A Systematic Study of the Subtribe Espeletiinae. Heliantheae (taxonomic keys) - Cuatrecasas 2013

CHAPTER 13. CHARACTERIZATION OF SUBTRIBE ESPELETIINAE CUATRECASAS Espeletiinae Cuatrec., Phytologia 35: 48. 1976.

Pachycaulous, perennial, and perennifoliate plants mostly with xerophytic structures. Trees, woody shrubs, and caulirosulas, monocarpic or polycarpic.

Stems solidly woody or mostly with a broad central pith. Leaves usually more or less crowded at the distal ends of the stem or branches or many congested in sessile or stipitate rosettes (or caulirosulas).

Leaves spirally arranged, the laminae rather large, entire (or minutely denticulate), rather oblong, pinnately veined, markedly xeromorphic, coriaceous or subcoriaceous, with areolar cavities, with hypodermis, and usually covered at least abaxially by dense indument. Foliar rays with compact files of procumbent cells.

Synflorescences monotelic, either dichasial or monochasial, terminal or axillary, bearing capitula varying in number from 1 to a few hundred.

Capitula heterogamous. Involucres pluriseriate; phyllaries subherbaceous or subchartaceous; outer ones sterile, 5-20(-50), ovate to oblong or linear, acute or obtuse, subequal or usually the external ones larger size and decreasing gradually inwards, abaxially more or less densely hairy; fertile phyllaries becoming less herbaceous inwards , indurate at base at maturity, embracing the achenes, at least distally hairy. Receptacle paleaceous, flat, convex, or conic, rarely semiglobose, glabrous or hirtulous; pales persistent, scarious or subscarious, embracing the flowers, distally barbate.

Ray flowers female in 2-6 rows, ligulate, seldom eligulate; corolla tubes usually hairy; ligules basically white or yellow; style branches with 2 parallel, discrete, marginal, stigmatic lines. Disc flowers pseudohermaphroditic, functionally male, with rudimentary ovary; corollas pentamerous, tubular with the limb tubular, campanulate, or slightly infundibuliform, shortly lobed or dentate, hairy or glabrous, usually yellow; anthers shortly sagittate, endothecial cells oblong, nodular at the transverse walls, apical appendages ovate with a median keel and no glands; styles distally papillose-pilose with subapical crown of longer papillae, nonstigmatic, apex conical, emarginate, base immersed in a tubular nectary (sometimes called a disc).

Achenes obovoid-pyramidal or oblong, sharply triangular to prismatic-quadrangular; apex rounded or truncate, base mostly acute or subacute, callose; abaxial side Âą convex, the other sides flat; surface

glabrous, smooth with no striations; cells of subepidermis with amorphous traces of a black pigment ("carbonized"); no pappus except for 3 sublanceolate squamellae in Tamania. .

Schizogenous resiniferous ducts and cells distributed generally throughout the plant except in the woody tissue; the ducts usually occurring along the major veins of the leaves and corollas. Gland-tipped hairs particularly rich in tetracyclic diterpenoid compounds of the kauranoid molecular structures.

Growth rings of the wood more or less marked or completely lacking, reflecting climatic oscillations in degree of dryness rather than seasonal periods. Tracheids present in arboreal species. Pore diameters larger in arboreal species (50-80 J.tm) than in rosulate species (28-61 J.tm). Vessel elements with simple vestigially bordered perforations, reticular unbordered perforations, and sometimes a few scalariform perforations with conspicuously bordered bars. Vessel lengths from 126-473 J.tm. Intervascular pitting mostly alternate, also scalariform and transitional, rarely opposite. Tyloses scarce, thin-walled, rarely abundant and with thick walls (Carramboa). Libriform fibers 2x longer than vessel elements, seldom septate. Woody uniseriate rays absent or scarce, multiseriate rays up to 10 cells wide. Ray cells usually 8- 90 x 10-25 J.tm in arboreal species, 90- 125 X 25-40 J.tm in caulirosulan species.

Pollen grains tricolporate, spheroidal prolate to spheroidal oblate with 22- 42 J.tm polar diameter and 22- 38 J.tm equatorial diameter. Colpi usually narrow, rather shallow, with acute ends and finely granulate membrane. Germ pores well defined, rarely circular, usually elongate, rather constricted at the polar axis, and equatorial ends acute or rounded. Exine thinner between spines. Sexine thick. Tectum spiny, foraminate, baculate. Spines 1.5-7.0 J.tm long, conical, acute or obtuse, with an internal cavity near the base; perforations near the base in l-3 circles, larger at the spine bases than between them. Columellae thin, circular, appearing capitate in thin section, united at base into a more or less homogeneous membrane. Infratectum lacking, replaced by a continuous cavus, the nexine united to the sexine only at colpi. Nexine apparently homogeneous, sometimes slightly differentiated into nexine 1 and 2, the internal surface granular.

Chromosome number n = 19.

The features of the Espeletiinae described above separate them readily from other subtribes of the

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114

A SYSTEMATIC STUDY OF THE SUBTRIBE ESPELETIINAE

TAMAN/A

RUILOPEZIA LIBANOTHAMNUS PARAMIFLOS

A

ESPELETIOPSIS ESPELETIA COESPELETIA

CARRAMBOA FIG. 13-1. Phylogenetic diagram of Espeletiinae genera: A. Prototype, a shrubby Heliantheae developing with spirally alternate leaves in primitive, pre-Andean area of Venezuela, probably in late Pliocene. B. Branched woody plants with tracheids , terminal reproductive meristem, monochasial synfloresences, achenes with vestigial pappus. C. Woody plants basically without tracheids, with lateral reproductive meristem , vestigial dichasial synftorescences, vestigial pappus lost.

D. Broadly branched trees with solid wood in stem, close tubular leaf bases, vestigial pappus lost. E. Branching stems with tracheids and solid wood, vestigial pappus retained. F. Woody stems with broader central pith, tracheids lost, branching reduced to monocaulous caulirosula, large apical synftorescences, vestigial pappus lost. G. Mostly monocaulous caulirosulas, a few vestigially branched trees, sy nflorescences reduced to monochasial type. H. Opposite branching retained (dichasial).

I. Branched tree form with thick woody stems retained, leaves with close annular bases, basic dichasial synftorescences retained. J. Reduction to monopodia] caulirosulas. K. Synftorescences permanently dichasial. L. Reduction of synftorescences to monochasial-racemiform type. M. Structure of involucre subtly changed. N. Basic involucra] structure retained.

tribe Heliantheae: the Polymniinae, Melampodiinae, and Ecliptinae. The major distinguishing features are: • The spiral leaf phyllotaxis.

• The obpyramidal to prismatic shape of the achenes, which also are smooth, without striations, carbonized, glabrous, and epappose (with one exception).

• The fertile, female ray flowers and functionally male disc flowers .

• The pluriseriate involucre and persistent pales of the receptacle.

• The thick and woody stem.

• The general xeromorphic structure and organization of the plants and various specialized life-forms.

• The chromosome number n = 19.

Figure 13-1 is a phylogenetic diagram showing changes in characteristics during the evolution of the 8 genera of the Espeletiinae, which can be distinguished by the following key.

Key to Genera 1. Synftorescences terminal, monochasial (with alternate branches), with many capitula.

2. Leaf bases tubular. Trees, rarely caulirosulas. Ray corollas white or creamy. Achenes epappose.

Receptacles hairless. Venezuela, Colombia ............................................................................ Libanothamnus 2. Leaf bases open, amplexicaulous or fiat.

CHARACTERIZATION OF SUBTRIBE ESPELETITNAE CUATRECASAS

11 5

3. Trees. Achenes with a pappus of 3 pales. Rays corollas yellow. Receptacles sparsely hirtulous.

Venezuela, Colombia ... ...... ... ... ... ..... ...... ... ....... .. ... ... ... ....... ......................... . ........ ... ... ............. . ....... Tamania 3. Caulirosulas, monocaulous and monoca rpic or slightly branched, each rosette monocarpic.

Achenes epappose. Ray corollas white, yellow, or rarely reddish or purpli sh. Receptacles hairless or spa rsely hirtulous. Venezuela................... ............................................................................ .......

Ruilopezia l. Synflorescences ax illary (lateral). Achenes epappose.

4. Leaf bases tubular or annular. Trees. Leaves usually large, rather broad, green. Ray corollas bright yell ow. Receptacles mostly hirsute. Synflorescences basically dichasial, bea ring many capitu la.

Venez uela .......... .................... .................................. ............. ..... ....... ................................................. .......... Carramboa 4. Leaf bases open, flat. Caulirosulas, monocaul ous, very tall to short, subsess ile or sessi le, seldom slightly branched. Ray corollas yellow, rarely white or purplish. Receptacles hirsute to hairless.

Synflorescences with number of capitu la vary ing from many to l.

5. Synflorescences dichasial, with branches, leaves, and bracts opposite, at least in the proximal part. Ray corollas yellow, rarely obsolete. Cau lirosulas normally monocaulous. Pollen grains with ll - 18 spines, 4 -7~-Lm long on equator (rarely with 18- 2 1 spi nes, 2-2.5 IJ.m long). Colombia, Ecuador, Venezuela ... Espeletia 5. Synflorescences monochasial, their branches, leaves, and bracts spira lly alternate.

6. Involu cres with gradual tran sition from sterile, outer phyllaries to inner, fert ile phyllaries and receptacular pales.

7. Synflorescences corymbiform-panicul ate. Ray corollas yellow, occasionally white or purplish, rarely obsolete. Caulirosulas, usually monocaulous, occasionally slightly branched. Pollen grains with 12-18 spines, (3.5 - )4- 7 IJ.m long on equator. Colombia, Venezuela .................. Espeletiopsis I. Synflorescence racemiform. Ray corollas yellow or orange, occasionally somewhat purplish. Caulirosulas, monocaulous. Pollen gra ins with 16-2 1 spi nes, 1.5-4 IJ.m long on equator.

Venezuela .................... ............ .......................................................... ........... .................... ......... Coespeletia 6. Involucres with sterile, outer phyll aries sharply separated from inner, fertile ones, followed by receptacular pales. Poll en grains with about 12 spines, 4.5-61J.m long on equ ator. Colombia ..... Paramifios

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A SYSTEMATIC STUDY OF THE SUBTRIBE ESPELETIINAE

straight, slightly thickened walls or thin sinuous walls; uniseriate xylem rays essentially absent, multiseriate xylem rays to 10 cells wide, ray cells 90- 125 f.Lm high, 25-40 f.Lm wide; mature pith rays broader with cell walls thinner and nonlignified; foliar rays present, procumbent cells prominent. Midrib with simple pattern, crescent-shaped with (1- )3- 9 vascular bundles; leaf lamina with areolar cavities, with hypodermis. (See Pls. 7-2, 7-4D, E, 7-5C, 7-VIB, 7-7C, 7-SA.) The key that follows relies on characters summarized in the preceding generic descriptions. The key

by Torres de Ricardi (1979) differed primarily by its initial emphasis on straight vessel elements 160 to 600 f.Lm long, radial arrangement of the pores and fibers, and vascular rays up to 100 f.Lm wide versus sinuous vessel elements 66 to 360 f.Lm long, nonradial arrangement of the pores and fibers, and vascular rays up to 300 f.Lm wide. This resulted in keying Ruilopezia into the group with the arborescent genera. His

key distinguished Libanothamnus from Tamania by the linear-oblique versus ample apertures of the intervascular pits, the abundant versus few or no procumbent cells, and the obsolete versus abundant tyloses .

Espeletiopsis was separated from Espeletia by vascular rays always heterocellular, marginal cells of the rays as high as the fibers, and vessel elements to 360 f.Lm long versus rays homocellular and heterocellular, marginal cells of the rays shorter than the fibers, and vessel elements to 180 f.Lm long. Some of the di stinctions at the generic level in both the Torres de Ricardi key and the following key may prove unreliable as more species are studied. This is particularly true in the case of the characters presented by Torres de Ricardi for Espeletiopsis, which were based on one species. Paramiflos is omitted. The genus is rosulate, and the leaf traces are multiple, but the vessel elements and fibers have not been observed adequately to allow further comparisons.

Anatomical Key to Genera Arborescent form s with pith small in compari son to surrounding wood , pith not tea ring when dry; fibers rather obviously radial in arrangement, mostly with straight, slightly thickened walls, sometimes septate; uni seriate xylem rays random and infreque nt, ray cells 80-90 f!m high , 10- 25 ~-tm wide, lignified.

2 Wood without distinct termini on growth rings, without tracheids at termini ; pore di ameter 79-97 ~J,m ; numerous thick-walled tyloses; multiseri ate xylem rays to 1276 f!m high ................................ ........ ....... Carramboa 2 Wood often with distinct termini on growth rings, with tracheid s at termini ; pore diameter 50-80 f!m ; tyloses variable, lacking or thinner-wall ed; multiseriate rays to 680 IJ.m high .

3 Vessel elements length averaging 380 ~J,m; intervascular pit size 3 -5~-tm .............. ..... ...... ...... ...... Libanothamnus 3 Vessel element length 235 - 255 ~J,m; intervascular pit size 4- 7 IJ.m ............................ ............................ Tamania Rosul ate form s or nonrosulate forms with pith large in comparison to surrounding wood, pith often tearing whe n dry; fib ers often not radial in arrange ment, often with sinuous walls, not or rarely septate; xylem ray cells 90-1 25 IJ.m high , 25 - 40 ~-tm wide, not or only partly lignified.

4 Vessel element length 159- 473 ~J.m , mostly over 250 ~J,m ; fibers rather obviou sly radial in arrangement ................................. ... ... ... ... ... ... ... .............. .............. ... ... ... ... .. .. ... ... ...... .......... .. ....... .. ..... Ruilopezia 4 Vessel element leng th 128- 337 IJ.m , averaging 200 f!m; fibers scarcely radial or ta ngential in a rrangement.

5 Fibers rather tangential in arrangement; nucleate fibers thin-walled and sinuou s; foliar rays with central procumbent cells very thin-walled; intervascular pit size 5- 8 ~-tm ......................................... Coespeletia 5 Fibers not or scarcely radial or tangenti al in arrangement; some fibers with strai ght, thickened walls and some with thin , sinuous wall s; intervascular pit size variable.

6 Midrib with intermediate, multiple arrangement of vascular bundles ................ .......... .............. Espeletiopsis 6 Midrib with simple arrangement of vascular bundles .................................. ............ ........ ................. Espeletia

ESPELETIA

133

COLOMBIAN POPULATION BRANCH: CORDILLERA OCCIDENTAL

Only a few scattered populations of Espeletia are found in this generally lower and very broken Cordillera, mainly in its northern part (Figs. 14-5, 14-6). Probably during the Last Glacial period, the lowering of the paramo limit established a direct link between the paramos and their vegetation from the Central to Occidental Cordilleras, like that between the paramos from the Cordillera Oriental to the Cordillera Central. Presumably, Espeletia could have traveled from Paramo de Las Papas along a chain of paramos following the irregular line dividing the watersheds of the Cauca River (north) from those of Rio Timbfo and Patfa (south) at about 2000m and above along the known hills of Almaguer and Sotara. Thus, it could have connected westwards with the paramos around Munchique in the Cordillera Occidental. Espeletia, probably a form of E. hartwegiana, extended from Munchique (north of the Pat fa break) to the north end of the Cordillera Occidental, producing E. praefrontina and E. frontinoensis, which remained scattered throughout isolated populations found on the highest isolated peaks of Paramo de Frontino, Paramo de Tatama, and Paramo de La Soledad (Cauca, Mpio. Argelia).

