–0.3 mm long; stipules deciduous, free, lanceolate 1.9–2.1 × 0.9–1 mm, pubescent with appressed golden trichomes 0.2 mm long, margin entire; lamina elliptic, 3.2–7.2 × 1.6–3.6 cm, adaxially pubescent on mid-vein with appressed golden trichomes 0.2–0.3 mm long, abaxially pubescence on mid-vein, secondary and tertiary veins with appressed golden trichomes 0.2–0.3 mm long; with 5–6 major secondary vein pairs, semi-craspedodromous; secondary veins with irregular spacing, and veins angle smoothly decreasing proximally between them; symmetrical base, cuneate, margin subserrate, apex acuminate, acumen 5–6 mm long, mucronate, leaf domatia present. Inflorescence axillary, lateral or terminal, racemose, 1.5–5 cm long, 0.5–1 cm diam., axis pubescent with erect golden trichomes 0.2 mm long; pedicels 1.7–3.3 mm long, articulated near the middle, pubescent with erect golden trichomes 0.2 mm long,; bractlets persistent below articulation, subopposite, ca. 1 × 1 mm, herbaceous, costa pubescent with appressed golden trichomes 0.2 mm long, 3–5 veins; peduncle bracts deciduous, narrowly triangular 1–1.4 × 1–1.5 mm, herbaceous, pubescent with appressed golden trichomes 0.2 mm long, 3–5 veins, margin ciliolate. Flowers 3.2–3.5 × 3–3.3 mm; sepals subequal in size and shape, triangular, 1.5–1.7 × 1.7–1.9 mm, 5–7 veined, costa pubescent with appressed golden trichomes 0.2 mm long, margin ciliolate, apex apiculate; petals lanceolate, 3–3.3 × 1.4–1.6 mm, pubescent with erect golden trichomes 0.2 mm long near the apex, margin entire, apex acute; stamens 2.6–3 mm long, filaments free, 0.7–0.9 mm, dorsal gland glabrous, completely covering the filament; anthers elliptic, 1–1.1 × 0.3–0.5 mm, glabrous, apex rounded, connective 0.6–0.7 mm long, dorsal connective scale lanceolate, 2.2–2.4 × 0.8–1 mm, scarious, margin subentire, brown-orange; ovary globose, 0.7–0.8 × 0.8–0.9 mm, pubescent with appressed golden trichomes 0.4 mm long; style erect, filiform, 1.7–2 × ca. 0.1 mm, glabrous, stigma acute. Fruit unknown. (Fig. 5)

Figure 5. 

Rinorea chiribiquetensis Hoyos-Gómez A habit B detail of young twig with pubescence C detail of young twig showing lenticels D detail of leaf showing pubescence, adaxial surface E detail of leaf showing domatia, abaxial surface F leaf architecture G infructescence H flower I sepals, abaxial surface J petals, abaxial surface K gynoecium L stamens, showing adaxial and abaxial surfaces, with a detail of the dorsal gland (left). (A–L: M. Velayos 6509 [MO)].

Rinorea chiribiquetensis is endemic to Caquetá Department, Colombia, an area belonging to the Biogeographical Province of Imerí in the Boreal Brazilian Dominion (sensu Morrone (2014)). It occurs in lowland tropical rainforest from rocky substrates at an elevation of 630 m. (Fig. 6)

Figure 6. 

Distribution of Rinorea chiribiquetensis (circle), R. ovalifolia (triangles), R. stevensii (stars).

Rinorea chiribiquetensis is named for the Chiribiquete National Park in Colombia.

The species is only known from one flowering specimen collected in December.

The region encompassing the vast and remote Chiribiquete National Park is relatively little explored as evidenced by the scarcity of collections in the Tropicos database (https://www.tropicos.org; accessed 25 February 2023), GBIF (https://www.gbif.org; accessed 25 February 2023) and other Colombian Herbaria (e.g. COAH, COL, HUA). There are insufficient data to assess the risk of extinction of Rinorea chiribiquetensis and, therefore, the species is preliminarily assigned to the IUCN (2012) category of “Data Deficient” [DD].

Rinorea chiribiquetensis is sympatric with R. macrocarpa, but it can be differentiated by the presence of domatia (vs. domatia absent in R. macrocarpa), smaller lamina, 3.2–4.2 × 1.6–3.6 cm (vs. larger lamina, 5.5–21.5 × 2–9.2 cm), filaments free (vs. filaments fused into a tube) and style filiform (vs. style conical). Rinorea chiribiquetensis can be differentiated from R. stevensii by the smaller lamina, 3.2–4.2 × 1.6–3.6 cm (vs. larger lamina, 4–12.5 × 2.5–7 cm in R. stevensii) and less dense pubescence on the abaxial lamina surface, with trichomes 0.2–0.3 mm long (vs. denser pubescence on the lamina, with trichomes 0.4–0.5 mm long).

Colombia. Dept. Vichada: Parque Nacional “El Tuparro,” piedra canal near the south end of airstrip at Centro Administrativo, 5°17'N, 67°52'W, ca. 100 m elev., 5 Mar 1985, J. L. Zarucchi et al. 3577 (holotype: NY [barcode 04205808]!; isotypes: COAH [cat. 24597]!, COAH [cat. 55474]!, FMB [cat. 4307]!, FMB [cat. 21506]!, MO [acc. 3499269; barcode MO-1592746]!, U [barcode U1766360]!).

