Aliso: A Journal of Systematic and Floristic Botany Aliso: A Journal of Systematic and Floristic Botany Uncinia (Cyperaceae) of Ecuador Uncinia (Cyperaceae) of Ecuador

Nine species of Uncinia (Cyperaceae: Caricoideae) are recorded from Ecuador, one of which, U. ecuadorensis, is newly described and illustrated here. Descriptions, illustrations, distribution maps, and both artificial and vegetative keys are provided for the nine species, and for some uncinias additional taxonomic, phytogeographic and ecological comments are made. A lectotype is designated for the name U. lenuis.


INTRODUCTION
All of the 60 to 70 species of Uncinia Pers. (Cyperaceae: Caricoideae) occur south of the Tropic of Cancer, with about 25 growing in the mountains and cooler regions of South America (Wheeler and Goetghebeur 1995). Of the ten Uncinia taxa (nine species and one variety) previously reported from the northern half of the continent (Kunth 1837;Kiikenthal 1909;Steyermark 1951;Wheeler and Goetghebeur 1995;Wheeler 1995), we recognize eight species as occurring in Ecuador; in addition, U. ecuadorensis G. A. Wheeler & Goetghebeur is newly described and illustrated here. This paper represents the first comprehensive treatment of the Ecuadorian species of Uncinia. Based on early collections made by William Jameson, Aloysius L. Sodiro, and Richard Spruce, Clarke (1883) and Kiikenthal (1909) reported two species and one variety, U. hamata (Sw.) Urban, U. phleoides (Cav,) Pers. and its var. nux-nigra C. B. Clarke, whose type was collected in Ecuador. In 1988, Lregaard (Cyperaceae of Ecuador, unpublished) recorded an additional species, U. tenuis Poeppig ex Kunth, and also made reference to two unknown species. Based on many new collections, four new species have recently been described from northern South America, U. lacustris G. A. Wheeler, U. paludosa G. A. Wheeler & Goetghebeur, U. subsacculata G. A. Wheeler & Goetghebeur, and U. tenuifolia G. A. Wheeler & Goetghebeur, all of which whose types come from Ecuador (Wheeler and Goetghebeur 1995).
All Ecuadorian Uncinia are montane or paramo plants (Table 1), with essentially none growing below 1200 m. The paramo uncinias, all of which are diminutive in stature, grow in moist to wet places on high, wind-swept plains, which are dominated by low shrubs and herbs, particularly grasses and sedges. Indeed, U. ecuadorensis, U. lacustris, U. paludosa, and U. macrolepis Decne. seldom occur below 3500 m and the last-named species is known to reach as high as 4600 m. The remaining five species grow at lower elevations, mostly in montane rain forest, although U. tenuifolia seems to be confined to steep, calcareous cliffs. By far the most frequently collected species are U. hamata and U. phleoides, both of which grow in forests and in partially disturbed sites. Of the nine species recorded from Ecuador, four are known only from that country: viz., U. ecuadorensis, U. lacustris, U. subsacculata, and U. tenuifolia, with the last two known thus far only from their types. Parenthetically, no members of the subfamily Caricoideae, such as Uncinia, occur in the nearby Galapagos Islands (Koyama 1971).
The members of Uncinia are characterized by perigynia (transformed prophylls) that contain a greatly exserted rachilla, the latter being invariably terete and tipped by a retrorse, inrolled scale (a transformed glume of a reduced male flower) that forms an "un- Table 1. Schematic of the relative occurrence, at increasing altitudes, for nine Uncinia species in Ecuadorian montane rain forest and paramo. cus" or hook (Snell 1936;Kukkonen 1967). Although it has been debated, pro (Savile and Calder 1953) and con (Hamlin 1959), whether the rachilla of Uncinia developed de facto as a dispersal mechanism from a simple seta, it is abundantly clear that the rachilla does indeed serve as an agent of dispersal. According to most authors (Croizat 1952;Nelmes 1952;Hamlin 1959), Uncinia most likely originated in the Southern Hemisphere. Because it is believed that the Cyperaceae originated in the late Cretaceous or early Tertiary, with rapid diversification of the major groups after that (Ball 1990), differentiation within the genus probably took place sometime in the Tertiary. Uncinia species may be either protandrous or protogynous (Edgar 1970) and most, if not all, species are wind-pollinated (anemophilous). However, very little is known about the reproductive biology of South American uncinias and no chromosome number based on Ecuadorian material has been reported. Although both natural and artificial hybrids have been reported from New Zealand (Hamlin 1959;Edgar 1970), essentially nothing is known about the hybridization of Uncinia in South America.

