Monograph |
Corresponding author: Nancy Hensold ( nhensold@fieldmuseum.org ) Academic editor: Reyjane Patricia Oliveira
© 2016 Nancy Hensold.
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:
Hensold N (2016) The Andean Paepalanthus pilosus complex (Eriocaulaceae): a revision with three new taxa. PhytoKeys 64: 1-57. https://doi.org/10.3897/phytokeys.64.6864
|
A herbarium-based revision is provided for Paepalanthus pilosus and allies, five commonly confused species of cushion plants native to Andean paramo. These are placed in the recircumscribed Paepalanthus subsect. Cryptanthella Suess. The group includes P. pilosus, P. dendroides, and P. lodiculoides. An additional two species and one variety are newly described: Paepalanthus caryonauta, P. huancabambensis, and P. pilosus var. leoniae. The latter two are Peruvian endemics, while P. caryonauta is known from four countries, and has long been confused with other species. An additional, possibly undescribed taxon is noted from the Serrania de Perijá, Colombia. Five new synonyms and three lectotypes are proposed, and the common misapplication of some names is noted. Within the P. pilosus complex, species differences were found in timing of peduncle elongation, sex ratio, and leaf, perianth, diaspore and nectary morphology. Ecological differences are suggested by specimen data and a review of ecological literature. Descriptions, photographs and maps are provided for all species, as is a key to the groups of eriocaulaceous cushion plants from Andean South America.
Andes, cushion plants, diaspores, leaf anatomy, nectaries, new species, paramo, taxonomy
The Eriocaulaceae (Liliopsida: Poales) are a family of 10 genera and about 1200 species, about 800 of which are Neotropical (
The Paepalanthus pilosus complex includes seven of the currently recognized Andean species: P. barkleyi Moldenke, P. dendroides (Kunth) Kunth, P. dennisii Moldenke, P. karstenii Ruhland, P. lodiculoides Moldenke, P. pilosus (Kunth) Kunth, and P. schultesii Moldenke. Members of this complex are characteristic elements of wet peaty sites in climatically humid páramo and subparamo from Costa Rica to Bolivia. All exhibit the cushion plant growth form, or “pulviniform” habit, found among diverse flowering plant families of high-elevation páramo (
These species are also easy to confuse with other Andean cushion plant species, including Eriocaulon microcephalum Kunth and the three Andean taxa of Paepalanthus subsect. Dichocladus Ruhland (P. dichotomus var. glabrescens Moldenke, P. ferreyrae Moldenke, and P. muscosus Körn.), but can be distinguished by floral and seed morphology and other microcharacters, as detailed below. Some of the 16 species of Paepalanthus subg. Platycaulon Mart. endemic to the Andes also proliferate from the base and have been described as cushion plants (“polsterförmig”;
This study was originally focused on resolving taxonomy of the Peruvian members of the P. pilosus complex, but later expanded to a herbarium study of all Andean material at hand. While I attempted to describe all material available to me, including that of Colombia and Venezuela, a detailed study of North Andean material was outside the scope of this work. Both P. dendroides and P. pilosus exhibit more complex variation over their range than is found in Peru and Ecuador, and the descriptions may not entirely reflect populations of northern South America. Results from more intensive field or molecular studies may add much to our understanding.
Specimens were examined from the herbaria F,
Specimen localities were interpreted according to the label description and in some cases, published collecting records, and mapped manually using the Google “My Maps” application. Coordinates of the manually placed markers were retrieved as KML data. Final distribution maps were generated using ArcGIS. Estimated coordinates used for mapping are included in the Suppl. material
The key characters used to differentiate Paepalanthus subsect. Dichocladus from P. subsect. Cryptanthella in the key were based on examination of the following vouchers. Paepalanthus dichotomus var. glabrescens: Peru. Amazonas: Chachapoyas, A. Sagástegui 7454 (F), H. van der Werff 14940 & 16912 (F), Wurdack 1388 (isotype, F). Paepalanthus ferreyrae: Peru. Cajamarca: Cutervo, J. Mostacero 1594 (F), Peru. Amazonas: Bongará, J. Wurdack 1081 (F), I. Sánchez Vega 10020 (F). Paepalanthus muscosus: Colombia. Norte de Santander, Linden 1330 (isotype, F); Venezuela. Táchira, Maas & Tillett 5282 (F).
Leaf anatomy of the P. pilosus complex was studied from median hand sections of dried herbarium specimens. Leaves were boiled, stored in 70% alcohol, then rinsed, hand-sectioned in water, and mounted in 50% glycerine. Anatomical vouchers are as follows. Paepalanthus caryonauta Hensold: Barclay 5176 (F), Boyle 4219 (F), Fuentes 13574 (F), Monteagudo 16143 (F), Valenzuela 8117 (F); P. dendroides: Leon 2243 & 2683 (F), Vásquez 29038 (F); P. huancabambensis Hensold: Sagástegui 16799 (F); P. pilosus var. pilosus: Barbour 3427 (F), Jørgensen 1817 (F), 2209 (F); P. pilosus var. leoniae Hensold: León 1597 (
Floral measurements were made by ocular micrometer in a stereomicroscope. Except for the Colombian, Venezuelan and Central American material of P. dendroides and P. pilosus, flower samples of all available specimens were measured. Photos of dry flowers, seeds and some capitula were taken with a Dino-Lite AM-413T USB Digital Microscope (Figs
Conservation status could be assessed with confidence only for taxa of very broad and very narrow distribution. Confident evaluation according to the
All species in the group are perennials with a compact, flat to rounded (pulviniform) cushion habit, formed by short densely branched sympodial stems covered with congested subulate leaves about 1−2 cm long. Inflorescences are solitary and terminal on young shoots, but soon overtopped by erect lateral shoots, and then appear either axillary or borne between two dichotomous branchlets. The cushions may reach up to 1 m in diameter (P. caryonauta) and 10 cm high (P. pilosus), with a hemispherical shape (Fig.
Cushion habit of Paepalanthus pilosus var. pilosus in Venezuela, November 2012. A P. pilosus cushions with bunchgrasses, Páramo Batallón B Individual cushion, with graminoids emerging, Páramo Los Conejos. Photos by Serge Aubert, Station Alpine Joseph Fourier, France. Used by permission (www.cushionplants.eu).
Leaf morphology is similar among species, with subtle differences found in color, thickening, and pubescence. Paepalanthus huancabambensis is distinguished by a deep blue-green leaf color, while P. dendroides has leaves distinctly pale green in drying. In other species, leaf color is variable. The upper leaf surface is persistently pubescent in P. huancabambensis and in P. dendroides of Huánuco and Cuzco. In P. pilosus of Peru and Ecuador, appressed pubescence commonly occurs on the upper surface near the apex, while robust scattered cilia extending to the distal margin are typical of plants from the northern part of the range. Paepalanthus caryonauta has a rounded leaf apex (Fig.
Capitula and leaf tips of Paepalanthus pilosus and relatives A−E Fruiting capitula with diaspores; sepals detaching (A, D) or intact (C, E) A P. dendroides B−C P. caryonauta DP. pilosus var. pilosusE P. lodiculoides F−JP. pilosus var. pilosus, involucral bract variation. Arrows = tips of the torn peduncle sheath. (ADavidse 28991BBarclay 5176, Ecuador CDudley 11060, Peru DCuatrecasas 25574EØllgaard 9717FCuatrecasas 25882GCuatrecasas 5553HØllgaard 9557ICano 16840JJørgensen 2366.)
The 13 specimens sampled represent all taxa of the P. pilosus complex except P. lodiculoides. Their anatomy may be characterized as follows: Epidermal layer one cell thick of large rounded cells, with those of the upper epidermis slightly larger than the lower epidermis; leaf margin rounded, with smaller usually thicker-walled cells. Stomata abaxial only. In P. dendroides, the epidermal cells larger and mostly thinner-walled than in the other samples, partly collapsed and deformed in section. In other sampled taxa, the outer epidermal wall thickened, and sometimes heavily cutinized on one or both surfaces. Hypodermis absent. Mesophyll of dense to moderately loose short-armed chlorenchyma, with an adaxial palisade layer sometimes discernible. Veins mostly 5–7. Vein buttresses (bundle sheath extensions) commonly absent in median section (Fig.
Some unusual anatomical features are present which may relate to the daily freeze-thaw cycles of high elevation paramo. First, the mesophyll often detaches cleanly from the epidermis in an intact layer enclosing the vascular bundles. This was conspicuous in the thick leaves of P. caryonauta and observed to some extent in all sampled taxa except P. huancabambensis. It can be seen in dry broken leaves, as well as in hydrated sections. Separation of mesophyll from the epidermis is known from petioles and leaves of various frost-resistant species of other families in connection with extracellular ice formation (
Unusual deformation of the bundle sheaths is also observed in some species. In P. caryonauta, P. huancabambensis, and most P. pilosus, cells of the bundle sheath (“endodermis” sensu
Paepalanthus pilosus var. pilosus is distinguished, especially in Peru and Ecuador, by the capitula usually subsessile and partly contained within the peduncle sheaths at flowering time (Fig.
Most species have protogynous capitula, with pistillate flowers at the periphery, and staminate flowers in the center. The capitula are about 4-24-flowered and usually strongly determinate, with no central floral primordia found at the start of anthesis. In capitula with relatively more flowers (P. huancabambensis, some P. pilosus) the ratio of pistillate to staminate flowers tends to be greater (up to 3:1) whereas in smaller capitula it approaches equality. The related taxon, treated as “Paepalanthus sp. A” below, differs by the flowers more numerous (40 or more) and the pistillate and staminate disposed in alternating whorls, with staminate flowers sometimes found in the outer whorl (Fig.