I postulated (Cuatrecasas 1986b: 296) that past volcanic acti vities had formed temporal bridges between Cordilleras Central and Occidental across the Cauca valley, allowing Espeletia to traverse from Cundinamarca to Antioquia at Santa Rosa. This is also a possibility, especially for E. occidentalis var. antioquensis, a taxon close to E. grandijlora. The other species in the Cordillera Occidental, E. praefron.tin.a and E.frontinoensis, are related more closely to E. hartwegiana. Later, with the considerable raising of the timberline during the Holocene, the paramos and populations of Espeletia were reduced again to the few isolated, high-altitude spots (Alto del Chaquiro, Paramo de Frontino, Cerro Tatama), which were broad enough to support a population along this otherwise very broken, narrow, low, mountain range.

ENVIRONMENT Espeletia develops exclusively in the microthermallife zo ne with small incursions into the cryothermal zone (see Chapter 4). It seems to be favored by every extreme physical condition of the habitat, in this case, a constantly or intermittently cold, humid environment with humic acid soil mainly in wet or marshy paramos. Under these circumstances, gregarious, monospecific populations have formed and cover extensive ecological areas, such as swampy fl at depressions; wet or humid valleys; and sometimes rocky, humid clefts. These populations may cover several squ are kilometers of paramo, but are often smaller because of topographic or ecological barriers that limit continued dispersal. The dominant communities of Espeletia that form around the apparently drier, upper part or peaks of mountains usually are surrounded by and limited at their base by the Andean forests covering the slopes below. In this way, populations on emerging mountains, hill s, or peaks become isolated. Other preferred habitats also are limited by fores ts. The bottoms of small to large, wet valleys and flat or moderately inclined grasslands quickly become covered by Espeletia communities, like islands completely surrounded by heavy Andean forest. Espeletia is the most representative genus of the paramos. Most of its species develop best in the proper paramo zone (about 3400-4200 m), but some also are found in the subparamo zone (about 32003400m), especially in mosiac distribution or in ecotonic communities and even in the microthermal forest. Espeletia is rather rare in the superparamo zone (above 4200 m), only approaching its lower border. A few species, e.g., E. cleefii and E. lopezii fma. alticola, may thrive in rocky habitats in the superparamo slopes of the Sierra Nevada del Cocuy (Colombia), as well as E. semiglobulata and E. schultzii in the Cordillera de Merida (Venezuela), defy ing the characteristic frost and sol ifluction of the ground.

Figures 14-7 and 14-8 show the altitudinal ranges of all species.

Key to the Sections of Espeletia I. Outer phyllari es, ovate, elliptic, triangular or broadly oblong, rarely narrower, length to width ratio 1-3:1.

Receptacles variously hairy or g labrous.

2. Tndume nt of leaf laminae patent-tomentose, subvelvety. Receptacles hirsute. Venezuela: Trujillo. Species No. I ....... ........ .......................................................................... . .............. ................................. I. Sect. Aristeguietana 2. Tndument of leaf lam inae sericeou s or lanate. Receptac les sparsely hirsute or glabrous.

3. Indument on both sides of lami nae sericeous, lustrous, the hairs stra ight, strongly appressed. Colombi a: Boyaca, C undinamarca. Species Nos. 2 and 3 ................................ .. ..................................... 2. Sect. Bonplandia 3. Indument on leaves lan ate.

134

A SYSTEMATIC STUDY OF THE SUBTRIBE ESPELETIINAE

ALTITUDINAL RANGE OF ESPELETIA SPECIES 1-29 ALTITUDE

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4. Receptacle mostly hirsute. Venezuela.

5. Capitul a sem iglobose short-radiate, the ray-corollas 3- 4.5mm long. Synflorescences consistently dichasia l. Indument lanate, appressed. Sp ines of po llen g rain s 2-2.5 1-lln, 18-21 on the equator.

Venezuela: Merida. Species No. 4 .. .................................................................................. 3. Sect. Badilloa 5. Capitul a campa nul ate, the ray co roll as 10-17m m long. Sy nflo rescences predominantly monochasial but at least basal leaves or bracts always opposite. Indument lanate shaggy.

Venezuela: Merid a. Spec ies No. 57 a nd some hybrids .. .. .. .. ...... ... .. ....... ..... ..... .. ... ...... 6. Gro up Aberrantes 4. Receptacles glabrous or subglabrous. Colombi a and Venezuelan border. Spec ies Nos . 5- 48 ....................................... .. .. .... .. ............................... .. .. .... .. .. .... ... ...... .. .. .. ............ .. 4. Sect. Espeletia 1. Outer phyll a ries li near, narrowly triangular, lanceolate, or narrowly oblong, acute, length to width ratio usually 4-1 0:1 , rarely ovate or broadl y lanceolate with a ratio of 3- 2:1, but always long-acuminate and acute.

Receptacles us ually densely hirs ute, rarely glabrous. Leaf sheaths oblong, rectang ular, not broadening .. .. .. ................ 5. Sect. Wedde/!ia towards the base. Venezuela: Meri da, Tr ujillo, La ra. Species nos. 49- 56 .....

Key to Sections and Species of Espeletia Section 1. Aristeguieta Indument of leaves densely erect-pate nt, tome ntose-subvelvety. Leaf lamina sublanceolate-oblong, thick, fi rm, sess ile.

Sy nflo rescences large panicles. Capitul a subglobose 10- 14m m in d iameter. Disc coro ll a lobes glabrous. Rosettes large, sessil e or stipitate. Sp ines of po ll en grai ns 4.0- 4.5 fl111 long, about 12 on equator. Venezuela: Trujillo .. .. ................................. .... ... ... .... .. ... ... ... .. .... ... ...... .. .... .. ... .... .. .. . .. 1. E. aristeguietana

ESPELETIA

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ALTITUDINAL RANGE OF ESPELETIA SPECIES 30-57 <ll

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Section 2. Bonplandia lndument of leaves appressed, sil very sericeous. Synflorescences large panicles, indument yellow-brown at distal part. Capitula semiglobose, 12-20mm diameter. Disc corolla lobes barbate. Spines of pollen grains 4.5- 6.5 J..Un long, 12- 15 on equator.

I. Leaf lamin ae linear or ensiform , flat , soft , and fl ex ible, not narrowed , slightly broadened or sometimes slightl y restricted at bases; costa weak; sheath s usually the same width as lower pa rt of laminae; indument of leaves dense and much appressed as if ironed, sericeous, silvery shining. Ligules of ray corollas well developed or rarely lacking. Colombia: Cu ndin a rn a rca .............. . ............................................................... 2. E. argentea I. Leaf la minae narrowly elliptic, thick, firm , contracting into na rrow bases that sudden ly broaden into a sheath , rounded on top; costa robust; indument of laminae very dense, thi ck, subappressed or appressed , sericeous. Limb of ray corollas well developed. Colombi a: Boyaca ..................................................... 3. E. boyacensis

Section 3. Badilloa Capitul a small, se miglobose, 15-20 rnm in diameter, short-radiate; ray corollas 3.0- 4.5 mm long, barely exceedin g involucres and with very conspicuous, adaxially lin gulate appendages; lobes of di sc coro ll as glabrous; outer phyllaries tri angul ar, 10.0-6.5 mmlong; synflorescences up to 2m long with thinner arched branch lets and pedicels; capitu la noddin g or

136

A SYSTEMATIC STUDY OF THE SUBTRIBE ESPELETIINAE

pendulous. Leaf blades narrowly oblanceolate, long-petiolate; indument lanate, appressed. Rosettes very large, sessile.

Spines of pollen grains 2.0~2.5 !-(m long, 18 ~21 on equator. Venezuela: Merida .................................. 4. E. semiglobulata

Section 4. Espeletia Capitula large, subglobose or campanulate, 20~70mm in diameter, long-radiate, rarely pseudodiscoid , ray corollas 6~24mm long, usually exceeding involucres, adaxial appendages on tubes dentiform or linear or lacking; outer phyllaries of involucres 12~30mm long, ovate, suborbicular, elliptic or oblong-ovate, ratio 1 ~3:1; only rarely narrowly oblong lanceolate; leaf laminae various; indument lanate. Spines of pollen grains 4~7 !-(ill, 11 ~ 18 on equator.

I. Proximal, vegetative parts of synflorescences each with from several to 1 pair of sterile, opposite leaves.

2. Leaf laminae narrowed toward bases into conspicuous pseudopetioles.

3. Synflorescences each with 1 ~3(~5) large, nodding capitula, 35~70mm in diameter; involucre thickly golden lanate; tubes of ray corollas 3.0~4.5 mm long.

4. Leaf laminae firm , coriaceous, elliptic-oblanceolate, 26~35 em x 2.0 ~3.2 em (including pseudopetiole of 4~8 em), indument appressed, yellowish when dry. Synflorescences each with 1~3 capitula (rarely 5), 30~55 mm in diameter; tubes of ray corollas glanduliferous; disc corollas 8.5~11.5 mm long, glanduliferous; anther appendages ovate, 0.4~0.7 mm long. Colombia: Norte de Santander ........................................................................................................................ 5. E. roberti 4. Leaf laminae flexible, subcoriaceous, linear-oblanceolate, l8~26cm x 1.0~1.5 em (including pseudopetiole of 3 ~5 em); indument rather lax but thickly softly lanate and superficially barbate, white or cinereous when dry. Synflorescences monocephalous; capitula 40 ~70mm in diameter; tubes of ray corollas subglabrous; disc corollas 11 ~ 12mm long, subglabrous; anther appendages ovate-elliptic, 1.0~ 1.5 mm long. Colombia: Santander, Norte de Santander ................................................................................................................................. 6. E. estanislana 3. Synflorescences generally with numerous capitula, (16~)20~35 mm in diameter; indument of involucres white, seldom yellow; tubes of ray corollas varying in length (1.2~5.5 mm).

5. Leaf laminae sublinear, narrowly elliptic-oblong, or eventually oblanceolate-oblong, acute, (21~) 30~40 x 3 ~ 6cm, length to width ratio 7~ 12:1 , densely white or cinerous lanate, secondary nerves perpendicularly spreading, the deviation angle (70~)80~90°; pseudopetiole 3~7(~10)cm long, 7~ 10mm wide at base. Capitula 20~25mm in diameter; ray corollas 8.5~13.0mm long, tubes 1.2~2.0mm long; disc corollas 6~7 mm long; lobes glanduliferous, eventually with a few hairs.

Colombia: Cundinamarca ................................................. ............................................................ 7. E. chocontana 5. Leaf laminae narrowly or broadly 1anceolate, oblanceolate, or subobovate; secondary nerves diverging at 40~50(30~60) angle.

6. Leaves rather flexible, weakly subcoriaceous; laminae narrowly elliptic-lanceolate, acute, long-cuneate at base, (16~)20 ~24cm x 1.8~2.3 em, length to width ratio (6.6 ~)8 ~ 11 :I; pseudopetioles 3 ~4cm long; sheaths narrow, 7~8cm x 1.8 ~2.2cm. Synflorescences 2~3x longer than leaves, each with 14~20 capitula, of25~28mm in diameter; tubes of ray corollas 2mm long; disc corollas 7.5 ~ 8.0mm long with a few hairs and glands at the middle; lobes with only sparse, stalked glands. Colombian-Venezuelan border: Sierra de Perij a; Zulia ...................... 8. E. tillettii 6. Leaves rigid and robust, coriaceous or subcoriaceous.

7. Leaf laminae oblanceolate and acute or subobovate and subobtuse or shortly apiculate, 26~34cm x 3.5~5.0cm, length to width ratio 5.8 ~7.5(~9.0):1; pseudopetioles 2~5 em long, 5~8mm wide at bases; sheaths 7~9cm x 4~5cm. Synflorescences 2~3x longer than rosettes, each with 24~7 capitula of 16~24mm in diameter; proximal fertile internodes and branches very long, 18 ~ 44cm, branches all rather thin, erect, and parallel-ascending (fastigiate); tubes of ray corollas 2.2~3.0 mm long; disc corollas 8~9 mm long, lobes glanduliferous, without hairs. General indument white or greenish white cinereous. Venezuela: Tachira, Apure ........................................................................................................................... 9. E. steyermarkii 7. Leaf laminae narrowly to broadly lanceolate, elliptic-lanceolate, or sublanceolate, sharply acute or acute; pseudopetioles very long to short. Synflorescences without an extra long, lower, fertile internode and without particularly long, vertically ascending, proximal branches.

8. Tubes of ray corollas 3 ~5.5 mm long.

9. Disc corollas copiously long-pilose and 8.0~8.5 mm long; lobes usually with sparse hairs and glands. Tubes of ray corollas densely long-pilose and 3 ~4 mm long.

Leaf laminae elliptic-lanceolate, acute, 19~32cm x 4~8cm ; pseudopetioles 2.0~3.5 em x 6~9 mm. Capitula 25~35 mm in diameter. Colombia: Boyaca ............................................ 13-3. E. grandiflora ssp. boyacana 0

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9. Disc coroll as glabrous or subglabrous, 9.0-11.5mm long; lobes glabrous. Tubes of ray coroll as subglabrous.

10. Leaf la minae broad-ell iptic-lanceolate, sharply acute, rather thick, 30-35cm x 9.5- 10.5 cm; pseudopetioles 2.5-5.0 cm long. Ray coroll as 20-23 mm long, tubes 3-4 mm long. Disc coroll as 9 mm long; lobes glabrous; phyllaries densely villous and long-barbate. Synfl orescences 1/J or 112 longer than rosettes, sterile leaves as well as subtending bracts acuminate, greatly exceed ing internodes and peduncles. General indument, especially on sy nfl orescences, thickly, laxly long-villous and cottony barbate. Colombia: Cundin amarca ..................... 10. E. cayetana 10. Leaf la minae oblong-elliptic or oblong-elliptic-lanceolate, ac ute, thick and firm , 35-38 cm x 8.0-8.5cm; pseudopetioles 8-J I em long. Ray corollas not exceeding the involucre, 16-17 mm long; tubes 4.5-5.5 mm long. Disc coroll as 11.0- 11. 5 mm long, subglabrous; lobes with a few clavate glands.

Phyllaries densely crisped-lanate; synflorescences 2x longer than rosettes, subtending bracts equaling or shorter than internodes of peduncles. General indument thick, closely compressed cri sped-lanate. Colombia: Cundinamarca ........................ .. .. .... .. ........................ .. ...... .. ...... 11. E. miradorensis 8. Tubes of ray coroll as 0. 8- 2.5(- 2.8) mm long.

11. Synflorescences shorter than or equaling rosettes; indu ment rather shaggy, barbate, yellowish. Pseudopetioles winged, about 1 em x 9- 11 mm. Tubes of ray corollas 0.8-1.2mm long. Colombi a: Valle del Cauca, Tolim a.26-3. E. hartwegiana ssp. barragensis 11. Synflorescences l/J to 3x longer than rosettes, usually great ly surpassing them.

12. Phyllaries ellipti c-oblong, obtuse, 30-19mm x 15-11 mm, densely yellowish cri sped-l anate and somewhat villous above. Involucres exceeding ray corollas.