Rinorea stevensii is similar to R. ovalifolia by the lamina base symmetrical, style erect and two glabrous seeds per valve, but it differs by the lamina abaxial surface with golden pubescence (vs. the lamina abaxial surface glabrescent in R. ovalifolia), domatia present (vs. domatia absent), inflorescences cylindrical, 2–5 cm long (vs. inflorescence conical, 4–12 cm long), costa of petals and sepals pubescent (vs. costa of petals and sepals glabrescent), filaments free (vs. filaments fused), and capsule pubescent (vs. capsule glabrescent).

Trees 1.5–4 m tall, terminal branchlets pubescent with erect golden trichomes 0.4–0.5 mm long. Leaves opposite, petiolate; petiole 2–6 mm, pubescent with long golden trichomes 0.3–0.4 mm long; stipules deciduous, free, lanceolate, herbaceous, 2.5–5 × 1–1.5 mm, pubescent with appressed golden trichomes 0.4–0.5 mm long; margin entire; lamina elliptic, 4–12.5 × 2.5–7 cm, adaxially pubescent on mid-vein and abaxially pubescent on mid-vein and secondary veins with erect golden trichomes 0.4–0.5 mm long; semi-craspedodromous, with 6–9 major secondary vein pairs, secondary veins with irregular spacing and vein angles smoothly decreasing proximally between them, base rounded, symmetrical, margin crenate, ciliate, apex acuminate, acumen 0.5–1 cm long, mucronate, domatia present. Inflorescence axillary, lateral or terminal, racemose, 2–5 cm long, 1–1.5 cm diam., axis pubescent with erect golden trichomes 0.4–0.5 mm long; pedicels 2.5–6.5 mm, pubescent with erect golden trichomes 0.4–0.5 mm long, articulated near the middle; bractlets persistent below articulation, subopposite, ca. 1 × 1 mm, herbaceous, costa pubescent with appressed golden trichomes 0.4–0.5 mm long; peduncle bracts persisting, broadly triangular, 1–1.5 × 1–1.5 mm, herbaceous, pubescent with appressed golden trichomes 0.4–0.5 mm long, margin ciliolate. Flowers 2.8–3.8 × 3–3.4 mm long, sepals subequal, triangular, 2–2.5 × 1–1.6 mm, 7–11-veined, pubescent with appressed golden trichomes 0.5 mm long, margin ciliolate; petals lanceolate, 2.8–4.3 × 1.2–1.9 mm, costa pubescent with appressed golden trichomes 0.3–0.4 mm long, margin entire, cream to yellow in vivo, pale brown when dry; stamens 2.5–3 mm long, all filaments free, 0.4–0.6 mm, dorsal gland covering completely the filament, glabrous; anthers elliptic, 1.4–1.7 × 0.3–0.8 mm, apex obtuse, connective 0.8–0.9 mm, dorsal connective scale lanceolate, 2.4–2.5 × 0.9–1.0 mm long, margin subentire, scarious, orange-brown; ovary globose, 0.8–1.5 × 0.7–1.3 mm, pubescent with appressed golden trichomes 0.5 mm long; style erect, filiform, 1.8–2.4 × ca. 0.1 mm long, pubescent proximally with appressed trichomes 0.5 mm long, stigma acute. Fruit a symmetrical, subligneous capsule dehiscent along three sutures, ellipsoid, 1.5–2.3 × 0.5–0.6 cm, acuminate at the apex, veined, pubescent with erect golden trichomes 0.5 mm long, green when fresh, brown when dry. Seeds two per valve, globose, 4–5 mm diam., glabrous, without maculae, brown when dry. (Fig. 7)

Figure 7. 

Rinorea stevensii Hoyos-Gómez A habit B detail of young twig showing lenticels and pubescence C detail of leaf showing domatia, abaxial surface D detail of leaf showing margin ciliate, adaxial surface E branchlet with inflorescences F leaf architecture G infructescence H flower I sepals, abaxial surface J petals, abaxial surface K stamens, showing adaxial and abaxial surfaces L gynoecium M fruit N detail of fruit showing two seeds per valve O seed. (A–D: B. M. Boom 6564 [MO]; E–O: J. L. Zarucchi 3577 [MO]).

Rinorea stevensii occurs in Colombia in the border region with Venezuela near the Orinoco River and in El Tuparro National Park. It also occurs in Venezuela in the vicinity of Puerto Ayacucho. This distribution corresponds to the Biogeographical Provinces of Pantepui and Sabana, in the Boreal Brazilian and Pacific Dominions, respectively (sensu Morrone (2014)). The species grows on granitic hills and slopes in lowland semi-deciduous tropical rainforests and savannahs, at elevations of 85–520 m. (Fig. 6)

Rinorea stevensii is named in honour of Dr. Peter Stevens, Professor of Biology at the University of Missouri Saint-Louis and curator of the Missouri Botanical Garden. Prof. Stevens was the major advisor for SEHG’s master’s degree.

Aruni yó (B. M. Boom et al. 6564).

Rinorea stevensii has a geographic range in the form of an EOO of 5,070 km² and AOO of 56 km². It is represented by two locations within the El Tuparro National Park in Colombia and eight locations outside of protected areas in Venezuela where it is threatened by deforestation, illegal resource extraction and uncontrolled fires. Based on its limited AOO, the number of locations and the projected continuing decline in the quality of habitat, the species is preliminarily assigned to the “Vulnerable” category [VU B2ab(iii)].

