Research Article |
Corresponding author: Andrés Orejuela ( aaorejuelar@gmail.com ) Academic editor: Leandro Giacomin
© 2023 Andrés Orejuela, Stacey D. Smith, Boris Villanueva, Rocío Deanna.
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, Smith SD, Villanueva B, Deanna R (2023) A new species of Iochroma Benth. (Solanaceae) from the eastern Andes of Colombia. PhytoKeys 232: 133-144. https://doi.org/10.3897/phytokeys.232.108474
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Iochroma orozcoae A.Orejuela & S.D.Sm., sp. nov. (Solanaceae) is described from the Andean forests of Cundinamarca in the eastern cordillera of Colombia. Iochroma orozcoae was first collected by the eminent Spanish priest and botanist José Celestino Mutis in the late part of the 18th century, but the specimens have lain unrecognised in herbaria for over 200 years. The species shares many features with its closest relative, Iochroma baumii S.D.Sm. & S.Leiva, but it differs from it in having small flowers with five corolla lobes and few inflorescences per branch, located near the shoot apex with 1 to 4 (–8) flowers, fruits that are greenish-yellow when ripe and its restricted geographic distribution. A description of I. orozcoae is provided, along with a detailed illustration, photographs of live plants, a comparison with closely-related species and a key to all Colombian species of Iochroma Benth. In closing, we emphasise the value of historical collections for the knowledge of biodiversity.
Iochroma orozcoae A.Orejuela & S.D.Sm., sp. nov. (Solanaceae) se describe a partir de los bosques andinos de Cundinamarca, en la cordillera oriental de Colombia. Iochroma orozcoae fue recolectada por primera vez por el eminente sacerdote y botánico español José Celestino Mutis a finales del siglo XVIII, pero los especímenes han permanecido sin ser reconocidos en herbarios durante más de 200 años. La especie comparte muchas características con su pariente más cercano, Iochroma baumii S.D.Sm. & S.Leiva, pero difiere de esta en sus flores más pequeñas con cinco lóbulos de la corola y pocas inflorescencias por rama ubicadas cerca del ápice de la rama, con 1 a 4 (–8) flores por inflorescencia, frutos verde-amarillentos al madurar, y su distribución geográfica restringida. Se presenta una descripción de I. orozcoae, junto con una ilustración detallada, fotografías en vivo, una comparación con las especies cercanamente relacionadas y una clave para las especies colombianas de Iochroma Benth. Finalmente, enfatizamos el valor de las colecciones históricas para el conocimiento de la biodiversidad.
Andes, Colombia, Iochroma, Iochrominae, IUCN Red List, Solanaceae
Iochroma Benth. (Solanaceae) is a neotropical genus that comprises approximately 30 species distributed from Colombia to Peru, with the highest concentration of species found in the Peruvian Andes (
All specimens of the genus Iochroma from the Colombian herbaria
For phylogenetic analysis, DNA was extracted from silica gel dried leaf material and three nuclear markers were sequenced (internal transcribed spacer region (ITS), the granule-bound starch synthase (GBSSI or waxy) gene and the second intron of LEAFY (LFY)), following
To map the distribution of the new species and its close relatives, 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 (
Colombia. Cundinamarca: vía Ubate-Lenguazaque, carretera sin pavimentar, antes del sector conocido como las balsas, 5°20'2.5"N, 73°43'23"W, 2600 m elev., 27 August 2017, A. Orejuela & J. Castillo 2942 [holotype:
Iochroma orozcoae A.Orejuela & S.D.Sm. A flowering branch B inner corolla surface, showing the stamens C flower. Detail is shown for the simple and branched hairs on the corolla D gynoecium E, F details of the stamens and anthers G fruit with persistent calyx H seed. Drawn by Omar Bernal from Orejuela & Castillo 2942 & Orejuela et al. 3407.
A Iochroma baumii S.D.Sm. & S.Leiva affinis, sed paucarum inflorescentiae in ramum prope apicem germinis dispositae sunt, quae 1 ad 4 (–8) floribus, flores minores 1.5–2 cm longi; corolla quinque lobis constat, et fructus viridis-flavus colore maturo differt.
