Corresponding author: Eric. J. Tepe (
Academic editor: M. A. Jaramillo
We describe
Describimos la nueva especie
Tepe EJ, Rodríguez-Castañeda G, Glassmire AE, Dyer LA (2014)
Documenting plant and phytochemical diversity is recognized as an important component of understanding interactions among plants, herbivores and predators, and for understanding the evolution of biodiversity (
Morphological data and locality information were taken from the personal collections of EJT, GRC, and AEG, the ecological studies mentioned above, and from specimens sent to EJT for identification. Although the new species occurs in Ecuador and Peru, the distribution model was restricted to Ecuador since the majority of collection data is from the northeast and southeast Andean slopes of Ecuador. Collection density was very high in Napo Province, Ecuador (where much of the project activity was focused) relative to other parts of the species range, and to avoid biased results due to this imbalance, locations were filtered so that the model was based on an even sample from across the species’ range. The distribution model was calculated using the maximum entropy method (Maxent software package;
Study sites were located in the northeastern Andes of Ecuador in the provinces of Napo and Sucumbíos along an elevational transect from Amazonian lowland rain forest (200 m) to Andean páramo (3200 m). Mean temperature along this gradient drops linearly with elevation at a rate of approximately -0.5°C per 200 m elevation gain, and precipitation ranges from 3900–4500 mm yr-1 (
To document trophic interactions associated with the new
Piperi imperiali
Shrub to small tree, 1.5–15 m tall, moderately branched; trunk of flowering individuals 5–8 cm d.b.h.; some individuals with prop roots. Stems glabrous, the nodes moderately to densely tuberculate, density of tubercules increasing distally along each internode, the internodes 6.5–9.6 cm long and 0.5–1.1 cm in diam.; shoot apex emerging from the sheathing leaf base. Prophyll minute and hidden by the sheathing leaf base. Leaves more or less distichous on flowering branches, with petioles 6–14 × 0.7–1 cm at flowering nodes, vaginate, and with persistent margins, the margins extending to the leaf base or projecting 1–10 mm beyond the insertion of lower leaf lobe, glabrous, sparsely to moderately tuberculate; laminae 25–50 × 15–42 cm, broadly ovate, the apex obtuse, rounded, the base oblique, cordate, the lobes equal or more commonly somewhat unequal, extending 2–8 cm below the petiole attachment, divergent and never overlapping the petiole, the sinus open and the apices of the lobes nearer the leaf margin than the petiole, the sides of the lamina arising 7–8 mm apart on the petiole, the lamina drying thickly chartaceous, densely glandular-dotted (usually blackish on dried specimens, 3–5 per mm2, increasing in density along the leaf margin), glabrous above, glabrous to sparsely pubescent on the lamina below and moderately to densely pubescent on the veins below with much shorter trichomes (< 0.2 mm long; these more conspicuous on the secondary and tertiary veins, often lacking on the midvein), the 4–6 pairs of major secondary veins arising from the lower 2/3 of the midvein, arcuate-ascending, primary–tertiary veins somewhat impressed above, prominent below. Inflorescences free from the leaf base of the same node, pendulous, 40–72 × 0.4–0.8 cm in flower and 0.8–1.1 cm in diameter in fruit, the flowers densely crowded and +/- banded; peduncles 2.5–6 × 0.25–0.5 cm, white to green in fruit, glabrous; floral bracts 0.7–1.0 mm broad, triangular to triangular-rounded, nearly glabrous with upper margin white ciliate to densely pubescent throughout; stamens 4 per flower, the stamens maturing asynchronously such that only one or two are apparently visible per flower at one time, white, the anthers 1–1.5 mm long, the filaments clavate, the connective somewhat broadened between thecae and these divergent at ca. 45°, dehiscing laterally; fruits rounded or rectangular from above by compression, 1.5–2 × 1.2–2 mm, the apex truncate, glabrous, gland-dotted, stigma lobes 3(–4), 0.5–0.8 mm long, sessile or on a very short style, caducous; seeds rounded-square, flattened, 1.5–2 mm.
Distribution of
Flowering specimens have been collected in Jan–Mar and Dec; fruiting specimens have been collected in Mar, Jul, and Sep.
There are three basic patterns of sympodial stem tip and leaf primordial (STLP) enclosure and protection in
According to the IUCN Red List Categories (
Ecological niche model performance was high (avg. train AUC: 0.99; avg. test AUC: 0.99; max. probability range: 0–0.86). Results predict that suitable habitat for
Plot data resulted in herbivory and parasitism rates, and measures of herbivore and parasitoid richness. Mean herbivory rates for the plant are close to 20%, and the only herbivores that are regularly found on
Project collaborators have reared nearly 30,000 caterpillars and parasitoids from over 100
The parasitism rate for all of the
Thus, to date,
A colony of
Empirical evidence has been accumulating that supports the hypothesis that plant biochemistry helps drive the evolution of biodiversity, not only among herbivores that feed directly on plants, but among higher trophic levels as well (
Our description of
We thank the Museo Ecuatoriano de Ciencias Naturales, the Ministerio del Ambiente, and Tom Walla for assistance with permits in Ecuador, and Segundo Leiva for assistance with permits and logistics in Peru; Harold Greeney, José Simbaña, and the rest of the staff at the Yanayacu Biological Station; Elicio Tapia, Wilmer Simbaña, Luchito Alberto, Brittany Bernik, Heidi Connahs, Jessica Hogue, Annie Kalson, Lauren Loue, Maria Paz Moreno, Colin Morrison, Elia Pirtle, Aleksandra Pokrovskaya, Kevin Salls, Joe Sexton, Stephanie Short, Stephen Stern, and Toni Walters for invaluable assistance in the field; and we thank Jürgen Homeier, Nixon Cumbicus, and Malte Ebinghaus for additional collections of the new species. We also acknowledge Aparna Palmer and three anonymous reviewers who provided useful information and important comments on the manuscript, and Lauren Ginsberg for providing much-needed help with our Latin grammar. This work was supported by NSF grant DEB 1020509, DEB 0614883, and DEB 1145609 to LAD; EJTs fieldwork was funded by W.S. Turrell Herbarium (MU) Fund (#214) and the PBI: