Research Article |
Corresponding author: Andrés Orejuela ( aaorejuelar@gmail.com ) Academic editor: Leandro Giacomin
© 2022 Andrés Orejuela, Boris Villanueva, Clara Inés Orozco, Sandra Knapp, Tiina Särkinen.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Orejuela A, Villanueva B, Orozco CI, Knapp S, Särkinen T (2022) Monograph of Doselia (Solanaceae), a new hemiepiphytic genus endemic to the northern Andes. PhytoKeys 202: 73-96. https://doi.org/10.3897/phytokeys.202.82101
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A new genus, Doselia A.Orejuela & Särkinen, gen. nov., is described in the tribe Solandreae (Solanaceae) consisting of four species of hemiepiphytic lianas endemic to the premontane forests of the Colombian and Ecuadorian Andes. The genus is distinguished based on the membranous leaves, usually sparsely pubescent with eglandular simple trichomes, pseudo-verticillate leaf arrangement, and elongated, pendulous, and few-flowered inflorescences with showy flowers and conical fruits. Three new combinations are made to transfer species to the new genus previously described as part of the polyphyletic genus Markea Rich. (Doselia epifita (S.Knapp) A.Orejuela & Särkinen, comb. nov., D. huilensis (A.Orejuela & J.M.Vélez) A.Orejuela & Särkinen, comb. nov. and D. lopezii (Hunz.) A.Orejuela & Särkinen, comb. nov.). One new species is described from the western slopes of the eastern cordillera of the Colombian Andes, known only from three localities in the Boyacá, Santander, and Tolima departments (Doselia galilensis A.Orejuela & Villanueva, sp. nov.). The new species is unique in the genus in having glabrescent adult leaves, green-purplish calyces and long, greenish-white, infundibuliform corollas with delicate purplish veins and large lobes tinged with purple, and pubescent styles. Here we provide a revision of Doselia with a distribution map of all species, an identification key, photographs, preliminary conservation assessments, and line drawings of all four species.
Un nuevo género, Doselia A.Orejuela & Särkinen, gen. nov., es descrito en la tribu Solandreae (Solanaceae), conformado por cuatro especies de lianas hemiepífitas endémicas de los bosques premontanos de los Andes de Colombia y Ecuador. El género se distingue por sus hojas membranáceas con tricomas simples no glandulares, el arreglo pseudoverticilado de las hojas y sus inflorescencias paucifloras, largo pedunculadas y péndulas, flores vistosas y frutos cónicos. Se proponen tres nuevas combinaciones para acomodar especies previamente descritas en el polifilético género Markea Rich. (Doselia epifita (S.Knapp) A.Orejuela & Särkinen, comb. nov., D. huilensis (A.Orejuela & J.M.Vélez) A.Orejuela & Särkinen, comb. nov. and D. lopezii (Hunz.) A.Orejuela & Särkinen, comb. nov.) Una nueva especie es descrita aquí proveniente de la vertiente occidental de la cordillera Oriental de los Andes colombianos, y conocida únicamente de tres localidades en los departamentos de Boyacá, Santander y Tolima (Doselia galilensis A.Orejuela & Villanueva, sp. nov.). La nueva especie se caracteriza por sus hojas glabrescentes cuando adultas, cáliz verde-púrpura, corolas infundibuliformes grandes, blanco-verdosas con una reticulación fina de venas púrpuras, lóbulos corolinos grandes y teñidos de púrpura en su lado ventral y estilos pubescentes. El nuevo género es formalmente descrito incluyendo un mapa de distribución, una clave taxonómica y fotografías e ilustraciones para sus cuatro especies.
Colombia, Doselia, Ecuador, endemism, hemiepiphytes, Juanulloeae, lianas, Markea, new species, Solandreae
Colombia, Doselia, Ecuador, endemismo, hemiepífitas, Juanulloeae, lianas, Markea, nuevas especies, Solandreae
The tribe Solandreae Miers (Solanaceae) contains ca. 80 species of mainly epiphytic or hemi-epiphytic lianas and shrubs in a number of genera currently being recircumscribed (
The tribe Solandreae is a unique clade within Solanaceae in that many of its component taxa are epiphytic and hemiepiphytic plants with a great diversity of floral forms, pollinators, and ant associations. Epiphytes are rare in Solanaceae, with only ca. 90 species with this growth form across the family in three distinct tribes (Solandreae 80 spp.; Capsiceae 4–5 spp.; and Solaneae 4–5 spp.), with Solandreae containing most of the epiphytic species (ca. 90%;
Within Solandreae, there is notable morphological variation in corolla shape, size, and colour. Corollas vary from large infundibuliform or campanulate, long tubular, hypocrateriform to minutely campanulate and include pale or dull-coloured to brightly coloured forms. This remarkable variation suggests a diverse coevolutionary history with pollinators; bats, hummingbirds, and bees have all been observed to visit these flowers (
Variation in floral form has been used as the basis of previous taxonomic classifications of the tribe. Molecular phylogenetic studies have shown, however, that many of the previously recognised genera in Solandreae are para- or polyphyletic and in dire need of taxonomic revision (
Here we focus on the morphologically distinct Markea lopezii clade (Figs
Comparison of the new genus Doselia with the morphologically most closely related genera and groups in the tribe Solandreae (Solanaceae).