Based on several morphological differences and an allopatric distribution, Rinorea stevensii is segregated from R. ovalifolia (as circumscribed by Hekking (1988)). Hekking noted the variable morphology of some specimens now representing R. stevensii, but he was unsure of their placement: “The leaves of Rinorea ovalifolia are variable in character, e.g. the underside of the leaves varies from densely hispidulous to glabrescent and domatia may be present or not”. Hekking invoked introgressive hybridisation between R. ovalifolia and R. pubiflora var. pubiflora to explain the morphological variability, but we disagree with his opinion. We hypothesise that the distinctive morphological characteristics and the discrete biogeographic distribution suggest that R. stevensii is a separately evolving lineage (sensu De Queiroz (2007)) making it worthy of recognition at the rank of species. Key morphological differences amongst R. stevensii, R. ovalifolia and R. pubiflora var. pubiflora are presented in Table 1.

In addition to an allopatric distribution, Rinorea stevensii can be differentiated from R. chiribiquetensis (the other species segregated from R. ovalifolia) by several morphological characters. It has longer and more abundant pubescence on the lamina, with trichomes 0.4–0.5 mm long (vs. shorter and less abundant pubescence on the lamina, with trichomes 0.2–0.3 mm long in R. chiribiquetensis), larger lamina, 4–12.5 × 2.5–7 cm (vs. smaller lamina, 3.2–4.2 × 1.6–3.6 cm) and larger petals 2.8–4.3 × 1.2–1.9 mm (vs. 3–3.3 × 1.4–1.6 mm). Rinorea pubiflora var. pubiflora co-occurs with R. stevensii, but it is differentiated by having three pubescent seeds per valve, whereas R. stevensii has two glabrous seeds per valve.

Colombia. Dept. Vichada. Mpio. de Puerto Carreño: Cerro al N del Centro Adm. Inderena, 5°21'17.0"N, 67°51'40.6"W, 4 Apr 1995 (fr), M. P. Córdoba et al. 1266 (COAH, COL, FMB); base del cerro Rocoso, 100 m elev., 8 Oct 1979 (fl), P. Vincelli 1047 (COAH, COL, FMB). Venezuela. Estado Amazonas. Mpio. De Puerto Ayacucho: 35 km. south of Puerto Ayacucho, at the “Tobagón”. Large igneous outcrop bordering forest on slope, 85 m elev., 4 May 1977 (fr), J. Steyermark & O. Huber 113844 (COL, MO). Mpio. de Atures: 23 km NE of Puerto Ayacucho and 10 km E of the highway, hills and base of hills, near Cachama, 90 m elev., 5°51'N, 67°24'W, 90 m elev., 19 Apr 1978 (fr), G. Davidse et al. 15300 (MO, NY, U, US); 14–15 km NE of Puerto Ayacucho, along road to “El Burro,” 5°47'N, 67°32'W, 85 m elev., 15 Apr 1978 (fr), G. Davidse et al. 15070 (MO, NY, US); alrededores de Puerto Ayacucho +/- 11 km N, 5°44'N, 67°30'W, 15 Apr 1978 (fl), O. Huber et al. 1450 (U, US); ibid., 5°43'N, 67°30'W, 29 Jan 1978 (fl), O. Huber et al. 1502 (K, NY, U, US); Piedra el Berraco, laja granítica que conduce a Provincial, 10 km al NE de Puerto Ayacucho, 5°47'N, 67°34'W, Apr 1997 (fl, fr), Á. Fernández-Pérez et al. 10800 (US); E of río Parguaza, 125 km N of Puerto Ayacucho, 6°17'N, 67°5'W, 11 Sep 1985 (st), J. A. Steyermark et al. 131751 (MO, U); Parguaza, 22 Apr 1946 (fr), I. Velez 2448 (US). Estado Bolivar. Mpio. Cedeño: vicinity of Panare village of Corozal, 6 km from Manipure towards Caicara, 6°55'N, 66°30'W, 24 Sep 1985 (st), B. M. Boom et al. 6083 (NY); ibid., 6°55'N, 66°30'W, 90 m elev., 19 Apr 1986 (fr), B. M. Boom & M. Grillo 6564 (MO, F, U, US); carretera Caicara–El Burro, 16 Apr 1984 (fl), B. Stergios et al. 8494 (MO); Caicara 100 m elev., 10 Jun 1940 (st), L. W. Williams 13255 (F). Mpio. Foráneo La Urbana: cerca a la desembocadura del río Orinoco, 6°46'N, 67°00'W, 31 Jan 1989 (fl), N. Cuello 718 (MO, U, US); Los Pijiguaos, afloramiento granítico 1.5 km al N del campamento Bauxiven, 6°35'N, 66°47'W, 7 Aug 1993 (st), A. Gröger et al. 1077-B (MO); E slopes of cerro Pijiguao (N end of serrania Suapure) above Pijiguao, ca. 70 km from mouth of Río Suapure, 110–520 m eelv., 19 Jan 1956 (fl), J. J. Wurdack et al. 41295 (F, K, MG, MO, NY, U, US).

Colombia. Dept. Antioquia: Mpio. de San Luis, Cañón del Río Claro, sector norte, margen izquierda, 5°53'N, 74°39'W, 340–500 m elev., 24 Dec 1983, A. Cogollo 1075 (holotype: HUA [032011]!; isotypes: COL [299700]!, MO [acc. 3737288; barcode MO-1590440]!).