Leaf laminae 27-33 x 5.5- 5.8 em; pseudopetioles 2-3 em x 7- 8 mm ; indument white, appressed, thickly crisped-lanate. Rosettes whi te or greenish cinereous.

Synflorescences yell owish. Colombia: Cauca .......... .. .... .. .. ...

12. E. idroboi 12. Phyll aries ovate, 28- 16 mm x 20-8 mm. Involucres much surpassed by ray corollas, rarely equaling them. General appea rance white; indument on synflorescences rather shaggy-wooly.

13. Leaf lamin ae conspicuously lanceolate or elliptic-lanceolate, contracted above bases, 24- 48 em x 4.0-9.5 em, length to width ratio 4- 7: I; pseudopetioles 2-7(- 11) em. Capitula 30-35(-40) mm in di ameter; ray corollas 16- 24 mm long; tubes 1.0- 2.5 mm long, de nsely hairy; disc coroll as 7.5-9.0 mm long, usually hairy at middle, lobes glabrous.

Synflorescences 2- 3x longer than leaves. Colombia: Cundinamarca .............. ...... .. .... ........ .. .... .. ......... '13- l a. E. grandifiora var. grandijlora 13. Leaf lamin ae moderately lanceolate or lanceolate-linear, gradually na rrow ing to bases, 21-38cm x 2.2-4.7 em, length to width ratio 7-11:1 ; pseudopetioles short to obsolete (l-3cm). Capitul a 20-30mm in diameter; ray coroll as 10-17 mm long, tubes l.0-2.5 mm long, densely hairy; disc corollas, 8.0- 8.5 mm long, copiously hairy at middle, lobes with sessile or subsessile glands . Syn florescen ces 12- 2x longer than leaves. Colombia: Cundi namarca ...... ............................................... 13-lb. E. grandifiora var. attenuata 2. Leaf laminae somewhat narrowed toward bases, but not obviously pseudopetiolate.

14. Synflorescences 2- 3x (or 12x) longer than rosettes.

15. La minae narrowly oblanceolate-oblong, acute, narrowed gradually to bases, rigid-coriaceous, 45- 48cm x 4.0- 4.5 cm, length to width ratio 9-10:1 ; indument whiti sh, appressed-lanate. Phyllaries and subtending bracts loosely antrorsely villous and conspicuously glanduli ferous. Disc corolla lobes glanduliferous, not hairy. Colombia: Cundinamarca, Meta ............ .. .............. .. ....... ! 4. E. tapirophila 15. Without the above combination of features; subtending bracts not conspicuously glanduliferous.

16. Indument throughout pl ants densely, thickly crisped-lanate. Leaf lam inae thickly felt-li ke; venation totally concealed by indumenta. Lobes of disc corollas usuall y with some hairs.

17. Capitula in groups of 3-7, sessile or subsessile, compressed into la nate, compact clusters (glomerules), nodding or drooping at tops of rigid, thick, synflorescence axes; ray corollas long and narrow, 20-30 mm; tube 7- 9mm long. Rosette appearance golden or yellowi sh.

Leaf laminae narrowly lanceolate, attenuate at both ends, (21- )28- 45cm x 3.0- 4.6cm; length to width ratio 7-12: 1. Colombia: Boyaca .......... .. ...... .. .... .. .. ................... 15. E. congestifiora

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A SYSTEMATIC STUDY OF THE SUB TRIBE ESPELETIINAE 17. Capitula distinctly pedunculate, lanate, subglobular, 20-30(- 35) mm in diameter; synftorescences paniculate, yellow ish with spreading, stout, thick branches and peduncles; ray corollas 10- 13 mm long; tubes 1.6-2.2(-3) mm long. Rosette appearance whiti sh.

Leaf laminae elliptic-sublanceolate, 27-46(-50) x 3.3- 6.5(- 7.3)cm, length to width ratio (4.2-)6- 11: I. Co lombi a: Santander, Norte de Santa nder .... ....... ... ..... . ...... 16. E. standleyana 16. Indument superficially vill ous-sericeous and white throughout plant or leaf laminae with secondary nerves abax ially very conspicuous. Lobes of di sc coroll as usually without hairs.

18. Synftorescences each with 5- 7 pairs of sterile leaves, the 3 or 4 prox imal ones crowded at bases; branches rather spreading. Leaf laminae narrowly elliptic, narrowly oblong, or narrowly suboblanceo late.

19. lndument of leaves compactly lanate with a looser, subappressed or appressed villous-sericeous layer on the surface; secondary nerves concealed or only noticeable abax ially by thin , linea r, sericeous traces in indument. Sy nft orescences very large (up to 1.7 m), erect, spreading-paniculate with rather slender, terminal branches and pedicels; subtending bracts longer than the internodes, peduncles, and pedicels; capitula 14- 18(- 20) mm in di ameter. Indument white th roughout. Leaf laminae 24- 38 cm x 3.5-6.0 cm. Disc coroll as 5.5-6.0mm long, tubes copiously glanduliferous; lobes glabrous with a few glands. Tubes of ray corollas 1.2-2.0 mm long. Colombia: Boyaca ........................................ ... ...... .... ... ... ... .... ... ................. 17. E. tunjana 19. Indument of leaves dense and appressed adaxially, only in juvenile stage with arachnoideous or villous-sericeous layer on the surface; seco nd ary nerves very conspicuous abax ially. Synftorescences robu st, 64-90 c m long, rather divergent from rosettes with rather thick, terminal branchlets and ped icels; subtending bracts equ aling or longer than internodes, peduncles, and ped icels. Capitula about 25 mm in diameter. lndument white or cinereous on leaves, yellowish on panicles. Leaf lamin ae 23- 32cm x 3.8-5.5 cm. Disc coroll as 6.5-7.5 mm long, tubes with sparse glands, lobes glabrous or with rare ha irs. Thbes of ray corollas 2.2-3.0mm long. Colombia: Boyaca .............. .. ... ... ... ... ..... ..... ... .... ... .. ...... ..... . 18. E. jaramillo 18. Synftorescences eac h with 2- 5 pairs of sterile leaves, only l or 2 close to bases; internodes and peduncles longer than bracts.

20. Leaf lamin ae narrowly oblong, 26-30cm x 3.2- 3.5cm, 1.4-1.7 em wide above bases. Synftorescencess erect, subfastigiate, subcory mbiform, yellowi sh lanate, about 48cm long with about 17 capitula, 20- 24mm in diameter. Colombia: Boyaca ........................................................ ....... ...... ... ... ... ... ... ... ... ........ ... ........ 19. E. azucarina 20. Leaf laminae more or less markedly lanceolate or obl anceolate.

21. Leaf lam inae 14-31 em x 2.3-5.0 cm, narrowly elli ptic or subl anceolate; indument white lanate, dense and appressed adax ia ll y, looser abax ially; reticul ar alveolae deep, densely pilose inside; secondary nerves spreading at an angle of 50- 75°. Sy nft orescences slender, erect, loosely white lanate, each with 5- 3 capitula 18- 22 mm in diameter. Colombi a: Antioqui a, C6rdoba .. ... .. ...... ............. .. ............................. ... ... ... ... ..... ... ........ ........ 20. E. occidentalis 21. Leaf lamin ae 34-70 cm x 5.4- 9(-13) cm, greenish or cinereous with white lanate indument and superficial flu shes of appressed, long clusters of hai rs; second ary nerves and costa more conspicously marked by co ncentrations of longer, stra ight, vill ous, white hairs. Synftoresce nces large, each usually with more than 13 capitula.

22. Leaf laminae cin ereous or whitish with more densely cri sped indument abax ial ly; venular reticulum prominulous, rather obtuse, concealed by indument, alveolae rather shallow with bottoms hairy and granulatepapillose th roughout. Synflorescences 12 or 2x longer than rosette, subcorymbi fo rm , densely branched, each with (3- )4- 6 pairs of branches (paraclad ia) and 22- 80 capitula about 30mm in diameter; disc corollas 7.5-lO.Omm long, glabrous or almost so; tubes of ray coroll as 2- 3 mm long with copious, subacute hairs 0.25- 0.60 mm long, with sparse glands abaxially. Colombia: Boyaca, Arauca ... ... .... ... .... ... 21. E. curialensis 22. Leaf lam inae greenish with a soft, thi ck, but rather lax, white, crispedlanate indument abaxially; venular reticulu m prominent and sharp, conceal ed by its thick indument, alveolae concave, deep, with clear, glabrous, nitid , green bottoms. Synflorescences 2- 3x longer than rosettes,

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thyrsoid, each with 2-4 pairs of branches (paracladia) and 15- 23 capitula 15-22mm in diameter; disc corollas 6.5- 8.0mm long, copiously hairy at middle; tubes of ray corollas 1.2- 2.0mm long, with copious, thick, obtuse hairs 0.2- 0.3 mm long and with shortly pediculate glands.

Colombia: Boyaca, Arauca ........................................................... 22. E. oswaldiana 14. Synflorescences shorter than, equaling, or slightly surpassing (to 1/4) rosette.

23. Capitula semiglobose, 18-20(-23) mm in diameter; involucres shallow-cupular or subpatelliform, cinereous, antrorse-villous , slightly surpassing the convex, protuding discs; ray corollas barely surpassing involucres. Leaf laminae gray-greenish with short, lax, lanate indument adaxially and whitish, lanate indument abaxially, elliptic-lanceolate or oblong-elliptic, abruptly acutate, 35- 52 em x 7.0-13 em; bases 1.2-2.0cm wide, rarely contracted to 1 em. Synflorescences robust, pyramidal, each with 1- 2 pairs of sterile leaves; peduncles rather straight; capitula suberect or drooping. Colombia: Cundinamarca ........................................................................................... 23. E. uribei 23. Capitula sub globose or campanulate; involucres usually distinctly surpassing di scs and with crisped-lanate or lanuginous, yellowish indument; ray corollas conspicuously much longer than involucres. Adult leaf laminae densely ian ate on both sides.

24. Synflorescences corymbiform, obconical; branches ascending, rigid , appearing thick and straight, proximal ones reaching or nearly reaching tops of distal ones; each with 1-4 pairs of sterile leaves; indument thick, yellowish. Capitula 15- 20(-22)mm in diameter, erect or inclined; outer phyllaries broadly ovate-acuminate, acute, densely yellowish lanate; disc corollas 5-6 mm with some long hairs at middle, rarely scarce hairs on lobes. Leaf laminae oblanceolate, very acute, 26-34cm x 3.5- 6.0cm, thickly and softly, white or cinereous lanate. Colombia: Boyaca ........................................................ ............................................... 24. E. cleefii 24. Synflorescences pyramidal, proximal branches not reaching tops of distal ones. Capitula 20-40mm in diameter, campanulate or deeply cupular, drooping or nodding; outer phyllaries obovate-oblong or elliptic, obtuse or abruptly cuspidate; disc corolla lobes without hairs. Leaf laminae thickly lanate, indument often shaggy and copiously barbate, particularly in young leaves and synflorescences. General appearance yellowish.

25. Phyllaries thinly membranaceous, loosely, thinly wooly; pales 6.5-8.0mm long with sparse hairs at apices; disc corollas 6.5- 7.0 mm long with sparse or rare hairs or glands at the middle. Leaflaminae 36- 58cm x 8.0- 12.5 em, elliptic or rather obovate-elliptic, abruptly narrowed , shortly acute or apiculate, gradually tapering without or with a slight contraction to bases (15- 23mm wide). Capitula small, 20-28(- 30)mm. Colombia: Narifio-Putumayo ............................................................. ..................... ... ......... 25. E. schultesiana 25. Phyllaries thick, firmly herbaceous, densely lanate, and distally barbate; pales distally, densely, brownish l:iarbate. Leaf laminae narrowly elliptic-oblong or elliptic, attenuate and acute, attenuate towards bases, usually with some contraction.

26. Leaf laminae (20-)26-40cm x 4.5-9.0(- lO)cm, narrowly elliptic-oblong or elliptic and gradually narrowing, or wide with some contraction above broad bases, these (1.0-)1.5-2.8 mm wide. Synflorescences each with 5- 17(-23) capitula, (22-)25-35 mm in diameter; outer phyllaries 20- 30mm x I0- 18mm; pales 8- 9 mm long, distall y, densely, brownish barbate; disc corollas 8- lOmm long. Colombia: Cordillera Central, Cauca, Valle del Cauca, Huila, Caldas ...................................................... 26. E. hartwegiana 26. Leaf laminae 20-25 em x 4-5.5 em, narrowly oblong, attenuate, acute at apices, 13-18 mm wide above base, not markedly contracted. Synflorescences usually each with 3-5 capitula, 22-25mm in diameter; outer phyllaries 17-20mm x 10- 11 mm; pales 5.5-6(-8)mm long, distally densely barbate and glanduliferous; disc corollas 6.5-8 mm long. Colombia: Cordillera Occidental, Antioquia .................. 27. E. praeji-ontina I. Proximal, vegetative parts of synflorescences naked, completely lacking sterile leaves, occasionally above proximal parts bearing a few leafy bracts originating from fertile parts and becoming sterile.

27. Leaf laminae lanceolate in outline, narrowly elliptic or linear, gradually attenuate toward the acute apex, also narrowing moderately to the base.

28. Leaf laminae sessile, narrowly oblong or linear (or occasionally slightly lanceolate), usually very long and thick, 34- 52cm x 3- 6cm, length to width ratio 7-13:1, gradually narrowing to 23-13(-lO)mm width at bases. Synflorescences erect, slightly surpassing rosettes, each with 3-7(-11) capitula 3-6cm in diameter.

29. Phyllaries linear-lanceolate or narrowly oblong, acuminate, very acute, 30- 53 mm x 6- 12 mm, length to width ratio 4-6.2:1, surpassing the ray corollas. Syntlorescences each with 3 capitula 4- 6cm wide; ray corollas 16- 22mm long, green-yellow or pale yellow; tubes 2.0- 2.8 mm long;

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A SYSTEMATIC STUDY OF THE SUBTRIBE ESPELETIINAE disc corollas 9.0-11.5 mm long; lobes sometimes with glands, without hairs. Plants snow-wh ite wooly. Colombia: Cundinamarca, Paramo de Sumapaz ............................... ............ ... .. 28. E. summapacis 29. Phyllaries ovate or oblong-ovate, 18- 35 mm x 13-19(-23)mm, length to width ratio 1.7-2.2:1.

Synflorescences each with (3-)5-7(-11) capitula 3-4cm in diameter; ray corollas bright yellow; plants light yellowish wooly.

30. Ray corollas 12- 16 mm long; tubes 3-6 mm long; disc corollas 8.0- 10.5 mm long; lobes with few hairs. Leaf laminae narrowly oblong, gradually narrowed to 13-23 mm minimum width above bases. Colombia: Cundinamarca, Bogota Massif ........................................ 29. E. killipii 30. Ray corollas 20-26mm long; tubes 1.5- 2.0(- 3.0)mm; disc corollas 9-l2mm long, lobes sparsely glanduliferous with no hairs. Leaf laminae narrowed towards bases to 10-16mm minimum width with some trend toward a lanceolate shape. Colombia: Cundinamarca, Bogota Massif ..... ....... ...................................................................... 29-2. E. killipii ssp. chisacana 28. Leaf laminae conspicuously broadly lanceolate, oblanceolate, or oblong-elliptic; pseudopetiolate or sessile.