Doselia | Solandra | Schultesianthus | Markea antioquiensis clade | |
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Habit | Hemiepiphytic lianas | Hemiepiphytic lianas | Hemiepiphytic lianas | Terrestrial/epiphytic shrubs |
Leaf texture | Membranous | Chartaceous | Coriaceous | Membranous |
Leaf arrangement | Clustered on adult branches | Alternate | Alternate | Clustered on adult branches |
Trichome type | Simple, not glandular | Simple or branched, glandular or not glandular | Simple, glandular or not glandular | Simple, not glandular |
Inflorescence branching | unbranched to forked | None; flowers solitary | Unbranched, forked or multi-branched | Unbranched |
Open flowers per inflorescence | 1–2 | Solitary | (1)6–10 (>100) | 4–6 |
Peduncles | Long (1–50 cm), hanging & slender | Absent | Short (0–3 cm), stout & woody | Intermediate (4–12 cm), hanging & slender |
Pedicels | Unwinged or distally winged | Unwinged | Unwinged | Unwinged or distally winged |
Floral bract number and size | 0–2 | Absent | Absent | 2 |
5–6 cm long × 1–2 cm wide | 0.5–2 cm long × 0.1–0.3 cm wide | |||
Corolla symmetry | Actinomorphic | Weakly zygomorphic | Weakly zygomorphic | Actinomorphic |
Corolla lobes | Entire | Fimbriate | Fimbriate | Entire |
Corolla shape | Infundibuliform or hypocrateriform to tubular-campanulate | Infundibuliform to cyathiform | Infundibuliform to cyathiform | Infundibuliform |
Corolla length | 8.5–15 cm | 15–40 cm | 2.5–21.5 cm | 7–12 cm |
Fruit shape | Conical | Conical | Globose, rarely conical | Conical or globose |
Fruit locule number | 2 | 4 | 2 | 2 |
Doselia A.Orejuela & Särkinen A pseudoverticillate leaves in D. epifita B infundibuliform cream-purplish corolla of D. epifita C infundibuliform greenish-white corolla with subtle purple veins of D. galilensis D tubular-campanulate yellowish-green corolla of D. huilensis with strong purple-tinged reticulation along major and minor veins E developing fruit in D. huilensis F hypocrateriform orange corollas in D. lopezii G pendent long flowering branches in D. lopezii with clustered leaves and orange corollas in frontal view H developing fruit in D. lopezii with a nectariferous disc at the base (Vouchers: A Orozco et al. 3876 (COL), B no voucher, C Corrales et al. 917 (JBB, TOLI), D Coral 34 (HEEA), E Orejuela & Vélez-Puerta 112 (COL) F Orejuela et al. 727 (JBB) G no voucher H Orejuela & Calderon 170 (COL): photos by Alistair Hay, Andreas Kay, Andrés Orejuela, Boris Villanueva, Brayan Coral, and Eduardo Calderon.
All specimens of the tribe Solandreae from 25 Colombian and Ecuadorian herbaria were revised for the study to understand morphological variation across the group (acronyms follow Index Herbariorum http://sweetgum.nybg.org/science/ih/): ANDES, CAUP, COAH, COL, CUVC, FAUC, FMB, HECASA, HUA, HUAZ, HUQ, JAUM, JBB, JBGP, LLANOS, MEDEL, PSO, QCA, QCNE, TOLI, UDBC, UIS, UPTC, UTMC, and VALLE; as well as major herbaria that hold representatives from countries across the Andes (BM, E, F, K, MO, US). Herbarium material of the new species was collected in 2018 and deposited at
Universidad del Tolima herbarium (
Specimens with coordinates were mapped directly, and those lacking coordinates were located using Google Earth, GeoNames gazetteer (http://www.geonames.org), and GEOLocate Web service (https://www.geo-locate.org/default.html). Distribution maps were created using QGIS (
Like Solandra Sw. and Schultesianthus Hunz., but differing from both in having membranous leaves (vs. chartaceous or coriaceous), lacking glandular trichomes, having pendulous inflorescences with long slender peduncles (vs. solitary flowers or short erect inflorescences with thick woody peduncles), distally winged pedicels, and actinomorphic corollas (vs. weakly zygomorphic); differing from Solandra in its smaller corollas (8.5–15 cm vs. 15–40 cm long), flowers borne in groups (vs. solitary), straight filaments and styles (vs. strongly curved and declinate), and 2-locular ovaries (vs. 4-locular); differing from Schultesianthus Hunz. in the chartaceous fruiting calyx with visible venation (vs. a thick, shiny and coriaceous fruiting calyx with no visible venation).
Doselia lopezii (Hunz.) A.Orejuela & Särkinen comb. nov. (basionym Markea lopezii Hunz.)
