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Darwiniana, nueva serie

versión On-line ISSN 1850-1699

Darwiniana v.45 n.1 San Isidro ene./jul. 2007

 

Notes on the genus Caiophora (Loasoideae, Loasaceae) in Chile and neighbouring countries

Markus Ackermann1 & Maximilian Weigend

Institut für Biologie, Systematische Botanik und Pflanzengeographie, Freie Universität Berlin, 14195 Berlin, Germany; ackermal@zedat.fu-berlin.de (1author for correspondence)

Abstract. Ackermann, M. & M. Weigend. 2007. Notes on the genus Caiophora (Loasoideae, Loasaceae) in Chile and neighbouring countries.
   This is the first revision for the representatives of the genus Caiophora (Loasaceae) in Chile. The genus is widely distributed in the Andes from Argentina/Chile in the South to Central Ecuador in the North, and comprises approximately 60 species. In Chile only five species are present, Caiophora chuquitensis, C. cirsiifolia, C. coronata, C. deserticola sp. nov. and C. rosulata. Caiophora rosulata is here subdivided into two subspecies: western Andean C. rosulata subsp. rosulata (present in northern Chile and southern Peru), and eastern Andean C. rosulata subsp. taraxacoides, stat. and comb. nov. Furthermore C. superba syn. nov. and C. macrocarpa syn. nov. are placed into synonymy under C. chuquitensis, and C. rahmeri syn. nov. is synonymized to C. rosulata subsp. rosulata. These five species comprise the complete range of growth forms known for the genus, i.e., subshrubs, cushionforming herbs, acaulescent, rosulate herbs and vines. For all taxa a key and full morphological descriptions and synonymy are provided, including illustrations, notes on habitat, distribution, floral biology and chromosome numbers.

Keywords. Caiophora; Chile; Chromosome number; High Andean flora; Loasaceae; Morphology; Pollination; Taxonomy.

Resumen. Ackermann, M. & M. Weigend. 2007. Notas sobre el Género Caiophora (Loasoideae, Loasaceae) en Chile y países limítrofes.
   Esta es la primera revisión del género Caiophora (Loasaceae) en Chile. El género comprende alrededor de 60 especies, y está ampliamente distribuido en los Andes, desde Argentina/Chile en el Sur, hasta el centro de Ecuador en el Norte. Se conocen solamente cinco especies de Chile: Caiophora chuquitensis, C. cirsiifolia, C. coronata, C. deserticola sp. nov. y C. rosulata. El material de Caiophora rosulata se divide entre dos subespecies, C. rosulata subsp. rosulata comb. nov., de los Andes Occidentales (presente en el norte de Chile y el sur de Perú, y C. rosulata subsp. taraxacoides stat. y comb. nov., de los Andes Orientales. Los nombres C. superba syn. nov. y C. macrocarpa syn. nov. son sinonimizados bajo C. chuquitensis, y C. rahmeri syn. nov., sinonimizado bajo C. rosulata subsp. rosulata. Estas cinco especies comprenden el rango completo de hábitos conocido para el género: sufrútices, hierbas perennes en forma de cojines, hierbas rosuladas acaules, y hierbas trepadoras. Proporcionamos una clave, descripciones morfológicas y la sinonimia completa para todos los taxones, incluyendo ilustraciones, notas sobre la distribución, el hábitat, la biología floral y números cromosómicos.

Palabras clave. Caiophora; Chile; Flora Alto Andina; Loasaceae; Morfología; Número cromosómico; Polinización; Taxonomía.