Iochroma orozcoae A.Orejuela & S.D.Sm. A young branch B stem detail C, D leaf detail showing the abaxial and adaxial sides E floriferous branch with flowers in lateral view F floriferous branch with a flower in frontal view G fruits showing the accrescent appressed calyx. Photos by Andrés Orejuela.
Shrub 1.5–3 m tall; young stems pubescent with branched hairs, older stems with corky bark. Leaves alternate, simple, (2.1–) 5.3–11.6 × (1.2–) 2.2–5.5 cm, elliptic; adaxial surface glabrescent; abaxial surface densely pubescent with branched hairs; base cuneate; margins entire; apex acute to attenuate; petioles (0.4–) 0.8–2.5 (–3) cm. Inflorescences axillary on young branches near the shoot apex, 1 to 4 (–8)-flowered; pedicels 1.1–1.7 (–2) cm in flower, 1.6–2.4 cm in fruit, terete, pendulous, densely pubescent with many-branched hairs. Calyx 6–7 × 5–6 mm, tubular to slightly urceolate, green, with few to many-branched hairs, with five broadly triangular lobes, ca. 0.8 × 2–2.5 mm in flower, shallowly divided in flower, deeply divided in fruit, in fruit, the calyx accrescent 9–10 × 10–11 mm, lobes 4–5 × 7–8 mm; corolla 15–20 × 4–4.5 mm at anthesis, tubular, flaring at the mouth, the exterior deep blue-purple, with many simple or occasionally branched hairs, the pubescence increasing towards the mouth, the interior deep blue-purple, glabrous, the lobes 5, 2.6–2.8 × 2.2–2.6 mm, acute to the apex and cucullate; stamens 5, included; filaments with simple and branched hairs, fused to the corolla at 3.8–5.2 mm from the base, with the free portion 7–11.4 mm long; anthers 3.5–5.2 × 1.3–1.7 mm, oblong, cream, longitudinally dehiscent; ovary 3.6–4.8 × 1.7–2.5 mm, pyriform, glabrous, with a yellow nectariferous disc, style 13–15 mm, the stigma green, clavate, bilobed. Berry 12–14 × 9–11 mm, slightly ovoid, greenish-yellow at maturity with 20 to 30 sclerosomes, the basal 3/4 enveloped in the accrescent fruiting calyx; seeds 110 to 170 per berry, 2–2.1 × 1.7–1.9 mm, yellow, reniform.
This species is named in honour of Clara Inés Orozco Pardo, an Associate Professor of botany at the Instituto de Ciencias Naturales of the Universidad Nacional de Colombia. Her dedication to the understanding of Colombian flora, particularly in the fields of taxonomy and systematics of the Brunelliaceae and Solanaceae families, has been remarkable. She has also played a crucial role in mentoring several Colombian botanists, including the first author of this paper.
Iochroma orozcoae is found in the Municipality of Lenguazaque, Cundinamarca Department, in the eastern Andes of Colombia, in the surroundings of the rock formation known as the Farallones de Lenguazaque, which is situated at an elevation of 2600 m (Fig.
Iochroma orozcoae belongs to the “F” clade (
Maximum Likelihood phylogeny of Iochrominae including the new species, I. orozcoae. Relationships based on three nuclear markers with taxon sampling following
Iochroma orozcoae is classified as a data-deficient species (DD) due to the limited information on its abundance and distribution that is inadequate for comprehensively assessing its conservation status. The species has been collected only on four occasions, suggesting probable local rarity. The initial collection was made by José Celestino Mutis in 1783 from an unknown locality, followed by another by Oscar Haught in 1947 from Lenguazaque, Cundinamarca and a couple of recent collections by the first author from the same locality as Haught’s collection. Despite conducting an extensive search through specialised literature, the drawings of the Royal Botanical Expedition of the New Kingdom of Granada (1783–1816) and Mutis’ journals, the precise location of Mutis’ first collection remains unknown. We infer that the distribution of the species may be highly restricted, based on the small number of known collections from the same locality. Assumptions regarding its scarcity and restricted distribution, however, require further studies. Living plants of I. orozcoae originated from seeds, collected at the type locality and are conserved ex situ in the living collections of the Jardín Botánico de Bogotá.