Montane Rain Forest Paramo
Like most genera of the Cyperaceae, mature fruit is sine qua non for the positive identification of Uncinia species. The primary taxonomic characters useful in the systematics of the genus are: deciduous versus persistent scales: perigynia vestiture or the lack of it; and staminal filament width compared to anther width. Other useful features are spike shape and density and variations in perigynia, and include size, shape, color, position and density of marginal hairs, and beak characteristics. Still other useful features are achenes, rachillae, rhizomes, anther length, and scale and leaf characters. On the other hand, some features are unreliable as taxonomic characters. For example, a sterile bract may be present or absent from different spikes of the same plant. Also, some features of the perigynium, such as differences in texture, veination, winging of the margins, and orifice shape, are only of limited importance in the taxonomy of the genus (Hamlin 1959). Moreover, in the Ecuadorian plants coloration of the basal sheaths vary little among species.
Two keys to the Ecuadorian Uncinia are provided below. The Artificial Key is based primarily on flowering and fruiting material and thus provides the most reliable means of identifying the material at hand. The second key is based primarily on vegetative characteristics, although habitat information has also been utilized. The Vegetative Key was constructed to provide a means of identifying Uncinia material when no flowering or fruiting parts are available. For instance, the latter key has been helpful in identifying sterile plants mounted on herbarium sheets. Issued as a caveat, however, vegetative material of Uncinia is often difficult to distinguish from other cyperaceous genera, particularly Carex L. (Wheeler 1994). Nevertheless, it is apposite to note that while many Carex species have long internodes with conspicuous cauline leaves, most Uncinia species have short internodes so that the leaves are basal (Hamlin 1958;Kukkonen 1967). Also, the basal sheaths of many Carex species are strongly reddishor purplish-tinged, whereas all Ecuadorian Uncinia have pale brown to brown or, more often, dark reddish brown basal sheaths. Although the keys are largely self-explanatory, explanations for some characters are as follows. In the Artificial Key: 1) rachilla length was measured unidirectionally from the point of attachment (at the achene base within the perigynium) to the "end" of the terminal hook (i.e., the length of the descending portion of the hook was excluded, though that dimension is given separately); and 2) spike width was measured from the tip of the perigynium (or pistillate scale) on one side of the rachis to that on the opposite side; hence, rachilla length (and its concomitant width) was excluded. In both keys, the measurements of the widest leaves include dead leaves and other leaf remnants at the base of the plant.
Although this species (Fig. 13) resembles Uncinia erinacea (Cav.) Pers. and somewhat less so two newly described South American uncinias (Wheeler 1997), all of which occur in Chile, it differs from them by having smaller and differently shaped perigynia and achenes, shorter rachillae, narrower leaves, and acute pistillate scales. Moreover, whereas the culms of the three above-mentioned Chilean species have long internodes and conspicuous cauline leaves, the internodes of U. ecuadorensis are short so that the leaves are basal. If one assumes that U. erinacea and U. ecuadorensis are closely related, then the robust U. erinacea, with its leafy culm, broad leaves, and longer rachillae, seemingly is the more "primitive" of the two. In this regard, Hamlin (1958, p. 85) considers U. erinacea to be "the most primitive living member of the genus." Like U. erinacea, the new species belongs in section Platyandrae. Plants cespitose, from short-to long-creeping rhizomes. Fertile culms 20-60 cm tall, 0.5-1 mm thick, erect or somewhat curved, from shorter than to exceeding the leaves, obscurely trigonous, smooth, with glabrous, brownish sheaths. Leaves 