Rinorea cogolloi is similar to R. hirsuta by the elliptic lamina with ferruginous trichomes and lacking domatia, but it differs by the asymmetrical lamina base (vs. lamina base symmetrical in R. hirsuta), inflorescence thyrsoid (vs. inflorescence racemose), petals 2–3 × 0.7–1.3 mm (vs. 4–4.3 × 1.5–1.8 mm), stamens 1.7–2 mm long (vs. 3–3.3 mm long), dorsal gland pubescent (vs. dorsal gland glabrous), seeds pubescent (vs. glabrous) and capsule asymmetrical and pilose (vs. capsule symmetrical and velutinous).

Shrubs or trees 2–6 m tall, terminal branchlets pubescent with erect ferruginous trichomes 0.4–0.6 mm long. Leaves opposite, petiolate; petiole 1.9–5 mm, pubescent with erect ferruginous trichomes 0.3–0.4 mm long; stipules deciduous, free, lanceolate, membranous, 1.9–2.5 × 0.7–1 mm, pubescent with appressed ferruginous trichomes 0.2–0.3 mm long, margin ciliolate; lamina elliptic, 7–17 × 2.5–7 cm long, herbaceous, adaxially pubescent on mid-vein with erect ferruginous trichomes 0.2–0.3 mm long, abaxially pubescent with erect ferruginous trichomes 0.4–0.5 mm long, with 5–8 major secondary vein pairs, semi-craspedodromous, secondary veins with regular spacing and angled to the mid-vein, base rounded, asymmetrical, margin crenate, apex acuminate, acumen 0.5–1 cm long, mucronate, leaf domatia absent. Inflorescence axillary, lateral or terminal, thrysoid, 4–8 cm long, 0.5–1 cm diam., axis pubescent with erect ferruginous trichomes 0.2–0.3 mm long; cymules 3–5 flowered; common peduncule 1–1.5 mm, pubescent with erect ferruginous trichomes 0.2 mm long; pedicels 0.5–1 mm, articulated at the base, pubescent with erect ferruginous trichomes 0.2 mm long; bractlets alternate, persistent below articulation, ca. 0.5 × 0.5 mm, herbaceous, costa pubescent with appressed ferruginous trichomes 0.2 mm long, margin ciliolate; peduncle bracts persistent, broadly triangular, 1–1.5 × 1–1.5 mm, herbaceous, costa pubescent with appressed ferruginous trichomes 0.3–0.4 mm long; margin ciliolate. Flowers 2.5–2.8 × 2.5–2.8 mm, sepals subequal in size and shape, triangular, 1–1.5 × 0.5–0.7 mm, pubescent with appressed ferruginous trichomes 0.2 mm long, margin ciliolate; petals elliptic, 2–3 × 0.7–1.3 mm long, pubescent with appressed ferruginous trichomes 0.2 mm long, margin ciliolate, cream to yellow in vivo, brown when dry; stamens 1.7–2 mm long, filaments free, glabrous, 0.9–1 mm, dorsal gland covering the filament, pubescent with spreading ferruginous trichomes 0.2 mm long; anthers elliptic, 0.6–1 × 0.4–0.6 mm, pubescent with spreading ferruginous trichomes 0.2 mm long, apex obtuse, connective 0.6–0.7 mm, pubescent with appressed ferruginous trichomes 0.2 mm long, dorsal connective scale lanceolate, 1.2–1.6 × 0.7–0.9 mm, scarious, margin subentire, orange-brown; ovary globose, 1–1.5 mm diam., pubescent with erect ferruginous trichomes 0.3–0.5 mm long; style erect subclavate, filiform, 1.5–1.8 × 0.1 mm, stigma acute. Fruit an asymmetrical, subligneous capsule dehiscent along three sutures, ellipsoid, 0.8–1.5 × 0.5–0.6 cm, apex acuminate, veined, pubescent with erect ferruginous trichomes 0.2–0.3 mm long, green when fresh, brown when dry. Seeds one per valve, globose, 5–7 mm diam., with maculae, pubescent with erect ferruginous trichomes 0.3 mm long, brown when dry. (Fig. 8)

Figure 8. 

Rinorea cogolloi Hoyos-Gómez A habit B detail of leaf pubescent, abaxial surface C leaf architecture D infructescence E cymule with flowers F sepals, abaxial surface G petals, abaxial surface H stamens, showing adaxial and abaxial surfaces I gynoecium J infructescence K detail of fruit showing one seed per valve L seed. (A–L: A. Cogollo 786 [MO]).

Rinorea cogolloi is narrowly distributed in Antioquia Department, Colombia, an area that coincides with the Biogeographical Province of Magdalena in the Pacific Dominion (sensu Morrone 2014). The species grows in lowland tropical rainforests in a region characterised by a karstic topography, occurring at elevations of 30–500 m. (Fig. 9)

Figure 9. 

Distribution of Rinorea cogolloi (squares), R. galeanoae-bernalii (bolts), R. hirsuta (triangles).

Rinorea cogolloi is named in honour of Professor Alvaro Cogollo who has made contributions to the knowledge of Magnoliaceae taxonomy and the flora of Colombia, especially the middle Magdalena River Valley, where the species is endemic.

The species flowers between August and December; fruiting specimens were collected in February and August through October.

