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. 2025 Sep 17;263:37–47. doi: 10.3897/phytokeys.263.159645

Ceratostema limonensis (Ericaceae), a new species from the Province of Morona-Santiago, Ecuador

Marco M Jiménez 1,✉, Gabriel A Iturralde 1, Diego Gutiérrez del Pozo 2,3, Nadia Lapo-González 3,4, J R Kuethe 5, Henry X Garzón-Suárez 3,6,7
PMCID: PMC12461215  PMID: 41017859

Abstract

A new species of Ceratostema (Vaccinieae) from south-eastern Ecuador is here described and illustrated. Ceratostema limonensis is distinguished by the singular, subflexuous young branches. The new species is compared to the morphologically similar C. gualaquizensis, which differs by the subterete, longer rachis and shorter corollas with narrower monochromatic corolla lobes. A brief history of the species from the tribe Vaccinieae in Ecuador is provided. Further taxonomic discussions, distribution and conservation status for the new species are provided in this paper.

Key words: Amazon, Andes, premontane forest, rainforest, taxonomy

Introduction

Members of the heather family (Ericaceae Juss.) can be found on all continents (except Antarctica), where it reaches its greatest diversity within the tropical latitudes, such as the mountains of the tropical Andes. Currently, this family is divided into eight subfamilies (Kron et al. 2002), with Vaccinieae being the most represented across Ecuador. This group is foremost characterised by having inferior ovaries and edible fruits and, for this reason, are of local economic importance, being cultivated as small fruit crops, but also in ornamental value (Luteyn 2021). Seventeen genera of Vaccinieae are currently known for Ecuador, with the largest number of species present within the genus Ceratostema Juss. (Luteyn 2021). Initially encompassing 28 species (Luteyn 1996), targeted studies conducted during recent years have increased this number to more than 40, making Ecuador amongst the most biodiverse countries for this genus (Jiménez et al. 2021; Cornejo and Luteyn 2024; Cornejo et al. 2024; Doucette et al. 2024; Jiménez et al. 2024a, 2024b, 2024c; Doucette et al. 2025a, 2025b, 2025c).

Ceratostema was originally established by de Jussieu (1789) and a few attempts have been made since the 20th century to divide the genus into independent genera, namely Englerodoxa Hoerold and Periclesia A.C.Sm. (Hoerold 1909; Smith 1942). Previous writings, including Smith (1952), recognised 16 species of Ceratostema during the mid-1900s; the number was increased to 23 by Luteyn (1986) three decades later. In 1996, Luteyn published a national survey focusing on the Ceratostema present in Ecuador, including a key for the species and introducing nine new species present within this country. Subsequently, Pedraza-Peñalosa et al. (2015) used molecular studies to suggest that Ceratostema is a polyphyletic group, but did not propose an infrageneric classification to better establish an intrageneric understanding of Ceratostema as a whole.

Ceratostema are typically represented by terrestrial plants or epiphytes that frequently arise from lignotubers with alternate, petiolate leaves. Flowers are sympetalous, bearing a lower ovary and stamens that are usually as long as the corolla. The long anther tubules and calyx are articulated with the pedicel and the basal corolla is divided into lobes that are proportionally more elongated than the rest of its length (Luteyn 1986; Luteyn 1996; Cornejo and Luteyn 2024).

Recent fieldwork carried out by the authorial team in the south-eastern Andes of Ecuador has uncovered yet another new species of Ceratostema for Ecuador. This new species was found during botanical studies carried out in the Morona-Santiago Province. It was seen growing in the evergreen forests of the superhumid regions of the eastern slope of the Andes, a region classed as one of the 25 biodiversity hotspots exhibiting some of the richest biotas of flora and fauna on Earth (Myers et al. 2000; Killeen et al. 2007). Unfortunately, a high degree of deforestation and land degradation poses questions about the long-term survival of this species and conservation efforts should be undertaken to prevent further decline of these important habitats. Notes regarding the distribution, natural habitat and conservation status of this new species have been provided in this paper.

Materials and methods

This new species of Ceratostema was discovered by the authors (MJ and HG) during botanical field explorations conducted between 2021 and 2025 in the south of the Morona-Santiago Province. The species was encountered during ongoing floristic surveys focused on poorly-known montane ecosystems present within the region.