INTRODUCTION

   The genus Caiophora C. Presl comprises about 60 species. It ranges from Central Argentina/Chile to central Ecuador with a single species in Uruguay and southern Brazil [C. arechavaletae (Urb.) Urb. & Gilg]. The genus is largely High Andean, and most taxa are restricted to elevations above 3000 m, ranging as high as 5000 m. The genus Caiophora forms a monophyletic group with clades of Loasa Adans. and Scyphanthus D. Don (Hufford et al., 2005; Weigend et al., 2004). Sleumer (1955) and Weigend (1997, 2000) made the first and preliminary attempts to organize and evaluate the numerous names published in the genus Caiophora since the studies by Urban & Gilg (1900, 1911), while some additional names were clarified in Weigend & Ackermann (2003). Many of the taxonomical problems in the genus stem from the fact that Urban & Gilg (1900) enthusiastically described new species on the basis of single, fragmentary specimens. However, a fullscale revision of the genus is still pending, and classification in the genus remains extremely problematical, due to the relative scarcity of clear morphological characters identifiable in herbarium specimens, the high degree of heterogeneity between populations of individual taxa as now recognized, and the abundance of interspecific hybrids (Sleumer, 1955). These complications are particularly true for Peru, where Caiophora is most diverse, and to a lesser extent for Bolivia. For Peru only one revision has been published (Macbride, 1941), which has been evaluated for the Checklist of that country (Schatz, 1996). These studies are based on a very superficial review of herbarium material only. Especially in Peru some species groups such as the C. cirsiifolia-group and C. carduifolia-group (Weigend & Ackermann, 2003) comprise numerous undescribed, often locally endemic taxa. In Chile the situation is less complex, with only a handful of relatively well defined species known. However, Caiophora is largely restricted to the North of Chile in the Region I, an area not well represented in herbaria due to the seasonality of its flora and the inaccessibility of the High Andean vegetation. Furthermore, the transport of organic material into the neighbouring Region is severely restricted for phytosanitary reasons, which provides a further inhibition for the collection of botanical specimens. Distribution limits are therefore not entirely clear. Another complication arises from the very incomplete locality data and absence of reliable numbering on the type collections by Philippi and Steinmann. Especially Philippi collected and described some specimens with nearly the same or without precise locality information (C. superba Phil. and C. rahmeri Phil. from Tarapacá) leading to confusion in both the literature and herbaria, as well as making the unequivocal identification of iso- and holotypes next to impossible. These problems are here addressed as best as possible.
   In the present study, evaluation of species limits, i.e., morphological divergence typically present within relatively homogeneous species (and determined by ontogeny or ecology) versus morphological divergence sufficient to warrant the recognition of different taxa, is mostly derived from more than 15 field trips, mainly to Peru. Also, ca. 20 accessions of Caiophora taxa were taken into cultivation in Munich or Berlin to compare the variability of morphological characters from natural habitat with different ecological conditions, and an extensive amount of herbarium material was studied.
   This study aims to provide well-defined names for the project of the "Checklist of the Southern Cone" (Weigend et al., forthcoming) and we here redefine the relevant taxa present there, reduce several names to synonymy and improve descriptions and illustrations. The following taxa are here reported for the flora of Chile: Caiophora chuquitensis (Meyen) Urb. & Gilg, C. cirsiifolia C. Presl, C. coronata (Gillies ex. Arn.) Hooker & Arn., C. deserticola Weigend & Mark. Ackermann, sp. nov. and C. rosulata (Wedd.) Urb. & Gilg subsp. rosulata comb. nov. and C. rosulata (Wedd.) Urb. & Gilg subsp. taraxacoides (Killip) Weigend & Mark. Ackermann stat. and comb. nov.

MATERIAL AND METHODS

   The present study is based on field studies in southern Peru and from cultivation of several accessions at Munich and Berlin (vouchers at BSB, M, MSB). Specimens of the following herbaria were revised: B, BA, BM, BOLV, BR, BSB, CORD, CUZ, E, F, FR, G, GB, GOET, HBG, GH, HUSA, HUT, IBBA, K, L, LIL, LPB, LPZ, M, MA, MICH, MO, MSB, NY, OXF, P, PR, PRC, S, SI, SGO, TRIER, TUEB, UC, UMSS, U, US, USM, W, WU, Z, plus the online type collections of the Field Museum of Natural History (F, fm1.fieldmuseum.org/vrrc/index.php), New York Botanical Gardens (NY, www.nybg.org/bsci/herb/), Smithsonian Museum of Natural History (US, rave nel.si.edu/botany/types/) and the Herbarium of the University of Vienna (WU, herbarium.univie.ac.at/database/collections.htm). For identification and description of the different species we used the morphological characters of growth habit, leafsize, flower-size, petal-size, nectar scale and staminode size and shape, and also the density of setae and trichomes (scabrid and glochidiate).
   Chromosome counts were made from embryonic roots of germinating seeds (Petri culture dishes, on moist filter paper) or obtained from potted seedlings that were pre-treated in hydroxyquinoline for 1.5 hours, fixed in 95% ethanol-acetic acid (3:1 V/V), stained with aceto-orcein, and then squashed and counted. Additional counts were compiled from the literature.

Key to the Chilean species of Caiophora:

1. Acaulescent, rosulate plant. Capsule up to 16 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . .5. C. rosulata
1. Plants with well developed aerial shoot, shoots erect, decumbent or winding. Capsule over 20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2.
2(1). Plants decumbent but leaves stiffly erect. Flowers on long, lax pedicels, geoflorous. Petals mostly white, cream, greenish-white (very rarely yellow or orange) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3. C. coronata
2. Plants stiffly erect or winding. Flowers deflexed or pendulous. Petals orange, red or pale pink . . . . . . . .3.
3(2) Plants winding. Petals and nectar scales orange. Nectar scales keeled and never with three dorsal filaments (rarely one or two, and then inserted at scale apex) . . . . . . . . . . . . . . . . . . . . . . . . . .2. C. cirsiifolia
3. Plants erect. Petals orange, red or pink. Nectar scales white, usually not keeled and with three dorsal filaments inserted on middle or upper half of scale . . . . . . . . .4.
4(3) Flowers penta- to heptamerous. Pedicel 20-30 (-70) mm (in anthetic and postanthetic flowers). Petals 15-25 mm long, pink. Abaxial leaf-surface nearly esetulose. Fruit conical . . . . . . . . . . . . . . . . . . .4. C. deserticola
4. Flowers penta- to nonamerous. Pedicel 3-15 (-50) mm (in anthetic and postanthetic flowers). Petals 20-30 (-40) mm long, pink, bright orange or red, rarely white or yellow. Abaxial leaf-surface setose, fruit ovoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1. C. chuquitensis

1. Caiophora chuquitensis (Meyen) Urb. & Gilg, Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 76: 301. 1900. Loasa chuquitensis Meyen, Reise Erde 1: 483. 1834. Blumenbachia chuquitensis (Meyen) Hook.f., Bot. Mag. 51: Tab. 6143. 1875. TYPE: Bolivia, Depto. La Paz, Prov. La Paz, Vic La Paz, 10000 ft, M. Bang 171 p.p. (neotype NY! designated by M. Weigend, Sendtnera 4: 232. 1997; isoneotypes BM!, E!, F!, GH!, MO, US!, W!). Figs. 1, 2.