Rinorea cogolloi has a geographic range in the form of an EOO of 1,641 km² and AOO of 32 km². It is currently known from eight locations outside of protected areas. The region, underlain by karstic rocks, is heavily impacted by limestone extraction for concrete production and uncontrolled deforestation for grazing, agriculture and logging. Another location in north-eastern Antioquia Department is threatened with gold mining and deforestation. Given the high demand for cement products, the generally poor management of forest resources in the area and the number of locations, we project a continuing decline in the extent and quality of habitat for the species. Rinorea cogolloi is preliminarily assigned to the “Vulnerable” category [VU B1+B2ab(iii)] in accordance with the IUCN Red List Categories and Criteria (IUCN 2012).

Hekking (1988) noted the aberrant lamina pubescence of several specimens of Rinorea lindeniana (e.g. A. Cogollo 786, 1075, 1483). Rinorea cogolloi and R. lindeniana share several characteristics, such as leaf venation architecture and asymmetrical lamina bases, but R. cogolloi is readily separated from R. lindeniana by the densely pubescent indument on the abaxial lamina surface (vs. the abaxial lamina surface pilose to glabrescent in R. lindeniana). Rinorea cogolloi occurs sympatrically with R. galeanoae-bernalii (see below), but R. cogolloi has densely pubescent indument on the abaxial lamina surface (vs. abaxial lamina surface glabrescent in R. galeanoae-bernalii), pubescent fruit with spreading, erect ferruginous trichomes 0.3–0.4 mm long (vs. densely tomentose fruit) and the seeds pubescent with erect ferruginous trichomes 0.3 mm long (vs. seeds glabrous). Rinorea cogolloi and R. hirsuta have been confused by the similarity of the leaves and indumenta, but they can be differentiated by the asymmetrical lamina base, thyrsoid inflorescence, dorsal gland pubescent, pubescent seeds and capsule asymmetrical and pilose. In addition, these two species have different distributions, with R. cogolloi occurring in the Magdalena Province and R. hirsuta restricted to the Chocó-Darien Province (sensu Morrone (2014)). Rinorea cogolloi is also similar to R. pubiflora var. pubiflora, but it can be differentiated by its valves containing one seed (vs. valves containing three seeds in R. pubiflora) and the thyrsoid inflorescence (vs. racemose inflorescence). Table 2 highlights the key morphological differences amongst these five species.

Colombia. Dept. Antioquia. Mpio. de Caucasia: Vda. la Arenosa, km 45 vía al Bagre, Finca la Natalia, 7°47'46"N, 74°53'9"W, 70–100 m elev., 26 Aug 2022 (st), S. E. Hoyos-Gómez et al. 5184 (HUA, UBDC); ibid., 7°47'59"N, 74°52'45"W, 80 m elev., 10 Nov 2022 (fr), M. Montoya M9004 (HUA); Mpio. de Nechí: Vda. El Catorce, entre la mina el 14 y la roca del camino, 8°94'60"N, 74°46'14"W, 30–40 m elev., 6 Mar 2010 (fl), W. Rodríguez 6599 (COL, MEDEL); Mpio. de Puerto Triunfo: alrededores de la Gruta “El Condor,” 5°56'N, 74°50'W, 12 Oct 1983 (fl, fr), A. Cogollo 786 (COL, HUA, MO); ibid., 22 Oct 1989 (fr), J. G. Ramírez 4263 (JAUM); Río Claro carretera al Cairo, 17 Sep 1982 (fl, fr), E. Rentería 2722 (JAUM, MO). Mpio. de Sonsón: vereda Jerusalén, 5°55'00"N, 74°51'00"W, 200 m elev., 24 Dec 2010 (st), S. E. Hoyos-Gómez et al. 1145 (HUA, JAUM, MO); ibid., 5°55'00"N, 74°51'00"W, 200 m elev., 24 Dec 2010 (fl), S. E. Hoyos-Gómez et al. 1152 (HUA, JAUM); ibid., 5°55'00"N, 74°51'00"W, 200 m elev., 24 Dec 2019 (fl), S. E. Hoyos-Gómez & G. A. Wahlert 3780 (HUA, JAUM); ibid., 5°53'30"N, 74°50'58"W, 200 m elev., 23 Sep 2018 (fl, fr), J. P. Tobón et al. 2798 (HUA, JAUM); vereda Jerusalén, 5°53'30"N, 74°50'58"W, 380 m elev., 23 Sep 2018 (fr), J. P. Tobón et al. 2799 (HUA, JAUM); ibid., 5°53'39"N, 74°51'08"W, 380 m elev., 24 Sep 2018 (fl, fr), J. P. Tobón et al. 2803 (HUA, JAUM); ibid., 5°53'25"N, 74°50'50"W, 24 Sep 2018 (fl, fr), J. P. Tobón et al. 2806 (HUA, JAUM); ibid., 5°53'44.1"N, 74°51'18"W, 1 Feb 2019 (fr), J. P. Tobón et al. 2897 (JAUM); vereda Jerusalén, Reserva Natural Cañón del Río Claro, “El Refugio”, margen derecha, 5°53.759'N, 74°51.219'W, Apr 2013 (fr), L. Cano et al. 43 (HUA); corregimiento Jerusalén, vía Medellín-Bogotá, predios de Sumicol, cuenca del Río Claro, en cercanías de la torre de energía T 66, 5.915756°N, 74.849186°W, 4 Aug 2020 (fl, fr). J. M. Velez et al. 7388 (MEDEL, COL); ibid., 5.916104°N, 74.849473°W, 17 Sep 2020 (fl), J. M. Velez et al. 7511 (JAUM); ibid., 5.916182°N, 74.850481°W, 16 Feb 2021 (fr). J. M. Velez et al. 7604 (JAUM).