Our phylogenetic analyses offer strong support for the placement of I. orozcoae in the northern Andean “F” clade from
A morphological and geographical comparison of Iochroma species in the “F” clade (sensu
Iochroma orozcoae | Iochroma baumii | Iochroma gesnerioides | Iochroma fuchsioides | Iochroma calycinum | |
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Geographical distribution | Eastern Colombia | Ecuador and central Colombia | Ecuador and southern and central Colombia | Peru, Ecuador and southern Colombia | Ecuador and Peru |
Habitat | Partially altered high Andean forests and roadsides to 2600 m | Disturbed cloud forest habitats, such as pasture hedges, forest gaps and roadsides from 2600 to 3300 m | Common in middle to high elevations in cloud forests, pastures hedges and roadsides from 1800 to 3300 m | Common in middle to high elevations in cloud forests, pastures hedges and roadsides from 2400 to 3500 m | Common in some areas of the wet cloud forest from 2000 to 3350 m |
Leaf size (cm) | (2.1–) 5.3–11.6 × (1.2–) 2.2–5.5 | (6–) 9–17 × (2.5–) 3–6 | 12–18 (–30) × 5–7.5 (–12) | 3.5–7 (–9) × 1.5–3 (–5) | 12–23 × 5–9 |
Leaf shape | Elliptic | Elliptic to lanceolate | Elliptic to lanceolate | Obovate to elliptic | Elliptic to lanceolate |
Inflorescence | Inflorescences axillary on young branches near the shoot apex, 1 to 4 (–8)-flowered | Inflorescences axillary, typically on older, often leafless branches, rarely near the shoot apex, 6 to 12-flowered | Inflorescences axillary, in clusters with masses of flowers (with up ca. 120 flowers) along older or upper leaf nodes | Inflorescences axillary, in small clusters of flowers (1 to 12 flowers) on young branches near the shoot apex or upper leaf nodes | Inflorescences axillary, in small clusters of flowers on young branches near the shoot apex (5 to 12 flowers) or older branches (1 to 6 flowers) |
Calyx in flower | Calyx 6–7 × 5–6 mm, slightly urceolate, green, with densely branched hairs | Calyx 4–9 × 3.8–6 mm, tubular to slightly urceolate, purplish-green, with few to many branched hairs | Calyx 3–5 × 4–5 mm, cup-shaped, green, with densely branched hairs | Calyx 7–13 × 4.5–5.5 mm, tubular to campanulate, green, hairless or with a few scattered branched hairs | Calyx 25 × 8–17 mm, inflated, elliptic, purplish, hairless or with a few branched hairs |
Corolla | Corolla 15–20 × 4–4.5 mm, deep blue-purple, 5-lobed | Corolla 28–45 × 4–6 mm, deep blue-purple, 10-lobed | Corolla 25–40 × 5–6 mm, red, orange-red, orange to salmon, 5-lobed | Corolla 20–30 × 6–7 mm, red to orange-red, 10-lobed | Corolla 50–65 × 4–8 mm, deep blue to purple, 5-lobed |
Fruit | Berry slightly ovoid ca. 1.2–1.4 × 0.9–1.1 cm, greenish-yellow when ripe | Berry markedly ovoid 1.2–1.7 × 1–1.5 cm, green and purple when ripe | Berry markedly ovoid, 1.3–1.8 x 0.9–1.3 cm green, reddish-brown or purple when ripe | Berry markedly ovoid, 1.6–2.5 x 1.2–1.7 cm, yellow, green or brown when ripe | Berry markedly ovoid to conical, 2.3–3.5 x 1.4–2 cm, white when ripe |
Species within the “F” clade of Iochroma that are closely related to I. orozcoae A flowering branch of I. fuchsioides observed at Parque Recreacional y Bosque Protector Jerusalem, Malchingui, Ecuador B flowers of I. gesnerioides photographed at the Jardín Botánico de Bogotá, Colombia C flowers of I. calycinum at the reserva Otonga, Sigchos, Cotopaxi, Ecuador D flowers and fruits of I. baumii observed close to the type locality in Papallacta-Baeza road, Quijos, Napo, Ecuador E flowering branch of I. baumii from populations from La Pastora, Caldas, Colombia F fruits of I. gesnerioides photographed at the Jardín Botánico de Bogotá, Colombia G fruits of I. fuchsioides photographed close to Pasto, Nariño, Colombia. Photos by Andrés Orejuela (B, C, F, G), Hal Mitchell (D), Nathaly Obregón (A) and Juan David Tovar (E).