Fig. 1. Caiophora chuquitensis. A, growth habit. B, E, nectar scale, dorsal view. C, G, apical view. D, lateral view. F, ventral view. H, petal, dorsal view. I, K, apical view. J, lateral view. L, flower (drawn by H. Lünser, Berlin). M, lateral view.. N-O, staminodes. P, sepal. Q, fruit. A and M, drawn from cultivated plants in Munich M. Weigend 3681 (MSB); B-D, K, L, O, P, from M. Ackermann et al. 274 (BSB, HUSA, M); E-J, N, Q, from F. Luebert 1720 (BSB, SGO).


Fig. 2. Caiophora chuquitensis. A, natural habitat in Chile (Photo from F. Luebert) B, natural habitat in Peru. C-F, two different morphotypes, cultivated in Berlin. C, D, flower, lateral. E, F, nectar scales and staminodes. A, C, E, F. Luebert 1720 (BSB, SGO): B, M. Weigend & K. Weigend 2000/203 (HUSA, MSB, NY); D, F, M. Ackermann et al. 274 (BSB, HUSA, M).

   Caiophora superba Phil., Anal. Mus. Nac. Chile 1891: 23. 1891, syn. nov. TYPE: Chile, I Región de Tarapacá, Tarapacá, R. A. Philippi s.n. (holotype SGO!; isotypes K!; WU!).
   Caiophora horrida Urb. & Gilg, Mem. Torrey Bot. Club 3/3: 36. 1893. TYPE: Bolivia, near La Paz, M. Bang 171 p.p. (lectotype NY! designated by M. Weigend, Sendtnera 4: 232. 1997; isolectotypes BM!, E!, F!, GH!, MO, US!, W!).
   Caiophora albiflora (Griseb.) Urb. & Gilg, in Engler and Prantl, Nat. Pflanzenfam. 3/6a: 119. 1894. Caiophora heptamera Wedd. var. albiflora Griseb., Symb. Fl. Argent. 139. 1879. TYPE: Argentina, Prov. Catamarca, Andalgalá, near Negrilla, F. Schickendantz 149 (lectotype GOET! designated by M. Weigend, Sendtnera 4: 232. 1997; isolectotypes B destroyed, photo F, neg. nr. 10140, CORD).
   Caiophora heptamera (Wedd.) Urb. & Gilg, in Engler and Prantl, Nat. Pflanzenfam. 6a: 119. 1894. Loasa heptamera Wedd., Chlor. And. 2: 218. 1857. TYPE: Bolivia, Depto. Potosi, H. A. Weddell 4095 (holotype P!, photo F!, neg. nr. 38479).
   Caiophora angustisecta Urb. & Gilg, Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 76: 300. 1900. TYPE: Argentina. Prov. Salta: Cafayate, Cuesta del Arca, 3090 m, C. Spegazzini 102321 (holotype B destroyed, photo F!, neg. nr. 10142; isotype LPS).
   Caiophora lorentziana Urb. & Gilg, Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 76: 289. 1900. TYPE: Argentina, Prov. Salta, Caldera, near Nevado del Castillo, P. G. Lorentz & G. H. Hieronymus s. n. (lectotype WU! designated by M. Weigend Sendtnera 4: 234. 1997).
   Caiophora macrocarpa Urb. & Gilg, Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 76: 285. 1900, syn. nov. TYPE: Argentina, Prov. Salta, Caldera, near Nevado del Castillo, P. G. Lorentz and G. H. Hieronymus 49 (lectotype K! designated by M. Weigend, Sendtnera 4: 234.1997; isolectotypes B destroyed, photo F!, neg. nr.10156; G, GOET!).
   Caiophora orbignyana Urb. & Gilg, Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 76: 302 1900. TYPE: Bolivia, Potosí, A. D'Orbigny 1436 (holotype BR!; isotypes G!, photo F!, neg. nr 24169, P!).
   Caiophora sphaerocarpa Urb. & Gilg, Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 76: 296. 1900. TYPE: Bolivia, La Paz, Larecaja, near Sorata, Arrilaya, Chuchu, 3800-4200 m, G. Mandon 619 p.p. (holotype P!, photo F!, neg. nr. 38498).
   Caiophora fiebrigii Urb. & Gilg, Bot. Jahrb. Syst. 45: 470. 1911. TYPE: Bolivia, Tarija, Prov. Avilez, Puna Patanca, 3800 m, K. Fiebrig 2603 (lectotype BM! designated by M. Weigend, Sendtnera 4: 234. 1997; isolectotypes B destroyed, photo F!, neg. nr. 10151; E!, G, HBG!, K!, L, M!, P!, U, US!, W!).

