(Cactaceae) in Chile A New Taxonomic Treatment of the ...

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A New Taxonomic Treatment of the Genus Trichocereus(Cactaceae) in ChileAuthor(s): Sofía AlbesianoSource: Haseltonia, 18():116-139. 2012.Published By: Cactus and Succulent Society of AmericaDOI: http://dx.doi.org/10.2985/026.018.0114URL: http://www.bioone.org/doi/full/10.2985/026.018.0114

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Haseltonia 18: 116–139. 2012 116

INTRODUCTION

The genus Trichocereus belongs to the subtribe Trichocereinae Buxbaum, which in turn belongs to the tribe Trichocereeae Buxbaum, subfamily Cactoi-deae. Columnar or sometimes globular stems char-acterize this subtribe. Its flowers are zygomorphic (Schick 2011), ranging in shape from campanulate to infundibuliform, with large tepals, usually white or whitish, or brightly colored in some taxa; they may or may not have nectaries, but the stamen inser-tion begins either at the base of the receptacle or on the nectaries (Buxbaum 1958).

Trichocereus is morphologically defined by cylin-

drical stems with shallow ribs, large flowers (8–30 cm), wide ovaries (2.0–2.5 cm) and floral tube cov-ered with subulate scales, presenting an abundance of black, brown, gray or white hair in their axils (Kiesling 1978, Kiesling & Ferrari 2005). Riccobono (1909) recognized Trichocereus as a genus based on the columnar stems and flower pilosity, and the ety-mology of the name Trichocereus refers to those two characteristics. This genus comprises about 45 spe-cies in the Andes of Ecuador, Peru, Bolivia, Argen-tina and Chile, occurring in the latter country from the regions of Arica and Parinacota to Maule.

Trichocereus is a monophyletic genus, if Harrisia hahniana and Harrisia earlei are included in it (Al-

Abstract: The genus Trichocereus is represented by 19 taxa (12 species, 5 subspecies and 2 varieties) in Chile, 13 of them endemic to this country. These occur from the regions of Arica and Parinacota on the border with Peru, to the region of Maule, the IV Region of Coquimbo presenting the highest taxonomic richness. Its species are a dominant feature of the landscape and define phytogeographic regions. This tax-onomic study has yielded iconographies, original descriptions, detailed descriptions, taxonomic comments, information on distribution and habitat, exsiccata, nomenclature, and three keys for the identification of spe-cies, subspecies and varieties. This treatment provides an updated synonymy including eight new synonyms; a new combination, Eulychnia coquimbana (Molina) S. Albesiano; three taxonomic level changes: (1) Tricho-cereus chiloensis subsp. australis (F. Ritter) S. Albesiano, (2) Trichocereus chiloensis subsp. eburneus (Phil. ex K. Schum.) S. Albesiano, and (3) Trichocereus chiloensis subsp. panhoplites (K. Schum.) S. Albesiano; three new names: (1) Trichocereus faundezii S. Albesiano, (2) Trichocereus pectiniferus S. Albesiano, (3) Trichocereus undu-losus S. Albesiano; and thirteen neotypifications.

Keywords: Cactaceae, Chile, distribution, endemism, taxonomy, Trichocereus.

Resumen: El género Trichocereus en Chile se encuentra representado por 19 taxones (12 especies, 5 sub-especies y 2 variedades), 13 de ellos endémicos de este país, y distribuidos desde las regiones de Arica y Pari-nacota; en límites con Perú hasta la región del Maule, destacándose la IV Región de Coquimbo por la mayor riqueza taxonómica. Sus especies constituyen elementos dominantes del paisaje y definen provincias fito-geográficas. Como resultado del estudio taxonómico se proporcionan iconografías, descripciones originales, descripciones detalladas, distribuciones y hábitats, comentarios taxonómicos y nomenclaturales, exsiccatas, y tres claves de especies, subespecies y variedades. Se provee una sinonimia actualizada con ocho nuevos sinóni-mos, una nueva combinación: Eulychnia coquimbana (Molina) S. Albesiano; tres cambios de rango: (1) Tricho-cereus chiloensis subsp. australis (F. Ritter) S. Albesiano, (2) Trichocereus chiloensis subsp. eburneus (Phil. ex K. Schum.) S. Albesiano, (3) Trichocereus chiloensis subsp. panhoplites (K. Schum.) S. Albesiano, tres nombres nuevos: (1) Trichocereus faundezii S. Albesiano, (2) Trichocereus pectiniferus S. Albesiano, (3) Trichocereus un-dulosus S. Albesiano y trece neotipificaciones.