In Paepalanthus and other genera of the Paepalanthoideae, modified style branches may function as nectaries (
In P. lodiculoides (Fig.
Flowers of Paepalanthus caryonauta (A−O) and suspected hybrid (P). A−G Pistillate flowers in anthesis A Involucral bract subtending flower B Whole flower C Sepal (abaxial) D Petal, adaxial E Petal, abaxial F Flower with one petal and two sepals removed G Gynoecial nectaries H−I Staminate flowers in anthesis H Floral bract I Flower J Staminate flower showing post-anthesis thickening, two anthers fallen K−L Mature diaspores M Diaspore with sepals removed, with thickened petals N−O Seeds P Mature flower of probable hybrid (Roldán 402), with seed. (A−I, NValenzuela 8117J−KDudley 11060L, OBoyle 4219MDudley 11194).
In P. dendroides, P. huancabambensis, and P. pilosus var. pilosus the basal half of the fruiting sepals thickens along the midvein at maturity and recurves hygroscopically upon drying, presumably pushing the detached corolla and fruit upward to the capitulum surface (Figs
The staminate flowers of P. caryonauta also incidentally thicken with age (Fig.
Smut fungus infection was observed in both P. dendroides (Nuñez 7773, Tupayachi 50) and P. pilosus var. pilosus (Sagástegui 12242). In these specimens the compact black spore masses swelling the ovary locules simulate mature fruit.
Paepalanthus pilosus and allies occur strictly in wet and very wet paramo and subparamo formations of the Andes, often associated with Sphagnum. Most taxa, except for P. dendroides, occur at about 3100-4000 m on wet but probably not inundated sites. Of these, P. pilosus and P. caryonauta are the most similar in habit, both forming dense mats or cushions 30 cm or more in diameter and reported as locally abundant. Paepalanthus pilosus is the more commonly collected species, noted for its colonization of disturbed sites, and often cited in ecological studies of paramo as “P. karstenii.” Further detail is found in the species discussions.
The principal distributions of P. pilosus and P. lodiculoides, on one hand, and P. caryonauta on the other, form an allopatric mosaic with respect to each other (Figs
Parallel disjunctions may be noted in other pulviniform Andean Eriocaulaceae. The Andean species of P. subsect. Dichocladus have a collective distribution similar to that of P. pilosus, occurring in the northern part of the eastern Cordillera of Colombia (P. muscosus), and in the Amotape-Huancabamba Zone (P. dichotomus var. glabrescens, P. ferreyrae). On the other hand, Eriocaulon microcephalum, which occurs in the Central Cordillera of Colombia and northern Ecuador, as well as in central Peru has an Andean distribution paralleling that of P. caryonauta, but also touching the edges of the Amotape-Huancabamba Zone. In Ecuador, it reaches to southern Azuay (specimen data from
Paepalanthus dendroides has a wide patchy range that overlaps that of both P. caryonauta and P. pilosus (Fig.
A number of suspected hybrids were detected in areas of sympatry. These include a sterile intermediate between P. dendroides and P. caryonauta in Pasco, Peru, discussed under P. dendroides. In addition, material morphologically intermediate between P. caryonauta and P. pilosus has been collected in Colombia in areas marginal to the distribution of both species. At the Paramo de Frontino in the western Cordillera, an apparently fertile intermediate plant was found sympatrically with normal P. pilosus, and a similar intermediate, with abortive flowers, was collected at the south end of the eastern Cordillera. Typical P. caryonauta is not known from either locality. (See P. caryonauta discussion.) Finally, as discussed under P. pilosus, long-term introgression with P. dendroides, or perhaps another local taxon, is suspected as a factor contributing to the unusually broad, and perhaps bimodal pattern of variation observed in P. pilosus in disturbed paramos near Bogotá, Colombia, but more study is needed.
A convenient alternative placement for these plants is available in the overlooked subsection Paepalanthus sect. Paepalanthus subsect. Cryptanthella Suess., described to accommodate P. kupperi Suess. (syn. of P. pilosus) from Costa Rica.
The species of P. subsect. Cryptanthella formally treated here are all Andean cushion plants. However, P. pilosus is very similar to a robust long-pedunculate taxon from páramo in the Serrania de Perijá, Colombia, discussed below under Paepalanthus species A. Further affinities of the group are not clear. It was not sampled in recent cladistic studies (
Due to the frequent confusion and misidentification of all the Andean cushion plant taxa of Eriocaulaceae, a key distinguishing major groups is provided.
1 | Roots white, spongy and compressible, easy to tear or cut, septate (with transverse partitions); leaves fenestrate at base (“windowed,” with transverse tissue partitions between the veins); leaf tips acuminose, membranous, pale or discolored; peduncles and outer involucral bracts glabrous; anthers deep black; petals with a black gland on inner apex | Eriocaulon microcephalum Kunth |
1' | Roots cream to brown, wiry, with persistent fibrous core, not easily broken by hand nor septate, nor leaves fenestrate; leaf tips thickened, rigid, rounded to aristate; peduncles and/or outer involucral bracts pubescent; anthers cream-colored; petals eglandular | 2 |
2 | Peduncle sheaths lacking or present, and then the tips ciliate around the whole mouth, sometimes glabrate with age, occasionally splitting with age; pistillate flowers with sepals linear-ligulate, the tips recurving or recoiling in fruit; petals tufted at upper margin only, not pilose on outer surface; seeds smooth and shiny even when wetted, pseudotrichomes absent. Plants of sandy and rocky soils of jalca or montane forest openings, ca. 2000–3000 m elevation | Paepalanthus subsect. Dichocladus (Andean taxa) |
2' | Peduncle sheaths always present, the tips papery-thin and scarious, often splitting deeply into two or three segments before anthesis, glabrous or minutely tufted at apex, never ciliate around mouth; pistillate flowers with sepals elliptic to spatulate, the tips not recoiling in fruit, and petals prominently pilose on outer and often inner surface; seeds with a whitish covering of matted or distinct pseudotrichomes after wetting. Plants of wet paramo or subparamo often associated with Sphagnum, ca. 2000–4000 m | Paepalanthus subsect. Cryptanthella |
1 | 1.7–6.0 mm long, less than 0.4 mm wide in the middle; peduncles glabrous at apex; floral sex ratio very variable, ranging from capitula with flowers all staminate to flowers mostly pistillate | 4. P. lodiculoides |
1' | Leaves (5.5–) 6-22 mm long, 0.6–2 mm wide in the middle; peduncles with a dense collar of hairs at the apex surrounding the base of the capitulum; sex ratio less variable, the capitula with the number of pistillate flowers subequaling to three times more than the staminate in number | 2 |
2 | Leaf apex narrowly obtuse to rounded, if acute, the tip curved under and not evident from above; leaves glabrous in the distal half or nearly so; sepals of the female flowers thickening uniformly at maturity, enclosing the fruit and dispersed with it | 1. P. caryonauta |
2' | Leaf apex acute to sharp-apiculate, never deflexed; leaves commonly appressed-pilose on upper surface near tip or ciliate, rarely glabrous; sepals of the female flowers thickening only in a narrow strap-like zone in the basal half of the midrib, and separating from fruit at dispersal (except P. pilosus var. leoniae, Peru, San Martín) | 3 |
3 | Peduncles mostly < 7 mm, usually shorter than the sheaths at time of anthesis, often elongating in fruit (rarely up to 50 mm at anthesis, Cano 16840, Piura); petals of the female flowers oblanceolate, mostly 2.2–6 times longer than wide; style base (below insertion of style branches) 0.5–1.05 mm long; nectaries dark, rigid | 4 |
3' | Peduncles 20–130 mm at anthesis; petals of the female flowers broadly spatulate, ca. 1.6–2.3 times longer than wide; style base less than 0.35 mm long; nectaries various | 5 |
4 | Fruiting sepals of the female flowers with basal half of midrib thickened, otherwise remaining chartaceous | 5a. P. pilosus var pilosus |
4' | Fruiting sepals of the female flowers uniformly thickened throughout and enclosing corolla and fruit at maturity. Plants of San Martín, Peru | 5b. P. pilosus var. leoniae |
5 | Leaves pale green; peduncle sheaths 11–23 mm, closely appressed, often hidden among the leaves; involucral bracts paler at least along the midvein; nectaries colorless to pinkish or light brown, those of the staminate flowers usually only half-equalling the corolla sinuses (except Vasquez 29038, Pasco) | 2. P. dendroides |
5' | Leaves dark blue-green; peduncle sheaths 25-30 mm, lax and open, much exsert from leaf mat; involucral bracts blackish brown throughout; nectaries dark brown, rigid, exsert in fruit, those of the staminate flowers equalling the corolla sinuses | 3. P. huancabambensis |
Paepalanthus subsect. Cryptanthella Suess., Bot. Jahrb. Syst. 72(2): 293. 1942.
Paepalanthus kupperi Suess.