5-7, the uppermost ones cauline with conspicuous sheaths and long internodes; blades (4.5-)8-50 cm long, 2.4-6.4 mm wide, flat, membranaceous, flaccid, glabrous, the margins antrorsely scabrous, terminating in a scabrous attenuate tip; leaf sheaths up to 6.5 cm long, more or less tightly enveloping culm, glabrous, green; inner band glabrous, greenish below and whitish green or pale reddish brown above, mouth slightly thickened and often dark reddish brown, the apex concave; ligules 2-6 mm long, rounded to acute. Inflorescence a solitary, androgynous spike, (4.5-)7-16 cm long, 2-4 mm wide, linear to cylindric, subclavate. Staminate part 6-16 mm long, 1O-30-flowered; scales 1.8-3.2 mm long, 0.8-1.8 mm wide, obovate, obtuse to subacute, coriaceous, glabrous, greenish center and stramineous or pale brown margins, with an inverted V-shaped reddish brown strip near the apex, 1-3veined, the tips with very narrow hyaline margins and ciliolate. Pistillate part tightly flowered, with ca. 25-100 perigynia; scales persistent, 3.3-5.8 mm long, 1.5-2.8 mm wide, shorter than to about equaling the perigynia, oblong-obovate, obtuse to subacute, coria-ceous, glabrous, green or greenish brown, with an inverted V -shaped reddish brown strip near the apex, 9-13-veined, the tips with very narrow hyaline margins and ciliolate. Perigynia 4-6.2 mm long, 1.3-1 .8 mm wide, oblong-obovate or elliptical, appressed hispid distally, smooth or sparsely hispid proximally, the margins ciliate from apex to near the base with the longest hairs distally, strarnineous or pale brown to brown, 2 veins prominent and the rest faint, tapered to a stipitatelike base; perigynium beak conical, appressed hispid, the margins densely ciliate. Achenes 3.2-3.8 mm long, 1.2-1.6 mm wide, compressed-trigonous with more or less flat, oblong sides, tightly enveloped by the perigynium, brownish, sessile. Rachilla 6-11.5 mm long, terete, projecting beyond orifice of perigynium, the exserted portion up to 6.5 mm long, but sometimes the uppermost ones mostly hidden by the scales, usually geniculate in the distal half, stramineous or brownish, the hook 1.3-1 .6 mm long and whitish green or pale brown to brown (particularly the descending part). Stigmas 3; style base somewhat thickened. Anthers 3, 0.8-1.6 mm long, ca. 0.2 mm wide; filaments linear, dilated (0.2-0.3 mm wide), as wide as or wider than the anthers.
Uncinia hamata (Fig. 3) occurs from northern Argentina (Barros 1947) northward to Mexico and the Caribbean islands of Jamaica and Hispaniola (Ktiken-thaI 1909;Foster 1965;Mora-Osejo 1966;Chater 1994; Wheeler in press); as such, it is the northernmost-occurring Uncinia in the Western Hemisphere. In Ecuador (Fig. 14), where it is quite common, it grows in loose to dense tufts in montane rain forest, at elevations from about 1200 to 3400 m, growing primarily in shaded places, such as on the forest floor and in ravines; however, it also grows in partially shaded sites, such as along river banks and adjacent to trails. Specimens with ripe fruit have been collected from February through October. The epithet refers to the hooked fruits of this species.
Uncinia lacustris is known only from two sites in north-central Ecuador (Fig. 14), where it grows in paramo at elevations from about 3900-4100 m (Wheeler and Goetghebeur 1995). At the type locality, it was growing on the margins of a small lake with other cyperaceous plants, such as Carex and Rhynchospora. At Volclin Cotocachi (see Fig. 15) it grows in pajonal with the rarity U. ecuadorensis. Plants with ripe fruit have been collected in late March and early June. The epithet refers to the occurrence of this species on the wet margins of lakes.
Uncinia macroLepis occurs in northern South America, in Patagonia and Tierra del Fuego, and on the south-Atlantic islands of South Georgia and Tristan da Cunha (Wheeler 1994(Wheeler , 1995 Wheeler (1995) has demonstrated that the populations from northern South America, South Georgia, and Tristan da Cunha are morphologically indistinct from the Patagonian-Fuegian populations of U. macroLepis, whose name has priority.
