Paepalanthus lodiculoides Moldenke

Hensold, Nancy, 2016, The Andean Paepalanthus pilosus complex (Eriocaulaceae): a revision with three new taxa, PhytoKeys 64, pp. 1-57 : 28-32

4. Paepalanthus lodiculoides Moldenke Figs 2E , 10

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: NY [NY00103700]; isotype: F).

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: LL; isotypes: COL [COL000006924], US [US00088389]).

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: COL [COL000006910]; isotype: LL, U [U0007809], US [US00088365]).

Type.

Colombia. Boyacá: Nevado de Cocuy, Alto Valle de las Lagunillas, 4000 m, 12 Sept 1938, J. Cuatrecasas & H. García Barriga 1537 (holotype: US [ US 00088366]; isotypes: BC [ BC638454 ], COL [COL000006909], F, NY [NY00102897], P [P00741963]) .

Description.

Plants compact densely branched mosslike cushions, reportedly up to 23 cm in diameter (Soderstrom 1262) and 2 cm high ( Pedraza-Peñalosa et al. 2005). Leaves 1.7-5 (-6.5) mm long including broad basal sheath, the sheath often com prising half or more of the leaf length, the lamina subulate, dark to pale green, ca. 0.25-0.35 mm wide in the middle when mature, apex sharply acute (Loja) to subacute or minutely rounded; leaves densely congested along the short stems, and often half buried in deep woolly pubescence of the stem and lower leaf cilia, the tips glabrous. Peduncles (2-) 5-11 mm long, usually exsert 1 mm or more from sheaths and leaf mat at anthesis, obscurely 3-costate and often scurfy-pilose in lower half, rigid, subterete, and glabrous at apex. Peduncle sheaths (1.7-) 3-5 mm long, with an oblique scarious, sharp-acuminate to irregular or bifid apex, glabrous or obscurely tufted apically, margins eciliate (but cf. Syngonanthus steyermarkii type, see below). Capitula (1.4-) 2.5-3 mm in diameter. Involucres about equaling flowers; involucral bracts ovate to orbicular-ovate, dark black-brown throughout or paler brown along midvein, tufted at apex with clavate to linear, smooth to slightly tuberculate trichomes. Flowers ca. 4-6 per capitulum, sex ratio of capitula varying widely, from flowers all male to mostly female to some mixture of the two, even on same plant, the few flowers mostly peripheral, subtended by broad upper involucral bracts; receptacular bracts only rarely produced, these narrower and more oblong than involucral bracts, and carinate at base. Pistillate flowers: Pedicels sclerified, blackish, 0.1-0.15 mm, persisting on receptacle as bumps. Sepals broadly elliptic to suborbicular, strongly rounded-cymbiform in fruit, 1.2-1.7 mm long by ca. 0.65 mm wide at middle, 0.35-0.45 mm wide at base, deep blackish brown, sometimes with a pale medial streak, tufted with trichomes at apex. Petals oblong-obovate to broadly spatulate, acute-erose to acuminate, 1.1-1.6 mm × 0.4-0.7 mm, cream to nearly black, the distal half moderately pilose on both surfaces in two submedial or submarginal bands. Gynoecium at anthesis with ovary ca. 0.3 mm, style column 0.3 mm, nectaries ca. 0.35-0.6 mm, the glandular portion about equaling the stalk, clavate, the papillae soft and membranous, concentrated at apex but scattered along outside, colorless or tinged orange-brown at base, style branches 0.7-0.9 mm, brownish. Seeds 0.55-0.6 mm long, reticulate with short pseudotrichomes; locule wall thin, dehiscent or in some specimens observed adhering to the seeds, and the locules splitting apart without dehiscing (perhaps only in dry material). Staminate flowers: Pedicels 0.15-0.25 mm, brown. Sepals broadly spatulate, blackish toward apex, tufted; corolla with narrow membranous anthophore and broad tube with acute to acuminate lobes; anthers cream, exsert; the nectaries only half-equaling the corolla sinuses.

Phenology.

In Peru, flowering May-June (early in dry season); in Ecuador, September; in Colombia and Venezuela, July-Sept and Nov-March. Pedraza-Peñalosa et al. (2005) reports flowering times similar to Paepalanthus pilosus at Chisaca, corresponding to the "little dry season."

Distribution.

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. 8 )

Habitat.