Plants cespitose or pulviniform, forming densely branched clumps with erect terminal shoots ca. 1-4 cm; inflorescences solitary, terminal, but soon overtopped by one or two sympodial branches; peduncle sheaths scarious, swollen at apex, usually splitting into two or three triangular segments; involucral bracts pale greenish to gold or blackish-brown, pilose, not or barely surpassing flowers. Trichomes of the involucral bract and sepal apices subacute to rounded at apex, tuberculate; apical trichome tuft of bracts and sepals relatively short, surpassing perianth tip by less than 0.2 mm. Pistillate flowers peripheral, the outer subtended by the broad inner bracts of the involucre; staminate flowers central or rarely (P. lodiculoides) the whole capitulum staminate; the inner flowers subtended by linear receptacular bracts, these often with sub-cucullate tips and carinate-clasping bases; pistillate flowers pedicellate; the petals usually long pilose abaxially with tuberculate trichomes disposed in submarginal bands in the upper 1/2 to 2/3 of petal, and also in two dense tufts just inside the upper margin either side of the apex and securing the stigma, the petal tips not involute after anthesis. Gynoecium with stigmas simple; stigmatic nectaries colorless to reddish or dark brown, usually with a distinctly broadened large-papillate upper rim, the papillae in the basal two-thirds or more usually indistinct and scattered. Staminate flower corolla with anthophore usually at least half the total corolla length, membranous or fleshy, corolla tube with three subacute lobes, non-involute after anthesis; staminal filaments prominently fleshy below the corolla lobes and often (not always) adnate to corolla, flat and membranous distally, the tips sometimes turning red-brown with age; anthers whitish, exsert above the lobes, not retracted after anthesis, and often deciduous. Seeds with longitudinal rows of hygroscopic pseudotrichomes.
Cushion plants with linear glabrescent leaves 1–2 cm long, rounded at the tip, or if cuspidate, the tip strongly deflexed. Peduncles ca. 9–25 mm, solitary, terminal at initiation, sheaths lacerate at apex, eciliate. Pistillate flowers in outer whorl of capitulum, the sepals broad, persistent, together with the petals enclosing the mature fruit at dispersal, nectaries uniformly dark brown, rigid. Staminate flowers central, with sepals fused ¼–¾ of their length.
PERU. Cuzco: Dist. Huayopata, sector San Luis, bosque primario intervenido, 13°04'S, 72°23'W, ca. 3000–3500 m, 24 Nov 2006, L. Valenzuela et al. 8117 (holotype: F; isotypes:
Densely branched cushion plants, the cushions reported to reach one meter in diameter (Dudley 11194), and 15 cm high (Barclay 5176). Branchlets 1–5 cm, densely and uniformly leafy. Leaves linear-subulate, 10–20 mm long including the open basal sheath 3–4 mm long, 1.2–1.7 mm wide at midpoint, ca. 2.5 mm wide at ampliate base; apex narrowly obtuse to rounded, if sharp-cuspidate the tip deflexed downward and not evident in adaxial view; inconspicuously appressed-pilose (rarely ciliate) in juvenile state, glabrate at maturity except for the irregularly ciliate basal sheath; “dark green and glossy” when fresh (Dudley 11060), chartaceous to subcoriaceous, the adaxial surface smooth, the abaxial surface often with margins prominently thickened in older leaves and 1–3 veins more or less salient. Inflorescences solitary and terminal, with one to four produced in succession along a sympodially branched axis, the lower appearing axillary. Peduncles (7–) 9–25 (–35) mm long, ca. 3-costate, obscurely angled, pale, glabrous to obscurely appressed-puberulent, usually with a collar of longer silvery hairs at apex investing base of involucre. Peduncle sheaths (9–) 10–17 (–25) mm long, equaling or surpassing the leaf mat by up to 5 mm, the lamina (tip) of the sheath 3–4 mm long, inflated, scarious, somewhat cucullate, almost closed at the apex in bud before emergence of capitulum, tufted-ciliolate at apex when young, often glabrate, frequently splitting into 3 triangular segments with emergence of capitulum. Capitula 3–4 mm in diameter. Involucres subequaling flowers at anthesis; involucral bracts 2–3-seriate, the outer bracts triangular-ovate to broadly ovate, dull gold, tinged gray or occasionally blackish on shoulders; the inner bracts usually more strongly pigmented except for the paler midvein; bracts bearded apically especially along upper midvein, the upper margin short-ciliate with proximal cilia slightly longer, often early glabrate abaxially. Bract and floral trichomes obtuse to clavate, subhyaline, obscurely ornamented within and very obscurely tuberculate. Flowers about 8–14 per capitulum, the pistillate flowers peripheral, the staminate central, equalling to subequalling the pistillate in number; the receptacle sparingly pilose or pilose only toward center. Receptacular bracts equaling to subequaling flowers, broadly linear-subspatulate, ca. 6 times longer than wide, the apex often slightly cucullate and pubescent as the sepals, the base distinctly carinate, clasping. Pistillate flowers: Pedicels ca. 0.1–0.35 mm, thick, glabrous, often becoming callose-thickened in fruit (Peru), leaving characteristic ‘stumps’ on the empty receptacle after abscission of flowers. Sepals broadly obovate, strongly cymbiform, with apex convex-acute to obtuse (1.35–) 1.5–2.0 mm long × 0.6–1.0 mm wide (width variable within a flower), (0.25–) 0.35–0.4 (–0.5) mm wide at the base; black-mottled on shoulders, the midvein area paler brown; short-ciliate along upper margin and bearded with longer appressed hairs on upper dorsum; membranaceous to chartaceous at anthesis, enlarging slightly and becoming uniformly thickened, and often rigid in fruit, husk-like and non-hygroscopic, enclosing the corolla and fruit and dispersed with it. Petals oblanceolate-spatulate, obtuse to often emarginate or truncate-emarginate, (1.2–) 1.35–1.7 (–1.85) mm long × (.35–) 0.4–0.7 (–0.8) mm wide, ca. 2.2–3.2 (–4.9) times longer than wide, cream to brownish-tinged, with scale-like staminode at base, pilose abaxially near margins of distal half with more or less tuberculate trichomes, also densely tufted subapically within in two patches either side of the apex, which enfold the style branch; like the sepals becoming more or less rigid-thickened in fruit, the staminode also thickening and tightly adherent to both petal and ovary base. Gynoecium with ovary 0.5 mm at anthesis, ca. 0.85 mm in fruit; the style base 0.3–0.5 mm long; the nectaries with stalks 0.25–0.65 mm long, glandular portion 0.2–0.35 mm long, very dark brown or dark reddish, subclavate, with a ring of brown membranous papillae ca. 2–3 rows thick at the apex, the whole structure maintaining its shape after anthesis; style branches 0.6–1.0 (–1.2) mm long, usually ca. 0.2 mm longer than nectaries though often developing unequally, the stigmas simple, dark red-brown, non-involute after anthesis. Seeds subglobose to ellipsoid, mostly 0.6–065 mm long, 0.4–0.55 mm wide, red-brown, reticulate with short weak pseudotrichomes, sometimes glabrate. Staminate flowers: Pedicels 0.15 mm to obsolete. Sepals (1.2–) 1.35–1.75 mm long × 0.5–0.85 mm, usually strongly and unequally fused at base for ¼–3/4 their length, obovate-navicular above, the apex obtuse-angled to broadly rounded, the tubular base often rigid and obconic at maturity; color and pubescence as in the pistillate flowers. Corolla including anthophore 1.35–1.9 mm; the anthophore 0.65–1.35 mm long, usually ca. 60–70% the corolla length, ca. 0.2–0.35 mm diameter at base, fleshy and columnar at maturity; the corolla tube 0.35–0.75 mm deep, fleshy towards base especially opposite the filaments, the corolla lobes hyaline to brownish, obtuse, 0.15–0.35 mm, not involute after anthesis; intermediate lobes lacking. Stamen filaments with the basal half fleshy, terete and adnate to the corolla, abruptly narrowed and loosely adhering to the lobes above, exsert 0.2–0.5 mm beyond the lobes, the exsert portion dark reddish-brown especially at tip; anthers cream-colored, ca. 0.3–0.35 mm long, usually deciduous after anthesis and not present in fruiting capitula. Nectaries similar to those of pistillate flowers, reaching or slightly surpassing the corolla sinuses.
The epithet is taken from the Greek caryonaute (nom. sing.), the name given to the “nutshell sailors” in Lucian of Samosata’s tale True Stories. It refers to the diaspores enclosed by the thick buoyant perianth.
In Peru and Bolivia, collected in early anthesis in November and December, and with older inflorescences in all months from February to September. The dry season here is May to August but mitigated at higher elevations by cloud cover (
Colombia (Central Cordillera): Cauca, Nariño. Ecuador: Carchi, probably Sucumbíos. Peru: Cuzco, Junín, Pasco. Bolivia: La Paz. In addition, some atypical specimens or hybrids (see below) are known from the Western and Eastern Cordilleras of Colombia, in Antioquia, Meta, and Norte de Santander. (Fig.
In Peru and Bolivia, this species is restricted to a narrow band of wet paramo-like habitat on the high eastern slope of the Andes, while in Ecuador and Colombia it is found in open wet páramo. It is reported from boggy wet bunchgrass meadows (pajonal) with Calamagrostis Adans. or Festuca procera Kunth, in shallow waterlogged soil of ridgetops and rocky slopes, and in cloud forests and páramo degraded by fire. In Ecuador and Colombia also reported from depressions in Espeletia páramo.