In Ecuador (Fig. 14), this species occurs near the upper limit of paramo, at elevations between 4000 and 4600 m, where it grows on wet rocks and with cushion plants. particularly along rocky lake shores and on rock ledges. It is probably best known from Cerro Antisana (see Fig. 15) on the border between Pichincha and Napo provinces. Plants with mature fruit have been collected essentially throughout the year. The epithet refers to the broad scales of this species (Fig. 2).
Because paramo plants are typically dwarfed and compact. different species may physiognomically look very similar. For instance, Uncinia macroLepis ( Fig. 1) resembles U. paLudosa and somewhat less so U. Lacustris, but differs from both of them in several morphological features. Indeed, U. macroLepis belongs in section Uncinia (Wheeler 1995), whereas the other two species belong in section PLatyandrae. In regard to differences that can be observed in the field (with the aid of a hand-lens), the culms of U. macroLepis are typically roughened (i.e .• scaberulent) beneath the inflorescence. whereas those of U. Lacustris and U. paLudosa are smooth throughout. However. in post-mature culms of U. macroLepis this character is sometimes difficult to observe. Also, U. macroLepis has slender rhizomes and short leaves that are wide spreading and often strongly curved, whereas both U. Lacustris and U. paLudosa have thickish rhizomes and leaves that are longer and mostly erect and little curved. It is also noteworthy that U. macroLepis is the only Ecuadorian Uncinia with a conspicuously enlarged style base, though that of U. hamata is somewhat thickened.
Uncinia paLudosa is known from many sites in Ecuador (Fig. 14) and several specimens have been seen from Arauca and Boyaca provinces in northeastern Colombia (Wheeler in press); a single collection is also known from Rio Abfesco National Park in Peru (Leon & Young 1663 [NY]). This species occurs in paramo, at elevations from about 3500 to 4400 m (Wheeler and Goetghebeur 1995), growing primarily in bogs and swamps dominated by low shrubs and herbs, particularly grasses and sedges. Flowering plants have been collected in February and those with mature fruit from April through early September. The epithet refers to the occurrence of this species in marshy habitats.
Uncinia phleoides (Fig. 8) ranges from northern Patagonia (Barros 1947) northward to Colombia (Kiiken-thaI 1909;Foster 1965;Wheeler in press) and is also reported from Mexico (Gonzalez 1983). In Ecuador (Fig. 15), where it is common, it occurs at elevations from about 2500 to 4200 m and grows in large, dense tufts in scrub forest and clearings and in disturbed sites (e.g., in roadside ditches and along trails). Among other localities, this species is well known from the slopes and environs of Chimborazo, Cotopaxi, Pasochoa, and Pichincha volcanoes (see Fig. 15). Specimens with ripe fruit have been collected essentially throughout the year. The epithet, which means "phleumlike," refers to the timothylike spikes of this species (Fig. 9).
Uncinia subsacculata is known only from the type locality (Fig. 15). It seemingly is a scotophilous (shade-loving) species that grows in Polylepis forest, at about 3800 m. Plants with well-developed perigynia have been collected in early June. The epithet refers to the very short, saccate appendages that persist after the deciduous scales have fallen.