Taxonomic evaluation was conducted through detailed comparisons with the known described species of Ceratostema from eastern Ecuador, using specialised literature for the Ecuadorian taxa (e.g. Luteyn (1996); Cornejo and Luteyn (2024); Cornejo et al. (2024); Jiménez et al. (2024b); Doucette et al. (2025c)) and a comparative study utilising online herbarium databases including Tropicos, JSTOR Global Plants and SpeciesLink. Morphological terminology follows Beentje (2016).

Photographs were taken in situ using a Nikon® D3100 camera, equipped with an AF-S VR Micro-Nikkor 40 mm f/2.8G IF-ED lens and a Raynox® DCR-250 mm Super Macro lens, supported by two Nikon® SB-700 AF speed-light flashes. Herbarium specimens were collected, pressed and deposited at the Herbarium of the Universidad Técnica Particular de Loja (HUTPL). Floral structures were dissected manually and macro-photographed in the field, including metric scale references and then preserved in 70% ethanol with 1% glycerine. ImageJ software was applied for obtaining measurements from fresh specimens and high-resolution photographs (https://ij.imjoy.io/). Illustrations were prepared using Adobe® Photoshop CS6. A distribution map was generated in ArcGIS Desktop 10.3.

All fieldwork and specimen collection were conducted under permit No. MAATE-DBI-CM-2022-0248, issued by the Ecuadorian Ministry of Environment and Ecological Transition (MAATE).

Taxonomic treatment

. Ceratostema limonensis

M.M.Jiménez & H.Garzón sp. nov.

A2628450-5379-5E5F-AF1D-340F78C7D82E

urn:lsid:ipni.org:names:77369307-1

Figs 1, 2, 4

Figure 1.

Figure 1.

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Lankester Composite Dissection Plate (LCDP) of Ceratostema limonensis. A. Lignotuber; B. Fertile branch with a close-up of the node (B1) and rachis of the inflorescence (B2); C. Complete flower with a ventral view of the corolla lobes (C1); D. Flower with a longitudinal section of the corolla showing stamens and the corolla without stamens (right); E. Calyx, pedicel and style with a close-up of the ovary (E1) and the bracteole (E2); F. Stamens with a close-up of the pores of the tubule (F1), anther thecae (F2) and external surface of the filament (F3). Prepared by N. Lapo-González from photographs of the holotype.

Figure 2.

Figure 2.

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Leaves of Ceratostema limonensis with close-ups of the indumentum. A. Adaxial view with a close-up of the margin (A1) and mid-vein (A2); B. Abaxial view with a close-up of the basal portion of the leaf (B1). Prepared by N. Lapo-González from photos by H.X. Garzón-Suárez.

Figure 4.

Figure 4.

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Ceratostema limonensis in situ. Photos by Henry X. Garzón-Suárez.

Diagnosis.

Ceratostema limonensis is morphologically similar to C. gualaquizensis, but differs by the longer (4.8–5.1 mm vs. 1 mm long), subterete (vs. obconical) rhachis; the shorter (1.4–1.8 mm vs. 3.4–4.7 mm long), ovate-deltate (vs. narrowly lanceolate) calyx lobes; the shorter corolla (2.9 cm vs. 4.5–4.7 cm long) with narrower lobes (2.3 mm vs. 3.7–4.0 mm wide), that have the same colour as the rest of the corolla (vs. black at the apex and inside); and the shorter (3.1–3.3 cm vs. 4.6–4.7 cm long), pilose (vs. glabrous) filaments.

Type.

Ecuador • Morona-Santiago: Cerca de Limón, 1298 m alt., 28 April 2025, H. Garzón 293 (holotype: HUTPL!).

Description.