This species is known from two disjunct paramo zones, one about 475 km long in the northern Andes and one 950 km long in the central Andes. However, unlike related P. pilosus, it is not reported from disturbed areas, and rare outlying populations in Colombia show signs of introgression with P. pilosus. In the event of climatic drying or warming this species would be vulnerable, especially in the southern part of its range where suitable habitat is narrowly restricted to the eastern slope.
Paepalanthus karstenii f. corei sensu
This species is most similar to P. pilosus, with a similar cushion-forming habit and similar habitat, and is often confused with that species (or “P. karstenii”). In his later annotations, Moldenke frequently identified this species as P. karstenii f. corei (Moldenke) Moldenke. It has also frequently been distributed as P. muscosus (subsect. Dichocladus), which also has rounded leaf tips. The report of P. muscosus from northern Ecuador by
Paepalanthus caryonauta is readily distinguished from typical P. pilosus and P. dendroides by the obtuse leaf tips, and by the sepals uniformly thickened and persistent in fruit, to form an ovoid-ellipsoid diaspore. Even in anthesis, the sepals of P. caryonauta are about twice as broad at the base as those of P. pilosus and P. dendroides. Paepalanthus caryonauta can also be recognized by eye due to subtle differences in aspect and leaf orientation, with the leaves commonly flatter and more ascending, i.e., less conduplicate and recurved than is commonly seen in P. pilosus. Boeke, who collected P. pilosus and an intermediate form of P. caryonauta at the same locality (see below), noted that in P. pilosus, the cushions were “easy to separate” and in P. aff. caryonauta, “difficult to separate.” In the Cordillera Vilcabamba Dudley reported cushions up to 3 feet in diameter (Dudley 11194). However in disturbed roadside páramo at Acjanaco,
Paepalanthus dendroides | Paepalanthus pilosus | Paepalanthus caryonauta | |
---|---|---|---|
Habitat | Terrestrial or emergent aquatic; 1900–3200 m; to 3900 m in Cuzco | Terrestrial, not in standing water; (2900–) 3100–4000 (–4300) m. | |
Leaf apex shape | Convex-acute to acute, cuspidate | Acute, cuspidate to short-aristate | Narrowly rounded; if cuspidate, tip sharply deflexed |
Leaf pubescence at adaxial apex; at distal margins | Glabrous to persistently hirsutulous; not ciliate |
Glabrous (N*) to pilose (S); often prominent long scattered cilia (N) | Early glabrate; not ciliate |
Peduncles | 20–130 mm | 1.5–8 mm (fl); 20–100 mm (fr) |
7–35 mm |
Capitulum diameter | 3–5 mm | 3–6 mm (S); 3–8 mm (N) |
3–4 mm |
♀ Sepal length | 1.2–1.9 mm | (1.4–) 2.0–2.7 mm | 1.3–2.0 mm |
Sepals in fruit | Hygroscopic thickening along basal midline; detaching from fruit at maturity. (Except P. pilosus var. leoniae) | Thickened throughout; non-hygroscopic, enclosing fruit | |
♀ Petals: Length/Width |
L/W = 1.6–2.3 Broadly spatulate, densely pilose along upper margin |
L/W = (2–)3–6 Oblanceolate, usu. acute, sparingly tufted |
L/W = 2.2–3.5 Oblanceolate-spatulate, obtuse; sparingly tufted |
Nectaries | Colorless (N) to pale pink-brown (S); weak, partly collapsed in old flowers | Dark red-brown, rigid, erect and exsert in old flowers. | |
Nectary position ♂ flower |
Only about half-reaching the corolla sinuses. | Reaching the corolla tube sinuses. | |
Seeds | Pink to orange-brown; pseudotrichomes weak | Red-brown; pseudotrichomes separating when wet, remaining erect |
Paepalanthus caryonauta has a more uniform morphology throughout its range than its close relatives. However, in Colombia and Ecuador the plants have less thickening in the leaves and flowers, the presence of a short pedicel in the staminate flowers, and peduncles often shorter at flowering time, approaching those of P. pilosus in length.
Paepalanthus caryonauta and P. pilosus are mostly allopatric, but in Colombia there are points of contact where intermediates occur. Typical P. caryonautais common in the Nudo de los Pastos and Central Cordillera of Colombia, from which only one historical collection of P. pilosus is known, collected ca. 1842. However at the isolated Paramo Frontino in the Western Cordillera, typical P. pilosus occurs sympatrically with a morphologically intermediate form of P. caryonauta. These two elements were treated as “P. karstenii” and “P. karstenii var. corei,” respectively, by
In the eastern Cordillera of Colombia, P. pilosus is abundant, while only two specimens suggesting atypical P. caryonauta were confirmed, these from opposite ends of the eastern Cordillera, on east-facing slopes. At the south end, an intermediate plant, similar to that of Paramo Frontino, but with flowers mostly abortive, was collected from the eastern slope of Sumapaz National Park (S. Diaz-Piedrahita 2608). This location is just south of the southernmost confirmed Colombian collection of P. pilosus. To the north, Cuatrecasas 10302 (F), collected at the “extreme east” of Paramo Santurban (Norte de Santander) may represent P. caryonauta or a hybrid intermediate, differing by the light gold bracts. Sympatric taxa in this area include P. dendroides, typical P. pilosus, and the taxon treated below as Paepalanthus sp. A.
In southern Peru, typical P. caryonauta occurs in mixed populations with P. dendroides at the entrance to Manú National Park (Abra Acjanaco, Cuzco). Both taxa have been abundantly collected and are readily distinguished in the field (A.Cano, pers. comm.) However, in Pasco (Oxapampa), where both species also occur, a sterile intermediate between the two has been collected. (See discussion of P. dendroides.)
(paratypes). COLOMBIA. Cauca: Purace – La Plata, 3200 m, 22 Aug 1957, Barclay 5176 (F,
(not paratypes). Intermediates with P. pilosus: COLOMBIA. Antioquia: Paramo de Frontino, near Llano Grande, 3450 m, 27 Oct 1976, Boeke 269 (
Paepalanthus dendroides (Kunth in H.B.K.) Kunth, Enum. Pl. 3: 507. 1841.
Eriocaulon dendroides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 251, t. 59, fig. 2. 1815 [1816].
Type: Colombia. Cundinamarca: “Crescit in frigidis montanae planitiei Bogotensis inter Suba et Suacha, alt. 1340 hex.,” Jul 1801, Bonpland & Humboldt s.n. (lectotype, here designated: B [B 10 0243900]; syntypes: B [B-W 2366],
Paepalanthus barkleyi Moldenke, Phytologia 3: 114. 1949. Syn. nov.
Type: Colombia. Antioquia: 1 km N of Santa Rosa de Osos, 2600 m, 25 Sep 1948, S. Posada S., M. Torregrosa & F. Barkley 18A100 (holotype:
Paepalanthus karstenii var. corei Moldenke, Phytologia 29: 386. 1975. Syn. nov.
Type: Colombia. Cauca: Above Purace, 11,000 ft, 19 Feb 1944, E. L. Core 272a (holotype:
Paepalanthus karstenii f. corei (Moldenke) Moldenke. Phytologia 45: 296. 1980. Syn. nov.
Type: Based on P. karstenii var. corei Moldenke.
Based on Eriocaulon dendroides Kunth in H.B.K.
Plants terrestrial or partly submerged, forming densely leafy cushions or mats reported up to 30 cm in diameter (Fassett 25929), the erect branchlets ca. 1–5 cm. Leaves linear-subulate, 9–22 mm long × 0.85–1.6 mm wide, tip cuspidate-acute, persistently pilose to hirsutulous adaxially with white roughened hairs, sometimes nearly glabrous (Peru, Cajamarca and Pasco), mostly eciliate, consistently paler green than P. caryonauta or P. pilosus, with veins slightly salient below. Peduncles 20–130 mm long, pale, rigidulous (Peru) or often soft and compressible and then noticeably constricted at apex when dry, glabrous except for an apical collar of sericeous appressed hairs; peduncle sheaths 13–23 mm long, tightly enclosing the peduncle, equaling or scarcely exsert from leaf mat, tufted at apex otherwise mostly glabrous. Capitula 3.5–5 mm wide. Involucral bracts about equaling flowers, ovate, similar to P. caryonauta but the outer bracts more hyaline, paler, sometimes uniformly gold and glabrous except at margins. Trichomes of bract and sepal apices obtuse to clavate, strongly tuberculate. Flowers ca. 12–17 per capitulum, pistillate peripheral, the staminate equaling to subequaling the pistillate in number. Pistillate flowers: Pedicels ca. 0.25–0.45 mm long, fine, not thickened at maturity. Sepals obovate-spatulate, 1.15–1.85 mm long × 0.65–0.85 mm wide, 0.15–0.25 mm wide at base, blackish-brown, short-ciliate along upper margin and bearded with longer appressed hairs on upper dorsum, the basal half of the midrib hygroscopically thickened, and spadiceous-brown in fruit, the broad distal half of the sepal remaining chartaceous, suberect, detaching from diaspore upon dispersal. Petals broadly spatulate, 1.15–1.75 mm long, 0.55–1.0 mm wide, ca. 1.6–2.3 times longer than wide, cream-colored and densely long-pilose with tuberculate trichomes on the abaxial surface flanking the midvein, not thickening, dispersed with fruit. Gynoecium with style base 0.15–0.25 mm long, the nectaries 0.55–0.7 (–0.85) mm long, the glandular portion colorless to pale pink to red- or yellow-brown, penicillate, slightly curved after anthesis, the apical ring of papillae colorless (white), thin- to thick-walled; styles 0.75–0.9 mm, mostly thinner and less pigmented than in P. pilosus or P. caryonauta. Seeds 0.6–0.75 mm long, orange-brown, the pseudotrichomes weak, erect upon wetting but collapsing soon after (few seeds observed). Staminate flowers: Pedicels (0.35–) 0.4–0.6 mm long, fine, membranous, nearly glabrous. Sepals 1.1–1.9 mm long × 0.35–0.5 mm wide, narrowed to base and very shallowly fused or if fused up to half the sepal length, not thickening, the calyx base not obconic at maturity, color and pubescence as in pistillate flowers. Corolla (1.0–) 1.4–1.7 (–1.95) mm long; the anthophore alone (0.35–) 0.55–0.95 mm long, comprising ca. 40–60% of the corolla length, membranous and ca. 0.1–0.2 mm in diameter at base, 0.3 mm at apex, the lobed tube (0.55–) 0.65–0.9 (–1.0) mm long. Filaments often unpigmented, slightly less exsert than in P. caryonauta, the base of the anther rarely exsert more than 0.3 mm beyond lobe tips. Nectaries well-included within tube, usually only half-equaling the sinuses.