This species (Fig. 12) resembles Uncinia tenuis but differs in several features, the most salient of which are given in the keys. It is noteworthy that these two South American uncinias are readily identified by their distinctive inflorescences, which, although greatly (or entirely) divested of perigynia, continue to display few to numerous saccate appendages, structures which are persistent portions of the otherwise deciduous scales (Wheeler and Goetghebeur 1995). However, in U. tenuis the proximal one-third of the scale persists as a conspicuous appendage (about 1 mm long), whereas in U. subsacculata the saccate appendage is appreciably shorter (less than 0.5 mm long . ".  0.3-0.5 mm thick, erect or slightly curved, from shorter than to exceeding the leaves, obscurely trigonous, smooth, with glabrous, brown basal sheaths. Leaves 3-7, basal; blades 4-17 cm long, 0.6-1 .5 mm wide, more or less spreading, flat or channeled (especially in the proximal half), membranaceous, glabrous, the margins antrorsely scabrous distally, terminating in a long, scabrous attenuate tip; inner band of leaf sheaths hy-aline or pale brown, glabrous, the apex concave; ligules 0.4-0.8 mm long, rounded. Inflorescence a solitary, androgynous spike, (7-)12-22 mm long, 1.5-2 mm wide, narrowly linear. Staminate part 4.5-8 mm long, 3-11-flowered; scales 2-2.8 mm long, 0.8-1.6 mm wide, obovate, obtuse to subacute, glabrous, brownish, 1(-3)-veined, the tips with hyaline margins and ciliolate. Pistillate part more or less tightly flow-ALISO ered with 3-15 perigynia; scales persistent, 2.5-3.4 mm long, 1.2-2 mm wide, from shorter than to about equaling the perigynia, oblong to obovate, obtuse to acute, subcoriaceous, glabrous, pale brown to brown, 5-7-veined, the tips with hyaline margins and ciliolate, the lowermost one infrequently with a scaberulent awn up to 9 mm long. Perigynia 2.5-3.3 mm long, 0.8-1.2 mm wide, elliptical, appressed hispid distally, smooth or sparsely hispid proximally, the margins ciliate from apex to near the base with the longest hairs distally, strarnineous or brownish, 2 veins prominent and the rest faint, tapered to the base; perigynium beak: conical, appressed hispid, the margins densely ciliate. Achenes 1.8-2.2 mm long, 0.9-1.1 mm wide, compressed-trigonous with slightly convex, oblong sides, tightly enveloped by the perigynium, brownish, sessile. Rachilla 5-7.6 mm long, terete, projecting beyond orifice of perigynium, the exserted portion (1.8-)2.5-4.6 mm long, smooth, strarnineous or pale brown, the hook 0.7-1.3 mm long and stramineous or brownish (particularly the descending part). Stigmas 3; style base little thickened. Anthers 3, 1-1.4 mm long, ca. 0.2 mm wide; filaments linear, dilated (ca. 0.2 mm wide), as wide as or wider than the anthers. This scopulicolous . (or cliff-growing) species is known only from the type locality in southern Ecuador (Fig. 15). It seemingly requires special edaphic conditions, growing in very thin soils on precipitous cliffs, where it forms small, dense cespitdse clumps. The type collection was made from a calcareous rock face near a cave. Plants with mature fruit have been collected in early March. The epithet refers to the narrow leaves of this species.
Uncinia tenuis ranges from Cape Hom in Tierra del Fuego (Wheeler 1994) northward to west-central Argentina and central Chile (Barros 1969;Munoz-Schick 1980) and is disjunct on the Juan Fernandez Islands (Skottsberg 1922); further north it occurs in Ecuador (Fig. 15) and Colombia (Wheeler in press) and is also reported from Costa Rica in Central America (Chater 1994). In Ecuador, this species (Fig. 10) grows at elevations from about 3100 to 4000 m with several collections seen from Cerro Sumaco (see Fig. 15) in Napo Province. It grows in loose mats in moist, shaded places on the forest floor and also grows epiphytically on fallen trees. Specimens with ripe fruit have been collected from April through November, though by late November the majority have been shed. Even then, however, this species is easily identified by its distinctive inflorescence axes (Fig. 11), which although greatly (or entirely) divested of perigynia continue to display whitish or pale brown saccate appendages, structures which are the persistent portions of otherwise   Fig. 15. Distributions of Uncinia ecuadorensis. U. phleoides. U. subsacculata. U. tenuifolia. and U. tenuis in Ecuador; selected localities in Ecuador mentioned in the text. (The two paratypes of Uncinia ecuadorensis [small closed circles] were examined and mapped after the maps were originally constructed.) deciduous scales (Wheeler 1994). The epithet, which means "slender" or " thin," refers to several features of this species (e.g., its narrow rhizomes, cuims, and spikes).
This species resembles U. subsacculata but differs by having shorter perigynia, achenes, rachillae, anthers, and scales. Also, the perigynium beak of U. ten-uis is shorter and broader than the beak of U. subsacculata. Notably, these are the only two South American uncinias that have deciduous scales and persistent saccate appendages. Like U. subsacculata, this species belongs in section Uncinia.