Pendant, epiphytic shrubs; indumentum consisting of short, white, almost persistent, eglandular trichomes of 0.2–0.7 mm long, trichomes arranged unevenly, sparsely to densely on younger branches, petioles, leaf blades, inflorescences and flowers, including stamens and style; axonomorphous roots with well-developed lignotubers, lignotubers fusiform, 6.0–7.3 × 2.1–2.8 cm. Stems terete to subterete, 5–8 cm long, glabrous, slightly arching, arising from the lignotuber, the older stems dark brown, cracking longitudinally and exfoliating; younger branches subterete, up to 18 cm long, 0.8–1.3 mm wide, pendant, subflexuous, dark brown, pilose, becoming glabrous and striate when old or after exfoliation; axillary buds emerging 1 mm below the leaf node. Leaves spirally arranged, subsessile, pendulous to almost horizontal; petioles subterete, 0.9–1.9 × 1.0 mm, pilose, pale green; blades lanceolate, 5.5–7.8 × 1.8–2.0 cm, thinly-coriaceous, convex with the basal margins folded to conceal flowers and fruits, base cordate to obtuse, apex acuminate, dark green adaxially, pale green abaxially, lustrous and pilose adaxially, dull and pilose abaxially, becoming glabrous with age, 5–7-plinerved from near the base, the mid-vein raised along almost its length adaxially, thickened in the proximal 12 mm, impressed and hollow abaxially, the secondary veins raised adaxially, weakly impressed abaxially, branching, anastomosing distally with reticulate veinlets, slightly raised adiaxally, obscure abaxially. Inflorescence axillary, 1–2-flowered, sessile; rachis subterete, constricted in the middle, 4.8–5.1 mm long, 1.4 mm thick, subverrucose, covered at the base by several bracts; bracts deltate, 0.4–0.6 mm long, minute, persistent, whitish-green, pilose, acute; floral bracts narrowly ovate, ca. 0.6 × 0.2 mm, suffused with pink, pilose, attenuate; pedicel subclavate, 5.1–5.5 mm long, 2.6–2.8 mm thick, subverrucose, slightly arcuate, pale green, pilose, articulate with the calyx; bracteoles 2, ovate-triangular, 0.5–0.7 × 0.3–0.5 mm, minute, whitish-green, pilose, attenuate, located near the base and opposite. Flowers pentamerous rarely tetramerous, descending to pendulous; calyx pilose, 5.1–6.0 × 4.5–5.0 mm, green; hypanthium obconic, 2.7–3.8 × 4.6–5.0 mm, obscurely 5-ridged, truncate; limb open, 1.9 × 4.2–4.5 mm, erect; lobes 5, ovate-deltate, 1.4–1.8 × 1.8–2.1 mm, small, shortly acuminate, the sinuses acute. Corolla tubular, slightly narrowing distally, 2.9 cm long (including the lobes), thick-carnose, bistratose, pubescent and bluntly 5-angled along its length, pubescent in the internal apical half, 5 mm in diameter at the base, 4 mm in diameter at the throat, magenta, paler at the angles; lobes 5, narrowly linear-triangular, 14.0–15.7 × 2.3 mm, spreading, acuminate, slightly incurved, glabrous, channelled and subverrucose internally. Stamens 10, nearly equalling the corolla in overall length, each pair unequal, 3.1–3.3 cm long; filaments connate forming a tubular staminal tube, slightly dilated distally, 8.9–9.6 mm long, white with a touch of pink, pilose externally in the proximal half; anthers 2.3–2.7 cm long, thecae conspicuously granulose, 6.2–6.8 mm long, prognathous, each pair of thecae unequal, 4.8–5.0 mm long; tubules distinct, 1.8–2.2 cm long, slightly unequal, seemingly connate in almost its length, straight, glabrous, dehiscing by terminal pores, 0.6 × 0.3 mm. Style exserted, 2.9–3.4 cm long, sparsely pilose at the middle, pale green, stigma truncated. Fruits not seen.

Distribution and habitat.

Ceratostema limonensis is currently known only from the vicinity of Limón in southern Morona-Santiago Province, where it was found on the south-eastern slopes of the Ecuadorian Andes (Fig. 3). It has been observed at elevations between 1100 and 1400 m in habitats characterised by mature, lower montane forests (Fig. 4). These forests are part of the “Bosque siempreverde piemontano del sur de la Cordillera Oriental de los Andes” (code BsPn04), as defined by the Ecuadorian Ministry for Environment (Ministerio del Ambiente del Ecuador 2013). The forest at the type locality is multi-stratified with a semi-open forest canopy and few lower forest glades with a maximum height of ca. 20 m. The most dominant species are Alchornea grandis Benth., Ficus cf. trapezicola Dugand, Metteniusa tessmanniana (Sleumer) Sleumer and Otoba parvifolia (Markgr.) A.H.Gentry. Specimens of C. limonensis were found sympatric with other flora, such as Evodianthus funifer Lindm., Maxillaria mapiriensis (Kraenzl.) L.O.Williams, Ronnbergia campanulata Gilmartin & H.Luther and Stenospermation sp.