   Perennial herbs to subshrubs (15-) 40-80 (-100) cm tall, with persistent basal leaf rosette from thick tap-root. Stems basally (for ca. 10-20 cm) lignescent and perennating, 4-20 mm thick, densely covered with setae 3-5 mm long, white, yellow or brown, and scabrid (0.2-0.5 mm long) and glochidiate trichomes (0.1-0.4 mm long). Basal leaves 10-15 (-25) cm long; laminas narrowly ovate, 70 x 30 to 150 x 45 (to 200 x 60) mm, pinnatisect (lower lobes generally free) with (6-) 9-12 (-14) lobes on each side; lobes narrowly ovate, up to 30 x 20 mm; lobe margins grossly serrate to pinnatifid to lobulate, with 5-8 serrations or lobules on each side; lobules up to 8 mm long, distal lobules usually recurved; adaxial leaf surface sparsely setose with stinging hairs 3-5 mm long and covered with scabrid trichomes up to 1 mm long; abaxial surface setose, very densely set with scabrid (up to 1 mm long) and glochidiate trichomes (up to 0.3 mm long). Inflorescences frondose, terminal, mono- or asymmetrical dichasia, rarely thyrsoids; with 3-7 flowers, with internodes 4-9 cm long; pedicels 3-15 (-50) mm long during anthesis. Flowers horizontal to deflexed, penta- to nonamerous. Calyx lobes spreading or deflexed, narrowly ovate-oblong to ovate-triangular, 9 x 2 to 11 x 3 mm, densely setose and covered with scabrid trichomes; margin coarsely serrate or dentate with 3-5 teeth on each side. Corolla widely balloon-shaped; petals erect, deeply cymbiform and sharply keeled, 20-30 (-40) mm long and 10 mm deep, dorsally setose and covered with scabrid and glochidiate trichomes; petals bright orange, red, pinkish, rarely yellow or white; apex blunt or acuminate. Stamens in 5-9 epipetalous fascicles, ca. 25 in number per fascicle each; filaments ca. 12-17 mm long, white; anthers ovoidal, pale yellow to orange, ca. 1 mm long. Nectar scales deeply cymbiform, white, hemispherical or keeled in dorsal view, ca. 5 x 6 to 7 x 8 mm; scale neck triangular or rounded, slightly thickened; with (rarely without) three filiform dorsal filaments up to 5 mm long, these sometimes basally widened, white with orange or red tip, inserted in the middle or in the upper third of scale back. Free staminodia Lshaped, 10-20 mm long, dorsally with spoonshaped appendage ca. 5 x 2 mm, appendage sometimes papillose; apex of staminodia filiform and hook-shaped. Style terete, up to 25 mm long (towards the end of anthesis); ovary inferior, conical, with 3-5 placentae and numerous ovules. Fruit horizontal or deflexed; pedicels up to 50 mm long; capsule ovoidal, 20 x 10 to 30 x 25 mm, sometimes protracted into a short beak, opening with 3- 5 longitudinal slits only; style persistent, up to 15 mm long (during anthesis), not accrescent in fruit; sepals accrescent, up to 20 mm long. Seeds numerous; testa deeply pitted, brown.

Geographical distribution and habitat. The species is distributed in Chile in Region II. (Marticorena et al., 1998), in Argentina in the provinces of Catamarca, Jujuy, Salta and Tucumán (Brücher, 1986, 1989; Sleumer, 1955), in Bolivia in the departments of Cochabamba, La Paz, Oruro, Potosi and Tarija (Weigend & Ackermann, forthcoming) and in Cuzco, Peru (Schatz, 1996). The elevational distribution ranges from (2500-) 3000-4500 m. This High Andean plant species is found on scree slopes, in corrals and at the base of dry stone walls, between rocks and in open grassland (Fig. 2A-B). Caiophora chuquitensis is pollinated by long-tongued bees and hummingbirds (Ackermann & Weigend, 2006; Harter, 1995; Coccuci & Sersic, 1998; Schlindwein, 2000).

Chromosome number. Sporophytic: 2n = 16 (incl. C. macrocarpa, Brücher 1986, 1989; own counts: M. Ackermann et al. 274).