31. Capitula 4-6(-7)cm in diameter. Leaf laminae broadly elliptic-lanceolate or sublanceolate, 32- 36cm x 6-7 em, very thick, thickly crisped-lanate, attenuate towards bases almost forming pseudopetioles 2-6cm x 10- 14 mm. Ray corollas 20-25 mm long; tubes 6- 8m m long. Disc corollas 12.5-14mm long; lobes with sparing or rare, very short hairs. Plants appearing yellowish. Synflorescences 2x longer than rosettes.

32. Synflorescences each usually with 5 capitula and a pair of steri le, upper leaves above the middle or with 3 capitula and 2 pairs of sterile bracts; capitula 4.0- 4.5(- S)cm in diameter, nodding. Colombia: Boyaca ...................................................................... 30-1. E. rositae ssp. rositae 32. Synflorescences monocephalous, each with 3 pairs of sterile, subtend ing bracts, the distal ones close to involucres, capitula 5- 7 em in diameter. Colombia: Boyaca ........................................................................................... 30-2. E. rositae ssp. macrocephala 31. Capitula 2-3cm in diameter.

33. Leaves with conspicuous, narrow pseudopetiole 4-8cm long. Synflorescences 2x longer than rosettes, large, each with many capitula or smaller with few capitula and 1-2 sterile, leafy bracts at the upper l/2 of each axis. Capitula 20- 25 mm in diameter; lobes of disc corollas with sparing hairs. Rosettes white.

34. Leaves very long, laminae 47-SOcm x 4.0-4.5cm, length to width ratio 11-12:1, rigid with robust middle nerve; pseudopetioles 6-8 em long. Synflorescences with broad, spreading branches each with as many as 43 capitula; branch lets lax and flexuous, distally golden lanate. Outer phyllaries oblong, slightly attenuate, abruptly acutate or subacute, 2.3- 3.0x longer than wide; ray corollas 12- 23 mm long; tubes 0.7-0.8 mm long. Colombia: Cundinamarca, Paramo de Sumapaz ..................................... 31. E. cabrerensis 34. Leaves smaller and more flexible; laminae 24-32cm x 2.2- 3.4cm; pseudopetioles 4.0-6.5(-8.0) em long. Synflorescences each with 3-5(- 6) capitu la, usually glomerate or congested at the tops of axes, upper 1/2 of axes often with I single or 2 opposite, additional , pedunculate capitula below and with I or 2 sma ll, sterile leaves; all white lanate; outer phyllaries ovate, acute or acuminate, 1.2-1.4x longer than wide; ray corollas 11.5-16.0mm long; tubes 1.5-2.5 mm. Sierra de Perij a. Colombia: Cesar; Venezuela: Zulia .......................................................... ....................... ............... 32. E. perijaensis 33. Without the combination of features above, with pseudopetioles or with shorter synflorescences or with yellowish appearance.

35. Leaf laminae tapering gradually to broad bases 20- 13(- 10) mm wide (not including indument), with no apparent pseudopetioles, broadly lanceolate, 20-30cm x 4.0-6.5 em.

Synflorescences usually equaling or only slightly surpassing rosettes. Lobes of disc corollas without hairs.

36. Indument yellow. Synflorescences equaling rosettes , each with 3-5 capitula.

Leaf laminae 20-30 cm x 4.0-5.6cm; bases 11-15 mm wide, length to width ratio 4.7-5.5:1. Ray corollas 10-13 mm long, tube 1.5- 2.2 mm long; disc corollas 7-8 mm long. Colombia: Cordillera Occidentalis, Antioquia ........ ........ 33. E.frontinoensis 36. Indument white. Colombia: Boyaca and adjacent Santander borders.

37. Branch lets or lateral pedicels not concaulescent with axes, aris ing directly from the axils of subtending bracts; these equaling or longer than the pedicels.

38. Synflorescences usually shorter than rosettes, each with 3 capitula (rarely 4); ray corollas 8.0-ll.Smm long; tubes 3-4mm long; disc

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corollas 8-9 mm long; lobes glabrous or seldom with I hair. Leaf laminae 18-32cm x 4.0-4.7 em; bases 16-!0mm wide; length to width ratio (2.7-)4- 6:1. Colombia: Santander, Boyaui ................................ 34. E. incana 38. Synftorescences surpassing rosettes, each with 7-9 capitula; ray corollas 9.5- 10.5mm long; tubes 1.0-1.5 mm long; disc corollas 7-8 mm long; lobes barbate. Leaf laminae 20-26 em x 4.0-7.4cm; bases 16-20mm wide; length to width ratio 3.5-5.0:1. Colombia: Boyaca ................ 35. E. pescana 37. Branch lets and lateral peduncles concaulescent with axes apparently arising several em higher than subtending bracts; these shorter than peduncles.

Synftorescences each with 5 capitula (rarely 7); ray corollas 4.5-8.0mm long; tubes 1.5-2.3 mm long; disc corollas 6.5-7 mm long; lobes rarely with a very few hairs. Leaflaminae 21-29cm x (4.3-)5.0- 6.5 em, bases (10- )13-20mm wide, length to width ratio 4.0-5.2:1. Colombia: Boyaca, Santander ..................................................................................... 36. E. brachyaxiantha 35. Leaf laminae tapering gradually to apparent pseudopetioles 4- lOmm wide.

39. Synftorescences with 5-11(-17) capitula and at least 2x longer than rosettes; branchlets and pedicels long, straight, subparallel, erect, proximal ones longer, almost reaching tops of corymbiform panicles. Capitula 18-22(- 25) mm in diameter, erect or drooping. Plant appearance yellow or greenish gray with yellowish synftorescences.

40. Leaf laminae rigid, thickly coriaceous, adaxially with fine, white, evanescent indument when young, soon becoming glabrous, smooth, green; abaxially cinereous or slightly ochraceous lanate; reticular venation very prominent in about 3 layers, alveolae deep, ovate or oval; outer phyllaries ovate-cordate, wider than long or as wide as long, soon glabrate. Lobes of disc corollas hairy; ray corollas 7- 11 mm long; tubes 1.2- 2.0mm long. Colombia: Boyaca ................................................................................................... 37. E. nemekenei 40. Leaf laminae ftexible-coriaceous, softer, with permanent yellowish indument on both sides; reticular venation prominent, forming obtusely polygonal alveolae. Outer phyllaries ovate, longer than wide; lobes of disc corollas glabrous or with very few hairs; ray corollas 6-9mm long, tubes 1.0-2.2mm long. Colombia: Boyaca ................................................................................. 38. E. murilloi 39. Synftorescences 1/2-2x longer than the leaves, with only 3 or occasionally 4 capitula; indument white.

41. Capitula pseudodiscoid, with ray corollas shorter than the emerging styles, reduced to tubes 0.2- 0.5 mm long each with a lobate or dentiform, obsolete, apical projection l-2mm long. Synftorescences usually shorter than or equal to the leaves, each always with 3 capitula; lateral peduncles concaulescent to some extent with axes, apparently arising at some higher point than the subtending bracts, much shorter than peduncles; the latter straight and erect; capitula 18- 20(- 22)mm in diameter; phyllaries suborbicular, obtuse, not surpassing the flowers. Colombia: Boyaca, S of Paramo de Guantiva ... 39. E. disco idea 41. Capitula distinctly radiate, with ray corollas ligular, 5.5-7.5 mm long; tubes 2.0-2.7 mm long. Synftorescences 1/2-2x longer than leaves and each with 3 or occasionally 4 capitula; peduncles never concaulescent with axes, arising directly from axils of the bracts, usually curved, supporting nodding capitula; bracts straight, erect, usually topping the panicles like a pair of ears; capitula 20-28 mm in diameter; phyllaries broadly ovate and acute.

Colombia: Boyaca .................................................................................. 40. E. arbelaezii 27. Leaf laminae oblong, obovate-oblong, elliptic, ovate-oblong, or spathulate, not gradually attenuate towards apex, subobtuse or shortly angulate.

42. Leaves conspicuously contracted into pseudopetioles, sometimes very short; synftorescences surpassing the rosettes.

43. Synftorescences usually a single, 3-cephalous cyme, occasionally 2 or more additional capitula, all thickly lanate, 3.5- 5.0(- 6.0) em in diameter; outer phyllaries ovate; tubes of ray corollas 4-6(-9) mm long. Leaf laminae narrowly elliptic, oblong-elliptic, or oblong-ovate, obtuse or shortly acute at apices, 16- 38cm x 3- 11 em, length to width ratio 2.8-5.6:1; pseudopetioles 4-lOcm x 5- lOmm; indument yellowish, very thick and dense, curly lanate and concealing the veins. Colombia: Boyaca ............................................................................................................................................ 41. E.lopezii

142

A SYSTEMATIC STUDY OF THE SUBTRIBE ESPELETIINAE 43. Synflorescences usually pluribranched with (5-)7-13(-15) capit ula; capitula densely lanate, 30-35 mm in diameter; outer phyllaries oblong; tubes of ray coroll as 2.5-3.8 mm long. Leaf laminae elliptic with abruptly narrowed, obtuse or angulate at apices or so metimes ovatesublanceolate, gradually or abruptly contracted at bases, 19- 35 em x 4.5-9.0cm, length to width ratio 3-5:1 ; pseudopetioles 1.0-3.5 em x 7.0- 14 mm. lndument white, thickly shaggy lanate and fl occose on many parts. Colombia: Cundinamarca, Boyaca ................................................ 42. E. barclayana 42. Leaf laminae attenuate or contracted but not distinctly pseudopetiolate.

44. Leaf laminae narrow, 3.2- 6.0cm wide, length to width ratio (5- )6-9(-13):1, oblong-spathulate or ob long-elliptic, occasionally obtrullate, gradually attenuate to bases. Synflorescences much exceeding rosettes.

45. Plant appearance yellowi sh. Ray coro ll as 14-18mm long; tubes 4.5- 6.0mm long. Leaf laminae thick, coriaceous, usua ll y rigid, narrowly oblong, subobtuse, or somewhat elliptic, (26-)34-40cm x 4.3- 5.5(- 6.5)cm, indument compact. Capitu la usually 5 or 3, nodding, 3- 4cm in diameter; peduncles long or short, flexuous. Colombia: Santander, south of Paramo Almorzadero .......................... .. ... ... .. ......... ............. . ................... ........... ... ...... .... . ... 43. E. dugandii 45. Plant appearance white or cinereous. Ray corollas 7-11(-13) mm long; tubes 1.5- 2.5 mm long. lndument thi ck, softer, looser at surface. Capitula 2.0-3.2cm in diameter.

46. Leaflaminae more thinl y cori aceous, flexible , oblong, spathulate or narrowly obovate-oblong, obtuse or subacute at apices, 22- 32(- 38)cm x 3.2-5.5 cm. Capitula usually glomerate on curled, contracted peduncles (rarely peduncles almost straight); disc coroll as hairy and glanduliferous at the middle, at least some lobes with a few hairs. Colombia: Santander, Norte de Santander, and Tach ira .......... ........................................... 44. E. conglomerata 46. Leaf laminae rather thickl y coriaceous, obtrullate-oblong, angul ate and acute at apices, 30-32cm x 5.3- 6.3cm. Capitula drooping or nodding on straight, rigid peduncles and pedicels; disc corollas glabrous or subglabrous; lobes with onl y sparse glands. Col ombia: Norte de Santa nder, Paramo del Romeral ...... .................. ...... ... ................ .. ... ... .... ... 45. E. canescens 44. Leaf laminae (4.5-)7.0-14.0cm wide, length to width ratio 1.3-5.0(-6.0):1 , elliptic, obovate, oblongobovate or oblong-elliptic, abruptly or gradually narrowed at bases. General appearance white.

Synflorescences shorter than to 2x longer than leaves.

47. Sy nflorescences about 2x longer than leaves. Tubes of ray coroll as 3.5-5.0mm long, copiously pilose. Synflorescences with (5-)9-19 capitula; proximal branches separated by long internodes, very long, thin , erect, stra ight, para ll el-ascending, close to axes and each topped by 1- 3 congested, nodding capitula; ped uncles erect; outer phyllaries ovate; disc coroll a lobes glabrous or in ssp. rarely with a very few hairs. Leaf laminae 30- 40cm x 7- 14crn , broadly elliptic, subobtuse, obtuse or attenuate at apices, margins irregularly, coarsely undulate, gradually attenuate in proxim al part or contracted to 12-25 rnm width above bases, sometimes narrower. General indument den se, looser at surface. Colombi a: Norte de Santander; Venezuela: Tachira. .. .... ........... ............................. ....... ......... .. ..... .. ... .... ... ... .... ...... 46. E. brassicoidea 47. Synflorescences shorter than or equaling the leaves. Tubes of the ray coroll as 0.3-1.5 mm long, densely hairy.

48. Leaf laminae 22- 25 ern x 7.8- 8.5 ern , length to width ratio 2.6-3.0: 1, elliptic or obovate-elliptic, abruptly subacute at apices, narrowed toward s the bases and constricted into short, apparent pseudopetioles 12- 15 rnrn wide. Synflorescences about 20 cm long with 5 capitula, much shorter than rosettes; lobes of di sc coroll as without hairs. Colombia: Boyaca ................. .................................................

47-2. E. annemariana v. rupicola 48. Leaf laminae usually 30- 50crn x 7-13cm, gradually narrowed to broad sess ile bases 30-20(-15)mm wide.

49. Outer phyllaries oblong-ovate or triangular, acute, acuminate, 15- 20mm x 8-12mrn, usually shorter than fl owers. Leaf laminae narrow, elliptic-oblong, narrowed to acute, often acuminate apices, 32-43cm x 7.0-10.5cm, width at bases 18-21 mm. Lobes of di sc corollas barbate. Synflorescence with (5-)13- 19 capitula and proximal, vegetative parts shorter than fertile parts; subtending leaves (lower bracts) linear or oblonglanceolate, acute. Alveolae of the venular reticulum irregul ar, deeply concave with copious hairs. Colombia: Boyaca . ............. ............ ...... .... ................. 47. E. anne mariana 49. Outer phyll aries ellipti c, ovate-elliptic or obovate, usua lly obtuse or subobtuse (outermost subacute) 20- 30mm x 15- 20mm, longer than the fl owers. Leaf laminae obovate-oblong, obovate, elliptic or broadly elliptic, subobtuse and shortly apiculate or attenuate and subac ute, usually 36- 52cm x 8- 14cm, width above bases (14-)20- 26 (-30)mm wide (in var. galerana 22- 30cm x 4.5-8.0cm). Lobes of disc corollas

ESPELETIA

143 without hairs. Synflorescences with (5-)7- 17 capitula, vegetative parts longer than fertil e parts; subtending bracts of lower branches narrowly ellipti c, ac utate (occasion ally sterile). Alveolae of leaves moderately deep, flat or slightly concave, glabrou s, green, minutely papillose. Colombia: Nar ino-Putum ayo; Ecuador: Carchi , Tungurahua ............ .. .. .. ............................................................... ... 48. E. pycnophylla

Section 5. Weddellia Indument lanate, rarely sericeous. Prox imal, vegetative parts of the sy nft orescences usuall y with several to I pair of sterile opposite leaves or a monocephalo us scape with no leaves (E. tenorae). Outer phyllaries of involucres lin ear, subulate, narrowly triangu lar, or narrowly elli ptic-lanceolate, length to width ratio 4-10:1, rarely ovate-acuminate or broad ly lanceolate, with ratio 3.5- 2.0: 1. Spines of pollen gra in s 4-6.5 J.lm long, 11- 14 on equator.