Figure 3.

Figure 3.

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Distribution of Ceratostema limonensis in the south-eastern region of Ecuador.

Etymology.

This species is named after the Municipality of Limón in the Province of Morona-Santiago, the township near to which the new species was discovered.

Conservation status.

The very limited distribution of Ceratostema limonensis suggests that this species is a narrow endemic to south-eastern Ecuador. Its low abundance, combined with its restricted occurrence, does not allow for assessment of the AOO or EOO at this stage. Conservation threats to this species are excessive farming and mining activities which already occurring near the type locality. Due to the absence of more known specimens belonging to this species, a tentative classification of Data Deficient (DD) has been given (IUCN 2024).

Taxonomic discussion.

Ceratostema limonensis is primarily distinguished from all other species within the genus Ceratostema by its subflexuous young branches and axillary inflorescences, characterised by an entire, subterete rachis and pedicel. The inflorescences likely produce a second successive flower, a trait observed in related species, such as C. alexportillae A.Doucette, H.Medina & J.Portilla, C. gualaquizensis M.M.Jiménez & H.Garzón, C. loucianae Cornejo, G.Tello & Luteyn and C. rauhii Luteyn; however, further data on the floral production of this new species are necessary. These species also share several morphological features, including an epiphytic, pendant growth habit; spirally arranged, subsessile, plinerved, acuminate leaves; subsessile inflorescences with minute floral bracts and bracteoles; dark-coloured corollas with lobes that are narrowly triangular to narrowly linear-triangular, acuminate and recurved; and an exserted style (Luteyn 1992; Cornejo and Luteyn 2024; Cornejo et al. 2024; Jiménez et al. 2024b; Doucette et al. 2025a).

Amongst these, Ceratostema limonensis most closely resembles C. gualaquizensis, as both possess non-amplexicaul, plinerved leaves folded at the base to enclose magenta flowers. Their hypanthia are similarly obconic and subtly 5-ridged, adorned with acuminate calyx lobes. However, they differ in the characteristics set out in the diagnosis and, in addition, in that C. limonensis has foliar veins raised adaxially, whereas in C. gualaquizensis, the veins are impressed in the proximal 4 mm. Additionally, the corolla lobes of C. limonensis comprise approximately 50% of the total corolla length, compared to about 30% in C. gualaquizensis. C. limonensis also shares similarities with C. alexportillae, lianoid branches and notably convex, lanceolate leaves folded at the base (Jiménez et al. 2024b; Doucette et al. 2025a). A comparative summary of morphological differences amongst these species is provided in Table 1.

Table 1.

Morphological comparison of Ceratostema limonensis and similar species. References taken from: (1) presented herein, (2) Jiménez et al. (2024b) and Cornejo and Luteyn (2024); (3) Doucette et al. (2025a); (4) Jiménez et al. (2024b); (5) Luteyn (1992).