Observations. Caiophora chuquitensis is a very heterogenous species with a wide range of morphological variations (Fig. 2A-F). Sizes of plants, laminas and flowers vary considerably. Leaf margins are typically recurved in nature, but may be flat in moist situations and in cultivation. Petal colour is typically bright orange and also red or pinkish, but specimens with yellow or white petals have been recorded. Nectar scales are either hemispherical or keeled, sometimes lacking dorsal filaments (Fig. 2E-F). Cultivated plants (specimen: M. Ackermann et al. 274) in the greenhouses in Berlin show that many of the characters used for species delimitation by Urban & Gilg (1900) (plant size, presence or absence of dorsal filaments on nectar scales, degree of leaf dissection) vary between the wild collected plant and its offspring in cultivation and are thus not stable. Merosity, flower-size and fruit-size, which Sleumer (1955) and Urban and Gilg (1900) used to distinguish C. heptamera and C. macrocarpa are not fixed. Cultivated plants show that the first flower and fruit of the inflorescence is huge in comparison to the final ones and that merosity can decrease with the subsequent flowers. Density and occurrence of the different trichomes (scabrid and glochidiate) depend on seasonal and ecological factors. Leaves from plants cultivated in shade have few setae and flat laminas, whereas plants cultivated in sunshine have margins that are usually recurved. These observations lead us to synonymize C. superba and C. macrocarpa under C. chuquitensis. Caiophora andina Urb. & Gilg (Peru and Argentina) remains problematical and is only distinguished by its few-flowered inflorescences (one or two flowers) and the less deeply dissected leaves. Closely allied C. mollis (Griseb.) Urb. & Gilg (Bolivia and Argentina) can be distinguished by the lack of or scarcity of setae on the entire plant and the presence of long, white scabrid trichomes. Caiophora rusbyana Urb. & Gilg and C. boliviana Urb. & Gilg are characterized by capsules with a distinct conical beak.

Representative specimens examined

   ARGENTINA. Catamarca. Depto. Andalgalá: Mina de Capillitas, subida el Cerro Yutuyaco, 3600-3900 m, 3-III-1952, H. O. Sleumer 2739 (UC, US). Jujuy. Depto. Cochinoca: Cerro Jucahuasi, 4000 m, 5-III-1930, S. Venturi 10371 (S). Depto. Humahuaca: Palca de Aparzo, 3700 m, 23o 10' S, 065o 11' W, 16-II-1997, F. O. Zuloaga et al. 5948 (MO). Depto. Rinconada: Mina Pirquitas, 4200 m, III-1970, H. Fabris and F. O. Zuloaga 7693 (P); Depto. Tilcara: Chorru Valley, near Tilcara, 4000 m, 13-II-1939, E. K. Balls 6029 (E, K, UC, US). Depto. Yavi: Quebrada de Toquero, 3600 m, 21-XI-1963, A. L. Cabrera 15370 (M). Salta. Depto. Rosario de Lerma, 3200 m, S. Venturi 8131 (BM, GH, K, NY, US). Depto. Caldera: Potrero del Castillo, ascent to Nevado del Castillo via Mal Paso, 3700-3750 m, 15-III-1952, H. O. Sleumer and F. B. Vervoorst 2953 (US). Depto. Chicoana: Cuesta del Obispo, 2800 m, A. L. Cabrera et al. 22021 (K). Depto. Orán: Cerro La Escalera, 3800 m, A. Pierotti 1337 (GH). Depto. Poma: Cobres, 3500 m, 31-I-1944, A. L. Cabrera 8333 (GH). Depto. San Antonio de los Pobres: Quebrada de Urcuro, 3700 m, 12-II-1945, A. L. Cabrera 8687 (GH). Depto. Santa Victoria: near the village Santa Victoria, 2500-2800 m, 13-16- XII-1988, J. L. Novara 8355 (M, S). Tucumán. Depto. Chicligasta: Estancia Las Pavas, 3000 m, 28-XI-1926, S. Venturi 4655 (GH, US). Depto. Tafi: Cumbre de Chaquiril, 12-I-1945, D. Olea 256 (BM, S).

   BOLIVIA. Cochabamba. Prov. Quillacolllo: laderas de la cordillera sobre Tiquipaya y la comunidad de Laphia, 3750 m, 11-II-1990, G. Navarro 1103 (BOLV). Prov. Tiraque: Kaspiconcha alto, Millumayo, 3950 m, 7-XII-1989, R. Guillen 34 (BOLV). La Paz. Prov. Aroma: La Paz 75 km hacia al Sur y 10 km hacia al desvio a Sapahaqui, 17° S, 068° W, 4150 m, 18-I-1981, S. Beck 6021 (MO, MSB). Prov. Bautista Saavedra: Charazani, Chajaya, 7-IV-1992, P. Gutte 339 (LPB, LPZ). Prov. Inquisivi: 35 km de Caracollo-Leque Palca, 4 km hacia Cochabamba, sobre el camino nuevo asfaltado Tholopampa, 17° 35' S, 066° 57' W, 3950 m, 15-I-1995, S. Beck 21725 (M). Prov. Los Andes: 6,6 km NW of Batallas on the principal road along Lake Titicaca, 16° 15' S, 068° 33' W, 3850 m, 5-II-1984, J. C. Solomon 11442 (MO, US). Prov. Loayza: 9.8 km NW of Villa Loza on road towards Urmiri and Sapahaqui, 5-III-1993, P. M. Petersen et al. 12679 (LPB). Prov. Murillo: al NW de La Paz, entre Lago Challapata y Lago Incachaca, 4300 m, 28-XII-1990, S. Beck 17894 (MSB). Prov. Omasuyos: near Sircapaca, 3880 m, III-1982, F. Casa & J. Molero 6464 (NY). Prov. Pacajes: Corocoro, 1 km east of town, 4050 m, 24- XI-1982, T. Johns 82-58 (F, LPB, MICH, MO). Oruro. Prov. Sajama: de Turco 3 km hacia Curahurare de Carangas, 3880 m, 18-III-1992, S. Beck 21050 (MSB). Potosi. Prov. Nor Chichas: Quechisla, XII-1931, R. Cárdenas 49 (GH). Prov. Sud Chichas: 1 km before Macho Cruz, the pass across the Cordillera de Mochara from Tupiza to Tarija, 3700 m, 4-XII-1967, B. B. Vuilleumier 405 (F, NY). Prov. Tomas Frias: Cerrania del Khare- Khare, arriba de la Ciudad de Potosi, a orrillas de la Laguna Chalaviri, 4400 m, 4-II-1988, Schulte 162a (M). Tarija. Prov. Aviles: Tajzara cerca Patancas, 3650 m, 11-III-1986, E. Bastión 1063 (LPB, MSB). Prov. Mendez: Abra entre Iscayachi y Cieneguillas, 3500 m, 27-XII-1985, R. Ehrich 28 (LPB, MSB).

