﻿Monograph of Doselia (Solanaceae), a new hemiepiphytic genus endemic to the northern Andes

﻿Abstract 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. (Doseliaepifita (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 (Doseliagalilensis 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.


Introduction
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 (Orejuela et al. 2017;Orejuela et al. in prep). The group is restricted to the Neotropics, with species distributed from Mexico and the Caribbean to Bolivia and southern Brazil (Orejuela et al. 2017). A centre of endemism for the tribe lies in Andean Ecuador and Colombia, where ca. 60% of the species are found (Orejuela et al. 2017).
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%; Hunziker 2001). The tribe is also the only group of Solanaceae with known ant associations (e.g., Merinthopodium Donn.Sm., Markea Rich., and species of Hawkesiophyton Hunz.; Knapp et al. 1997;Hunziker 2001;Orejuela et al. 2017).
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 (Vogel 1958;Cocucci 1999;Muchhala and Jarrin-V 2002;Sazima et al. 2003;Knapp 2010).
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 (Orejuela et al. 2017). In addition to extensive recircumscription of genera, two new lineages have been identified within Solandreae based on nuclear and plastid Sanger sequences and whole plastome data that represent distinct morphological groups comprised of species previously described as members of Markea that are distinct at the generic level: the Markea lopezii and Markea antioquiensis clades (Orejuela et al. 2017;Orejuela et al. in prep).
Here we focus on the morphologically distinct Markea lopezii clade (Figs 1, 2; Table 1), a group of four species from mid-elevation moist Andean forests of Ecuador and Colombia. The group includes three previously described species, M. epifita S. Knapp, M. huilensis A.Orejuela & J.M.Vélez, and M. lopezii Hunz. The fourth was discovered in 2018 during fieldwork in Colombia in the Parque Natural Regional Bosque de Galilea in the municipality of Villarrica, Tolima, and is described here. The four species treated here were resolved as a monophyletic group, named the Markea lopezii clade, with strong branch support in a molecular phylogenomic study of Solandreae that included 95% of the species (76 spp.; Orejuela et al. in prep).
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 (QGIS Development Team 2021). Conservation assessments were made based on the IUCN Red List categories and criteria (IUCN 2012) and the most recent guidelines for using the IUCN Red List Categories and Criteria . For the conservation assessments, Extent of Occurrence (EOO) and Area of Occupancy (AOO) were calculated using GeoCat (www.geocat.kew.org; Bachman et al. 2011) with a 2 km 2 cell size. Herbarium material, field observations, and photos were all used to construct the identification key.
Etymology. 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. Distribution (Fig. 2). Mid-elevation moist Andean forests from 500 to 2,300 m in Ecuador (Provinces of Morona Santiago, Napo, Pastaza) and Colombia (Departments of Antioquia, Boyacá, Caldas, Caquetá, Huila, Putumayo, Risaralda, Santander, Tolima, Valle del Cauca).
Discussion. 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 1). Leaves of all Doselia species are highly clustered on branch tips in whorls of 4-6 similar to species in the Markea antioquiensis clade and some species of Markea (e.g., M. plowmanii Hunz.) and differ from all other genera and species of the tribe where leaves are more spread apart and clearly alternate (Table 1). Leaves in Doselia are membranous with simple eglandular trichomes on both surfaces, a character shared with some species of the Markea antioquiensis clade (e.g., M. pilosa S.Knapp; Table 1). In many other genera of Solandreae, the leaves are chartaceous (e.g., Hawkesiophyton Hunz., Juanulloa Ruiz & Pav., Merinthopodium Donn. Sm., Solandra and Trianaea Planch. & Linden) or subcoriaceous to coriaceous (e.g., Schultesianthus) and often have simple glandular and/or dendritic trichomes in addition to the simple eglandular ones (Table 1). 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 1). Such inflorescences are not typical in the tribe but are observed only in a few other species in Solandreae, including Markea coccinea Rich., Merinthopodium neuranthum (Hemsl.) Donn.Sm., Merinthopodium pendulum (Cuatrec.) Hunz., and Trianaea nobilis Planch. & Linden. Pedicels in some Doselia species are distally winged because the sutures of the calyx are winged and continue onto the pedicel. Distally winged pedicels are also known in some species of the Markea antioquiensis clade (e.g., Markea antioquiensis S.Knapp and Markea pilosa S.Knapp; Table 1). 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 1). The two groups also differ in their calyx lobes, where lobes have acute to long-acuminate tips in Doselia but are rounded in the Markea antioquiensis clade. Corollas in the two other morphologically closely related genera Solandra and Schultesianthus are slightly zygomorphic (Table 1).
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). Fruits in Schultesianthus appear similarly leathery but are globose in shape and covered only partially by an irregularly splitting calyx (Table 1). Chromosome number is not known for Doselia, but count numbers in other members of Solandreae, have shown a basic chromosome number ×=12 for Dyssochroma Miers (Piovano 1989;Acosta and Moscone 2000), Solandra (Campin 1924;Lepper 1982) and Trianaea (Chiarini et al. 2019). Similar chromosome counts might be expected for Doselia, but further research is necessary to confirm this assumption.
Distribution (Fig. 2). On the eastern slopes of the Andes in central Ecuador (Provinces Morona-Santiago, Napo, and Pastaza) and Colombia (Departments Putumayo and Caquetá).
Ecology. In premontane forest between 500-1,500 m elevation. Preliminary conservation status . Our data support the assessment of the species by Knapp et al. (2017) who considered D. epifita as vulnerable (VU) based on the criteria B1ab [iii]. Doselia epifita is known from a few collections in the Cordillera de los Guacamayos, the protected areas of Sumaco-Napo-Galeras and Sangay, areas near the city of Puyo in Ecuador, the Natural Reserve "La isla escondida" in Putumayo, and the surroundings of the Alto Fragua indiwasi National Park in Caquetá, Colombia. The biggest threat to the species is deforestation (Knapp et al. 2017).
Discussion. 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 (Knapp 1998). Recent collections have revealed that the inflorescences are axillary and long (18.5-45 cm long; Fig. 3B), as correctly predicted by Knapp (1998). The fruits of this species remain unknown.
Etymology. 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.   Description. 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. Distribution (Fig. 2). Doselia galilensis occurs in the western slopes of the eastern cordillera of the Colombian Andes and is only known from three localities in the municipality of Arcabuco (Department of Boyacá), the natural reserve "Reinita Cielo Azul" (Department of Santander) and the Parque Natural Regional Bosque de Galilea (Department of Tolima).