The rediscovery of I. orozcoae over two centuries since the first collection by José Celestino Mutis in 1783 underscores the immense importance of historical collections in documenting and describing botanical diversity and their potential contribution in setting conservation priorities within biodiverse regions. This is a widely-debated topic that is gaining increasing attention over time (
Colombia. Cundinamarca: Ubate – Lenguazaque Highway, 2600 m elev., 16 September 1947, O.L. Haught 6188 (
1 | Corolla 15–70 mm long, tubular, flaring at the mouth, red, orange-red, orange, deep blue or purple, calyx conspicuously accrescent in fruit, fruit a slightly to markedly ovoid to conical berry | 2 |
– | Corolla 8–11 mm long, campanulate-infundibuliform, white, greenish or greenish-cream coloured, calyx non-accrescent or accrescent to an insignificant degree in fruit, fruit a globose berry | Iochroma arborescens |
2 | Corolla red, orange-red or orange-coloured | 3 |
– | Corolla deep blue or purple-coloured | 4 |
3 | Leaves usually pubescent; inflorescences with 30 to 120 flowers; calyx 3–7 mm long, cup-shaped, densely pubescent; corolla pubescent; anthers usually included | Iochroma gesnerioides |
– | Leaves usually glabrescent; inflorescences with 1 to 15 flowers; calyx 7–13 mm long, tubular to campanulate, glabrous or with a few scattered hairs; corolla glabrescent; anthers usually exserted or partially exserted | Iochroma fuchsioides |
4 | Leaves (2.1–) 5.3–11.6 × (1.2–) 2.2–5.5 cm. Inflorescences few per branch, located near the shoot apex; flowers 1 to 4 (–8) per inflorescence; corolla 1.5–2 cm long, 5-lobed; berry greenish-yellow when ripe | Iochroma orozcoae |
– | Leaves (6–) 9–17 × (2.5–) 3–6 cm. Inflorescences many per branch, typically in massive clusters on older, often leafless branches, rarely near the shoot apex; flowers 6 to 12 per inflorescences; corolla 2.8–4.5 cm, 10-lobed; berry green and purple when ripe | Iochroma baumii |
We would like to express our gratitude to herbarium curators for their assistance and granting permission to access the collections under their care. We are also thankful to the following colleagues for their valuable support and expertise during our fieldwork in Colombia and Ecuador: Segundo Leiva, Gloria Barboza, Victor Luna, Juan David Tovar, Felipe Bohorquez, Jhoana Castillo, Diego Másmela and Humberto Mendoza. Special thanks go to Juan David Tovar and Felipe Bohorquez for their efforts in collecting and confirming the presence of I. baumii in Colombia. Lastly, we extend our appreciation to Hal Mitchell, Nathaly Obregón and Juan David Tovar for their live photographs, as well as to Omar Bernal for the excellent illustration accompanying this paper. We also thank Sandy Knapp, Gloria Barboza and Leandro Giacomin for their comments and suggestions, which improved this manuscript.
The authors have declared that no competing interests exist.
No ethical statement was reported.
AO and BV would like to acknowledge the Jardín Botánico de Bogotá (
AO wrote the original draft and prepared the figures; RD and SDS performed the molecular analysis; AO, SDS, BV, and RD reviewed and edited the drafts.
Andrés Orejuela https://orcid.org/0000-0002-3511-1478
Stacey D. Smith https://orcid.org/0000-0003-2010-4459
Boris Villanueva https://orcid.org/0000-0002-6929-3572
Rocío Deanna https://orcid.org/0000-0001-8753-7596
All of the data that support the findings of this study are available in the main text.