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 ( Madriñán and Zapata 2001, Pedraza-Peñalosa et al. 2005). However Cleef (1981) reports Paepalanthus lodiculoides var. floccosus from the "upper zone of atmospherically dry [bunchgrass] páramo,” on the west side of the Eastern Cordillera, and the typical variety from "lower bamboo páramo” on atmospherically wet (east) slopes.

Conservation notes.

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.

Discussion.

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 Paepalanthus caryonauta . According to Madriñán and Zapata (2001) this species forms larger and more compact cushions than Paepalanthus pilosus at Páramo Chingaza, Cundinamarca. Pedraza-Peñalosa et al. (2005) describe the cushions as less than 2 cm tall, or only about half as tall as those of sympatric Paepalanthus pilosus (" Paepalanthus karstenii "). The branch architecture is similar to the other species but the dense cushions are sometimes distinguished by the many soft, nearly upright stems tightly packed together, possibly on wetter sites. Other individuals have shorter, more rigid stems and shorter branchlets. There are also pronounced differences in the amount of stem pubescence and leaf coloration. A densely white-woolly form with long hairs on the stems and lower leaf margins was described as Paepalanthus lodiculoides var. floccosus , but treated as a synonym by Luteyn (1999). That synonymy is provisionally accepted here, since other characteristics are the same, and the difference in pubescence seems likely due to environmental variation. A photograph of mature ring-shaped cushions of the floccose form was published by Moldenke (1976, p. 50).

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. Pedraza-Peñalosa et al. (2005) describe capitula with staminate flowers peripheral, but this was not observed in herbarium material.

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 Luteyn (1999) at my suggestion.

Additional specimens examined.

COLOMBIA. Bogotá, D.C.: Sumapaz, Páramo de Chisacá, 3900 m, 9-11 Nov 1958, Barclay & Juajibioy 6113 (F, MO); 22 Mar 1959, Barclay 7183 (MO); Parque Natural de Sumapaz , La Union - La Pedregal 3350-3720 m, 13-15 Jan 1997, Betancur 6935 (MO); Páramo Chisacá, 3680-3700 m, 16 Sep 1961, Cuatrecasas & Jaramillo 25987 (F); Andabobos, 3720-3760 m, 8 Jan 1969, Cuatrecasas & Idrobo 27054 (F); Páramo Chisacá, 3596 m, 26 Sep 1966, Soderstrom 1262 (MO); Lagunas de Chisaca , 3600 m, 15 Feb 1964, Uribe 4672 (MO) . Boyacá: Nevado del Cocuy, valle de Las Lagunillas, 4000-4300 m, 12 Sep 1938, Cuatrecasas 1537 (F); Paramo del Pedrisco , km 270 Sogamoso - Pajarito, 3000 m, 27 Aug 1953, Langenheim 3589 (F, MO) ; Santander: Paramo de Almorzadero , 3600 m, 1 Jan 1960, Barclay 10412 (MO) . ECUADOR. Loja: Muletrack Amaluza - Palanda , 3350-3450 m, 22 Sep 1976, Øllgaard & Balslev 9717 (F) . PERU. Piura: Huancabamba, El Talanco, 13 Jun 1961, Acleto 572 (USM [photo]), 3400 m, 12 Jun 1961, Friedberg 256 (USM [photo]); Huancabamba, Huaringas, 3957 m, May 1984, H. Ochoa 13 (USM [photo]) .

Figures

Figure 2. Capitula and leaf tips of Paepalanthus pilosus and relatives A-E Fruiting capitula with diaspores; sepals detaching (A, D) or intact (C, E) A Paepalanthus dendroides B-C Paepalanthus caryonauta D Paepalanthus pilosus var. pilosus E Paepalanthus lodiculoides F-J Paepalanthus pilosus var. pilosus, involucral bract variation. Arrows = tips of the torn peduncle sheath. (A Davidse 28991 B Barclay 5176, Ecuador C Dudley 11060, Peru D Cuatrecasas 25574 E Ollgaard 9717 F Cuatrecasas 25882 G Cuatrecasas 5553 H Ollgaard 9557 I Cano 16840 J Jorgensen 2366.)

Figure 8. Distribution of Paepalanthus dendroides, Paepalanthus huancabambensis, and Paepalanthus lodiculoides.

Figure 10. Paepalanthus lodiculoides. A Habit B-H Pistillate flower and fruit B Petal, adaxial C Petal, abaxial D Gynoecium (ovary abortive, atypical.) E Mature diaspore F-G Diaspore (F) with sepals removed (G) H Seed. (A Cuatrecasas 25987 B-D Barclay 6113 E-G Ollgaard 9717 H Steyermark 57372)