I. Rosettes large, 40 - lOO cm in diameter; leaflaminae 25- 50cm long, oblong or linear, sess ile with no petiolar contraction; indument thickly lanate. Synflorescences large, up to 1.0-1.7 m long, highly branched; branch lets and peduncles sto ut, more or less ascending, the capitu la suberect or droop ing and num erous, subglobose or cam panul ate, 20-3 0mm in di ameter, long-radiate.

2. Leaf lam inae oblong or narrowly elliptic-oblong, eventua lly slightly subobovate-oblong, 25-45 em x 3- 6(-7)cm, length to width ratio 6- 10:1 (5-1 3: 1); apices obtuse or suddenly narrowed and subacute, not or slightly attenuate to bases, both surfaces covered by very thick, crisped-Ia nate indument, abaxial su rfaces obv iously strongly nervate. Lobes of disc corollas glabrous. Venezuela: Merida, Trujillo .......... .............. 49. E. schultzii 2. Leaf lam inae linear, 40-45cm X 2- 3cm, length to width ratio 13.5- 22: 1, attenuate di stally and acute at apices, grad ua ll y and slightly narrowed to bases; adax ially subappressed lanate and subser iceous longvillous chi efl y on the costa; abaxially nervation visibl e, indument densely crisped-l anate and also sericeous-v illous or barbate on the main vein s. Lobes of disc corollas with a few patent hairs and obovoid glands. Venezuela: Merida, Sierra de La Culata ........................................... .. .......................... 50. E. cuniculorum I. Rosettes small , 10- 30cm in diameter, sessile or subsess ile. Leaf laminae 3-18(-22)cm long. Synftorescences scapose, J0- 45(-75)cm long, with on ly 1 capitulum or 2-5 capitula, nodding or drooping.

3. Leaf sheaths a nd proximal parts of costae totally glabrous. Synflorescences monocep halo us.

4. Scapes aphyllous, with only I or 2 small, opposite or alternate, di sta l bracts, densely and thickly white lanate. Leaf laminae oblanceolate-oblong, spathulate, attenuate towards bases, densely floccose-cottony, the white indument very abundant and thicker at the obtuse, club-shaped tips. Involucres thick ly white cottony, rou nd in outline, 20- 30mm wide, surpass ing the rays. Venezuela: Merid a, Truj ill o .......... 5 1. E . tenorae 4. Scapes leafy and bracteose; 1- 3 pairs of opposite leaves near the base with long, tubular sheaths, upwards few to many small bracts; indument ochraceous, brownish or rusty lanate. Leaf laminae linear-ensiform , narrowly oblong or linear-obl anceolate, slightl y or not attenu ate to bases. Ray corollas usually surpassi ng the densely ochraceou s or brownish Janate involucres.

5. Leaf lamin ae densely, sil very sericeous, 4.0- 6.5cm x 1.4-0.5cm, the hairs appressed, straight, antrorse, 1.0-1.5 mm long. Capitula 20mm in diameter; involucres campanulate-turbinate Venezuela: Trujillo- Merida .. ............ ... .... .............. ................. ............. ................. ............. ........... .. .... ........... 52. E. mart/we 5. Leaf laminae white villous-lanate, 4- 13cm x 0.4-1.1 em, hairs rather flexuose or curled, 3-6 mm long.

Capitu la 20-26 mm in diameter; involucres rounded-cupular. Venezuela: Trujillo-Merida ................. 53. E. nana 3. Leaf sheaths abaxially den sely barbate; costae vested. Synfl orescences with 1-5 capitu la.

6. Leaf lam inae throughout softly and thickly cri sped-lanate, with cylindric outline (particularl y when young), n-like, entangled, softly thick, equal on both sides, in add ition, an arachnoid velum on the surface, becoming roseate or ru sty in drying. Synflorescences usually with I capitulu m, sometimes 2-3 capitu la, 24- 32 mm in di ameter, 1 pair (rarely 2 pairs) of opposite leaves above base, and 6-1 2 sparse, obtuse, li ne ar bracts, upwards becoming gradually smaller, bracts and scape thickly rounded-lanate.

Venezuela: Truji llo, Lara .... ...... .............................................. ................................ ................... ................. 54. E. ulotricha 6. Leaf laminae long-villous or lanate, linear or oblong-oblanceolate, obviously flat ; indument white, even when dry. Synflorescences with 2- 3 pairs of opposite leaves, with long, tubul ar sheaths.

7. Leaf lam in ae linear or slightly linear-oblanceolate, 5-12(-18) em x 0.3- 0.8(-l) cm (excluding the indument), rather acute; adaxially den sely subappressed antrorse-villous; abaxially loosely Januginou s, costa also subsericeous-villous. Scapes thin , white Janate, bearing 2- 4 small capitula, rarely monocepha lous. Capitula villous-lan ate, semiglobose, 14-20(-25) mm in diameter.

Venezuela: Merida, Trujillo ........................................ .. .................................................... ...... 55. E. weddellii 7. Leaf lami nae oblong or oblanceolate-oblong or spathulate, 6.5- 15cm x 0.8- 2.0cm (excluding the indument), obtuse or subobtu se, den sely and abundantly lanate and villous on both sides.

Synflorescences relatively thick, den sely wh ite lanate, di sta lly more thickly lanate, monocephalous. Capitula thickly lanate, subg lobose, 20-30mm in diameter (excluding the rays).

Venezu ela: Merida ..... .................................... ... ... ... ........... .... ................... .... .... ... ... ... ... ...... .......... .. 56. E. batata

144

A SYSTEMATIC STUDY OF THE SUBTRIBE ESPELETIINAE

Group Abberantes Synflorescences usually with alterna te branchlets, peduncles, and bracts, except the basal leaves or bracts always opposite. Capitula campanulate, long-radiate; ray corollas 10-17 mm long. Outer phyll ar ies usually wide, ovate, lanceolate, or elliptic, abaxially lanate or long villous; inner fertile ones ciliate and apically barbate . Receptacles hairy. Vegetative indument lanate. Venezuela: Trujillo, Merida ......................................................................................................... 57. E. jajoensis [Note: Some recent work is not accounted for in thi s monograph. Most importantly, S. Dfa z-P ied rahita et a!. (2006) described 8 new species and I new variety in Espeletia: E. ariana Rodrfguez-Cabeza & S. Dfaz, E. betancurii RodrfguezCabeza & S. Dfaz, E. curialensis var. exigua Rodrfguez-Cabeza & S. Dfaz, E. episcopalis Rodrfguez-Cabeza & S. Dfaz, E. formosa S. Dfaz & Rodrfguez-Cabeza, E. mutabilis S. Dfaz & Rodrfguez-Cabeza , E. pisbana S. Dfaz & RodrfguezCabeza, E. pulcherrima Rodrfguez-Cabeza & S. Dfaz, and E. soroca S. Dfaz & Rodrfguez-Cabeza.]

Section 1. Aristeguietana Cuatrec. sect. nov. [Type species: Espeletia aristeguietana Cuatrec.].

Phyllaria exteriora involucri ovata vel oblongoovata abaxiale subappresse villosa. Indumentum foliorum patulo-tomentoso, pilis patentibus rectis vel tantum apice breviter flexuosis. Receptaculum hirtum. Venezuela: Trujillo.

1. Espeletia aristeguietana Cuatrec., Phytologia 27(3): 174-176. 1973b. [Type: Paramo La Cristalina, La Canada 2500-2600m , Venezuela, Cuatrecasas, Ruiz-Terdn & L6pez-Figueiras 28194, holotype US].

Figs. 14-9A-14-ll ; Pl. 14-lA Caulirosula sessile or with a trunk up to lOOcm high, 5cm in diameter, densely covered by marcescent leaves. Rosettes rather large (70- 90cm in diameter) with loose, erect and patent, large leaves of yellowgreenish white appearance. Initial young leaves surrounding the terminal bud with very thick, dense, compressed, long-hairy villous-sericeous tomentum.

Leaves coriaceous, thick and soft to the touch, somewhat flexible when young, rigid when adult, sessile, total length 46-60cm. Laminae suboblanceolateoblong, attenuate and acute at apex, narrowing gradually to the base, 43-56 em x 7.5-12(- 14) em, length to width ratio (4-)4.6-5.7:1 , width above the base 2.3- 3.8cm, margins revolute and thickened, entire, or with remote, minute, callose teeth abaxially hidden; adaxially costa and secondary nerves conspicuous; abaxially, under indument, costa thick and prominent, secondary nerves prominent, 8-20mm apart, slightly curved ascending, the deviation angle 60-70(50- 75) 0 , more curved or furcate near margins and anastomosing, tertiary nerves raised, irregularly transverse, joining with minor vei ns into a prominent reticulum. Indument yellow-greenish white, soft and velvety to the touch, adaxially subvelvety-tomentose, the hairs patent, slightly ascendent, more curved when old, 0.5-l.Omm long, thickened at base, the end very thin, straight or curved or even hooked; above

the costa very dense, straight, antrorse appressedsericeous hairs, 3 mm long; abaxially thicker and softer tomentose, hairs 1.5- 3.0 mm long, proximally patent, distally very thin, curved or inclined, ascending, forming uniform covering, but on the costa much more dense, appressed or subappressed, sericeous.

Sheaths robust, subtrapezoidal, 3.0-3.5 x 4.0- 5.0cm, adaxially glabrous and conspicuously plurinerved, abaxially long, tawny, sericeous-barbate.

Synftorescences, 2x longer than rosettes, 7090cm x about 30cm. Axes robust near the base, up to 2.0-2.5 em in diameter, fistulose, cylindrical , sharply striate. Proximal, vegetative part about 1!3 of total length, with 2 pairs of sterile leaves; lower leaves near the base 26-28cm x 4.5cm, similar to rosette leaves but smaller, the base sheathed and tubular, medial leaves 13-20 em x 3.5 em. Distal, fertile part l/ 2 or 2/3 of total length, broadly paniculate, floribundous, with up to 90 capitula distributed on about 5 pairs of branches; lower opposite branches robust, 25- 40 cm long, with distal branching about 3x in opposite, corymbiform cymes; medial opposite branches similar but smaller, the upper branchlets opposite or alternate, carrying from 3 to 1 capitula; general branching gradually decreasing in size upwards, the compound panicle with a rather rounded top. Peduncles 1.53.0 em long, usually erect. Subtending bracts smaller than the respective branches, the proximal ones 8-6cm x 1.8-1.5cm, oblong, obtuse, the others progressively decreasing upwards from 4cm x I em to 1 em x 0.15 em, oblong, linear, acute. Indument on axis when young densely whitish-tomentose or villous-tomentose, the hairs at first straight, patentascending, 1-2(3- 5) mm long, more or less flexuou s, when old, shorter, crisped-lanate; branches, branchlets, and peduncles densely tomentose or tomentosevillous, the hairs 2- 3 mm long, patent at base, thin at the end, ascendent, somewhat intricate; subtending bracts densely whitish tomentose-villous.

Capitula radiate, medium-sized, subglobose, l014mm in diameter, each with 112-127 flowers; ligular circles 24-28 mm in diameter, the discs 10-11 mm

ESPELETTA

in which the var. mucurubana represents the extreme form.

Origin. The variations described above indicate instability and a probable hybrid origin. Espeletia jajoensis may be the result of a cross-pollination probably between E. schultzii and Espeletiopsis cristalinensis (currently considered possibly to be Libanothamnus liscanoanus). The fertile product may be stabilized partially with some fluctuating features. The broad base of the typical leaves of E. jajoensis is also a character of E. schultzii, whereas the narrower base in some specimens represents an E. cristalinensis character. The closer, parallel, secondary nerves of E. schultzii are present in some individuals of E. jajoensis, but the more separate and angular nerves in some others may represent an introgression from E. cristalinensis. The opposite elements of the synfiorescence may be inherited from E. schultzii, and the alternate from Espeletiopsis introgression. Until the taxonomic status of this species can be resolved, it is placed in this informal group.

EXCLUDED SPECIES This list shows names of species once included in Espeletia that now are considered not Espeletiinae or are in the genus Espeletiopsis, which is not treated in this monograph.

Espeletia amplexicaulis Nutt., J. Acad. Nat. Sci. Philadelphia 7: 38. 1834. =Wyethia amplexicaulis (Nutt.) Nutt.

E. angustifolia Cuatrec., Bol. Soc. Venez. Ci. Nat. 17(85): 80. 1956. = Espeletiopsis angustifolia (Cuatrec.) Cuatrec.

E. bogotensis Cuatrec., Revista Acad. Colomb. Ci. Exact. 3: 427. 1940. =Espeletiopsis bogotensis (Cuatrec.) Cuatrec.

E. caldasii Cuatrec., Revista Acad. Colomb. Ci .

Exact. 3: 431. 1940. = Espeletiopsis caldasii (Cuatrec.) Cuatrec.

E. colombiana Cuatrec., Revista Acad. Colomb.

Ci. Exact. 3: 249. 1940. =Espeletiopsis colombiana (Cuatrec.) Cuatrec.

E. corymbosa H.&B., Pl. Aequin. 2: 16. 1808 = Espeletiopsis corymbosa (Humb. & Bonpl.)

Cuatrec.

379

E. funckii Sch.Bip. ex Wedd., Chlor. And. 1: 64.

1856 = Espeletiopsis funckii (Sch.Bip. ex Wedd.)

Cuatrec.

E. garciae Cuatrec., Phytologia 23(4): 358. May 1972; Ciencias 27(6). Oct 1972. = Espeletiopsis garciae (Cuatrec.) Cuatrec.

E . guacharaca Diaz, Caldasia 11(53): 19. 1975. = Espeletiopsis guacharaca (Diaz.) Cuatrec.

E. helianthoides Nutt., J. Acad. Nat. Sci. Philadelphia 7: 39. 1834. = Balsamorhiza sagittata (Pursh.) Nutt.

E. insignis Cuatrec., Revista Acad. Colomb. Ci. Exact. 3: 432. 1940. = Espeletiopsis insignis (Cuatrec.) Cuatrec.

E. jimenez-quesadae Cuatrec., Revista Acad. Colomb. Ci. Exact. 3: 247. 1940. = Espeletiopsis jimenez-quesadae (Cuatrec.)

Cuatrec.

E. muiska Cuatrec., Revista Acad. Colomb. Ci. Exact 3: 429. 1940. = Espeletiopsis muiska (Cuatrec.) Cuatrec.

E. pannosa Standi., Am. J. Bot. 2: 480. 1915. = Espeletiopsis pannosa (Standi.) Cuatrec.

E. petiolata Cuatrec., Revista Acad. Colomb. Ci. Exact. 4: 338. 1941. = Espeletiopsis petiolata (Cuatrec.) Cuatrec.

E. pleiochasia Cuatrec., Revista Acad. Colomb.

Ci. Exact. 3: 432. 1940. = Espeletiopsis pleiochasia (Cuatrec.) Cuatrec.

E. pozoensis Cuatrec., Ciencia (Mexico) 6: 266.

1945. = Espeletiopsis pozoensis (Cuatrec.) Cuatrec.

E. purpurascens Cuatrec., Revista Acad. Colomb.

Ci. Exact. 5: 16. 1942. = Espeletiopsis purpurascens (Cuatrec.) Cuatrec.