In Central America, flowering from March to April, in the dry season, and from August to September, the wet season, punctuated by a short dry period or veranillo (
Costa Rica (Cerro de Talamanca): Limón, Puntarenas. Panama: Bocas del Toro. Colombia (Central and Eastern Cordilleras): Antioquia, Bogotá D.C., Boyacá, Cauca, Cundinamarca, Nariño, Santander, Norte de Santander. Peru: Cajamarca, Cuzco, Huánuco, Pasco, Puno. Brazil (Pico da Neblina): Amazonas. (Fig.
A terrestrial or partly submerged aquatic, in open marshy subparamo, low paramo or cloud forest margins, in bogs, wet meadows, seeps, commonly associated with Sphagnum or tussock grasses, sometimes shrubs or Blechnum. Elevation mostly 2300–3200 m, but as low as 1900 m in Antioquia, and up to 3800 m in Cuzco (Abra Acjanaco), Peru. In the high elevation Cuzco plants seed production does not appear abundant, abortive flowers are common, and smut fungus infection is observed.
The conservation status of this widespread species is presumed to be of Least Concern (
Paepalanthus dendroides was initially described by Kunth and later
Paepalanthus dendroides was placed in synonymy of P. pilosus by
Paepalanthus pilosus sensu
Paepalanthus dendroides is a variable species across its range, but may be distinguished from its close relatives by the character syndrome in Table
Of the Peruvian collections, those from Huánuco, Cuzco, and Puno share a similar morphology, with leaves relatively narrow and conspicuously pilose above, the outer involucral bracts dusky gray, and the nectaries somewhat pigmented. Leaf pubescence easily distinguishes these populations from sympatric P. caryonauta. In comparison, specimens from Cajamarca and Pasco have broader, glabrous leaves. The Cajamarca collections were examined only from photos, but the plants closely match Colombian material of P. dendroides and are from a similar habitat and elevation. The Pasco collection (Vasquez 29038) has floral characters somewhat intermediate with P. pilosus or P. caryonauta, as follows: petals 2.0–2.3(–3.0) times longer than wide; style base 0.4–0.5 mm; nectaries darker, 0.85–0.95 mm long; styles to 1.0 mm long; nectaries of male flowers almost reaching the mouth of the corolla.
The Pasco, Peru, specimen (Vásquez 29038) was collected very near a plant fully intermediate between it and P. caryonauta, with abortive locules and stigmas (Monteagudo 7938; full specimen citation under P. caryonauta). Typical P. caryonauta (Monteagudo et al. 16143) is recorded 20 km to the east.
In Colombia, introgression between P. dendroides and P. pilosus is suspected in the disturbed paramos east of Bogotá. Although P. dendroides was originally described from near Bogotá, I have only seen one other typical individual from this vicinity (Pennell 1997), collected at Quebrada Chapinero in 1917, the same locality where the type of P. schultesii (=P. pilosus) was collected in 1941. Paepalanthus pilosus is presently abundant near Bogotá, but seems particularly variable and with an unusual tendency to long peduncles, lax habit, and variably shaped bracts, suggesting intermediacy with P. dendroides (see P. pilosus discussion). Typical P. dendroides is currently found north of Bogotá, where it is mostly recorded from elevations 500–900 m lower than nearby collections of P. pilosus. In Panamá, however, typical P. dendroides and P. pilosus are sympatric at the Cerro Fabrega massif, without apparent intermediates.
(of 56 total). COSTA RICA. Limón: Cerro Kamuk, 9°14'30"-15'30"N, 83°03'30"-04'30"W, 2900–3100 m, 23–26 Mar 1984, Davidse et al. 25928 (F,
Plants forming loose cushions or mats; leaves blue-green, pilose above, older leaves strongly costate below; peduncle sheaths 25–30 mm, very lax, and strongly exsert from the leaf mat, the involucral bracts dark brown, the heads with over 20 flowers; pistillate flower petals spatulate, pilose; nectaries dark red-brown and rigidulous, those of the male flowers equaling the corolla sinuses.
PERU. Piura: Huancabamba, Jalca de Chiguelas, 5°8.2'S, 79°23.6'W, 3082 m, 19 Oct 2001, A. Sagástegui et al. 16799 (holotype: F; isotype:
Plants short-caulescent, with erect actively growing shoots to 4 cm, densely leafy, branching to form rounded mats. Leaves subulate, acute, 13–22 mm long × 1–2 mm wide at midpoint, tip cuspidate to apiculate, densely pilose to villous above with appressed to spreading white tuberculate hairs, upper margin eciliate, lamina dark blue-green, mature leaves prominently 3–5-costate below. Inflorescences solitary and terminal, soon overtopped by one or two erect lateral shoots which may flower in rapid succession, so that peduncles superficially appear fascicled. Peduncles 6.0–11.5 cm long at anthesis, perhaps continuing to elongate into fruiting, with peduncle of previous season up to 15 cm observed on same plant, ca. 3-costate, densely subappressed-villous especially above, with a dense sericeous collar of trichomes subtending involucre. Peduncle sheaths 25–30 mm, much surpassing the leaves, and strongly surpassing the leaf mat, scarious, very lax, nearly glabrous except for the tufted apex, the lamina cucullate, enclosing the bud when young and then splitting broadly into two or three triangular segments. Capitula 4–6 mm, depressed-hemispheric. Involucres subequaling flowers at anthesis, and opening broadly at maturity; involucral bracts 2–3-seriate, the outer bracts triangular-ovate, greenish to uniformly dark brown, shaggy-ciliate on margins and villous in two submedial bands. Floral trichomes obtuse to clavate, strongly tuberculate. Flowers ca. 20–24 per capitulum, the pistillate flowers peripheral, the staminate central, with 14–18 pistillate flowers to 6 staminate flowers (in two capitula sampled). Receptacle sparingly long-pilose with brownish hairs. Receptacular bracts subequaling flowers, linear-subspatulate, the apex slightly cucullate, pubescent as sepals, the base sharply carinate. Pistillate flowers: Pedicels 0.3–0.45 mm long, fine and membranous. Sepals broadly obovate-spatulate to subtruncate at apex, sometimes weakly cymbiform, 1.55–1.65 mm long × 0.65–0.8 mm wide at middle, 0.15–0.2 mm wide at base, blackish-brown, short-ciliate (apical cilia to 0.17 mm) along upper margin, and appressed-long-pilose in two bands flanking the upper dorsum, the basal half of the midrib hygroscopically thickened, spadiceous-brown in fruit, and recurving when dry, the broad upper half of the sepal remaining chartaceous, erect; sepals detaching from fruit on dispersal. Petals spatulate, obtuse, brownish at tip, 1.25–1.35 mm long, the widest in a flower ca. 0.65 mm wide, ca. 2.2 times longer than wide, bearing scale-like staminodes at base, densely long-pilose with long tuberculate hairs on abaxial upper half except for midvein, also tufted subapically within, the hairs enfolding the style branch, petals not thickening in fruit, dispersed with fruit. Gynoecium with style base 0.35 mm long; nectaries ca. 0.7 mm long, dark red-brown, penicillate to subclavate-infundibular, with fringe of colorless stiff-walled papillae at mouth, these rigidulous and maintaining shape after anthesis; style branches 0.85 mm long, thick and dark red-brown, non-involute. Only two slightly misshapen seeds seen, 0.6–0.63 mm long, pinkish to red-brown, the pseudotrichomes weak. Staminate flowers: Pedicels 0.35–0.4 mm, membranous, nearly glabrous. Sepals 1.6–1.7 mm × 0.5–0.6 mm, spatulate to obrhombic or subtruncate, color and pubescence as in the female flowers, narrowed toward base and shallowly fused. Corolla 1.8 mm long; the anthophore 1.15 mm long, comprising ca. 65% the length of the corolla, grading from 0.15 mm wide at base to 0.35 mm wide at apex; the tube 0.65 mm long including well-defined brownish-tinged lobes. Filaments brownish-tinged above; exsert not more than 0.3 mm beyond lobe tips, anthers persistent, cream to light brownish. Nectaries equaling corolla sinuses.
Collected in flower May, September, October.