Hemiepiphytic lianas adhering to the trees by adventitious roots. Stems terete when fresh, irregularly angulate when dry, pubescent with simple, uniseriate multicellular eglandular trichomes, older stems with pale brown and papery bark, often peeling, with broad circular foliar scars left by fallen leaves. Leaves alternate on young stems or tightly clustered appearing sub-opposite in adult branches, simple, broadly elliptic to obovate, membranous, concolorous, sparsely to densely pubescent both abaxially and adaxially with simple, uniseriate eglandular trichomes like those on stem; major veins 3–6 pairs; base attenuate to obtuse, sometimes asymmetric; margins entire to undulate; apex acute, acuminate, or mucronate; petioles well-developed, conspicuously articulate, green. Inflorescences axillary to sub-axillary monochasial cymes, simple to one-branched, occasionally bracteate, usually long-pedunculate and pendulous, 1–7-flowered, pubescent with trichomes as on the stems; peduncles (1.2–) 8.5–50 cm long; bracts absent or if present foliaceous and linear, 5–6 cm long, 1–2 cm wide; pedicels 0.5–3.0 cm long, conical, 5-ribbed, distally thickened and winged in some species, articulated at the base. Flowers 5-merous, actinomorphic, perfect, aestivation valvate in the calyx and cochlear in corolla. Calyx cupuliform, green to whitish-green with purple colouration sometimes on the veins or along the margins only; lobes flat to undulate, 2.4–5.2 cm long, 1–1.5 cm wide, long-triangular to lanceolate, apically long-acuminate to acute, pubescent with simple, uniseriate eglandular, transparent to brown trichomes. Corollas 8.5–15 cm long, the inner corolla diameter from 2.5–5.0 cm, infundibuliform, hypocrateriform to tubular-campanulate, orange, purple, white, yellowish-green with purple veins, sparsely pubescent abaxially with trichomes like those on the calyx; lobes 1.6–4.2 cm long, 1.6–4.3 cm wide, triangular to oblong, spreading to reflexed during anthesis, glabrous to sparsely pubescent, the margins entire to undulate to revolute, the apices acute to obtuse. Stamens 5, equal, included within corolla tube or fully exserted beyond the mouth; filaments 1.7–6.1 cm long, adnate to the base of the corolla, white to purple-tinged, glabrous to pubescent at the point of insertion; anthers 1.4–2.7 cm long, 1.3–1.8 mm wide, elongate, basifixed, not connivent. Ovary conical, 2.9–7.5 mm long, 2.9–7 mm in diameter, light yellow to brown, 2-carpellate, 2-locular, glabrous, with a well-developed 5-lobed light green to pale yellow nectariferous disc; style 5.9–8.8 cm long, straight, glabrous to sparsely pubescent with simple uniseriate trichomes 0.3–0.5 mm long; stigma 2-lobed, ca. 1 mm long and wide, usually clavate. Fruit a conical berry, 1.5–4.4 cm long, 1.5–4.4 cm in diameter, pale to dark green, chartaceous to coriaceous when dry, 2-locular, the exocarp 2–2.8 mm thick; fruiting calyx persistent, the lobes 3–5 cm long, 1.2–2.3 cm wide, appressed and enveloping the berry loosely, fully covering the fruit. Seeds numerous, 2.2–3.6 mm long, 1–1.7 mm wide, subreniform, the testa reticulate, the testa cells rectangular and straight in the outline, the embryo slightly curved, the cotyledons accumbent, slightly longer than the embryo rest, endosperm rather scanty. Chromosome number not known.
The generic name Doselia is derived from the Spanish word “dosel”, meaning canopy. It refers to the hemiepiphytic lianescent habit of all species of Doselia, with branches rising high up to the canopy to the top of tree crowns. The plants can be challenging to see because of their position on top of the tree canopy unless the plants have their showy pendulous flowers.
(Fig.
Doselia represents a morphologically distinct group of four hemiepiphytic lianas from mid-elevation moist Andean forests with very long branches extending to the forest canopy through adventitious roots. The combination of hemiepiphytic lianescent habit, membranous leaves arranged in tight clusters on adult branches, indumentum consisting of only simple eglandular trichomes, showy actinomorphic flowers arranged in elongated, pendulous, and few-flowered inflorescences, and conical fruits is unique within the tribe.
Within Solandreae, the lianescent hemiepiphytic habit is also known in Solandra and Schultesianthus, with the rest of the tribe mainly being epiphytic or rarely terrestrial shrubs (Markea antioquiensis clade; Table
Inflorescences in Doselia are long and pendulous (up to 50 cm long), with up to three flowers of which only one or rarely two develops at a time (Table
Corollas in Doselia are actinomorphic and showy, similar to species of the Markea antioquiensis clade, but these two groups can be distinguished based on other characters such as growth form, peduncle length, number of open flowers per inflorescence, and floral bract and calyx size (Table
Fruits in Doselia are conical, leathery, and fully covered by the calyx, like those of Solandra, but differ from the latter in being 2-carpellate and 2-locular, in contrast to the 2-carpellate and 4-locular fruits in Solandra (Table
1 | Sparse pubescence throughout the plant, on the leaves confined to the veins and margins only, leaves glabrescent with age; style pubescent along its entire length | D. galilensis |
– | Dense pubescence throughout the plant, on the leaves extending to the blade mesophyll, leaves persistently pubescent with age; style glabrous or with only a few trichomes at the very base | 2 |
2 | Anthers included within the corolla tube; corolla pale purple or purplish-cream; style with a few trichomes at the base | D. epifita |
– | Anthers partially to entirely exserted beyond the corolla tube; corolla yellowish-green, brownish-green or orange; style completely glabrous | 3 |
3 | Corolla tubular-campanulate, the tube yellowish-green with a conspicuous reticulum of purple veins on both surfaces; anthers completely exserted | D. huilensis |
– | Corolla infundibuliform, the tube orange with a conspicuous reticulum of purple veins at the base on the adaxial side only; anthers partially included | D. lopezii |
Markea epifita S.Knapp, Novon 8(2): 155–157, f. 3a, b. 1998. Type. Ecuador. Napo: Canton Archidona, carretera Hollín-Loreto km 25, sector Challua Yacu, faldas al S de Volcán Sumaco, 1°45'S, 77°38'W, 1,200 m, 21 Apr 1989 (fl), C. Cerón & F. Hurtado 6534 (holotype: QCNE; isotypes: MO! [MO-289398, acc. # 5343691], NY! [00214503]).