E. sagittata (Pursh.) Nutt., J. Acad. Nat. Sci. Philadelphia 7: 39. 1834. = Balsamorhiza sagittata (Pursh.) Nutt.

E. santanderensis A. C. Sm., Brittonia 1: 527. 1935. = Espeletiopsis santanderensis (A.C. Sm.) Cuatrec.

E. sclerophylla Cuatrec., Revista Acad. Colomb.

Ci. Exact. 3: 436. 1940. = Espeletiopsis sclerophylla (Cuatrec.) Cuatrec.

E. trianae Cuatrec., Revista Acad. Colomb. Ci. Exact. 5: 18. 1942. = Espeletiopsis trianae (Cuatrec.) Cuatrec.

CARRAMBOA

381

showy. The capitula are usually 6-9 mm in diameter, rarely 10 mm, but C. tachirensis differs from the other species by having capitula 10-5 mm in diameter.

Its ligular circle is 27-33 mm in diameter, whereas the usual diameters in the other species range from 12 to 22mm.

The sterile phyllaries are only 4-6 in about 2 rows, usually quincuncially arranged, ovate-acuminate pointed, and 2.0- 4.5 mm x 1.2-3.0mm; C. tachirensis is an exception, with sterile phyllaries 6.77.5 mm x 3.5- 4.0 mm. The fertile phyllaries, except in C. tachirensis, are 2.5-4.0mmx 1.3-2.2mm, and the pales 2.8-4.0 mm x 1.2 m m, all brownish to reddish or tawny barbate distally and with some sparse, minute glands interspersed.

The total number of flowers found in a capitulum of Carramboa ranges from 28 to 79 (to 140 for C. tachirensis), the lowest number for a genus of the Espeletiinae, except Tamania.

The number of ray flowers in a capitulum is always lower than the number of disc flowers. The number of female flowers in most species varies between 10 and 23 (up to 52 for C. tachirensis), whereas the number of male flowers ranges from 20 to 61 (up to 94 for C. tachirensis).

For most of the species, in capitula with a larger number of total flowers, the ratio of male to female flowers is 2.3- 3.3:1. In smaller capitula with fewer flowers , the relative number of functionally male flowers diminishes, so the ratio is 5-1.8:1. An exception is C. tachirensis, with larger capitula and greater numbers of flowers, but a ratio of 1.3-1.70:12 .

ORIGIN AND RADIATION The genus Carramboa probably is derived directly from a common Espeletiinae ancestor and adopted the tree form . It retains the basic, spirally alternate phyllotaxis with usually large or very large, green, and sparsely or moderately pubescent leaves.

The main axes and branches have indefinite, terminal growth. The synflorescences are axillary, with branching primarily dichasial; at least the lower branches and bracts are opposite, whereas the upper ones may be opposite or alternate. Carramboa prob-

ably was adapted originally to a humid, montane forest that spread throughout the Sierra Nevada de Merida and into valleys north and southwards at somewhat lower elevations. The isothermic and permanently humid conditions of the cloud forest probably favored retention of the dichasial inflorescences, which contrast with the marked spirally alternate, vegetative leaves and branches characteristic of the subtribe. In the humid periods of the Pleistocene, the genus probably spread throughout the region, but vertical oscillations of the vegetation caused by the alternating periods of glaciation and warmer, drier periods eliminated it from large areas (Fig. 15-1). The presently known localities may represent mainly refuge habitats, especially the region of Trujillo State and other deforested areas. But Carramboa persists in deep, less accessible valleys and along still-forested banks of small creeks. There is no doubt that more localities and perhaps even undescribed species will be found with future explorations of the rugged Venezuelan Andes.

Carramboa species are limited to forested areas in the Andes of Venezuela (Fig. 15-2). They are scattered in some parts of the state of Trujillo (C. trujillensis and C. wurdackii). However, the genus grows profusely in the southern part of the Cordillera de Merida, south from Merida and southwestwards to Tachira at the headwaters of the Rfo Uribante. In the latter area, between longitudes 71 o and 72°E, C. badilloi and varieties are especially abundant.

ENVIRONMENT Carramboa occurs only in the cloudy, humid, Andean forest, where shade and rain are constant, within or at the margins of forested areas and along deep creeks. It grows best between 2300 m and 2600 m altitudes but ranges up as high as 3000m and sometimes higher and down as low as 2000 and 1900m (Fig. 15-3). The Andean forest where the species of Carramboa grow usually is populated also by species of Bejaria, Beilschmiedia, Brunellia, Cybianthus, Drimys, Escallonia, Gaiadendron, Hedyosmium, Hesperomeles, Ladenbergia, Libanothamnus, Myrica, Nectandra, Ocotea, Paragynoxys, Podocarpus, Prunus, and Weinmannia.

Key to Species of Carramboa I. Leaf laminae sessile, rather large, broadly lanceolate, acute, green on both surfaces; bases broadly auriculate-

amplectant. Capitula variable in size.

2. Capitula large, 10- 15 mm; ligular circles (l8-)22-33mm in diameter. Ray corollas 8- 13 mm x .. ... l. C. tachirensis 3.0-4.5 m m.. ... ... ................................ ................................... ................... ..............................

2. Capitula small, 6- 9 mm; ligular circles 16- 21 mm in diameter. Ray corollas 7.0-9.5 mm x 3.0- 4.5 mm ............................................................................................................................................ 2. C. trujillensis

382

A SYSTEMATIC STUDY OF THE SUBTRIBE ESPELETIINAE

FIG.

15-1. Present distribution of Carramboa in relation to glaciated area.

I. Leaf laminae petiolate. Capitu la small.

3. Leaf lamin ae ovate, usually large and wide, length to width ratio 1.3-2.2:1, green on both surfaces, bottoms of in trareticul ar alveolae rather large, open, gree n, with minute, glandular papill ae, glabrou s, or with sca rce hai rs similar to those on venation. Sy nflorescences with proxim al, vegetative parts aphyllous. Involuc res glabrous or subglabrous ........................................ .. .............. ........ ....................... 3. C. badilloi 3. Leaf lamin ae thi ck, lanceolate, oblong, length to width ratio 3- 6:1 ; adaxia ll y green, abaxially densely villous, felt-]ike or velvety, whitish; bottoms of intrareticular alveolae with dense, minute, white, curled-lanate indument. Synflorescences with proximal, vegetative parts leafy. Involucres villous.

4. Disc coroll as hairy. Proximal, vegetative parts of sy nflorescences with usually 2 nodes each with a pair of steril e leaves and I or 2 additional nodes with alternate leaves; nodes with encircling sheaths.

Petioles 2.0-4.5 em long ....................................... .. .................................................. .... .4. C. rodriguezii 4. Disc corollas glabrous. Proximal, vegetative parts of synflorescences w ith 8-10 foliate nodes encircled by sheaths, 2 or 3 nodes with decu ssate, sterile leaves, the others bearing single, alternate leaves, occasiona ll y all alternate. Petioles 1.0- 2.5 em long. .. .............. ..... ........................................... 5. C. wurdackii

1. Carramboa tachirensis (Aristeg.) Cuatrec., Phytologia 52(3): 158, 1982.

Figs. 15-4, 15-5B, C; Pl. 15-lD Espeletia tachirensis Ari steg uieta, Compositae in Fl. Venez. 10( 1): 427-428, 1964. [Type: Tachira: Paramo del Batall6n, 26 Sep 1956, Aristeguieta 2523, holotype YEN, isotype US .]

Espeletiopsis tachirensis (A ri steg.) Cuatrec., Phytologia 35(1): 56, 1976.

Tree about 8 m tall with trunk 15 em in diameter; branches robust, elongate; branchlets erect, each ending with a cluster of leaves. Leafy stems terete, internodes short, densely and appressed antrorsely villous, the hairs 3 mm long. Young plants often growing as

LIBANOTHAMNUS 82

417

80

78

76

74

66

Li banothamn us

6

PA C I FIC

r------r)(ti~~~---1--~~~~~~~L-~~~~~~~~::== ~==~W 0 o; E A N

4. arboreus 5. banksiaefolius 5b. ssp. granatesianus 6.tamanus 7. grillini i 8. divisoriensis 9. parvulus 10. spectabilis

4

FIG. 17-7. Present distr ibution of Libanothamnus species, excluding 1 L. neriifolius and 11 L. occultus (numbers refer to order of presentation in keys and description s that follow).

Alternate Abridged Key to Species of Libanothamnus I. Lobes of the di sc corollas glabrous.

2. Capitu la radiate .. .... ... ... ... .............. ......... ........ ......... ........... ......... .......... ................... ....................... ........ !. L. neriifolius 2. Capitula pseudodiscoid (no ray ligules) ........................ ..... .. ... .... .......... .... ........ ...... ... ..... .... ....... .. ... ...... ... ... 3. L. lucidus I. Lobes of disc coro ll as with rather numerou s clavate hairs.

3. Leaf sheaths barbate at both sides.

4. Leaf laminae dentate and narrowly oblong or oblong-elliptic, sinuate dentate or almost entire, subobtu se or obtuse at both ends, long-petiolate ..................................................................... 5. L. banksiaefolius 4. Leaf laminae entire.

5. Leaves pseudopetiolate; tree ...... ......... ......... .... ................ .......... ..................... ... ........ ............. ....... 6. L. tamanus 5. Leaves sessile; caulirosula...................... ....... ..... ........ ... ...... ............ ......... ..... .... ........... 10. L. spectabilis 3. Leaf-sheaths on ly abaxially barbate.

6. Seconda ry nerves 5-lOmm apart ...... ... ... ... ... ....... ...... .. .... ........................ ..................... .............. 2. L. liscanoanus 6. Secondary nerves much closer together.

7. Leaves pseudopetiolate.

8. Leaf laminae oblong, attenuate at both ends or lanceolate, 14- 25 em x 2.8- 6.7 em. Capitula (12- )14-!S mm in diameter excluding ray flowers; involucres usu ally imbricate .. ... ... ......... 4. L. arboreus 8. Leaf lami nae elliptic. Capitula 8-11(-13) mm in diameter excluding ray flowers; outer phyllaries covering interior ones.

9. Leaves small, laminae 7.0-14.5 em x 2.4-4.6cm, subacute or obtuse at apex .................... 7. L. griffinii 9. Leaves larger, laminae 19- 34cm x 5- 8 em, subobtuse or acutate at apex, cuneate at base .......................................... ... ... .. .... ... ... ... ... .. ... ... .... .. ......................... .... ........ ................. 8. L. divisoriensis 7. Leaves sessi le or subsessile.

10. Leaves subsessile and small, laminae 6-10(-lS)cm x 2.3- 4.5 em ......................... .............. 9. L. parvulus 10. Leaves sessile and larger, laminae IS-50 em x 2.6-17 em .............. ... ... ...... ... ... ... . .... ... .... ... . 11. L. occult us

Key to Species of Libanothamnus I. Leaf margins , seen from above, more or less dentate or serrate, teeth acute, 0.5-1.0 mm long, mucronate, the mucron s antrorsely patent, or the sinuses shallow and the teeth patent or recurved; laminae oblong, 17-30cm x 4.5- 6.5 em, the base usually obtuse, long-petiolate. Secondary nerves 1.0- 2.2 mm apart.

418

A SYSTEM ATIC STUDY OF THE SUBTRIBE ESPELETllNAE

var. neriifolius var. cristamontis neriifolius var. boconensis var. turmalensis

..--

var. columbicus liscanoanus banksiaefolius ssp. granatesianus griffinii Libanotham nus

arboreus divisoriensis taman us lucidus parvulus ssp. humbertii

1.....--

occult us

ssp. occultus ssp. oroquensis ssp. glossophyllus spectabilis

FIG.

17-8. Phylogenetic diagram of species, subspecies, and varieties of Libanothamnus.

Leaf sheaths appressed sericeous-villous on both sides. Capitula 18-22mm in diameter ........................................... 5-1. L. banksiaefolius ssp. banksiaefolius I. Leaf margins, seen from above, entire, revolute (sometimes dentate in seed lings); laminae usually cuneate at base, pseudopetiolate or sess ile.

2. Seconda ry nerves usually (3 - )4-8(- 10) mm apart.

3. Leaves distinctly pseudopetiolate. Capitula 7-IOmm in diam eter, each with 47- 11 8 flowers; outer, steri le phyllaries 2.5-4.5 mm x 2-4 mm, deltoid-ovate, oblong-triangu lar, acute or subacute; ligular circles 14- 20mm in diameter, disc 6- lOmm in diameter; ray corollas 5-8(-lO)mm lon g; disc corol las 3.0- 4.5 mmlong, lobes glabrou s..... ...................................... ... .. .......... ....... 1. L. neri(folius 3. Leaves sessile. Capitula 15-20mm in diameter, each with 86- 153 flowers; outer, steri le phyll aries 13-20 x 6-9 mm , ovate-acuminate, oblong, very acute; ligular circles 25-30 mm in diameter; disc 10- 15 mm in diameter; ray corollas 8- 13 mm long; disc corollas 5.5- 6.5 mm long, lobes with clavate hairs .. ............................. .. .................................................................................... 2. L. liscanoanns 2. Secondary nerves l-4mm apa rt.

4. Leaves distinctly pse udopetiolate.

5. Ray corollas elig ulate, reduced to the basal tube. Lobes of disc corollas glabrou s ............. .. .. .... 3. L. lucidus 5. Ray corollas ligulate. Lobes of disc corollas with clavate hairs.

LIBANOTHAMNUS

419

6. Capitula 14-18 mm in diameter, each with 176-262 flowers. Sterile phyllaries ovate, usually gradually imbricate. Disc corollas 5-6mm long. Leaf laminae 14-25 em x (2.8-)4.5-6.7 em, oblong, subacute or lanceolate, cuneate at base ..................................................................... .4. L. arboreus 6. Capitula 8- 12mm in diameter, each with 54-130 flowers. Phyllaries not conspicuously gradate.

7. Leaf sheaths densely, appressed-barbate on both sides. Lamina 20- 47 em long.

Pseudopetioles 2-4(-6)cm long.

8. Leaflaminae elliptic-oblong or broadly oblong, obtuse or obtusely cuneate at base, 20-40x7- 14cm (in young plants up to 70x !Scm), length to width ratio 2.2-3.0(-3.4):1; margins often sinuate-denticulate; abaxially ashy or light ochraceous; secondary nerves 1.5-3.0mm apart. Capitula 10-12mm in diameter, each with 65-130 flowers; ligular circles 16-26 mm in diameter. Disc corollas 4.0-4.2 mm long ... ........... ............ ................................................................. S-2. L. banksiaefolius ssp. granatesianus 8. Leaf laminae narrowly oblong, gradually attenuate to the cuneate, acute base, 24- 47 em x 3.2-6.5 em, length to width ratio (5.4-)7-9:1; margins entire; abaxially ferruginous or ochraceous; secondary nerves 1-2 mm apart. Capitula 10-12 mm in diameter, each with 62-128 flowers; ligular circles 19-25mm in diameter. Disc corollas 3.7- 4.0mm long ............................................................................................................ 6. L. tamanus 7. Leaf sheaths adaxially glabrous.