Characters C. limonensis (1) C. gualaquizensis (2) C. alexportillae (3) C. loucianae (4) C. rauhii (5)
Indumentum on vegetative and floral parts Pilose, pubescent on the corolla Puberulous Glabrous Villose Puberulous
Branches Lianoid Filiform Lianoid Long-filiform Long-filiform
Leaf blades
Shape Lanceolate, convex with basal margins folded Elliptic, convex with basal margins folded Cordate, convex with basal margins folded Ovate, flat-canaliculate Narrowly ovate to linear-ovate, flat
Length (cm) 5.5–7.8 3.9–6.0 4.8–7.2 2.5–6.0 1.5–3.5
Width (cm) 1.8–2.0 2.2–2.5 Ca. 0.2 1.5–3.3 0.5–0.7
Venation 5–7-plinerved 5–7-plinerved Obscure 5–7-plinerved 3–5-plinerved
Base Cordate to obtuse, non-amplexicaul Obtuse, non-amplexicaul Cordate, amplexicaul Subcordate to truncate, non-amplexicaul Obtuse, non-amplexicaul
Rachis
Shape Subterete Obconical Obconical Obconical Not available
Length (mm) 4.8–5.1 Ca. 1.0 Ca. 3.8 Ca. 1.0 Not available
Pedicel
Shape Subclavate Subterete Terete-conical Subterete Terete
Length (mm) 5.1–5.5 5.0–6.0 4.9–5.0 6.0–11.0 Ca. 2.0–3.0
Hypanthium
Shape Obconical, obscurely 5-ridged Obconical, obscurely 5-ridged Broad obconical, 5-carinate Obconical, subterete to bluntly 5-costate Cylindrical to obconical, terete
Length (mm) 2.7–3.8 3.9–4.2 3.2–3.7 2.9–3.5 Ca. 2.0
Calyx lobes
Shape Ovate–deltate Narrowly lanceolate Broadly deltoid Broadly deltoid Narrowly triangular
Length (mm) 1.4–1.8 3.4–4.7 Ca. 1.0 0.7–0.8 3.5–4.5
Apex Shortly-acuminate Acuminate Apiculate Apiculate Long-acuminate
Sinuses Acute Acute Obtuse Rounded Acute
Corolla
Colour Magenta, paler at the angles Magenta Red-purple Red and black Bright red
Shape Tubular, bluntly 5-angled Tubular, obscurely to bluntly 5-angled Tubular, pentagonal Cylindrical-urceolate to tubular, bluntly 5-angled Tubular, bluntly 5-angled
Length (mm) Ca. 29.0 45.0–47.0 Ca. 29.0 18.0–28.0 28.0–30.0
Corolla lobes
Colour Magenta Magenta and black Purple-black Black Bright red
Shape Narrowly linear-triangular Narrowly triangular Narrowly triangular Narrowly linear-triangular Narrowly triangular
Length (mm) 14.0–15.7 12.1–17.0 Ca. 12.5 11.5–17.8 3.5–4.5
Apex Acuminate Acuminate Acute Acuminate Long-acuminate
Stamens
Length (cm) 3.1–3.3 4.4–4.6 2.3–2.4 2.6–3.1 Ca. 2.8
Filaments texture Pilose Glabrous Sparsely pubescent Glabrous Glabrous
Thecae length (mm) 6.2–6.8 Ca. 5.1 mm 3.8–4.2 6.0–10.0 Ca. 4.0
Tubules shape Straight Straight Sinuous Straight Straight
Tubules length (cm) 1.8–2.2 3.4–3.5 Ca. 1.6 1.2–1.5 Ca. 1.8
Style
Length (cm) 2.9–3.4 4.8–5.1 Ca. 3.0 3.3–3.4 2.8–3.0
Texture Sparsely pilose Glabrous Glabrous Glabrous Glabrous
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Supplementary Material

XML Treatment for Ceratostema limonensis

Acknowledgements

We would like to thank the herbarium staff of the Universidad Técnica Particular de Loja Herbarium (HUTPL) for their support in developing this research. We are also thankful to the Ministerio del Ambiente, Agua y Transición Ecológica (MAATE) of Ecuador for granting the research permit MAATE-DBI-CM-2022-0248 to carry out all necessary collections. The authors also acknowledge Alan Elliott for his input and suggestions in improving this manuscript.

Citation

Jiménez MM, Iturralde GA, del Pozo DG, Lapo-González N, Kuethe JR, Garzón-Suárez HX (2025) Ceratostema limonensis (Ericaceae), a new species from the Province of Morona-Santiago, Ecuador. PhytoKeys 263: 37–47. https://doi.org/10.3897/phytokeys.263.159645

Funding Statement

Universidad de Las Américas, grant No. 527.A.XV.24.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of AI

No use of AI was reported.

Funding

MMJ and GAI thank the Universidad de Las Américas (UDLA) for funding orchid research in Ecuador, grant No. 527.A.XV.24.

Author contributions

Resources: NLG, HXGS. Writing – original draft: MMJ, GAI, DGP. Writing – review and editing: GAI, JRK.

Author ORCIDs

Marco M. Jiménez https://orcid.org/0000-0002-9502-5651

Gabriel A. Iturralde https://orcid.org/0000-0003-2456-0929

Diego Gutiérrez del Pozo https://orcid.org/0000-0003-3611-7932

Nadia Lapo-González https://orcid.org/0009-0001-8863-1557

J. R. Kuethe https://orcid.org/0000-0001-5390-9654

Henry X. Garzón-Suárez https://orcid.org/0000-0003-4049-1652

Data availability

All of the data that support the findings of this study are available in the main text.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

XML Treatment for Ceratostema limonensis
phytokeys.263.159645-treatment1.xml (40.8KB, xml)

Data Availability Statement

All of the data that support the findings of this study are available in the main text.

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