Research Article |
Corresponding author: Laura Lagomarsino ( lagomarsino.l@gmail.com ) Academic editor: Clifford Morden
© 2016 Laura Lagomarsino, Daniel Santamaría-Aguilar.
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:
Lagomarsino LP, Santamaría-Aguilar D (2015) Two new species of Siphocampylus (Campanulaceae, Lobelioideae) from the Central Andes. PhytoKeys 58: 105-117. https://doi.org/10.3897/phytokeys.58.6973
|
Two species of Siphocampylus (Campanulaceae: Lobelioideae) from the Central Andes of Peru and Bolivia are described, illustrated, and discussed with reference to related species. One species, Siphocampylus antonellii, is endemic to high elevation grasslands of Calca, Peru, while the second, S. siberiensis, is endemic to cloud forests of Cochabamba, Bolivia. Both species are robust shrubs that produce tubular pink flowers that are likely pollinated by hummingbirds.
Andean biodiversity hotspot, Asterales , Bolivia, centropogonid clade, Peru, South America, taxonomy
While the subfamily Lobelioideae Burnett of Campanulaceae Juss. is cosmopolitan in its distribution, more than half of its ~1200 species are restricted to the Neotropics (
Here we describe two new species of hummingbird-pollinated Siphocampylus. Type specimens for both species were included in the most recent molecular phylogeny of the centropogonid clade, which is based on five plastid markers and includes relatively dense taxon sampling that spans all recognized taxonomic divisions, geographical occurrences, and morphological variation within Neotropical Lobelioideae (
Field collections focused on Campanulaceae were conducted in Bolivia in November–December, 2011 and in Peru in November–December, 2012. Species identification and description of new species resulting from this fieldwork utilized many taxonomic references (
Siphocampylus antonellii is similar to S. elfriedii, but differs in its smaller, linear-oblanceolate leaves, ventral corolla lobe >1.3 cm long, and pleasant, lemon-like odor emitted from living plants.
Peru. Cusco: Calca, Lares, Calle entre Amaparaes y Suyo, Arriba de Amaparaes, 12°58'902"S, 077°50'W, 3799 m, 10 December 2012 (fl), L. Lagomarsino, D. Santamaría, J. Wells, F. Farro 400 (holotype: A!; isotypes:
Shrub 1.5 m tall, branching 20 cm above the base, with soft wood; branches 0.2–0.6 cm in diameter, solid and fistulose, light brown to reddish purple in living material, glabrescent or white-tomentose; internodes 0.2–1.0 cm long; latex white. Leaves spirally arranged, distributed evenly along the branches, producing lemon-like odor; petiole 0.1–0.5 cm long, sometimes subsessile, villous, the trichomes whitish, adaxially canaliculate, abaxially rounded to triangular; lamina 3.8–5.2 × 0.3–0.55 cm, linear-oblanceolate, not rugose, appearing glabrous but densely pubescent with diminutive, whitish, stellate to echinoid trichomes; base attenuate to decurrent, sometimes with uneven sides; apex acuminate; margin sinuate, subentire, or diminutively dentate, 11–25 teeth per side, rounded to uncinate, sometimes appearing as a glandular callosity; venation reticulate, with 5–10 pairs of lateral nerves, ascending, impressed or indistinct adaxially, flat abaxially. Flowers solitary, axillary, generally towards the apex of branch; pedicel 3.0–5.0 cm long, straight for almost the entire length, but curved below the hypanthium, cylindrical or flattened, densely pubescent, bibracteolate; calyx lobes 5, 0.7–1.6 × 0.1–0.12 cm, linear-oblanceolate, margins diminutively dentate with 3–5 teeth per side, densely pubescent on both surfaces, straight, the apex acuminate; corolla (2.8–) 3.9–5.1 cm long, tube pink with yellow to light green lines parallel to the lobes, lobes light green-yellow externally, light yellow to cream colored inside, completely pubescent externally, pubescent internally with stellate to echinoid trichomes; tube 2.7–3.5 × 0.4–0.6 cm, constricted at the base and widening distally, straight at anthesis; corolla lobes 5, lanceolate to narrowly triangular, apex acute to acuminate, the two dorsal lobes 1.0–1.6 cm long, the two lateral lobes 1.3–1.5 cm long, the ventral lobe 1.4–1.9 cm long; staminal tube 3.5–4.5 × 0.1 cm, straight, glabrous, cream-colored to light green in living material, exserted between the two dorsal lobes; anther tube 0.5–0.7 × 0.2–0.21 cm, dark gray, glabrous, ventral anthers 0.4–0.6 cm long, penicillate at the apex, the trichomes white or yellowish gold, dorsal anther 0.45–0.6 cm long, glabrous. Fruits not seen.
Siphocampylus antonellii is endemic to Peru, where it grows on rocky slopes in puna habitat at ~3800 m in elevation. It is only known from the type collection.