Based on Markea epifita S.Knapp
Hemiepiphytic liana with adventitious roots. Stems sparsely pubescent with simple, uniseriate 2–4-celled hyaline trichomes ca. 0.5 mm long, giving the stems a tuberculate look. Leaves tightly clustered towards the branch tips, 11–25 cm long, 6–12 cm wide, obovate, sparsely pubescent with simple uniseriate 2–4-celled trichomes ca. 0.5 mm long, the trichomes denser and stiffer abaxially along the veins on both surfaces; major veins 4–5 pairs, not raised abaxially, drying dark brown; base attenuate, symmetric; margins entire; apex acute to acuminate; petiole 1–5 cm long, pubescent with stiff trichomes abaxially like those of the leaves. Inflorescence 18.5–45 cm long, axillary, unbranched, ebracteate, ca. 4–5-flowered, densely pubescent with simple, uniseriate trichomes like those of the stems; peduncle 8.5–35.2 cm long; pedicels 1.2–1.7 cm long, winged. Calyx 3.8–4 cm long, 1.4–1.5 cm wide, green tinged with purple, pubescent with simple uniseriate trichomes 0.5–1 mm long; tube 0.5–0.7 cm long; lobes undulate, 3.0–3.3 cm long, 0.8–1 cm wide, long-triangular, apically acuminate, densely pubescent abaxially with simple uniseriate trichomes 0.5–1 mm long, adaxially similar but also with tiny brownish papillate trichomes. Corolla 9–11 cm long, the inner corolla diameter 3–3.5 cm, infundibuliform, gradually widening from the base, glabrous; tube 7.5–8 cm, cream to green with lines or patches of purple; lobes 3.4–4.2 cm long, 2.5–3.3 cm wide, ovate, purple or violet, reflexed at anthesis, the margins slightly undulate, the apex obtuse to rounded, glabrous or with a few minute trichomes along the veins. Stamens 3.6–7.7 cm long, included within corolla tube; filaments 2–5 cm long, adnate to ca. 1 cm from the base of the corolla tube, white to purple-tinged, glabrous; anthers 1.6–2.7 cm long, 1.7–1.8 mm wide, cream. Ovary 3.3–3.4 mm long, 3.3–3.4 mm wide, colour unknown, glabrous; style 5.5–6 cm long, cream, glabrous except for a few simple uniseriate trichomes ca. 0.5 mm long at the very base; stigma clavate. Fruit not known. Chromosome number not known.
(Fig.
In premontane forest between 500–1,500 m elevation.
(
Doselia epifita is the only species of Doselia that reaches Ecuador and has the lowest elevational range within the genus. Doselia epifita is morphologically most similar to D. galilensis, and a detailed comparison is presented under the latter. The inflorescence morphology of D. epifita was unknown until recently because no complete specimens with entire inflorescences were known when the species was first described (
The specific epithet refers to the apparent epiphytic habit of the species, though, like other species in the genus, D. epifita is a hemiepiphyte rather than an obligate epiphyte.
Colombia. Caquetá: Belén de Los Andaquíes, camino Andaquí, via que comunica Acevedo con Belén de los Andaquíes, vegetación a orilla de quebrada, 12 Mar 2016 (fl), Cárdenas 45979 (COAH). Ecuador. Morona-Santiago: Parque Nacional Sangay, lagunas de Sardinayacu, 2°05'54"S, 78°09'19"W, 1,400–1,500 m, 18 Jan 2015 (fl), Pérez et al. 7875 (QCA). Napo: Canton Archidona, Carretera Hollin–Loreto, km 25, Sector Challua Yacu, Faldas al sur del Volcán Sumaco, 1°27'00"S, 77°22'48"W, 1,200 m, 26 Aug 1980 (fl), Cerón 6534 (MO, NY); Cantón Loreto, Parque Nacional Sumaco, Napo-Galeras, Matorral de Bambú, Bloque 19, línea sísmica 22, Compañía Triton, 0°47'00"S, 77°28'00"W, 500 m, 25 Mar 1996 (fl), Freire & Cerda 271 (MO, QCNE); Cantón El Chaco, Proyecto Hidroeléctrico Coca, Punto ST3, margen derecha del Río Quijos, 10 km al S de Reventador, 0°11'00"S, 77°39'00"W, 1,500 m, 3 Oct 1990 (fl), Palacios 5893 (MO). Pastaza: Cantón Mera, Carretera al Rio Ansu, 5 km al NE de Mera, 1°15'36"S, 78°03'36"W, 1,200 m, 16 Mar 1985 (fl), Palacios et al. 123 (MO); Cantón Mera, Colonia Pindo, Mirador en la Reserva Pindo, 1°27'23"S, 78°04'47"W, 12 Nov 2011, Orozco et al. 3876 (COL); Road Veracruz (Indillama) – Canelos, 1°35'00"S, 77°51'00"W, 25 Jun 1968 (fl), Lugo 75 (MO); Cantón Pastaza, Shell, Río Pindo, 1°29'59"S, 78°03'44"W, 1,050 m, 18 Aug 1992 (fl), Palacios 10380 (MO).