9. Leaf laminae 7.0-14.5cmx2.4-4.6cm, length to width ratio (2.5- )3.0 - 3.8:1; abaxially the indument ferruginous; secondary nerves l- 2mm apart. Pseudopetioles 1-3cm long. Disc corollas 4.5-5.3 mm long, tubes with sessile glands .................................... 7. L. griffinii 9. Leaf laminae 19-34cm x 5- 8 em, length to width ratio 3.8- 5:1; the indument abaxially whitish-ashy; secondary nerves 2-4mm apart. Pseudopetioles 2- 3cm long. Disc corollas 4-5 mm long, tubes with clavate hairs and pediculate-capitate glands .. 8. L. divisoriensis 4. Leaves sessile or subsessile.

10. Leaf lamina 6-18 em long, subsessile, abaxially thickly and densely lanate, covering nervation, the base contracted like a winged pseudopetiole 0-8 mm long. Capitula 8- 11 mm in diameter, each with 54- 105 flowers.

11. Leaflaminae 6- 10(-15)cm x 2.3- 4.5cm, elliptic or oblong-elliptic, obtuse or subotuse at apex, margins strongly revolute; secondary nerves l-2mm apart. Capitula 10- 11 mm in diameter, each with 68- 105 flowers. Sterile phyllaries 4.5-5.5 mm x 4.0-5.5 mm, broadly ovate. Disc corollas 4- 5 mm long, tubes pilose and glanduliferous, lobes clavate-pilose ........ 9. L. parvulus II. Leaf laminae 9- 15 em x 3-4 em, oblong-elliptic, acute at apex, margins inconspicuously revolute; secondary nerves 2-4mm apart. Capitula 8- IOmm in diameter. Sterile phyllaries 3.5- 4.0mm x 2.3-2.8 mm, ovate-triangular, subobtuse or subacute. Disc corollas 3.4-3.7 mm long, tubes distally somewhat pilose, lobes without hairs ................. l-2. L. neriifolius var. cristamontis 10. Leaf laminae surpassing !Scm in length, broadly sessile at base.

12. Leaf sheaths adaxially villous-barbate. Caulirosula, monocaulous and monocarpic.

Laminae 40- 65 x 7.5-IS.Ocm, minimal width above base 2.0- 3.5 em. Secondary nerves 1.5-3.0(-4.0)mm apart. Synflorescences large, usually showy. Capitula 16- 20mm in diameter, each with 180-264 flowers. Sterile phyllaries 8-6 mm x 4- 5 em, oblong-ovate, subacute. Disc corollas 5.2-6.5 mm long ........................................................................ 10. L. spectabilis 12. Leaf sheaths adaxially glabrous. Tree, many-branched Laminae 18-34(-50) x 2-10(- 17) em, elliptic oblong, linguiform, lanciform, obovate-oblong or oblong-ovate, acute or obtuse.

Secondary nerves 1-4(- S)mm apart.. ................................................................................. II. L. occultus

1. Libanothamnus neriifolius (Bonpl. ex Humb.)

Figs. 17-9- 17-14; Ernst, Vargasia 7: 186. 1870.

Pl. 17-1 Trixis neriifolia Bonp!. ex Hum b., Voy. Reg. Eguin. Rei. 1: 605. 1814. As T. nereifolia, typographic error.

[Type: "Syngenesia necessaria, Trixis neriifolia foliis lanceolatis integerrimis subtus laniculatis, Habitat in Caracas." "Incienso, Silla de Caracas (Humboldt)

Cass.," lectotype Herb. Willdenow 16672 in B, photo FM-16103. Isolectotype: "Caracas, Jan 1800, No. 652, incienso, Buphthalmum, Silla de Caracas," in other label "Andromachia? Trixis neriifolia Willd!

Espeletia neriifolia Sch. Bip. 19/10/55, Silla de Caracas, Herbier donne par M. Bonpland in 1833," P.] Baillieria? neriifolia H.B.K., Nov. Gen. Sp. Pl. 4: 289.

1820; Folio. 4: 227. 1818.

Clibadium? neriifolium DC., Prodr. (DC.) 5: 507. 1836.

RUILOPEZIA

479

center, in the opposite direction from subgenus Ruilopezia. Otherwise, the two sections are rather close.

The present species grows in depressions of rather swampy or wet areas in the region of Paramo de Las Coloradas at 2600- 2800m altitudes.

The major diversification within Ruilopezia has taken place along the southwestern hills of the Cordillera de Merida. One of the most remarkable, xeromorph ic species of the genus, R. jahnii, is well suited for the open, rocky, and stressful paramo environment.

This species probably spread through a more extensive region during the last glacial period or most likely through a dry environment during earlier glaciations.

The present distribution is reduced to the paramos of Zumbador and Batall6n, from 3000 to 3600m altitudes, at the southwestern end of the Cordillera. This multi leaved species forms compact rosettes with narrowly linear, recurved, stiffleaves; lanate indument on the stout panicles; and flowers concentrated in globose and rayless capitula. It represents an end point in the migration to open high zones that have been subjected to strong eli matic, seasonal, and diurnal oscillations.

A notable point of departure from the mainstream of Ruilopezia is the branch of the genus that extended into the hi ghest, open, grassy paramos of the Cordi!-

!era. This group, subgenus Argentea, adopted a sessile, caulirosular form usually with a large, tuberose corm and densely sericeous, linear leaves with a shiny, silvery appearance. The oldest apparent representative of the line, R. marga rita, with white rays occurs in the high paramos of Santo DomingoGranates. Along the line of migration to the southwest of the Cordillera is the widespread R. jloccosa with yellow rays, representing the genus in the central paramos of Sierras Nevadas, Apartaderos, and Sierra del Norte (Sierra de la Culata). The lack of species of this group in the immediate, southern hills of Merida is probably the result of extinction of older species, leaving a relictual population of the whiteflowered R. leucactina on the distant Paramo del Batall6n at 3300 to 3300m altitudes. Migration to the lower hills of the eastern front of the Cordillera produced R. vergarae and the most xeromorphic of the group, R. jabonensis, which grows on the open, upper slopes and tops of the windy hills, such as in the upper region of Paramo del Cende, Jab6n at 3400 m.

The latter species also invades deforested areas at lower altitudes.

Relationships of the subgenera and species of Ruilopezia are summarized in Figure 18-3.

Key to Subgenera of Ruilopezia J. Phyllaries ovate or broadly oblong, herbaceous, green, sparsely pilose, strongly nerved, glanduliferous.

Leaf laminae sessile, herbaceous-chartaceous, green on both sides or subcoriaceous and white vested.

Caulirosula, usually tall .................................... ... ................................................................... Subgenus 2. Ma crophyllaria J. Phyllaries linear, oblong or ovate, densely villous, sericeous, or lanate. Leaf lami nae coriaceous or chartaceous, densely la nate or sericeous, at least on the abax ial side.

2. Receptacles obovoid or ovoid, strongly protruding. Capitula globose. Leaf lam inae very narrowly linea r; ma rgins strongly revolute; ad ax iall y glabrous and green; abaxially venose-reticulate and lan ate.

Caulirosul a, densely leafy. Synflorescences obconica l, floribundou s, densely white lanate ....... .... ... ... ... ... ....... ................ .. ... ... ... ... ...... ... ... ......... ............. ............. ............ ... ... ... ... ...... ......... Subgenus 3. A/fredonia 2. Receptacles moderately convex or conica l.

3. Second ary nerves of leaf lamin ae para llel a nd longitudinal or ascending at an angle of up to 45°.

4. Leaf laminae oblong-obovate, broadly oblanceolate or elliptic-oblanceolate, rigidly coriaceous; secondary nerves diverging at an angle of 30- 45°; pseudopetioles long, narrow, rigid , rarely short. Phyllaries linear or shortly ovate ..................................................................... Subgenus I. Frailejonota 4. Leaf lami nae flat, narrow, linear, ens iform, or narrowly lanceolate, sess ile, usually g reen at least adax ially. Caulirosula, green with monocotyledonous aspect.

5. Median and proximal parts of the leaf laminae parallel-nerved; distally the nerves diverging at an angle of 5- 15°. Leaf sheaths very short and almost indistinct from the laminae.

Sy nflorescences narrowly paniculate, green. Capitul a semiglobose ............. .. ... .Subgenus 4. Bromeliaster 5. Secondary nerves of leaf la minae diverging up from the base at an acute a ngle of 5-40° ................. ... ... ......... ... ... .. ............. .. ... .... .................................................................... Subgenus 5. Ruilopezia 3. Second ary ne rves of leaf laminae diverging a t a n angle of 50- 90°. Lamin ae usually covered by a white, cine reous, or ochroleucous indument, sometimes adaxially glabrate and green when mature.

6. Tall caulirosula, with a trunk up to 3 m. Leaf laminae sessile or pseudopetiolate; adax ially whiti sh lanate or grayish, sometimes glabrate and green; abaxially la nate or villous-lanug inou s, white, gray, or ochroleucous ..... ................................................................................. Subgenus 6. Ruizonia 6. Sessile caulirosula with trunk usually somewhat tuberose. Leaf laminae sess il e, linear or sublin ear; adaxially si lvery ser iceo us, shiny ............................. ......... ... ....... ........ .. ......... Subgenus 7. Argentea

RUILOPEZIA

481

Key to Subgenera and Species of Ruilopezia I. Leaf laminae broadly obovate-lanceolate, oblanceolate, or elliptic, contracted into a long, strong

pseudopetiole or abruptly contracted at the base and sessile, rigid-coriaceous or chartaceous; secondary nerves diverging at a 30-45(-60) 0 angle; adaxially glabrous or sub glabrous at maturity. Rosettes sessile, with rhizomatous trunk producing replacement shoots or short branches or an emerging caulirosulan trunk ............ ........................... ..................................................................................................... Subgenus 1. Frailejonota 2. Leaf laminae rigidly, thickly coriaceous, strongly long-pseudopetiolate, 20-75 em x 2-9 em; obovate, obovate-lanceolate or broad-lanceolate. Rosettes sessile.

3. Capitula radiate. Ray corollas well developed, narrowly oblong, 6.5-13.0mm long, yellow. Leaf laminae densely, thickly lanate abaxially. Synflorescences 1- 2m long, corymboid-paniculate ............. 1. R. bracteosa 3. Capitula pseudodiscoid. Ray corollas reduced to short tubes, with dentiform or short obsolete appendages. Leaf laminae appressed sericeous abaxially. Synflorescences 1-2m long, thyrsoidpaniculate .. .............................................................................. ..................... ................................ 2. R. atropurpurea 2. Leaf laminae firmly, rather flexibly chartaceous, sessile or subsessile, 12-20cm x 3.0-3.3 em; lanceolate-elliptic, densely appressed-sericeous abaxially. Capitula radiate; ray corollas 6.0- 7.5 mm long, elliptical, white or with some purplish blotches. Synftorescences moderately laxly branched. Low caulirosula; stems sometimes branched .................... .......................................................................... 3. R. usubillagae 1. Leaf laminae not broadly obovate-lanceolate or elliptical and not strongly long-petiolate nor abruptly contracted at base; several shapes from linear to oblong or elliptic-oblong.

4. Outer phyllaries broadly ovate or ovate-oblong, 26-13 mm x 16-5(-3) mm , firmly herbaceous; nerves prominulous, reticulate; surfaces green, reddish, or purplish stained, copiously glanduliferous and sparsely pubescent or ciliate at margins. Leaf laminae sessile with lateral nerves diverging at an angle of 60-85° .......................................................................................................................... Subgenus 2. Macrophyllaria 5. Leaf laminae herbaceous-chartaceous, oblong or elliptic-oblong, 20-50(- 60)cm x 3.5-8.5(- 10.0) em, green on both sides, somewhat pubescent; margins flat or slightly revolute, sinuate-dentate with the teeth long mucronate, prominently nerved, abaxially and loosely reticulate; sessile, the base continued with lateral wings to the crisped-marginate somewhat undulate-auriculate sheaths.

Caulirosulae 2-6(-12)m tall, simple. Synflorescences corymboid-paniculate.

6. Indument of leaf laminae abaxially with rigid, straight or curved, acute hairs, thickened at base, patent or subpatent, l-2mm long; veins with very copious or scattered hairs; reticular areoles flat, glabrous, green. Axes and branches of synftorescences green, hirsute, the hairs firm, patent.

Peduncles hirsutulous and glanduliferous. Subtending bracts ovate, foliaceous. Leaf sheaths of synftorescence very short, green. Outer phyllaries ovate, reticulate. Ray corollas white or creamy or greenish white, 12-17mm long ........................................................................................... .4. R. rnarcescens 6. Indument of leaf laminae abaxially very fine, the hairs long, flexuous, ascendent, forming a sericeous veil over the costa, intricate and arachnoid on rest of the surface; reticular alveolae densely white lanate. Axes and branches of synflorescences whitish villous-lanuginous, the hairs appressed. Peduncles appressed silky-lanate, not glanduliferous. Subtending bracts oblong, distally linear, with sheathing bases. Leaf sheaths of synflorescences long and reddish. Outer phyllaries oblong or elliptical, acutate, parallel-nerved except distally reticulate. Ray corollas white-roseate, reddish, or purplish, 7-9 mm long ......................................................................................... 5. R. cuatrecasasii 5. Leaflaminae coriaceous, linear-oblong, or narrowly sublanceolate, 14-35(-56)cm x 1.6-4.0(-5.0)cm, adaxially greenish ashy, appressed-pilose, subsericeous, abaxially white lanate, the costa sericeous; margins revolute, apparently entire; sheaths coriaceous, ovate, neither winged nor auriculate.

Caulirosula with trunk up to 1m tall. Synflorescences thyrsoid-paniculate. Outer phyllaries ovate, acute, 20- 12mmx 12- 5mm. Ray corollas white or greenish white, 10- 14mmx2.0- 3.5mm ..... ........... 6. R.lindenii 4. Outer phyllaries linear-triangular, linear-lanceolate, narrowly oblong or ovate, when ovate, smaller than in sub g. Macrophyllaria and densely villous or lanate.

7. Leaf laminae green or greenish adaxially with secondary nerves diverging at a very acute angle (5- 30°) or extremely narrowly linear, strongly recurved without secondary nerves. Phyllaries linear or linear-triangular.

8. Leaf laminae 15-36cm x 1.5-4.0mm, extremely narrowly linear, with strongly recurved margins and rigidly pointed apex; abaxially thickly costate and reticulately veined, lanate.

Capitula globose, pseudodiscoid. Receptacles protruding, ovoid to subglose. Stems branching, the branches procumbent or prostrate. Synftorescences robust, ftoribundous, corymbiform ............................................................................................... Subgenus 3. A/fredonia: 7. R.jahnii 8. Leaflaminae with margins slightly or narrowly recurved. Receptacles conical.

482

A SYSTEMATIC STUDY OF THE SUBTRIBE ESPELETIINAE 9. Leaf laminae linear-triangular or ensiform, acute, with ascend ing secondary nerves parallel to the costa in the lower lfz, the nerves angular distally; adaxially glabrous. Capitula pseudodiscoid, subglobose. Synflorescences distally corymboid. Stems shortly branched, prostrate. Terminal rosettes green , usually dense, appearing sessile with a monocotyledonous aspect .......................................................................... Subgenus 4. Bromeliaster: 8. R. bromelioides 9. Leaf laminae abaxially with angular, secondary nerves from the base, conspicuous or obsolete and veins reticulate .................................. .......... ........................................ .Subgenus 5. Ruilopezia 10. Leaf laminae adaxially glabrous, bright green or with only the costa pilose.