Palabras clave: Cactaceae, Chile, distribución, endemismo, taxonomía, Trichocereus.

A NEW TAXONOMIC TREATMENT OF THE GENUS TRICHOCEREUS(CACTACEAE) IN CHILE

SOFÍA ALBESIANO1, 2, 3

1Universidad Pedagógica y Tecnológica de Colombia.

Tunja, Colombia

2Universidad Nacional de la Plata,

Buenos Aires, Argentina

3Instituto Argentino de Investigaciones de las Zonas Áridas-CONICET,

C.C. 507 (5500) Mendoza, Argentina

email: [email protected]

HASELTONIA VOL. 18. 2012 117

besiano & Terrazas 2012). The clade is supported by three synapomorphies and eight homoplasies. The basitonic habit with prostrate branches (branches are produced from the base, with no central trunk), imbricate scales along the floral tube, and subglobose fruit were recovered as synapomorphies, but not the growth form (columnar) and not flower pilosity (Al-besiano & Terrazas 2012). Trichocereus has a close affinity with Lobivia as they share two homoplasies: sharp ribs (45–90o) and closely positioned scales on the surface of the receptacle.

Albesiano and Kiesling (2012) determined the identity and neotypification of Trichocereus macrogo-nus (type species of the genus) and its synonyms and affinities, contributing to the nomenclatural stability of the genus. This species has been commonly culti-vated in Bolivia, Chile, Colombia, Ecuador and Peru for medicinal properties and for its use as a living fence, among other uses.

The complexity and diversity of the genera of the tribe Trichocereeae (Acanthocalycium, Chamaecereus, Echinopsis (s.s.), Hymenorebutia, Lobivia, Pseudolobiv-ia, Soehrensia and Trichocereus) present possibly the most interesting taxonomic problem at the generic level in the family Cactaceae, at least for the South American cacti. There are over 100 names for the species of Trichocereus alone, and there is no modern collective treatment for the genus; there are only par-tial treatments of regional floras, Chile (Hoffmann & Walter 2004; Ritter 1980b); Argentina (Kiesling 1978; Ritter 1980a); Bolivia (Ritter 1980a; Navarro 1996; Navarro & Maldonado 2002); Perú (Ritter 1981; Brako & Zarucchi 1993); and Ecuador (Mad-sen 1989). While there are species listed for Paraguay and Brazil, this information is either inaccurate or includes species currently assigned to other genera (Kiesling 1978). Based on the literature mentioned, it is estimated that the current number of valid Trichocereus species approaches 45; this number de-pends, of course, on the taxonomic treatment of the genus, which is in the process of being reviewed by Albesiano (in prep.).

The objective of this paper is to classify the spe-cies of Trichocereus with natural distribution in Chile, based on morphological characteristics, in addition to resolving nomenclatural issues in order to elu-cidate the taxonomy of a complex group in a little-known tribe that is of great biological and cultural importance.

MATERIALS AND METHODS

Two botanical explorations to the arid and semi-arid regions of Chile were conducted − one in No-vember 2008 and the other in October 2010. Data were compiled on the habitat (location and type of vegetation), and on the specimens (individual height and morphology of the ribs, areoles, spines, flow-ers, fruits and seeds). Fragments were collected and processed, then placed in the MERL and AGUCH Herbaria (Luis Faúndez collection, Santiago, Chile). In addition, exomorphological studies were comple-

mented by the observation of materials from the fol-lowing herbaria: LIL, LP, LPB, MERL, SGO, SI, U. These materials are denoted by the acronym of the herbarium, followed by an exclamation mark to de-note personal examination by this author. In many cases, neotypes had to be designated because the authors had based their species descriptions on live specimens which had never been documented as her-barium specimens. A data sheet was created for each taxon, which included the following information: synonyms, types, iconography, original description, extended description, distribution and habitat in-formation, taxonomic and nomenclatural comments, as well as exsiccata. Three keys are provided to de-termine (1) species, (2) subspecies of Trichocereus chiloensis, and (3) varieties of T. macrogonus. A list of excluded taxa that belong to another genus, or whose determination could not be achieved due to lack of information, is also included.