Differing from all other members of Doselia in its mature leaves with sparse pubescence of trichomes on the midvein and along margins only (vs. on entire lamina). Like D. epifita (S.Knapp) A.Orejuela & Särkinen but differing in the pale green to purplish-green calyces with flat lobes (vs. calyces green with purple patches with undulate lobes) and larger corolla 12–15 cm long (vs. 9–11 cm long).
Colombia. Tolima: Municipio Villarrica, Vereda Galilea, Bosque de Galilea, zona Campo Hermoso, junto a parcela permanente de Monitoreo de 1 ha, 03°46'21"N, 74°39'56"W, 1,543 m, 11 Jun 2018 (fl, fr), L. Corrales, B. Villanueva, K. Sánchez & H. Díaz 917 (holotype:
Hemiepiphytic liana with adventitious roots. Stems sparsely pubescent with simple, uniseriate 4–7-celled, hyaline trichomes 0.4–1.3 mm long, becoming glabrescent with age. Leaves tightly clustered towards the branch tips, 9.2–17.5 cm long, 6.4–8.4 cm wide, ovate to elliptic, sparsely pubescent with a few simple trichomes like those on the stems distributed along the margins and veins on both surfaces, especially on the young growth, glabrescent with age; major veins 3–4 pairs, slightly raised abaxially; base cuneate or obtuse, symmetric or rarely asymmetric; margins entire; apex acuminate to mucronate; petiole 0.8–1.8 cm long, sparsely pubescent with a few simple trichomes like those on the stems, glabrescent with age. Inflorescence axillary, simple, ebracteate, 11.5–17.2(–44) cm long, 1(–3)-flowered, sparsely pubescent with a few simple trichomes like those on the stems; peduncle 1.2–5.7(–32.5) cm long; pedicels 0.5–1.8 cm long, distally winged and thickened. Calyx 3.7–3.8 cm long, 1.7–1.8 cm wide, pale green with purple margins and reticulation along the veins, sparsely pubescent with simple, uniseriate trichomes like those on the stems; tube 0.5–0.7 cm long; lobes flat, 2.4–3.0 cm long, 1.0–1.2 cm wide, short-lanceolate, apically acute. Corolla 12–15 cm long, the inner corolla diameter 3.5–4.0 cm, infundibuliform; tube 8.3–9.5 cm long, with a narrow base 1.4–1.9 cm long, 0.8–0.9 cm wide and a wide distal portion 7.6–7.7 cm long, 3.6–3.8 cm wide, greenish-white with subtle purple veins, glabrous or sparsely pubescent with a few simple uniseriate trichomes like those of the rest of the plant on the tube externally; lobes 3.2–3.8 cm wide, 2.8–3.1 cm long, ovate, greenish-white with bright purple patches within, reflexed at anthesis, the margins revolute, the apex obtuse, glabrous. Stamens 4.1–4.2 cm long, included inside the corolla tube; filaments 3.1–3.4 cm long, adnate at ca. 1.4–1.8 cm from the base of the corolla, white, densely pubescent with simple, uniseriate 4–7(–12)-celled, hyaline trichomes at the insertion point; anthers 1.6–2.1 cm long, 1.4–1.5 mm wide. Ovary 3.7(–5.4) mm long, 6.2–6.3 mm wide, light brown, glabrous; style 5.9–6.5 cm long, cream, sparsely pubescent with simple short 2–4-celled uniseriate trichomes ca. 0.3 mm long; stigma clavate. Fruit ca. 4.4 cm long, ca. 2.9 cm wide, light green, the exocarp 2.1–2.4 mm thick, coriaceous and light yellow when dry; fruiting calyx persistent, accrescent and covering the fruit, enveloping the berry loosely, the lobes to 4–4.5 cm long, 1.3 cm wide. Seeds numerous, 3.3–3.6 mm long, 1.5–1.7 mm wide, ochre yellow when dry, the testa reticulate, the testal cells rectangular in outline, the embryo slightly curved, the cotyledons accumbent, slightly longer than embryo rest, endosperm rather scanty. Chromosome number not known.
(Fig.
Grows in Andean tropical cloud forest from 1,500 to 2,300 m elevation.
(
Doselia galilensis has been collected in flower in May, June and October and with fruits in June.