11. Capitula radiate. Ray coroll as yellow, 8-13 mm x 1.0- 1.8 mm. Leaf laminae abaxially pale green or ci nereous green, the costa glabrous, the venular reticulum conspicuous, the alveolae minutely, densely, appressed lanate. Peduncles subappressed long-villous. Rosettes sessile. Trunks cryptoligneous, undivided or with short, basal shoots. Synflorescences corymboid-paniculate ............ .. ........... ............... 9. R. viridis 11. Capitula pseudodi scoid. Ray corollas reduced to tubes, 0.8-1.5 mm long, truncate or with short, dentiform projections. Leaf laminae abaxially white or whitish; indument thickly lanate, hairs covering the whole reticu lu m; the costa glabrous and green. Pedicels sparsely glanduliferous and copiously lanugi nous, somewhat hirsutulous. Rosettes topping unbranched stem up to 2.3 m tall. Synflorescences thyrsoid-paniculate ......................... ......................................... .................. ............ lO. R. figueirasii 10. Leaf laminae adaxially pi liferous throughout.

12. Capitula long-radiate. Ray corollas 5.0-6.7 mm x 1.2- 2.5 mm, white or creamy white. Leaf laminae adaxially cinereous-green, appres sed sericeous, slightly substrigose, the hairs 0.7-1.0 mm long; abaxially whitish ochroleucous, densely sericeous, the hairs antrorse, somewhat appressed, rigidulous , intricate. Sheaths deltate-oblong, attenuate at top. Synflorescences floribundou s, multiflorous, thyrsoid-paniculate, !.0- 1.8 m x 0.8-!.5 m, pyramidal, rounded at top .......... II. R. lopez-palacii 12. Capitula short-radiate. Ray corollas 2.2-3.5 mm x 0.7-1.4 mm , pale yellow or greenish yellow. Leaf laminae adax ially green or light green , asperulous, shortly subsericeous-strigose, the hairs rigid and pointed, O.l-0.4mm long; abaxially appressed, silvery sericeou s. Sheaths ovate, obtuse or emarg inate at apex.

Synflorescences subcorymboid-paniculate, 30-45cm high, usually twice as wide, very crowded , branched from the base .................................... .. ....................... 12. R. paltonioides 7. Leaf laminae covered throughout with a white indument or ad axially glabrate at maturity with seco ndary nerves conspicuously parallel at deviation angle of 50- 90°.

13. Caulirosula stipitate; trunk erect, 1/z to several m tall. ............................. ...................... Subgenus 6. Ruizonia 14. Ray corollas white or whitish.

15. Leaf laminae adaxially permanently piliferous, pale green or gray-green, thinly villous, the hairs long and thin , somewhat flexuou s, appressed; abaxially greenish ashy, densely villous-lanuginous; secondary nerves weak but conspicuous, parallel with deviation angle of 50-70°. Ray corollas small , 4-6x0.5- l.5 mm , whitish. Disc corollas 5.0-5.5 mm long, glabrous or subglabrous except for the lobes .................... 13. R. grisea 15. Leaf laminae becoming glabrous and green adaxially.

16. Leaf laminae thickly coriaceous, rigid , long-linear, acute, 28-50cm x 1.5- 3.0cm., with thickly revolute margins; adaxia lly shiny green, somewhat viscid, abaxially densely, thickly, compressed curly-lanate, ochraceous, secondary nerves closely parallel, 1.5- 2.0 mm apart, patent, at deviation angle of 80- 90°; base attenuated into a pseudopetiole, 4- 8 em long and almost reduced to width of the costa.

Sheaths triangul ar, appressed-sericeous on both surfaces. Synflorescences robust, corymboid-paniculate, 60-100 em x 80-120 em . Ray corollas 7.5- 11.0 mm long, tubes 1 mm long. Disc corollas 4.1-4.5mm long, glabrous except for the barbate lobes .. ... ........................... ... ... ........ ....... ..... ........ ... ............................................... 14. R. cardonae 16. Leaf laminae coriaceous, obtuse, subobtuse or acute at apex , oblanceolate-linear, linear or lanceolate, adaxially not glutinous; abaxially thickly white lanate or canescent. Sheaths fully appressed-sericeous only abaxially.

17. Outer phyllaries ovate long-acum inate, 15.0-9.5 mm x 6.0-3.5 mm, apices conspicuously surpassing the ligular circles at full anthesis. Leaf laminae linear or narrowly oblanceolate, acute at apex , attenuated to base, sessile; adaxially glabrate, green; abaxially densely white lanate, secondary nerves

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with deviation angle of 80- 90°. Ray corollas 5.2-8.0mm x 2- 3 mm. Disc corollas 5.4- 6.0 mm long; tubes copiously glanduliferous ............. ... ........ .... .. 15. R. ruizii 17. Outer phyllaries not exceeding the ligular circles.

18. Leaf lamin ae sessile, linear, oblanceolate or sublanceolate-linear, rather narrow, 22- 43 em x 1.2-2.2cm, obtuse or subobtuse; adaxially olive-green in adult stage; abaxially canescent villous-lanuginous and overall subsericeous-barbate; secondary nerves with deviation angle of (75 - )80-90°. Outer phyllaries long villous-barbate, 17-11 mm x 5-3 mm.

Ray coroll as ll - 16mmx2.0- 3.5mm. Disc coroll as 6- 7 mm long; tubes sparsely pi lose ....... .. ....................................................................... 16. R. hanburiana 18. Leaf laminae pseudopetiolate, oblong-oblanceolate or lanceolate, acute, 24-41 em x 2- 6cm; in adult stage ad axially glabrous, green or yellowish green; abaxially densely, thickly lanate, wh ite, ci nereous or creamy greenish; secondary nerves with deviation angle of (60-)65- 75(-80) 0 • Outer phyll aries vi llous and sparsely glandul ar, 7- 5 mm x 4-3 mm. Ray corollas 4.0- 7.3 mm x 1.5-2.2 mm. Disc coroll as 3.8-4. 8 mm long; tubes pilose and glandu liferous .................... ........................................ . 17. R. coloradarum 14. Ray corollas yellow. Leaf lam inae lanceolate, acute.

19. Leaf laminae adaxially glabrous at maturity, sessile, although attenu ate to the base, 27-48cmx4.2-8.0cm; abaxia lly densely lanate, greenish ci nereous; secondary nerves with deviation angle of 65-80(-90t. Sheaths ovate or semiorbicular, 3.0-4.4 em x 3.58.0cm. Outer phyllaries ovate-acum inate 6-9 mm x 3.5- 5.0 mm , sparsely villous and densely glanduliferous. Ray corollas 6.5- 7.8mm x 2.0-2.6mm. Disc corollas 6.0-6.5 mm long; tubes somewhat glanduliferous and pilose ................................ 18. R. josephensis 19. Leaf laminae adaxia lly permanently, densely, appressed subsericeous, 3748cm x3.8- 5.3cm, conti nued at base by a pseudopetiole l-4cm long; abaxially densely whiti sh sublanate, secondary nerves with deviation angle of 45-66°. Sheaths oblong, trapezoidal, 4.8- 5.5 em x 2.0-3.4cm. Outer phyll aries triangular-lanceolate, 5.5-8 .0mm x 1.4-2.0 mm, densely vill ous and sparsely glanduliferous. Ray corollas 8-lOmm x 1.5- 2.2 mm. Disc corollas 4.2-5.0mm long; tubes adaxially sca rcely pilose ................ ........ .... ................................ ........................................................... 9. R. emmanuelis 13. Cau lirosu la sessile, not stipitate. Leaf laminae permanently covered throughout by a dense, white indument, at least adaxially appressed-sericeous. Phyllaries linear or linear triangular, villous .. ............ ...... ...... .............................. ....... .................. ........................ ... .............. .... Subgenus 7. Argentea 20. Ray corollas white.

21. Leaf laminae adaxially glandular-glutinous and densely sericeou s, narrowly linear-lanceolate, 26- 40cm x 0.8-1.3 em, length to width ratio 21.5- 31(- 41 ): 1; secondary nerves ascendi ng at deviation angle up to 25° or obsolete. Sheaths oblong 3-6cm x 1.5-3.0(- 3.5)cm. Ray corollas 13-18 mm x 3-4 mm; tubes 2- 3 mm long.

Disc corollas 5-6mm long ....................................................... ... .. ..... ................ ... ... 20. R. margarita 21. Leaf laminae adaxia lly thickly and densely appressed, silvery, shiny sericeous, not glutinous, linear-lanceolate or ensiform, 38-43(- 48)cm x 1.9-3.2cm , length to width ratio 13-20(-29):1 ; secondary nerves conspicuous, parallel, patent, the deviation angle (60-)70-80° Sheaths triangu lar 4- 5 em x 3-7 em. Ray corollas 8- 10 mm x 1.52.5 mm; tubes 1.5-1.8 mm long. Disc corollas 4.2-4.5 mm long ............ ... ............. 21. R. leucactina 20. Ray corollas yellow.

22. Ray corollas 18-23 mm x 3-4 mm; tubes 3-4 mm lon g. Disc coroll as (6.0-)6.5 -7.8 mm long. Capitula 15- 20mm in diameter; ligular circles 40-50mm in diameter. Leaf lam inae linear, linear-lanceolate or ensiform , 25-32cm x 1.3- 2.0cm, length to width ratio 15-20(-25):1 ; seco ndary nerves with deviation angle of 27-40(-50t . Axes of sy nflorescences blunt, very short, branches congested, subvertic ill ate in 1-3 rows, and up to 1m lon g, producing copious branchlets in racemose arrangement. .................. .22. R.jioccosa 22. Ray corollas 8-19mm x l.2- 2.0(- 3.0)mm; tubes l.0-2.2m m long. Disc corollas 4.5-5.5 mm long. Capitula 10-15 mm in diameter; ligular circles 24-35(- 40) mm in diameter. Axes of synflorescences well developed and equalling or longer than the basal branches, these and branchlets in corymboid arrangement.

23. Leaf laminae strictly linear or narrowly linear-lanc eolate, 10-30cmx0.3-0.7 (-0.9) em, length to width ratio 33- 42: I ; margin s strongly revolute and covering

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in diameter. All three species are widespread throughout the upper limits of the superparamo, although C. moritziana is the only one living successfully at the highest altitude, 4800 m, on rocky crests emerging from permanent snow blocks. Nevertheless, C. timotensis is more apt to thrive in this zone, in spite of the great difficulties that solifiution poses for seedling survival (the mortality rate may reach 100%). This species, helped by its unique structure and longevity, has managed not only to survive but to dominate large areas of this hostile habitat. Well-based estimates (A. P. Smith, personal communication 1978) indicated that adult individuals of C. timotensis greatly surpass 100 years in age. Relationships of these six species and one variety are summarized in Figure 19-2.

Coespeletia is the most characteristic genus of the Venezuelan superparamo landscape and the best adapted to the environment around and above the freezing line. The caulirosulan structure of Coespeletia is certainly a superb, morphophysiological design for protection against the extreme conditions. The massive columnar trunk consists mainly of a cover of marcescent, exceedingly tight leaves that is much thicker, more compact, and insulating than the vestiture of any Espeletiinae of the wetter Colombian paramos. These thick trunks give the impression of great stability and majesty. Considering the complexity of its morphology, the strategies developed by Coespeletia unquestionably represent a major, evolutionary success in the Espeletiinae.

Key to Species of Coespeletia 1. Capitula large, 35-50(-80) mm in diameter. Outer phyllaries (16-)20-32 mm long. Ray corollas much longer

than styles, tubes glanduliferous, the glands capitate-pediculate. Disc corollas glanduliferous or glabrous.

2. Synftorescences of single, large capitulum 40-50(-80)mm in diameter supported by a long, axillary scape. Tubes of ray corollas copiously glanduliferous; tubes of disc corollas glanduliferous; lobes and limbs glabrous or with a few glands at base of limbs. Leaf laminae strictly narrowly linear or lanceolate-linear, 0.6-1.5(-2.0)cm wide, length to width ratio (16-)23- 40:1. Caulirosula usually subsessile or short stemmed ................................................................................................................ I. C. moritziana 2. Synftorescences simple, determinate racemes of 8-18 monocephalous branchlets. Capitula 35- 55mm in diameter. Leaf laminae broadly linear or linear-oblanceolate, 1.2- 5.0cm wide, length to width ratio 9-18:1. Caulirosula usually tall.

3. Synftorescences usually with 8 to 18 monocephalous branch lets (peduncles); axes 85-48 em long.

Peduncles usually 3- 15(- 17)cm long, rather weak and ftexuose. Ray corollas 5.5-8.0(- 9.0)mm long; tubes copiously glanduliferous. Disc corollas sparsely glanduliferous. Leaflaminae widely linear or I inear-oblanceolate, (2.2-)2.5-4.0(-5.0)cm wide, length to width ratio 10-18:1 .......................... 2. C. timotensis 3. Synftorescences usually with 9- 11 monocephalous branchlets; axes 60-80cm long.

Peduncles 4- 7(- 10)cm long, rather thick, straight, erect, rigid, rarely curved. Ray corollas 11 - 13mm long; tubes copiously glanduliferous and also hirtellous. Disc corollas glabrous, sometimes sparsely glanduliferous. Leaf laminae linear oblanceolate, 2.8-5.0cm wide, length to width ratio 9-10:1 ................................................................................................................................. 3. C. albarregensis 1. Capitula smaller, (10-)15- 26mm in diameter. Outer phyllaries 9- 15 mm long. Ray-corolla tubes densely hirsute. Disc corollas somewhat hirtellous, nonglanduliferous, or occasionally with a few scattered, minute glands.

4. Synftorescences simply racemose; branchlets (peduncles) 14-38, all monocephalous. Ray corollas 1.3-4.5(-5.0)mm long, shorter than the styles. Leaf laminae narrowly linear, 1.2- 2.0cm wide.

Caulirosulas tall, usually 1.0-2.5 m ......................................................................................................... 4. C. spicata 4. Synftorescences narrowly racemose-thyrsoid; distal branchlets monocephalous, median and proximal branchlets polycephalous. Ray corollas 3- 5 mm long, longer than or equaling the styles. Leaf laminae broader, 3.5-8.0cm wide.

5. Leaf laminae broadly oblong or lanceolate-oblong, shortly attenuate or contracted at base, 26-47 em x 3.5-8.0cm, length to width ratio 4-10:1. Pseudopetioles, if any, 1-2cm x 1-3 em. Fertile part of synftorescences with usually 17- 33 branchlets, median and proximal ones each with 3- 7 capitula, 15-22 mm in diameter. Ray corollas 3-5 mm long, about equal to or exceeding the styles.

Caulirosula subsessile or with stem up to 50 em tall ................................................................... 5. C. thyrsiformis 5. Leaf laminae oblong-oblanceolate or lanceolate, progressively attenuate at the base into a long pseudopetiole, 49- 68 em x 3.5- 5.0cm, length to width ratio 9-18:1. Pseudopetioles 5-10cm x0.7- l.Ocm. Fertile part of synftorescences robust and elongated, with 30- 59 branch lets, median and proximal ones each with 2-4 capitula, 15-26 mm in diameter. Ray corollas always longer than the styles.

Caulirosula usually with long stem, 1.0-2.5 m ................................................................................. 6. C. elongata