Colombia. Dept. Caldas: La Dorada, vereda la Atarraya, finca los Achiles, relictos de bosque al margen trasera de la finca, 5°40'20.3"N, 74°44'26.4"W, 214 m elev., 3 Aug 2021 (fl), D. Sanín et al. 7966 (holotype: HUA [acc. 225716]!; isotype: FAUC!).

Rinorea galeanoae-bernalii is similar to R. hirsuta by the elliptic lamina with symmetrical base and lacking domatia, inflorescence racemose, capsule symmetrical and pubescent with velutinous trichomes and one glabrous seed per valve, but it differs by the abaxial surface of the lamina glabrescent (vs. abaxial surface of the lamina pubescent with ferruginous trichomes in R. hirsuta), filaments fused at the base forming a staminal tube (vs. filaments free not forming a staminal tube), anthers with pubescence between the thecae (vs. anthers glabrous between the thecae) and style proximally pubescent (vs. style proximally glabrous).

Treelets 1.5–4 m tall, terminal branchlets pubescent with erect golden trichomes 0.2–0.4 mm long. Leaves opposite, petiolate; petiole 4–13 mm, pubescent with erect golden trichomes 0.2 mm long, glabrescent; stipules deciduous, free, elliptic, 2.5–4 × 1–1.5 mm, pubescent with appressed golden trichomes, 0.3–0.5 mm long; lamina elliptic, 7–19 × 2.5–8 cm, adaxially pubescent on the mid-vein with appressed golden trichomes 0.1 mm long, glabrescent, abaxially pubescent with appressed golden trichomes 0.3–0.5 mm long, glabrescent, semi-craspedodromous, with 6–8 major secondary vein pairs, secondary vein spacing decreasing proximally between them, base symmetrical, cuneate, margin subcrenate, apex acuminate, acumen 0.5–1 cm, mucronate, leaf domatia absent. Inflorescence axillary or terminal, racemose, 3–10 cm, 1.5–3 cm diam., axis pubescent with erect golden trichomes 0.2 mm long; pedicels 3–6 mm, pubescent with erect golden trichomes 0.1–0.2 mm long, articulated at the base, peduncle bracts deciduous, triangular, 1.5–1.8 × 1.5–1.8 mm, herbaceous, pubescent with appressed golden trichomes 0.3–0.5 mm long, margin ciliolate, bractlets persistent, 1 × 1 mm, herbaceous, pubescent with appressed golden trichomes 0.1 mm long, margin ciliolate. Flowers 3–4 × 3–4 mm, sepals subequal in size and shape, triangular, 1.6–2 × 1–1.5 mm, 1–5-veined, pubescent on the distal part of the costa with appressed golden trichomes 0.1–0.2 mm long, margin ciliolate, apex apiculate; petals elliptic, 3–4 × 1.4–2 mm, costa pubescent with spreading golden trichomes 0.1 mm long, margin ciliolate, apex acute, white; stamens 2.5–3 mm long, all filaments united at the base forming a staminal tube, tube glabrous, filaments 0.4–0.5 mm, anthers elliptic, 1.1–1.4 × 0.4–0.6 mm, pubescent between the thecae adaxially with spreading golden trichomes 0.1–0.2 mm long, apex rounded, connective 0.7–0.8 mm, dorsal connective scale conspicuous, lanceolate, 2.3–2.5 × 0.8–1 mm, margin entire, orange-brown; ovary globose, 0.8–1 × 0.9–1 mm, pubescent with appressed golden trichomes 0.1–0.2 mm long; style filiform, curved, 1.5–2 × 0.1 mm, pubescent proximally with appressed trichomes 0.1 mm long, stigma acute. Fruit a symmetrical, subligneous capsule dehiscent along three sutures, ellipsoid, 1–2 × 0.5–0.8 cm, apex acuminate, tomentose with golden trichomes 0.1 mm long. Seeds one per valve, globose, 5–7 mm diam., glabrous, with maculae, brown when dry. (Fig. 10)

Figure 10. 

Rinorea galeanoae-bernalii Hoyos-Gómez A habit B detail of leaf, abaxial surface C leaf architecture D infructescence E flower F detail of filaments fused at the base forming a staminal tube G sepals, abaxial surface H petals, abaxial surface I stamens, adaxial and abaxial surfaces J detail of anthers showing pubescence between the thecae K gynoecium L fruit M detail of fruit showing one seed per valve N seed. (A–E, N: D. Sanin 7704 [HUA]; D. Sanin 7966 [HUA]).

Rinorea galeanoae-bernalii is apparently endemic to Colombia, where it occurs in the Departments of Antioquia, Caldas and Santander. Its distribution corresponds to the Biogeographical Province of Magdalena in the Pacific Dominion (sensu Morrone (2014)). It grows in lowland tropical rainforest at elevations of 192–500 m. (Fig. 9)

Rinorea galeanoae-bernalii is named in honour of the late Dr. Gloria Galeano Garcés (1958–2016) and Dr. Rodrigo Bernal, a husband-wife botanical team who contributed greatly to the knowledge of the Colombian flora and the taxonomy of Neotropical palms, Cyclanthaceae and Marantaceae. As professors and mentors, they inspired many of their students to pursue a botanical career studying the flora of Colombia.