The epithet “galilensis” is in honour of the recently created “Parque Natural Regional Bosque de Galilea”, where the type specimen was collected. The Galilea Forest is located between 3°53'36"N, 74°31'51"W and 3°40'32"N, 74°44'20"W in the municipalities of Villarrica and Dolores. We hope that the description of this new Colombian endemic species highlights the importance of the Galilea Forest and stimulates more researchers to explore this beautiful reserve. The Galilea Forest covers more than 26,000 hectares and occupies an elevational range from 1,480 to 3,080 m. It represents a mid-elevation Andean montane forest sandwiched between the lowland tropical rain forest and treeline. Besides the typical Andean cloud forest, the Galilea Forest comprises cushion mire wetlands known as “turberas” and white-sand forests with species adapted to grow in these highly specialised soil conditions (e.g., Utricularia L., Lentibulariaceae). The Galilea Forest is considered a strategic ecosystem for water regulation in the watershed area of the Negro River and the Aco and Lusitania ravines that feed the Hidroprado Dam (
In the area of Arcabuco, Boyacá, D. galilensis is sympatric with Merinthopodium vogelii (Cuatrec.) Castillo & R.E.Schult., a vegetatively similar species of Solandreae. Merinthopodium vogelii differs in having green campanulate corollas with strongly reflexed lobes at anthesis and partially exserted anthers, while D. galilensis has included anthers and to greenish-white, infundibuliform corollas with slightly reflexed lobes that are purple-tinged at anthesis.
Doselia galilensis A floriferous branch B branch with clustered and circular foliar scars C flower at anthesis D dissected corolla showing inserted stamens E detail of a stamen F gynoecium with a five-lobed nectariferous disc G fruit with an accrescent calyx H dissected fruit showing placentation (cross-section) I seed (A–I Corrales et al. 917: Drawing by Marcela Morales).
Doselia galilensis can be easily differentiated from other species of Doselia in its glabrescent mature leaf blades where pubescence is sparse and restricted to midveins and margins (Fig.
Morphological and geographical comparison of the four species assigned to Doselia.
D. epifita | D. galilensis | D. huilensis | D. lopezii | |
---|---|---|---|---|
Distribution | Napo, Pastaza (Ecuador), Putumayo, Caquetá (Colombia) | Boyacá, Santander, Tolima (Colombia) | Huila, Putumayo (Colombia) | Antioquia, Caldas, Risaralda, Valle del Cauca (Colombia) |
Elevation (m) | 500–1,500 | 1,500–2,300 | 2,200–2,300 | 1,700–2,100 |
Leaf length × width (cm) | 11–25 × 6–12 | 9.2–17.5 × 6.4–8.4 | 9.0–16.7 × 4.6–11.7 | 14–22 × 4–9.8 |
Leaf shape | Obovate | Ovate to elliptic | Elliptic to broadly elliptic | Elliptic to broadly elliptic |
Mature leaf pubescence | Sparsely pubescent on both surfaces with simple uniseriate trichomes | Sparsely pubescent on the main veins and margins with simple uniseriate trichomes, becoming glabrescent with age | Densely pubescent on both surfaces with simple uniseriate trichomes | Sparsely pubescent on both surfaces with simple uniseriate trichomes |
Peduncle length (cm) | 8.5–32.2 | 1.2–5.7(–32.5) | 8.5–39 | 6–24(–39) |
Corolla shape | Infundibuliform | Infundibuliform | Tubular-campanulate | Hypocrateriform |
Corolla length (cm) | 9–11 | 12–15 | 8.5–10 | 8–11 |
Corolla lobe length × width (cm) | 3.4–4.2 × 2.5–3.3 | 3.2–3.8 × 2.8–3.1 | 2.3–3.3 × 1.6–1.7 | 3.2–3.9 × 3.7–4.1 |
Anther position | Included | Included | Exserted | Partially exserted |
Anther length (cm) | 1.6–2.7 | 1.6–2.1 | 1.4–1.9 | 1.9–2.2 |
Doselia galilensis A habitat in type locality in Galilea forest, Villarrica, Tolima B mid-elevation moist forest habitat in Galilea forest C floriferous branch with a flower D floriferous branch with a dissected and opened corolla showing corolla colouration inside and the inserted stamens E female Euglossa bee visiting the flower F front view of the corolla showing the purple-tinged lobes and the inserted stamens G developing fruit covered by an accrescent calyx (A–G Corrales et al. 917: Photographs by Boris Villanueva).