Individuals were collected in flower in December; the rest of the phenology of this species remains unknown.
It is an honor to name this attractive species for Dr. Alexandre Antonelli (1978–), a biogeographer and phylogeneticist at the University of Gothenburg. Antonelli has made many important contributions to our understanding of Neotropical biodiversity through space and time, and to the evolution of various taxa, including Lobelioideae. His efforts in the latter brought the second author to the field in Costa Rica in 2005 in search of Campanulaceae, and helped to inspire the first author to study Centropogon, Siphocampylus, and Burmeistera.
Siphocampylus antonellii is endemic to a narrow stretch of high-elevation grassland (puna) in Calca, Peru, where it is locally abundant. Only a single population of this species is known, from which the type collection was made. Due to its small area of occurrence and the threat of future deforestation in its habitat, we tentatively consider this species to be Vulnerable (
Siphocampylus antonellii is most similar to S. elfriedii E. Wimm. (Fig.
Differences between Siphocampylus antonellii, S. elfriedii, and S. parvifolius.
S. antonellii | S. elfriedii | S. parvifolius | |
---|---|---|---|
Leaf shape | Linear-oblanceolate | Lanceolate | Lanceolate to oblanceolate |
Leaf size | 3.8–5.2 × 0.3–0.55 cm | 5.0–7.4 × 1.1–1.9 cm | 2.0 × 0.5 cm |
Leaf margin | Margin sinuate, subentire, or diminutively dentate | Denticulate | Weakly crenulate |
Sepal length | 0.7–1.6 cm | 0.7–1.4 cm | 0.3 cm long |
Sepal margin | Diminutively dentate | Denticulate | Entire or diminutively denticulate |
Corolla length | (2.8–) 3.9–5.1 cm | 3.9–4.6 cm | 3.4 cm |
Corolla indument | Pubescent | Pubescent | Glabrous |
Dorsal corolla lobe length | 1.0–1.6 cm | 0.9–1.2 cm | 1.1–1.4 cm |
Ventral corolla lobe length | 1.4–1.9 cm | 0.5–1 cm | 0.9–1.2 cm |
Reference sheets | L. Lagomarsino 400 (A) |
L. Lagomarsino 387 ( |
R. D. Metcalf 30469 (A) |
This new species would be placed near S. nobilis E. Wimm. and S. rosmarinifolius G. Don in the dichotomous key of genus in
Siphocampylus antonellii. A Flowering branch B Leaf, abaxially, including detail of leaf margin and stellate hairs that cover surface C Staminate-phase flower, including bibracteolate pedicel, with detail of sepal and stellate hairs that cover the outer corolla surface D Longitudinal section of a pistillate-phase flower, showing the insertion of staminal tube to corolla, style and stigma as situated relative to the stamens, and bilocular ovary with axile placentation E Corolla lobe detail F Detail of anther tube, including apical hairs on ventral anthers, and stigma. Drawing by Bobbi Angel from the type.
Siphocampylus antonellii. A Habit and high-elevation grassland (puna) habitat B Flowering branch C Cross-section of stem showing woody habit D Adaxial leaf surface E Abaxial leaf surface F Lateral view of flower in staminate phase G Anterior view of a flower, showing corolla aperture. All photos of the type collection, taken in the field by L. Lagomarsino.
Molecular phylogenetic analysis places S. antonellii in a well-supported clade that includes S. actinotrix E. Wimm. (Fig.
Closest relatives of Siphocampylus antonellii, based on molecular phylogeny of
Siphocampylus siberiensis is similar to S. boliviensis, but with a corolla tube lacking a constriction at its base and a hemispherical hypanthium.