Endemic to Peru, Piura, Cordillera de Huancabamba, District Carmen de la Frontera (Fig.
Grass páramo (or jalca), probably of anthropic origin, and “burnt cloud forest, growing under Pteridium aquilinum” (Weigend & Dostert 98/252). Elevation ca. 2900–3000 m.
Assessed as Critically Endangered, according to IUCN Criteria B1ab(iii) (
Paepalanthus huancabambensis is similar in habit and dimensions to P. dendroides, but differs by its very lax, elongate peduncle sheaths well exsert from the leaf mat, and the large capitula with more flowers. It also differs in the dark blue-green leaf color, compared to the consistently pale green leaves of the widespread P. dendroides, and preliminary anatomical study distinguishes it from that species by the presence of adaxial vein buttresses (bundle sheath extensions) in leaf median section. The broadly spatulate densely pilose female petals are similar to those of P. dendroides. However, the longer style base, the dark rigidulous nectaries with stiff colorless papillae fringing the rim, and the size of the nectaries relative to the corolla tube in the male flowers all suggest P. pilosus.
Except for the lax peduncle sheaths, this species lacks any strong distinctive features of its own but its mixture of critical characters prevent it from being easily placed in any related species, and do not immediately suggest hybrid origin. It is endemic to the Cordillera de Huancabamba near the border of Peru and Ecuador in the western part of the Andean chain. Notably, in the same vicinity are also found an atypical form of P. pilosus (Cano 16840, discussed under P. pilosus var. pilosus), and at higher elevations the only known populations of P. lodiculoides from Peru and Ecuador.
PERU. Piura: Huancabamba, Cordillera Chinguela (Sapalache el Cármen), 2900 m, 15 Sept 1981, Sagástegui et al. 10225 (
Paepalanthus lodiculoides Moldenke, Bull. Torrey Bot. Club 68: 68. 1941 [31 Dec 1940].
Syngonanthus steyermarkii Moldenke, Phytologia 2: 418. 1948.
Type: Venezuela. Táchira: Páramo de Tamá, near Venezuelan-Colombian border, 3045–3475 m, 15 July 1944, J. Steyermark 57372 (holotype:
Paepalanthus polytrichoides var. densus Moldenke, Phytologia 8: 392. 1962.
Type: Colombia. Cundinamarca: Paramo de Chisacá, around Laguna de Chisacá, 3650–3700 m, 29 Dec 1959, J. Cuatrecasas & R. Jaramillo M. 25737 (holotype:
Paepalanthus lodiculoides var. floccosus Moldenke, Phytologia 32: 47. 1975.
Type: Colombia. Boyacá: Paramo de la Sarna, entre Sogamoso y Vado Hondo, 5 km al NE de la Laguna de Tota, 3510 m, 30 Mar 1973, A. Cleef et al. 9214 (holotype:
Colombia. Boyacá: Nevado de Cocuy, Alto Valle de las Lagunillas, 4000 m, 12 Sept 1938, J. Cuatrecasas & H.García Barriga 1537 (holotype:
Plants compact densely branched mosslike cushions, reportedly up to 23 cm in diameter (Soderstrom 1262) and 2 cm high (
In Peru, flowering May–June (early in dry season); in Ecuador, September; in Colombia and Venezuela, July–Sept and Nov–March.
Colombia (Eastern Cordillera); Bogotá D.C., Boyacá, Cundinamarca, Santander. Venezuela (Paramo de Tamá): Táchira. Ecuador: Loja. Peru: Piura. This is the first report of the species from Peru. (Fig.
In paramo, at (3000–) 3300–4000 m. Cited by some authors as characteristic of very wet páramo of lake margins, bogs, and in grass paramo (
Known from two disjunct bands of paramo, one 420 km long in the north, and one about 100 km long in the south, under cool wet conditions.
Paepalanthus lodiculoides is easily distinguished from all other species in the group by the tiny moss-like leaves, delicate peduncles, and small capitula. The diaspores enclosed by persistent sepals and petals are similar to those of P. caryonauta. According to
This species is also unique in the complex for the variable sex ratios of the capitula, which may be all staminate to mostly pistillate on the same plant. Capitula with exclusively pistillate flowers were not observed.
The type of Syngonanthus steyermarkii Moldenke is the northernmost record of the species. It differs by the shorter peduncles (2–3 mm vs. 5–11 mm), shorter sheaths (1.7–2.2 mm vs. 3–5 mm) and capitula only about half the diameter of other specimens (1.4–1.5 mm vs 2.5–3 mm). The peduncle sheaths of this specimen are also abnormally developed, the apical lobe resembling an involucral bract in color and texture. Steyermark described the habitat as a limestone outcrop, which, if true, would be unusual for Eriocaulaceae, which almost universally occur on acidic substrates.
The three names cited in synonymy here were first cited as synonyms by
COLOMBIA. Bogotá, D.C.: Sumapaz, Páramo de Chisacá, 3900 m, 9–11 Nov 1958, Barclay & Juajibioy 6113 (F,
Paepalanthus pilosus (Kunth in H.B.K.) Kunth, Enum. Pl. 3: 518. 1841.
Eriocaulon pilosum Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 251–252. 1815 [1816].
Type: Colombia. Cundinamarca: “Crescit in frigidis montanae planitiei Bogotensis inter Suba et Suacha, alt. 1340 hex.,” Jul 1801, Bonpland & Humboldt s.n. (lectotype, here designated: B [B_10_0243899] in part, as to plants mounted on left and right side of sheet. The plant mounted in the center of the sheet is P. dendroides (Kunth) Kunth. Syntypes: B [B-W 2371],
?Eriocaulon selaginoides Benth., Pl. Hartw. 260. 1846.
Type: Colombia. Cauca: Popayán, near Laguna de Guanacas, 12,000 ft, [yr 1842], K. T. Hartweg 1445 (holotype: K [K000640107]; isotypes: K [K000640106], B,
?Paepalanthus selaginoides (Benth.) Körn., Fl. Bras. 3(1): 362. 1863.
Type: Based on Eriocaulon selaginoides Benth.
Dupatya pilosa (Kunth) Kuntze, Revis. Gen. Pl. 2: 746. 1891.
Type: Based on Eriocaulon pilosum Kunth in H.B.K.
Paepalanthus kupperi Suess., Bot. Jahrb. 72: 293, t. 3, fig. 8. 1942.
Type: Costa Rica. Cartago/Limón/San José: [Cerro] Chirripó Grande, 3450 m, W. Kupper 1315 (holotype: M [M0137218]; isotype:
Paepalanthus espinosianus Moldenke, Phytologia 2: 228. 1947.
Type. Ecuador Morona-Santiago: trail between Pailas and El Pan, 3400 m, 10 Sep 1943, J.A. Steyermark 54342 (holotype:
Paepalanthus loxensis Moldenke, Phytologia 2: 229. 1947.
Type: Ecuador. Loja: between Tambo Cachiyacu, La Entrada, and Nudo de Sabanilla, 2500–3500 m, 7 Oct 1943, J. A. Steyermark 54452 [‘54432' in protologue] (holotype:
Paepalanthus subsessilis Moldenke, Phytologia 2: 232.1947.
Type: Venezuela. Lara: between Buenos Aires and Páramo de las Rosas, 2285–3290 m, 11 Feb 1944, J. Steyermark 55495 (holotype:
Paepalanthus schultesii Moldenke, Bot. Mus. Leafl. 16(4): 65. 1953. Syn. nov.
Type: Colombia. Cundinamarca: Macizo de Bogotá, Quebrada de Chapinero, 9000 ft, 24 Sep 1941, R. E. Schultes 1024 (holotype:
Paepalanthus dennisii Moldenke, Phytologia 7: 88. 1959. ‘dennisi.’ Type citation corrected by Moldenke, Phytologia 7: 120. 1959. Syn. nov.
Type: Venezuela Merida: Mucubaji, Sierra de Santo Domingo, 3550 m, 26 Aug 1958, R. W. G. Dennis s.n. (holotype: K [K000640179]; isotype:
Paepalanthus karstenii var. minimus Moldenke, Phytologia 30: 15. 1975. Syn. nov.
Type: Colombia. Cundinamarca: Laguna de Verjón, 27 Jul 1917, Bro. Aristé Joseph A.73 (holotype:
Paepalanthus karstenii var. subsessilis (Moldenke) Moldenke, Phytologia 32: 47. 1975.
Type: Based on Paepalanthus subsessilis Moldenke
Based on E. pilosum Kunth in H.B.K.