   CHILE. II Región de Antofagasta. Prov. El Loa: Quebrada de Caspana, 5 km sur de Caspana, 3250 m, 18-II-2003, F. Luebert 1720 (BSB).

   PERU. Cuzco. Prov. Calca: road from Calca to Lares, after Rancal, 13° 10' 26'' S, 071° 57' 55'' W, 4000 m, 11-IX-2002, M. Ackermann et al. 274 (BSB, HUSA, M). Huancavelica. Prov. Castrovirrayna: near Apacheta Grande, 4500 m, 28-XII- 1974, T. C. Plowman & W. Davis 4646 (USM).

2. Caiophora cirsiifolia C. Presl, Reliq. Haenk. 2: 42, plate 56. 1831. TYPE: Peru. Depto. Junin, Tarma?, T. Haenke s.n. (holotype PR!, photo PR!, neg. nr. 919; isotype PR!, photo PR!, neg. nr. 920). Figs. 3, 4A-B.


Fig. 3. Caiophora cirsiifolia. A-C, leaves. D, sepal. E, nectar scale, lateral view. F, ventral view. G, dorsal view. H, staminode. I, petal, ventral view. J, lateral view. K, fruit. A, C-H, K, M. Ackermann et al. 420 (BSB, HUSA, M, USM, NY, F); B, I, J, M. Ackermann et al. 555 (BSB).


Fig. 4. Caiophora cirsiifolia. A, growth habit. B, flower. Caiophora coronata. C, growth habit. D, flowers (pictures from www.opuntiadelsur.de). Caiophora rosulata subsp. rosulata. E, habitat. F, growth habit. A-B, M. Ackermann et al. 420 (BSB, HUSA, M, USM, NY, F); E-F, M. Weigend & Ch. Schwarzer 7837 (BSB, HUSA, HUT, USM).

   Caiophora sepiaria (Ruiz & Pav. ex G. Don) J. F. Macbr., Candollea 8: 23. 1940. Blumenbachia sepiaria Ruiz & Pav. ex G. Don, Gen. Hist. 3: 62. 1834. Loasa sepiaria Ruiz & Pav., Fl. peruv, in Anales Inst. Bot. Cavanilles 16: 420, tab. 449. 1958. TYPE: Ruiz & Pav., Fl. peruv, in Anales Inst. Bot. Cavanilles 16: tab. 449. 1958. (Lectotype designated by M. Weigend, Sendtnera 4: 227. 1997). Peru, Depto. Lima, Huacho, cerca Juncal, Mayobamba, H. Ruiz & J. A. Pavón s.n. (epitype MA! designated by M. Weigend, Sendtnera 4: 227. 1997, photo M!, fragment F!).
   Caiophora preslii Urb. & Gilg, Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 1900: 306. TYPE: Peru. Depto. Lima, Matucana, cerca Matucana, "In vallibus cordillerum Peruvia", T. Haenke s.n. (holotype PR! nr. 24293).
   Caiophora contorta auct. non (Descr.) Presl.