Doselia galilensis
Ecology. Grows in Andean tropical cloud forest from 1,500 to 2,300 m elevation. Preliminary conservation status . Doselia galilensis is considered Data Deficient (DD) due to the small number of known populations. Based on our field observations, the biggest threat to the species is habitat loss due to agricultural expansion near the known localities. The situation has been alarming in the Galilea Forest during the last few years, with several direct threats to forest conservation such as agricultural expansion, unsustainable logging, and oil exploitation activities. Fortunately, the Galilea Forest has been recently declared as a protected area through the Corporación Autónoma Regional del Tolima ("CORTOLIMA" resolution 31 adopted on December 16, 2019). The Arcabuco oak forests in Boyacá do not, however, have any legal protection. It is unclear whether the new species remains in the area based on our unsuccessful attempt to collect D. galilensis in Arcabuco in 2019. The third population recently discovered in Santander is under the protection of the Proaves NGO in the natural reserve "Reinita Cielo Azul".
Phenology. Doselia galilensis has been collected in flower in May, June and October and with fruits in June.
Etymology. 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 (Quimbayo-Cardona et al. 2019).

Discussion.
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 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. 1; Table 2). Doselia galilensis is morphologically most similar to D. epifita; both species share several characters that are not present in other species of Doselia, such as infundibuliform corollas and included stamens with very short filaments ( Fig. 1; Table 2). Unlike D. epifita, D. galilensis is sparsely pubescent, with only a few trichomes along the main veins of the leaves and very few trichomes in other parts of the plant. In contrast, D. epifita has a dense and persistent pubescence covering the entire plant with persistent trichomes on both sides of the leaves. The calyx lobes in D. galilensis are flat and lanceolate compared to the long-triangular undulate calyx lobes in D. epifita. Doselia galilensis has slightly larger corollas with greenish-white tubes and purple-tinged lobes on the abaxial side ( Fig. 5C-F) compared to D. epifita with white to purplish corolla tubes with purple lobes on both surfaces (Fig. 1B). Styles are consistently pubescent in D. galilensis along their entire length, while D. epifita has glabrous styles except for a few simple uniseriate trichomes at the very base.  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. Distribution (Fig. 2). Doselia huilensis is known only from the Departments of Huila and Putumayo in southwestern Colombia. Ecology. Doselia huilensis is found in preserved or partially altered oak forests from 2,200 to 2,300 m elevation.
Preliminary conservation status . Doselia huilensis is reaffirmed (following Orejuela et al. 2014) here as an endangered species (EN) according to criteria B1ab [i, iii] based on the small EOO (~750 km 2 ), a small number of known populations, and the highly fragmented condition of the relictual forests where it occurs. The species is known from five collections from three localities. Two of these localities are in the Department of Huila 80 km apart, and one recent collection is known from the Valle del Sibundoy, Department of Putumayo, that extends the species distribution approximately 100 km to the south.
Ecology. Mid-elevation moist forests from 1,700 to 2,100 m elevation. Preliminary conservation status . Doselia lopezii is classified as vulnerable (VU) according to the B1a criterion with an EOO of ca. 6,000 km 2 . The area where it is distributed is severely fragmented and the species is known from fewer than ten localities..
Discussion. 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 2; Fig. 1F, G). Doselia lopezii has anomalous and apparently unique pollen in the genus with prominent spiny supratectal processes (Persson et al. 1994). Preliminary observations in D. huilensis (Orejuela et al. 2014) and specimens of D. epifita examined by Hunziker (1997, as M. lopezii) indicate that the pollen of these two species lack these spiny supratectal processes.  Hunziker (1985), reproduced with permission, the original drawing was edited by Omar Bernal and the fruit drawn by Humberto Mendoza). 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.