Eriocaulon microcephalumsensu
Paepalanthus pilosus var. pilosus
Densely branched cushion plants, the cushions reportedly reaching 30 cm in diameter (Molau 3223) and up to 10 cm tall (Berry 4366). Leaves subulate, recurved, 7–16 (–20) mm long × (0.5–) 0.7–1.3 (–1.5) mm wide at midpoint; apex cuspidate-acute to short-aristate, the sharp tip evident in adaxial view; usually appressed pilose adaxially near tip (in Peru and Ecuador), the hairs smooth to roughened, in northern part of range usually with conspicuous long coarse scattered cilia to apex (rare in Peru and Ecuador), often early glabrate, bright green, texture chartaceous to rigidulous, the abaxial surface smooth or with nerves salient. Peduncles appressed-pilose when young especially at apex, often only 1.5–8 mm long and surpassed by the sheaths at anthesis, but frequently and variably elongating up to ten or more times this length (20–100 mm) in fruit; in Peru and Ecuador only rarely up to 50 mm at anthesis and strongly exsert (Cano 16840); the sheaths 3–15 mm long, scarious and glabrous or minutely tufted at apex, splitting apically into 2 or 3 triangular segments. Capitula 3–6 mm in diameter, often borne among the leaves at anthesis, frequently exsert in fruit, more rarely exsert at flowering. Involucral bracts equaling the capitulum to slightly surpassing it, the outer bracts 1.6–4.2 mm long, ovate to often triangular, pale gold or greenish especially along midvein, pilose on upper back to merely tufted or glabrate, the inner bracts more broadly ovate to triangular, sometimes tinged grayish-brown on shoulders; receptacle long-pilose. Trichomes of bract and sepal apices subacute to bulbous, obscurely tuberculate. Flowers ca.10–16 per capitulum, the pistillate peripheral, usually 1–2 (–3) times as many as the staminate flowers. Pistillate flowers: Pedicels 0.35–0.5 mm, fine or rarely wide and spongy, not thickened-rigid in fruit. Sepals elliptic to obovate or oblanceolate, apex acute to rounded, the base linear-ligulate to narrowly cuneate, 1.4–2.65 mm long, 0.3–1.1 mm wide at middle, 0.15–0.3 (–0.4) mm wide at base; usually tinged dusky brown above, rarely uniformly pale cream (Sagastegui 12242), short-tufted at apex, ciliate at upper margins, and usually pilose either side of the dorsal midvein; the basal half of the midrib hygroscopically thickened, spadiceous-brown in fruit and recurving when dry, the margins and distal half of the sepal remaining chartaceous, erect; sepals detaching from diaspore before dispersal. Petals oblanceolate-spatulate, acute to obtuse to emarginate, rarely (Wurdack 1616) broadly cuneiform; apex truncate-emarginate, apiculate; 1.5–2.35 mm long × 0.3–0.75 (–1.0) mm wide, ca. 2–6 times longer than wide, cream to brownish-tinged, pubescence similar to P. caryonauta; usually not thickening in fruit, dispersed with the fruit. Gynoecium with style base 0.5–1.05 mm, nectaries with stalks 0.3–0.6 mm long, glandular portion dark red to brown, 0.3–0.4 mm long, penicillate with stiff whitish papillae at upper margin (northernmost Peru and northwards), or 0.4–0.5 mm long, clavate-infundibular with stiff whitish sometimes elongate papillae distributed along the outer surface, densest at the rim (Cajamarca: Celendín, Amazonas: Chachapoyas); papillae rigidulous after anthesis; styles 0.8–1.1 mm long (Barbour 3427: 1.4–1.6 mm long), dark red-brown. Seeds 0.55–0.8 mm long, red-brown, reticulate with abundant white erect to suberect pseudotrichomes. Staminate flowers: Pedicels 0.2–0.5 mm. Sepals 1.2–2.45 mm, fused at base unequally, from very briefly to about 1/3 sepal length, obovate to usually spatulate, acute to rounded, narrowed to base and not thickening with age. Corolla 1.2–2.5 mm long, the anthophore 0.5–1.5 mm long, comprising 30–65 % the corolla length, membranous and 0.1–0.2 mm diameter at base, broadening to 0.2–0.3 mm near apex; the corolla tube including lobes 0.5–1.65 mm long. Filaments similar to P. caryonauta, usually dark red-brown between apex and the base of the corolla lobe, exsert ca.0.2–0.5 mm beyond the lobes. Nectaries reaching the sinuses of the corolla tube.
In Central America, mostly collected January to May, and in July, coinciding with the main dry season, and possibly a shorter second dry season (veranillo), respectively (
Costa Rica (Cerro Talamanca): Cartago, Limón, San José. Panama: Bocas del Toro. Colombia (Eastern Cordillera, Sierra Nevada de Santa Marta, possibly Serrania de Perijá, also rare in Western and Central Cordilleras): Antioquia, Bogotá D.C., Boyacá, Cauca, Cundinamarca, Magdalena, Norte de Santander, Santander. Venezuela (Cordillera de Merida and Ramal de Guaramacal): Apure, Lara(?), Merida, Táchira, Trujillo. Ecuador: Azuay, Loja, Morona-Santiago, Zamora-Chinchipe(?). Peru (to 7° S lat.): Amazonas, Cajamarca, Lambayeque, Piura. (Fig.
Common and characteristic ground cover (rasante) or cushion species of wet páramo, rarely subpáramo, on boggy organic soils, or at pond and bog margins but not in standing water (
Paepalanthus pilosus is a commonly cited species in phytosociological analyses of North Andean paramo, usually under the name P. karstenii (
The status of this widespread variable taxon, reported as a colonizer tolerant of disturbance, is presumed to be of Least Concern (
No type material is found for E. pilosum Kunth in the herbarium P-Bonpl. in Paris (
Paepalanthus pilosus has suffered confusing taxonomic treatment over time.
Moldenke did not specify distinguishing characters for his other species here placed in synonymy of P. pilosus, nor were these names widely used. Of his new infraspecific taxa, Paepalanthus karstenii f. corei was said to differ from that species by the peduncles only 1–2 cm in length, and P. karstenii var. minimus by both leaves and peduncles shorter. The type of the former is in fact P. dendroides, though Moldenke most commonly applied the name to P. pilosus and P. caryonauta.
Paepalanthus espinosianus and P. loxensis were first treated in synonymy of P. pilosus by
Paepalanthus pilosus in Peru and southern Ecuador is characterized by the usually dwarf flowering peduncles with capitula subsessile at anthesis, and the outer involucral bracts greenish or with a green midvein and often longer than wide. In this area, soft hirsutulous pubescence of the upper leaf surface near the apex, similar to that found in P. dendroides, is common. From Costa Rica to Colombia and Venezuela, long robust scattered cilia along the distal margin are frequent and diagnostic when they occur (hence the species epithet; cf. Fig.
Habit varies from a dense compact mat pressed horizontally by collectors (as in the type of P. espinosianus Moldenke), to a rounded cushion with branch lengths of a few centimeters, pressed and mounted vertically (as in the type of P. loxensis Moldenke) The superficial difference in aspect of these two forms caused Moldenke to ally the former species with P. karstenii, but the latter species with P. glaziovii Ruhland (subsect. Dichocladus). This character almost certainly varies in response to habitat, as
Peduncle length is highly variable, as already noted by
Other characters showing wide variation in Peru and Ecuador are involucral bract length, capitulum size, and flower and seed size. For example, the type of P. loxensis (Loja) contrasts sharply with that of P. espinosianus (Azuay) by its much smaller involucral bracts, capitula, flowers, and seeds. Indeed, in most plants from Azuay, in the Cerro Fierro-Urcu (Loja), and in some localities in Peru (Jaén, Bagua, Celendín), outer involucral bracts are ca.2.6–4.0 mm, with the tips often surpassing the large capitula 4–6 mm in diameter (cf. Fig.
Paepalanthus pilosus in Colombia requires further careful study. In the Eastern Cordillera, particularly in disturbed paramo near Bogotá, there are more pronounced morphological extremes in the species than in other parts of its distribution. Here, leaf lengths range up to 3 cm, capitula from 3.5–8 mm in diameter, peduncles are occasionally strongly exsert at flowering, and involucral bract color and form are variable, ranging from nearly obovate-rounded to narrowly triangular, and pale gold to dark pigmented on the shoulders.
Hybridization may be a factor contributing to the chaotic variation in P. pilosus in the vicinity of Bogotá. Some material, such as the type of P. karstenii (see Doubtful Taxa) looks intermediate with P. dendroides, as to elongate peduncles and rounded involucral bracts. Typical P. dendroides was originally described from Bogotá, but has not been collected in the vicinity since 1917, while it is still widespread elsewhere. In addition, probable hybrids between P. pilosus and P. caryonauta have been collected at the margins of the range of P. pilosus, at Paramo Frontino (Antioquia), and on the eastern slope of Paramo Sumapaz, south of Bogotá (see P. caryonauta discussion).
A final complication in the taxonomy of Colombian P. pilosus is the presence in the Serranía del Perijá and vicinity of a closely related but more robust taxon, which may intergrade with it, discussed below under Paepalanthus sp. A.
(of 122 total). COSTA RICA. Cartago: Cerro Asunción, 3350 m, 19 Mar 1978, Wilbur 26123 (F). Limón: Parque Nac. Chirripó, Valle Las Morrenas, 9°29'24"N, 83°29'24"W, 3500 m, E. Alfaro 417 (
Habits of Paepalanthus pilosus varieties and “Paepalanthus sp. A.” A−DP. pilosus var. pilosus. (ASoderstrom 1346BSagástegui 12242CCano 16840DLarsen 237.) EP. pilosus var. leoniae (León 1597)F−G “Paepalanthus sp. A” (Cuadros 3732) F Habit G Capitulum. In D arrows indicate flowering (fl) and fruiting (fr) capitula.
Pistillate flowers and fruits of P. pilosus varieties. A−KP. pilosus var. pilosus (A−BJørgensen 2366C−FCano 16840GCuatrecasas 25574HMacDougal 4463IWurdack 1616JØllgaard 9557KBoeke 2133) A−B Flower in anthesis with detail of gynoecial nectaries C Flower in anthesis D Gynoecium E Petal F Seed G Diaspore with detached calyx H Sepal (fr) I Diaspore releasing seed J−K Seeds after wetting LP. pilosus var. leoniae. Mature diaspore, sepals and petals intact. (León 1597).