   Perennial, winding herbs up to 2-5 m, without basal leaf rosette, with taproot and short underground rhizomes. Stems basally 2-4 mm thick, sparsely covered with stinging hairs (2-3 mm long), trichomes scabrid (up to 1 mm long) and glochidiate (0.3 mm long). Basal leaves with petioles 15-25 mm long; lamina (narrowly) triangularovate, 60 x 35 to 90 x 40 mm, apex acuminate, with proximal pair of leaflets sometimes free (basal leaves), pinnate-pinnatifid to bipinnatifid with (4-) 6-7 (-9) lobes on each side; lobes narrowly ovate, triangular, 7 x 4 to 15 x 7 (to 24 x 15) mm; margins grossly serrate to pinnatifid with 2-3 lobules/teeth on each side; lobules up to 4 mm long; adaxial leaf surface sparsely setose with stinging hairs up to 3 mm long and set with scabrid trichomes up to 1 mm long; abaxial leaf surface esetulose or with scattered stinging hairs mainly on major veins, up to 2 mm long, very densely covered with scabrid trichomes (up to 0.6 mm long) and glochidiate trichomes (ca. 0.3 mm long). Inflorescences frondose, winding anthoclades, terminal monochasia or very asymmetrical dichasia; internodes 5-25 cm, pedicels 20-30 (-60) mm long during anthesis. Flowers pendulous, pentamerous. Calyx lobes spreading, apically reflexed, narrowly triangular-ovate, 6 x 1 to 10 x 2 mm, sparsely setose and densely covered with scabrid trichomes, with margins coarsely serrate with 1-3 teeth on each side. Corolla saucer-shaped; petals deeply cymbiform, 13-20 mm long and 6-8 mm deep, setose and covered with scabrid and glochidiate trichomes, orange, petals laterally winged towards the base. Stamens in 5 epipetalous fascicles, 20-25 in number per fascicle; filaments ca. 10-12 mm long; anthers ovoid, pale yellow to brownish, ca. 1 mm long. Nectar scales deeply cymbiform, ca. 5 mm high, 6 mm wide and 6 mm deep, same colour as petals (orange), keeled in dorsal view; generally without dorsal filaments, scale neck thickened. Free staminodia L-shaped, 8-10 mm long, dorsally with a spoon-shaped, papillose (up to 1 mm) appendage ca. 3 x 1.5 to 4 x 2 mm. Style terete, up to 7 mm long (towards the end of anthesis); ovary inferior, conical to cylindrical, with 3 placentae with numerous ovules. Fruit deflexed, pedicel 30-40 (-70) mm long; capsule conical, (15-) 20-35 mm x 8-11 mm, twisted, opening with 3 longitudinal slits; style persistent, not accrescent in fruit; sepals accrescent, up to 10 mm long. Seeds numerous; testa deeply pitted, brown.

Geographical distribution and habitat. The species is part of a complex of several closely allied taxa mostly on the western slope of the Andes ranging from North Peru (Depto. Cajamarca) into North Chile (Region I). The plant from ing it at infraspecific level. The collections from the southern part of the range generally have smaller flowers and more deeply dissected leaves than those from Central Peru, but these differences largely vanish in cultivation, indicating that they simply reflect the generally drier conditions for plants growing in South Peru and Chile. Caiophora cirsiifolia ranges from elevations of 2400 to 3700 m and is usually found in dry scrub forest, hedges (Fig. 4A), road banks and dry stone walls. The flowers of Caiophora cirsiifolia are largely visited by long-tongued bees of the genera Bombus and Centris (Ackermann and Weigend, 2006).

Chromosome number. Chromosome counts of the southern C. cirsiifolia form are not available, but northwards in the adjacent departments our counts result in 2n =16 [Depto. Apurimac: M. & K. Weigend 2000/392 (BSB, HUSA, M, USM), Depto. Ayacucho: M. & K. Weigend 2000/341 (BSB, HUSA, M, USM)].

Observations. Caiophora cirsiifolia is a poorly documented species for Chile. There are only a few collections from Region I, of which we have seen a single specimen. We do not know whether the species is truly rare, or only undercollected, since Region I of Chile is particularly poorly documented. It certainly is a very common species e.g. in Depto. Arequipa, Moquegua and Tacna in Peru. It is the only winding species of Caiophora on the western side of the Andes south of the city of Arequipa.

Representative specimens examined

    CHILE. I Región de Tarapacá. Prov. de Tarapacá: Belén, am Friedhof, ca. 3500 m, J. Grau s.n. (M).

    PERU. Arequipa. Prov. Arequipa: Environment of Chiguata, east from Arequipa, 16° 24' 20'' S, 071° 22' 38'' W, 3100 m, 1-X-2002, M. Ackermann et al. 420 (BSB, F, HUSA, M, NY, USM). Prov. Caylloma: Sibayo, 3500 m, IV-2002, F. Caceres 2494 (BSB, HUSA). Prov. La Union: Dist. Puyca, arriba de Puyca, S 15°34.00' W 72° 41.35', 3562 m, 18-IX-1999, V. Quipuscoa S. et al. 1591 (HUSA). Moquegua. Prov. Comás: Road from Arequipa to Puquina (Moquegua), between Arequipa and Pocsi, 16° 35' 23'' S, 071° 25' 52'' W, 3300-3400 m, 29-IV-2000, M. Weigend et al. 2000/557 (HUSA, NY). Prov. General Sanchez Cerro: Omate, village Challoguaya above Omate, S16°38'42,5'' W070°57'42,2'', 2520m. 09-XII- 2006, M. Ackermann & F. Caceres 680 (BSB, HUSA, USM). Prov. Mariscal Nieto: Puquina, 3400 m, 21-IV-1967, C. Vargas 19363 (BSB, CUZ). Puno. Prov. Moho: Dist. Moho, centro poblado de Huaraya, 3800-3900 m, 18-III-1997, G. Arenas T. s/n (HUSA). Tacna. Prov. Candarave: Volcan Yucamani, 3100-3400 m, 09-XII-1997, M. I. La Torre 1956 (USM). Prov. Tarata: 16 km above Candarave on Mazo Cruz road (196 km west of Llave) 3680 m, 9-X-1997, M. Weigend & H. Förther 97/797 (F, HUT, MO, MSB, NY, USM).