Description of the genus Trichocereus

Habit: Erect plants, pendulous or cespitose; soli-tary (up to 15 m high) or forming dense cespitose shrubs 2–3 m high or less than one meter. Stem:Species such as Trichocereus atacamensis and Tricho-cereus chiloensis subsp. litoralis have branchless trunks of 0.5 to 3 m high, and a diameter of 50 cm. Many individuals of some species lack trunks (e.g., Tricho-cereus deserticola, Trichocereus spinibarbis). Branch-es: Cylindrical, narrow, straight or slightly curved, branched at ground level or from a trunk, with acute apices (angle 45–90 degrees). The diameter (6–25 cm) and color (light green, dark green or gray) varies within and among species. Ribs 9–25(–40), wide (up to 5 cm), short (up to 2 cm depth), surface curva-ture predominantly obtuse. In cespitose species, such as Trichocereus nigripilis and Trichocereus spinibarbis,rib characteristics remain constant throughout the length of branches, while arborescent species, such as T. atacamensis, present differences in number as well as in size, between apical and basal ribs. In Tricho-cereus faundezii and T. undulosus ribs are bulbous in the area of the areola (i.e., tuberculate). Areoles: May be round, oblong or obovate, short (up to 10 mm high) and pannose with white, yellow, gray, brown or black hairs. Spines: Terete; radials short (2 cm), numerous (11–21), at a 75° angle to the axis of the stem (divaricate) or appressed to the surface of the branch (15° angle), intersecting or not intersecting with the spines of neighboring areoles; centrals long (10−20 cm), few (1–6), and perpendicular to the branch or divaricate. Their structure may be weak and flexible (bristly) or rigid and stiff (acicular or su-bulate). Spines on the apices of branches have brown tips and a light green base, while those toward the base are white with black apices or have black, gray, olive green, dark brown or black dots. Flowers: Api-cal and/or subapical, infundibular, 8–21 cm long; ovary 2–3 cm in diameter; floral tube 4–7 cm long, 2–3 cm in diameter distally, and covered with subu-late scales from 0.6 to 3 cm in length (shorter than

118 ALBESIANO — TREATMENT OF CHILEAN TRICHOCEREUS

those on the ovary), entirely green or green with brown tips, with abundant (20–40) gray, brown or black hairs, from 0.6 to 2 cm long, stemming from their axils; nectaries 3–4 mm in diameter, tubular; stamens white, 13–23 mm long, arranged in two se-ries; the first series distributed along the floral tube, the second series forming a ring around the neck of the floral tube; style light green, 5–11 cm long, 1–3 mm wide; style branches 13–16, light yellow, erect, 0.7−2 cm long; outer tepals linear acuminate, inner-most tepals spatulate, 3−7 cm long, 1.3−2.0 cm wide, yellowish green or yellowish white with pink or brown (rarely deep pink) longitudinal lines; perianth 6–18 cm in diameter. Fruits: Apical and/or subapi-cal, umbilicate berry, 3–4 cm long, 3–5 cm in diame-ter, covered with subulate scales, axils have abundant brown, gray or white hairs, 4–6 mm long; dark green exocarp, mesocarp and endocarp white. Seeds: 1–2 mm long, 0.8–1.0 mm wide, with black testa.

TAXONOMIC TREATMENT

1. Trichocereus atacamensis (Phil.) W.T. Marshall & T.M. Bock, Cactaceae: 94. 1941. (Fig. 1).

Cereus atacamensis Phil. Fl. Atacam.: 23. 1860. Helianthocereus atacamensis (Phil.) Backeb., Die Cac-taceae 2: 1315. 1959. Echinopsis atacamensis (Phil.) H. Friedrich & G. D. Rowley, I.O.S. Bull. 3(3): 94.

1974. Echinopsis atacamensis (Phil.) H. Friedrich & G.D. Rowley subsp. atacamensis, Lazaroa 17: 54. 1996. TYPE: Chile, II Región de Antofagasta, Prov. El Loa, San Pedro de Atacama, Mina de San Bartolo, 22°14' lat. m., 2438 m, R. A. Philippi 31. (Lectotype: SGO 052685! designated by Muñoz: 91. 1960.)