Colombia. Boyacá: Municipio de Arcabuco, La Cumbre, 2300 m, 22 May 1980 (fl), Pérez 01 (COL). Santander: Municipio de San Vicente de Chucurí, vereda Centro, sector Germania, parte alta, serranía de Los Yariguíes, camino de Lengerke entre Zapatoca y San Vicente de Chucurí, sector Reserva Proaves “Reinita Cielo Azul”, 6°50'46"N, 73°22'38W, 1672 m, 20 Oct 2021 (fl), D. Díaz-Rueda et al. 2272 (
Markea huilensis
A.Orejuela & J.M.Vélez, Phytotaxa 167(2): 156, Figs
Based on Markea huilensis A.Orejuela & J.M.Vélez
Doselia huilensis A floriferous branch B indument on the leaf blade C indument on young stems D leaf arrangement E flower bud F flower in anthesis G stamens H gynoecium I fruit with sepals removed J fruit in cross section K seed L embryo (A–L A. Orejuela & J.M. Vélez 112: Drawing by Marcela Morales, first published in
Hemiepiphytic liana with adventitious roots. Stems densely pubescent with simple, uniseriate (2–) 4–7 (–11)-celled, hyaline to ochre-brown trichomes 0.2–1.8 mm long, with a deciduous apex and a persistent multicellular base giving the surface a tuberculate appearance, stems glabrescent with age. Leaves tightly clustered towards the branch tips, 9.0–16.7 cm long, 4.6–11.7 cm wide, elliptic to broadly elliptic, densely pubescent with simple 4–9-celled uniseriate hyaline to dark olive-brown trichomes 0.3–2 mm long on both surfaces; major veins 4–6 pairs, slightly raised abaxially; base cuneate or obtuse, asymmetric; margins entire to undulate; apex usually acuminate, mucronate; petiole 0.4–3.8 cm long, densely pubescent. Inflorescence sub-axillary, simple to branched, bracteate, 18–50 cm long, ca. 2–7-flowered, surface tuberculate and densely pubescent with trichomes as on the stems; peduncle 8.5–39 cm long; bracts foliaceous and linear, 5–6 cm long, 1–2 cm wide; pedicels 1.5–2 cm long, distally winged and thickened. Calyx ca. 3.3 cm long, 1.5 cm wide, dark green with purple margins and reticulate along the veins, pubescent with simple 4–7-celled uniseriate white hyaline to brown trichomes; tube 0.5–0.7 mm long; lobes undulate, 2.7–5.2 cm long, 1.3–1.5 cm wide, lanceolate, apically acuminate with an acumen 0.6–0.9 mm long, green with the main vein and the margins purple-brown, pubescent with simple uniseriate trichomes on the abaxial side. Corolla 8.5–10 cm long, the inner corolla diameter 4.5–5 cm, tubular-campanulate; tube 6.2–6.7 cm long, scarcely pubescent with trichomes similar to those of the calyx, yellowish green with strong purple-tinged reticulation along major and minor veins both abaxially and adaxially; tube differentiated into a narrow base ca. 0.2 cm long and 0.8–1 cm wide and a wide distal portion 4.2–4.6 cm long, ca. 5 cm wide; lobes 2.3–3.3 cm long, 1.6–1.7 cm wide, oblong, reflexed during anthesis, colour similar to that of the corolla tube, the margins revolute, the apex obtuse, glabrous. Stamens 6.1–6.9 cm long, fully exserted beyond corolla tube; filaments 4.7–5 cm long, adnate at ca. 2 cm from the base of the corolla, purplish, densely pubescent with simple uniseriate trichomes at the insertion point like those on calyx; anthers 1.4–1.9 cm long, 1.3–1.5 mm wide. Ovary ca. 7 mm long, ca. 3.5 mm wide, light yellow, glabrous; style 7.3–8 cm long, cream; stigma clavate. Fruit ca. 4.2 cm long, ca. 2.5 cm wide, dark green, exocarp 2–2.8 mm thick when fresh, coriaceous, black when dry; fruiting calyx persistent, accrescent and covering the fruit, appressed at maturity, the lobes 4–5 cm long, 2.2 cm wide. Seeds numerous, 2.6–3.0 mm long, 1.2–1.4 mm wide, ochre when fresh, dark brown when dry, the testa reticulate, the testal cells rectangular in outline. Chromosome number unknown.
Doselia lopezii A branch with clustered leaves and an inflorescence axis B trichomes on leaves C trichomes on the calyx D trichomes at the filament insertion point E flower at anthesis F dissected corolla showing partially exserted stamens G stamen in lateral view H detail of a stamen showing the insertion of the filament I detail of the stigma J fruit with an accrescent calyx with two calyx lobes removed (A–I López Filgueiras 8208: Drawing by Nidia Flury, first published in
(Fig.
Doselia huilensis is found in preserved or partially altered oak forests from 2,200 to 2,300 m elevation.
Doselia huilensis differs from all other Doselia species in its tubular-campanulate corollas with fully exserted anthers (Table
Colombia. Huila: Municipio de La Plata,-vereda Agua Bonita, Finca Meremberg, 2,200–2,300 m, 21 Jul 1975 (fl), Díaz-Piedrahita et al. 793 (COL); Carretera Popayán-La Plata, km 100, Reserva forestal de Fauna y Flora Meremberg, 2,300 m, 12 Dec 1982 (fl), Murcia 09 (COL); Municipio San Agustín, vereda La Castellana, reserva privada Los Yalcones, interfluvio quebrada El Palmar-río Balseros, sitio El Palmar, 1°47´44"N, 76°21´5"W, 2,400–2,600 m, 15–20 Aug 2005, Mendoza & Robles 16548 (FMB). Putumayo: Valle del Sibundoy, Reserva Natural Yumartán, cuenca alta del río Putumayo, por la garganta del Balsayaco, 3 Mar 2021 (fl), Coral 34 (HEAA).
Markea lopezii Hunz., Lorentzia 5: 9. 1985. Type. Colombia. Valle: Cuenca del Rio Cali, cercanías de Peñas Blancas, 10–11 Jan 1963 (fl), M. López-Figueiras 8208 (holotype [two sheets]: US! [00385918, acc. # 2451166; 00385936, acc. # 24511165]).
Based on Markea lopezii Hunz.