3. Caiophora coronata (Gillies ex. Arn.) Hook. & Arn., Bot. Misc. 3: 327. 1833. Loasa coronata Gillies ex Arn., Edinburgh J. Nat. Geogr. Sci. 3: 274. 1831. TYPE: Argentina, Mendoza, Andes of Mendoza, above Puente del Inca, Aguas del Cerro Pelado, J. Gillies s.n., "anno 1821" (lectotype E! designated by M. Weigend, Sendtnera 4: 235. 1997; isolectotypes BM!, GH!, K!, OXF!). Figs. 4C-D, 5.


Fig. 5. Caiophora coronata. A-D, leaves. E-F, sepals. G, nectar scale, lateral view. H, ventral view. I, staminode. J, petal, ventral view. K, dorsal view. L, lateral view. M, fruit. A, Meyen 4659 (NY, UC); B, D, L, J. R. I. Wood 14626 (LPB); G-K, M, T. H. Goodspeed & Y. Mexia 4611 (GH, UC); C, E, M. Ackermann 60 (BSB); F, E. Budin 7437 (UC); (drawn by C. Becker, Berlin).

   Caiophora pycnophylla Urb. & Gilg, Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 76: 274. TYPE: Argentina, La Rioja, Famatina, Cueva de Pérez, 3700 m, G. H. Hieronymus & G. Niederlein 388 (holotype B destroyed, photo F!, neg. nr. 10164; isotypes CORD, K!).
   Caiophora absinthiifolia C. Presl, Reliq. Haenk.: 43, plate 57. 1831. TYPE: Chile, T. Haenke s.n. (holotype PRC, not located).

   Perennial cushion-forming herb with spreading, decumbent stems and stiffly erect leaves, ca. 15-30 cm tall, 20-50 cm in diameter, with persistent basal leaf rosette. Stems rarely basally lignescent 3-5 mm thick, sparsely to densely covered with stinging hairs 3-4 mm long, scabrid trichomes (ca. 0.5 mm long) and glochidiate trichomes (0.1-0.3 mm long). Leaves 6-18 (-22) cm long; lamina oblong/ovate, 50 x 15 to 90 x 40 mm, pinnate-pinnatifid to bipinnatifid with 5-9 lobes on each side, proximal pair of leaflets often free; leaf lobes up to 22 x 15 mm; lobe margins reflexed, grossly serrate to pinnatifid with 2-4 (-6) lobules/teeth on each side, triangular to linear; adaxial leaf surface densely setose with stinging hairs 3-4 mm long, sparsely covered with scabrid trichomes up to 0.4 mm long; abaxial surface densely covered with glochidiate trichomes ca. 0.2 mm long; major veins both setose from stinging hairs 3-4 mm long and scattered scabrid trichomes ca. 0.3 mm long. Inflorescences frondose, terminal monochasia up to 10 cm long and with 2-5 flowers; internodes 1- 3 cm, pedicels 2-12 cm long during anthesis. Flowers geoflorous, generally lying on the ground, oriented horizontally, pentamerous. Calyx lobes reflexed, narrowly triangular-linear, 8 x 1 to 15 x 2 mm, sparsely to densely setose and covered with scabrid trichomes; margin serrate to pinnatifid with (1-) 2-3 teeth/lobes on each side. Corolla bowl-shaped; petals deeply cymbiform, 20-30 mm long and 10-15 mm deep, dorsally setose and covered with scabrid and glochidiate trichomes, white, rarely cream, pale yellow, greenish or orange. Stamens in 5 epipetalous fascicles, 20-25 in number per fascicle; filaments ca. 15 mm long: anthers ovoid, pale yellow or brown, ca. 1.5 mm long. Nectar scales cymbiform to rectangular, white, ca. 4-5 x 5-8 mm, usually without dorsal filaments, sometimes with two or three white, filiform filaments up to 4 mm long, rising from the upper half of scale back. Free staminodia inflexed, 6-10 mm long, without appendage. Style terete, up to 12 mm long (towards the end of anthesis); ovary inferior, conical, with 3 placentae and numerous ovules. Fruit deflexed; pedicel 50-120 mm long; capsule globose to conical, slightly twisted, 20 (- 40) x 18 (-25) mm, opening with 3-4 longitudinal slits; style persistent, not accrescent in fruit, sepals accrescent, up to 15 mm long. Seeds numerous; testa deeply pitted, brown.

Geographical distribution and ecology. The species is distributed in Chile from Santiago up to the Regions I and II (Arroyo et al., 1982; Marticorena et al., 1998, 2001), in Argentina from Mendoza up to Jujuy (Brücher, 1986, 1989; Sleumer, 1955) and in Bolivia in the southern departments Potosi and Tarija (Weigend, 1997). The elevational distribution ranges from (2000-) 2500-4500 (- 5000) m, with elevation increasing from the South to the North. Caiophora coronata is found in at least seasonally dry habitats. It grows in corrals, at the base of rocks or in open habitats on scree slopes. (Fig. 4C). Caiophora coronata is pollinated by rodents, but is also visited by hymenoptera, hummingbirds and passerines (Coccuci and Sersic, 1998).

Chromosome number. Sporophytic 2n = 16 (Brücher 1986, 1989;