Differs from the typical variety by the capitula 2–3 mm in diameter, flowers 5-10 per capitulum, the pistillate flowers with sepals elliptic, uniformly thickened in fruit and dispersed with the fruit; petals of pistillate flowers oblong to oblong-spatulate with broadly rounded to truncate apex.
PERU. San Martín: Prov. Mariscal Caceres, NW sector Rio Abiseo National Park, grassland in Paredones, [07°40'16.73"S, 77°29'1.78"W], 3600 m, 16 March 1988, B. León & K. Young 1597 (Holotype:
Leaves 5.5–12 mm long × 0.6–1.4 mm wide at midpoint, sharply cuspidate, finely appressed-ciliate near apex when young, early glabrate. Peduncles 3–6.5 mm long, fruiting peduncles the same length in specimens observed, the sheaths 5–7 mm. Capitula 2–3 mm in diameter, the involucres obconic or cupulate in flower and fruit, not opening broadly, the bracts similar to typical variety. Flowers 5–10 per capitulum, the staminate equal in number to pistillate or in fewer-flowered capitula the proportion of staminate flowers may be reduced. Pistillate flowers: Pedicels 0.25–0.35 mm. Sepals elliptic, obtuse to rounded, 1.35–2.4 mm long × 0.65–0.8 mm wide at middle, 0.25–0.35 mm wide at base, pale brownish, tinged dusky-brown on shoulders, becoming uniformly thickened and cymbiform at maturity, enclosing the fruit and dispersed with it. Petals oblong to oblong-spatulate, with apex broadly rounded to truncate, retuse or apiculate, 1.5–2.0 mm long × 0.65–0.85 mm wide, 2.2–2.5 times longer than wide, pilose as in the typical variety or with trichomes restricted to subapical tufts, uniformly thickening in fruit, as the sepals. Gynoecium with style base 0.5–0.65 mm long, nectaries 0.65–1.0 mm long, papillae at upper margin colorless, stiff, globose (Leon 1597) to linear (Young 4368); style branches 0.75–1.7 mm. Seeds not seen but mature ovary locules 0.65 mm (Young 4368), 0.75 mm (Leon 1597). Staminate flowers: Pedicels 0.25–0.35 mm. Sepals 1.35–2.1 mm long, fused 35–55 % of their length. Corolla 1.55–2.05 mm long, with the anthophore 0.85–1.15 mm long, comprising 45–75% the length of the whole corolla, ca. 0.1–0.25 mm in diameter at base, the tube and lobes 0.4–1.05 mm long. Filaments brownish toward apex; nectaries reaching corolla sinuses.
Named in honor of Dr. Blanca León, of the Universidad Nacional Mayor de San Marcos and the University of Texas, a generous and accomplished botanist whose many contributions include ecological and taxonomic study of Rio Abiseo National Park.
Collected in March and July. A mild dry season occurs June to August (
Endemic to Peru, San Martín, Prov. Mariscal Caceres, Rio Abiseo National Park. (Fig.
From high-elevation grasslands, at 3450–3800 m.
Endangered, Criteria B1ab(iii) (
This variety is found at the southernmost and highest elevation station for the species in Peru. It occurs above 3400 m at the very south end of the Amotape-Huancabamba floristic zone as characterized by
The uniformly thickened fruiting perianth of this variety is similar to that of P. caryonauta, complicating the otherwise tidy distinction between the two species. However the size and shape of the leaves, the dwarf peduncles, the pale greenish involucres, and the large rigid nectary papillae, all suggest P. pilosus, and the type specimen in particular looks nearly identical to populations of typical P. pilosus from adjacent southern Amazonas. In fact, these populations are partly intermediate in floral morphology between the two varieties, as discussed under the typical variety. Given the variation in P. pilosus in Peru and the contiguity of distribution, it seems best for now to treat this taxon as a variety of that species.
The specimen Young & León 4368, collected ca. 10 km from the other two, is marked by the very small size (5.5–7 mm) of the dark green leaves. The flowers and seeds are also smaller. The habitats of the two localities differed, with the type locality a typical patchy wet páramo among outcrops, while the small-leaved plant was in a broad boggy area bordered by small trees, where cattle once pastured (B. León, pers. comm.)
Perú. San Martín: Prov. Mariscal Caceres. Puerta del Monte, northwest corner of Rio Abiseo National Park; high elevation grassland on bottom of u-shaped valley; [7°39'30.20"S; 77°28'13.67"W], 3450 m, 10 Jul 1987, K. Young & B. León 4368 (F); Pastizales de Empedrada, entre manojos de Calamagrostis, [07°40'17"S, 77°29'2" W], 3750–3785 m, 27 Jul 2000, B. León & K. Young 4579 (
Paepalanthus macarenensissensu
This robust taxon has a clumping habit structurally similar to P. pilosus but lacks the dense pulviniform aspect. Its other similarities include acute to aristate leaves, scarious splitting peduncle sheath tips, pale gold lanceolate involucral bracts, and a nearly identical flower and fruit morphology. It differs by the following characters:
Leaves longer, narrowly linear-lanceolate, 3–4 cm long, the peduncles 10–20 cm long at anthesis, and the capitula 6–7.5 mm wide, much more floriferous (> 40 flowers) than P. pilosus and sometimes globose at maturity, with alternating whorls of staminate and pistillate flowers. In addition, the capitula are “indeterminate,” with floral primordia found at the center of capitulum at the time of anthesis of the outer whorls. This contrasts with P. pilosus, in which pistillate flowers are limited to the outer whorl, staminate to the inner, and no floral primordia are found at the start of anthesis.
The most robust individuals are found in the northern part of Serrania del Perija (ca. 10°15'– 10°20' N), but similar smaller plants are also found at the north end of the main Cordillera Oriental, about 300 km to the south. These have leaf, peduncle and sheath lengths approaching those of the Perijá plants, and in spite of their small capitula, the flowers are more numerous than in typical P. pilosus (up to 40 per capitulum), and pistillate and staminate whorls alternate in the capitulum. It isn’t clear whether to treat these plants as small individuals of Paepalanthus sp. A, or intermediates with P. pilosus. Smaller individuals are also found at Sa. de Perijá (Cuatrecasas 25027, 25143,
The plants of Perijá had been distributed in part as P. macarenensis, a species otherwise only known from ca. 800 m in the Sierra de la Macarena (Meta). I have seen an image of the P. macarenensis type, and do not believe it is the same species or closely related, but pending closer examination treat the Perijá plants only provisionally here.
COLOMBIA. Cesar: [Serranía de Perijá], Paramo de Sabana Rubia, 3250 m, 22 Jul 1987, H. Cuadros 3732 (
COLOMBIA. Norte de Santander: de La Laguna a Nariz de Judío (Mutiscua), 19 Jun 1946, M. de Garganta 1209 (F); Santander: Paramo de Las Vegas, 3700–3800 m, 20–21 Dec 1926, E. P. Killip & A. C. Smith 15626 (F); Paramo de Santurban, between Tona and Mutiscua, 3800–4300 m, 18 Feb 1927, E. P. Killip & A. C. Smith 19557 (F).
Paepalanthus karstenii Ruhl., Pflanzenr. IV. 30: 155. 1903.
Type: Colombia. Cundinamarca: Páramo de Chipaque, [yrs 1848–1856], H. Karsten s.n. (holotype: W, destroyed [F neg. no. 29991]; lectotype, here designated: B [B_10_0243950]; isotype:
Dupatya karstenii (Ruhl.) Gleason, Bull. Torrey Bot. Club 53: 195. 1925.
Based on Paepalanthus karstenii Ruhl.
These unusual specimens with large capitula, broadly rounded bracts, coarsely ciliate leaves, and irregular leaf lengths appear intermediate between P. pilosus and P. dendroides as to bract characters and habit, and were collected at an elevation where distributions would potentially overlap. A topotype (Paramo de Chipaque, 3000 m, Dec 1855, J. J. Triana 1022-5,
Eriocaulon caulescens Willd. mscr., non Poir. nec Hook.f. & Thomson ex Thwaites. Based on: Humboldt & Bonpland s.n. (B-Willd. 2366 [BW02366000]), annotated as E. caulescens on sheet and folder. Presumed syntype of E. dendroides Kunth in H.B.K.
= Paepalanthus dendroides (Kunth) Kunth
Eriocaulon parvum Ruíz & Pav. nom. nud., non Körn. Cited by H. Ruíz (1940, p. 230). Annotated sheets: PERU. [Huánuco:] Saxiapata, Ruíz & Pavón s.n. (
= Paepalanthus dendroides (Kunth) Kunth
Paepalanthus pilosus var. microcephalus Moldenke, Phytologia 55(6): 372. 1984.
= Paepalanthus lamarckii Kunth, in P. ser Leptocephali (Ruhl.) Giul.
Synonymy according to
I thank the Field Museum Department of Botany and Keller Science Action Center for research space and use of equipment. I am especially grateful to Robin Foster for photos of specimens at
Paepalanthus pilosus complex, exsiccatae list
Data type: Text file
Paepalanthus pilosus complex, specimen database
Data type: Excel spreadsheet. List of examined specimens and geo-references of the localities
Explanation note: Contains complete label information of all examined specimens, as well as estimated geographic coordinates (in decimal degrees) for all specimens.