Hemipiphytic liana with adventitious roots. Stems sparsely pubescent with simple, uniseriate 4–8-celled, hyaline ochre trichomes 0.5–1.2 mm long, with deciduous apex and a persistent multicellular base giving the surface a tuberculate appearance, stems glabrescent with age. Leaves tightly clustered towards the branch tips, (7–)10–22 cm long, 4–9.8 cm wide, elliptic to broadly elliptic, sparsely pubescent with simple uniseriate 4–7-celled hyaline trichomes 0.8–1.2 mm long on both sides; major veins 4–6 pairs, slightly raised abaxially; base cuneate, slightly asymmetric; margins entire; apex acute to acuminate; petiole (7–)10–19(–25) mm long, densely pubescent with simple trichomes as on the leaves. Inflorescence axillary, simple, ebracteate, 7–35(–50) cm long, ~2–5-flowered, densely pubescent with simple, uniseriate trichomes like those on stems; peduncle 6–24(–39) cm long; pedicels 0.8–1.7(–3.0) cm long, distally winged and thickened. Calyx 3.8–5.6 cm long, 1.4–1.9 cm wide, green, sometimes tinged with purple, densely pubescent with simple, uniseriate trichomes like those on leaves; tube 8–9 mm long; lobes flat, 3.5–4.4 cm long, 1.2–1.4 cm wide, long-triangular, apically long-acuminate, green or green with purple margins, sparsely pubescent with simple uniseriate trichomes on the abaxial side. Corolla 8–11 cm long, the inner corolla diameter 2.5–2.8 cm, hypocrateriform; tube 7–10 cm long, orange and tinged with purple in the throat internally, glabrous; lobes 3.2–3.9 cm long, 3.7–4.1 cm wide, triangular, orange, spreading to slightly reflexed at anthesis, the margins flat and entire, the apex acute, glabrous or with a few minute trichomes along the veins. Stamens 7–8.6 cm long, partially exserted beyond the mouth of corolla tube; filaments 5.3–6.1 cm long, adnate at 1.0–1.3 cm from the base of the corolla tube, purple-tinged, densely pubescent with simple 6–10-celled uniseriate trichomes at the insertion point; anthers 1.9–2.2 cm long, 1.4–1.8 mm wide. Ovary ca. 7.5 mm long, ca. 2.9 mm wide, yellow, glabrous; style 7.9–8.8 cm long, glabrous; stigma clavate. Fruit 2.7–3.2 cm long, 1.5–3.3 cm wide, light green; fruiting calyx persistent, accrescent and fully covering the fruit, the lobes 3–3.3 cm long, 1.2–1.4 cm wide. Seeds numerous, 2.2–2.7 mm long, 1–1.3 mm wide, ochre when fresh, brown when dry, the testa reticulate, the testal cells rectangular in outline. Chromosome number not known.
(Fig.
Mid-elevation moist forests from 1,700 to 2,100 m elevation.
(
Doselia lopezii is the type species of the genus and the easiest species to recognise on account of its showy flowers with large orange corollas (Table
Colombia. Antioquia: Medellín– Puerto Triunfo, Cocorna, ca. 5 km E of Cocorna Peaje, Quebrada El Biadal, 6°N, 75°10´W, 1,830 m, 23 Nov 1983 (fl), Juncosa 1400 (JAUM); Pulperies 6000, Jul 1880 (fl), Kalbreyer 1638 (K, MO). Caldas: Municipio Samaná, corregimiento Florencia, Vereda San Vicente, sector río Claro - Sierra Morena, Parque Nacional Natural Selva de Florencia, 5°31'21"N, 75°3'40"W, 1,840 m, 2 Oct 2012 (fl), Betancur et al. 16698 (COL). Risaralda: Municipio de Pueblo Rico, en cercanías de los límites con el PNN Tatamá, Reserva Santuario-Tatamá, 5°11'03.1"N, 76°01'16.5"W, 1821 m, 18 Mar 2022 (fl), Orejuela et al. 3849 (
We thank herbarium curators at the herbaria whom we visited for their help and permission to consult the collections in their care, and the following colleagues for their support and expertise during fieldwork in Colombia and Ecuador: Gloria Barboza, Jhoana Castillo, Brayan Coral, Lina Corrales, Rocio Deanna, Daniel Mauricio Diaz Rueda, David Granados, Segundo Leiva, Alvaro Perez, Federico Roda, Gina Sierra, Juan David Tovar, Edwin Trujillo, Carlos Vargas and Jorge Mario Vélez. We extend our thanks to Marcela Morales, Omar Bernal, Nidia Flury and Humberto Mendoza for the excellent illustrations accompanying this paper. The study was funded by the Fundación CEIBA through the scholarship program “Rodolfo Llinás para la promoción de la formación avanzada y el espíritu científico en Bogotá”. Additional financial support was provided by the Darwin Trust and the Davis Fund from the University of Edinburgh, and the Royal Botanic Garden Edinburgh, the Systematics Association Fund and the GEME Max Planck Tandem Group (Agreement 566 from 2014 between the Universidad Nacional de Colombia (https://unal.edu.co/) and Colciencias (now called Minciencias https://minciencias.gov.co/). To Alistair Hay, Andreas Kay (deceased), Brayan Coral, and Eduardo Calderon for providing Doselia photos. We also thank Lynn Bohs, Gloria Barboza and Leandro Giacomin for their comments and suggestions, which improved this manuscript.