The species flowers between July and October; fruiting specimens were collected in January, April, July and September through November.

Rinorea galeanoae-bernalii has a geographic range in the form of an EOO of 9,409 km² and AOO of 48 km². It is known from 11 locations that are not included in any protected area. The locations in the middle Magdalena River Valley are heavily impacted by deforestation for grazing and agriculture, especially for cultivation of Elaeis guineensis (oil palm), while some known locations in Antioquia Department are threatened with limestone extraction for concrete production. Given ongoing habitat disturbance and projected continuing decline in the quality and extent of habitat, it is likely that the species will qualify in the near future for a threatened status and, thus, it is preliminarily assessed with a status of “Near Threatened” (NT).

Hekking noted the unusual glabrous lamina in at least one specimen which he had identified as Rinorea hirsuta (e.g. E. Rentería et al. 1779). Based on the discrete biogeographic distributions of R. galeanoae-bernalii and R. hirsuta (Magdalena Valley vs. Chocó-Darien, respectively) and the morphological characters that differentiate the two (e.g. the presence of a staminal tube and the patterns of pubescence on the anther connectives and style), we hypothesise that R. galeanoae-bernalii is a separately evolving lineage worthy of recognition at the rank of species.

Rinorea galeanoae-bernalii is sympatric with R. cogolloi, but it can be differentiated by the racemose inflorescence and glabrous seeds (vs. thyrsoid inflorescence and pilose seeds in R. cogolloi). Rinorea galeanoae-bernalii is also sympatric with R. pubiflora var. pubiflora, but it can be differentiated by the valves that each contain one glabrous seed (vs. valves that each contain three pubescent seeds in R. pubiflora var. pubiflora); see Table 2 for a comparison of key diagnostic characters differentiating R. galeanoae-bernalii, R. cogolloi and R. pubiflora var. pubiflora.

Colombia. Dept. Antioquia. Mpio. Puerto Triunfo: corregimiento Puerto Perales, hacienda Cerritos, 7.65833, -74.81861, 192 m elev., Nov 2009 (fr), L. Londoño et al. 938 (HUA). Mpio de Sonsón: Corregimiento de Jerusalén, Vía Medellín-Bogotá, predios de Sumicol, cuenca del Río Claro, 5.915969. -74.850459, 370 m elev., 16 Mar 2021 (fl), J. M. Vélez et al. 7621 (MEDEL, HUA). Mpio de Yondó: ciénaga de Barbacoas, Hacienda Java, bosque Catanga, 6°43'N, 74°19'W, 100–130 m elev., 18 Apr 2011 (fr), J. Betancur 15261 et al. (COL, HUA, NYBG, US). Dept. Caldas. Mpio. La Dorada: vereda la Atarraya, finca los Achiles, relictos de bosque de la finca, 5°41'03.9"N, 74°44'16.4"W, 30 Jan 2021 (fr), D. Sanín et al. 7704 (HUA, FAUC). Mpio. La Dorada: Vda. La Habana, 250 m elev., 16 Jan 2000 (fr), M. V. Bustos Giraldo 62 (COL); predio el palmar, 5°18.551'N, 74°47.839'W, 305 m elev., 4 Nov 2012 (fl, fr), J. M. Vélez 4303 (MEDEL). Dept. Santander. Mpio. Cimitarra: Potrero Quito, 6°28.285'N, 74°21.116'W, 3 Mar 1999 (st), Á. Idárraga et al. 902 (HUA); corregimiento Puerto Olaya, Hacienda el Bosque, Puerto Arturo, 6°27.576'N, 74°21.034'W, 4 Aug 1999 (fl), Á. Idárraga et al. 1491 (JAUM); corregimiento Puerto Olaya, Hacienda Piamonte, 6°26'17.8"N, 74°22'07.6"W, 29 May 2015 (fr), Á. Idárraga et al. 6034 (HUA); ibid., 6°26'25.3"N, 74°22'22"W, 123 m elev., 22 Jul 2015 (fl), Á. Idárraga et al. 6116 (HUA); ibid., 23 Jul 2015 (fl, fr) Á. Idárraga et al. 6138 (HUA); Puerto Araujo, 500 m elev., 18 Sep 1979 (fr), E. Rentería et al. 1779 (COL, HUA, JAUM, US); ibid., 700 m elev., 20 Sep 1979 (fr), E. Rentería et al. 1809 (COL); corregimiento Puerto Olaya, Hacienda el Bosque, fragmento Monte Cristo, 6°28'N, 74°21'W, 19 Sep 2001 (fl), A. Rivas et al. 152 (HUA, JAUM); vereda los Ranchos, Hacienda Monterrey, 6°15'42"N, 74°27'51"W, 123 m elev., 1 Oct 1998 (fl, fr), W. Rodríguez et al. 1070 (JAUM); ibid. (fl, fr), W. Rodríguez et al. 1505 (JAUM). Mpio. Girón: vereda Sogamoso, hacienda Trigueros, 7°05'27.1"N, 73°21'41.4"W, 332 m elev., 25 Feb 2011 (fl, fr), E. Y. Rodríguez-Ch. 1989 (COL).

Colombia. Dept. Caquetá: Mpio. de Solano, región de Araracuara, sector Chiribiquete, camino a Tepuí, 0°12'16"S, 72°29'14"W, 170 m elev., 10 Dec 2010, F. Castro 10919 (holotype: COAH [acc. 78339]!; isotype: NY [barcode 02691382]!).