Bolivia. Cochabamba: Carrasco, en la entrada para Sunchal, cerca al rótulo “Unidad Educativa Manuela Gandarillas”, 17°47'267"S, 064°47'669"W, 2668 m, 18 December 2011 (fl & fr), L. Lagomarsino, D. Santamaría & J. M. Mendoza 241 (holotype: A!; isotype:
Multi-stemmed shrub 3–4 m tall, branched, all branches arising from a single point at ground level, with soft wood, the bark suberose; branches 0.4–0.6 cm in diameter, fistulose, the youngest parts purple, brown when mature (greyish to whitish in dry material), glabrescent to tomentose; internodes 0.9–1.7 cm long; latex white. Leaves spirally arranged and generally clustered at the apex of branches, leaving prominent leaf scar after falling; petiole 0.3–0.7 cm long, glabrescent to tomentose with whitish trichomes, winged, adaxially canaliculate, abaxially more or less triangular with two ribs; leaf blade 10.5–19.5 × 2.9–4.6 cm, oblanceolate, adaxially tomentose and abaxially densely pubescent, the pubescence principally on the veins, trichomes simple, the base decurrent; apex acuminate; margin doubly dentate and ciliate, 75–95+ teeth per side, the teeth triangular; venation reticulate with 16–21 pairs of lateral nerves, lightly ascendant, impressed adaxially and elevated abaxially. Flowers solitary, axillary, generally towards branch apex; pedicel 4.5–9.4 cm long, straight, cylindrical, densely pubescent; bracteoles absent; hypanthium 0.5–0.8 × 0.3–0.4 cm, hemispherical, tomentose; calyx lobes 5, (0.8–) 1.0–1.4 × (0.2–) 0.3 cm, narrowly triangular, the margins ciliate, entire, pubescent on both surfaces, erect or recurved, the apex acuminate; corolla 3.5–3.8 cm long, completely pink, diminutively pubescent on both surfaces; corolla tube 0.9–2.2 × 0.7–0.8 cm, cylindrical for its entire length, a little wider apically than basally, straight at anthesis; corolla lobes 5, narrowly triangular, slightly falcate, the margins ciliate, the apex acute to acuminate, the two dorsal lobes 1.5 cm long, the two lateral lobes 1.4 cm long, the ventral lobe 1.4 cm long, staminal tube 3.0–3.8 × 0.1–0.2 cm, straight, sparsely pubescent, pink, exserted between the two dorsal lobes; anther tube 0.8–0.9 × 0.2–0.3 cm, gray in living material, glabrous except in the sutures between anthers, which are densely pubescent, the trichomes white, ventral anthers 0.6–0.8 cm long, penicillate at the apex, the trichomes white, the dorsal anthers 0.7–1.0 cm long, penicillate at the apex, the trichomes white. Fruit capsule, 5.0 × 1.0–1.2 cm, ca. 15-lobed, with external ridges, the calyx persistent; seeds not seen.
Siphocampylus siberiensis is endemic to Bolivia, where it has been collected at the edge of the road at ca. 2700–2900 m in elevation in cloud forest.
Individuals were collected in flower and fruit in December and in flower only in April; the rest of the phenology of this species remains unknown.
The specific epithet of this species refers to the type locality, the Serranía de Siberia, a mountain range at the limit between the Cochabamba and Santa Cruz departments in Bolivia.
Siphocampylus siberiensis is known only from a single population in Serranía de Siberia in central Bolivia; this population is represented by the two cited collections. This species appears to be locally rare, and only one individual was encountered during our fieldwork. Due to its small area of occurrence and the threat of future deforestation in its habitat, we tentatively consider this species to be Vulnerable (
Siphocampylus siberiensis can be recognized by its shrubby habit with multiple stems arising from a single point; leaves aggregated at the apex of branches (Fig.
Siphocampylus siberiensis. A Flowering branch, showing the persistent leaf scars and developmental procession of distal flower at anthesis to basal capsular fruit B Flower in pistillate phase, including detail of sepal with pubescence C Corolla lobe detail, including marginal pubescence D Longitudinal section of a pistillate phase flower, showing the insertion of staminal tube to corolla, style and stigma as situated relative to the stamens, and bilocular ovary with axile placentation E Anther tube in staminate-phase flower F Capsule with lobes and ridges. Drawing by Bobbi Angel from the type.
Siphocampylus siberiensis. A Habit B Detail of young stem C Detail of leaf margin and venation on abaxial leaf surface D Flower bud E Capsule F Lateral view of flower in pistillate phase G Anterior view of corolla, showing corolla aperture H Abaxial leaf surface I Adaxial leaf surface. All photos of the type collection, taken in the field by L. Lagomarsino; D. Santamaría-Aguilar is shown collecting the type in A.
Molecular phylogenetic analysis places S. siberiensis in a clade that includes S. tunarensis Zahlbr., S. tunicatus Zahlbr., and S. umbellatus (Kunth) G. Don; this clade is closely related to S. boliviensis Zahlbr. and S. sparsipilus E. Wimm. (
Closest relatives of Siphocampylus siberiensis, based on molecular phylogeny of
Even though they are not the most closely related species, the pink, narrow flowers of S. siberiensis most closely resemble those of S. boliviensis and S. sparsipilus. However, the latter two species can be easily distinguished by their corollas that are constricted at the base (vs. not constricted) and much rounder hypanthium (vs. flattened at top). The other species in the immediate clade that includes S. siberiensis differ in their dull reddish (S. tunarensis [Fig.
The species that is most superficially similar to S. siberiensis, S. boliviensis, is placed in the same couplet as S. macrostemon A. DC. in the dichotomous key to the members of the genus in
The measurements of the calyx and corolla in parentheses correspond to E. Fernández et al. 3583 (
Bolivia: Cochabamba, Carrasco, Siberia, 17°48'11"S, 064°46'12"W, 2900 m, 16 April 2005 (fl), E. Fernández et al. 3583 (
We would like to thank the directors and curatorial staff at the following herbaria for permitting the use their collections: A,