Rinorea betancurii is similar to R. macrocarpa by the elliptic lamina lacking domatia, lamina base symmetrical and capsule symmetrical with apex acuminate, but it differs by the abaxial lamina surface pubescent (vs. abaxial lamina surface glabrous in R. macrocarpa), lamina base obtuse (vs. lamina base cuneate), fruit smaller 2.5–3 cm long (vs. fruit larger, 3–5.7 cm) and valves containing two pubescent seeds (vs. valves containing three glabrous seeds).

Treelets 1.5–5 m tall, branchlets with puberulent erect ferruginous trichomes 0.1–0.2 mm long, glabrescent. Leaves opposite, petiolate; petiole 2–4 mm long, pubescent with erect ferruginous trichomes 0.2 mm long; stipules deciduous, free, lanceolate, 2.3–3 × 1–1.1 mm, herbaceous, pubescent with appressed ferruginous trichomes 0.2 mm long, lamina elliptic, 6.3–14.4 × 2.6–7 cm, adaxially glabrous, abaxially pubescent with erect ferruginous trichomes 0.2–0.4 mm long, semi-craspedodromous, with 5–7 major secondary vein pairs, secondary veins with spacing between them decreasing proximally, symmetrical, base obtuse, margin entire to subcrenate, apex acute, acumen 8–14 mm long, mucronate, domatia absent. Flowers unknown. Fruit a symmetrical, subligneous capsule dehiscent along three sutures, ellipsoid, 2.5–3 × 0.5–0.6 cm, apex acuminate, pubescent with curved golden trichomes 0.3–0.4 mm long, green in vivo, brown when dry. Seeds two per valve, globose, 7–8 mm diam., pubescent with spreading trichomes 0.1–0.2 mm long, with maculae, seeds brown when dry. (Fig. 11)

Figure 11. 

Rinorea betancurii Hoyos-Gómez A habit B detail of leaf, abaxial surface C leaf architecture D fruit and detail of pubescence E detail of fruit showing two seeds per valve F seed. (A–F: R. L. Liesner 15689 [MO]).

Rinorea betancurii occurs in Brazil, Colombia and Venezuela, an area that corresponds to the Biogeographic Provinces of Imerí and Pantepui in the Boreal Brazilian Dominion (sensu Morrone (2014)). The species grows in lowland tropical rainforest at elevations of 60–170 m. Rinorea betancurii has been recorded as growing in sandy soils near rivers and on slopes and hills. (Fig. 4)

Rinorea betancurii honours Julio Betancur, Colombian botanist and curator at the National Herbarium in Bogotá. Dr. Betancur has inspired a new generation of students to pursue a career in botany and has made many contributions to the taxonomic knowledge of the Bromeliaceae, Heliconiaceae and the flora of Colombia.

Fruiting specimens were collected in February through May and in September, November and December.

Rinorea betancurii has a geographic range in the form of an estimated EOO of 511,480 km² and an AOO of 36 km². The species is known from eight localities, none of which appear to be in protected areas. Based on satellite imagery from Google Earth, the locations in Colombia and Brazil appear to be heavily impacted by deforestation, while the four Venezuelan locations are in remote and vast primary forests with large buffers from disturbed areas. Even though we project continuing decline in the extent and quality of habitat for the Colombian and Brazilian locations, the undisturbed forests in southern Venezuela appear to be safe from large-scale disturbance for the foreseeable future. Nevertheless, it appears unlikely that the species will qualify for a threatened status in the near future and is assigned a preliminary status of Least Concern (LC).

Rinorea betancurii occurs with R. pubiflora var. pubiflora, but it can be separated by the valves that contain two seeds (vs. valves that contain three seeds in R. pubiflora var. pubiflora).

Colombia. Dept. Caquetá: cabeceras del río Masay, desembocadura del río San Jorge al río Cuñaré, 1–6 Mar 1980 (fr), M. Pabón 946 (COL). Dept. Meta: Mpio. Puerto López, vereda el Tigre, finca Pista, orilla del canal, 8 Feb 2008 (fr), F. Castro 4760 (UBDC). Dept. Vaupés: Mpio Taraira, comunidad Jotabeyá, 0°35'S, 70°11'W, 150–250 m elev., 27 Mar 2009 (fr), J. Betancur et al. 13854 (COAH, COL, HUA). Venezuela. Bolivar State: Mpio. Foráneo Aripao, margen derecha del caño Minchaquene (Hormiga), tributario del Alto Caura, entre Araguaña y Campamento, 4°45'N, 64°12'W, 2 May 1988–5 May 1988 (fr), G. Aymard 6810 (MO, NY, U). Amazonas State: Mpio. Atabapo, río Cunucunuma, raudal Mapaco, 3°37'N, 65°51'W, Apr 1990 (fr), W. R. Anderson 13343 (F, MICH); close to cerro Neblina base camp on río Mawarinuma, 0°50'N, 66°10'W, 6 Feb 1984 (fr), R. L. Liesner 15689 (BHO, MO, U, VEN); Cerro Yapacana, 3°45'N, 66°45'W, 125–400 m elev., 3 May 1970 (fr), J. Steyermark et al. 113015 (COL). Brazil. Amazonas State: Mpio. Barcelos, base cerro Aracá, 0°42'N, 63°22'W, 14 Jul 1985 (fr), I. Cordeiro 193 (INPA, NY).