Pilocereus pasacanus F.A.C. Weber, in Förster, Handb. Cacteenk. 2: 678. 1885. Cereus pasacanaF.A.C. Weber, Monatsschr. Kakteenk. 3: 165. 1893. Cephalocereus pasacanus (F.A.C. Weber) F.A.C. Weber, Index Kewensis 1: 89. 1906. Trichocereus pasacana(F.A.C. Weber) Britton & Rose, The Cactaceae 2: 133. 1920. Helianthocereus pasacana (F.A.C. Weber) Backeb, Cactac. Handb. Kakteenk. 2: 1314. 1959. Echinopsis pasacana (F.A.C. Weber) H. Friedrich & G.D. Rowley, I.O.S. Bull. 3(3): 96. 1974. Trichocer-eus atacamensis var. pasacana (F.A.C. Weber) F. Ritter, Kakteen Südamerika 2: 447. 1980. Echinopsis ataca-mensis subsp. pasacana (F.A.C. Weber) G. Navarro, Lazaroa 17: 54. 1996. TYPE: Argentina, Prov. Jujuy, Tilcara, 27 Oct. 1964, A. L. Cabrera 16340B. (Neo-type: LP, designated here.)

Cereus eriocarpus Phil., Anal. Mus. Nac. Chile 2: 27. 1891. TYPE: Chile, I Región de Tarapacá, Calc-alhuay, 3700 m, R. A. Philippi s.n. (Lectotype: SGO 052688! designated by Muñoz: 91. 1960.)Trichocereus cephalopasacanus Frič. Möller’s Deutsche Gärtn.–Zeitung (15) 44. 169. 1929, nom. nud.

Key for identification of species

1. Apical spines flexible, weak (bristly). ..................................................................................... 21 . Apical spines rigid, stiff. ....................................................................................................... 32 (1). Plants columnar, scarcely branched, usually unbranched.

Ribs numerous (24–40). .................................................................................. T. atacamensis2 . Plants branched, shrubby or arborescent. Ribs 12−17. ........................................ T. skottsbergii3 (1 ). Ribs few (5−)6−8(−9) and deep (2.5 cm).

Distance between areoles greater (2.5 cm).......................................................... T. macrogonus3 . Ribs numerous (10–21) and short (less than 2 cm deep).

Distance between areoles shorter (2 cm or less). .................................................................... 44 (3 ). Branches 5–10 cm in diameter........................................................................................ 54 . Branches 10–25 cm in diameter. .......................................................................................... 65 (4). Adult outer spines appressed to the stem (divergence angle ca. 15 degrees),

pectinate arrangement. ...................................................................................... T. pectiniferus5’. Adult outer spines divaricate to stem (divergence angle ca. 75 degrees),

radial arrangement. ............................................................................................................. 76. Ribs noticeably bulging in the area of the areoles (tuberculate). ............................................. 86 (4 ). Ribs not thickened in the area of the areoles. ................................................................. 97 (5 ). Radial spines 7–10, gray. Plants of Region XV. ......................................... T. uyupampensis7 . Radial spines 14–20, brown to greenish brown. Plants of Regions II, III and IV. ............... 108 (6). Plants short (80 cm high), cespitose and branched at ground level. .................. T. faundezii8 . Plants tall (2.5 cm high), shrubby and branched from the trunk...........................T. undulosus9 (6 ). Low rib height (0.6–1.0 cm), at middle point of stems. ............................. T. bolligerianus9 . High rib height (1.1–1.5 cm), at middle point of stems.........................................T. chiloensis10 (7 ). Spines entirely black, subapical. ..................................................................... T. nigripilis10 . Spines gray, with gray or brown tips, subapical. ............................................................... 1111 (10 ). Spines abundant (more than 10), bristly,

flexible and curved at apex, subapex and middle of stem. .................................... T. spinibarbis11 . Spines absent or scarce (less than 10), bristly. .................................................... T. deserticola


Trichocereus eremophilus F. Ritter, Kakteen Sü-damerika 2: 559. 1980. TYPE: Bolivia, Dpto. Potosí, Prov. Nor Lípez, Mpio. Colcha “K”, 3600–3900 m, F. Ritter 49a. (Holotype: U not seen; isotype: SGO 121151!.)

Iconography: Spegazzini (1925): 118. Mar-shall & Bock (1941), photograph 42: 93. Backeberg (1959), photographs 1261–1263: 1308–1310. Rit-ter (1980b), photograph 1080: 1226. Mauseth et al. (2002): 270–271. Hoffmann & Walter (2004), plate 14: 87. Kiesling & Ferrari (2005), photograph 40: 56.

Original description:

Extended description: Plant erect, up to 15 m high. Trunk 47 cm diameter, branched at heights of 1 to 3 m above the ground, 25 cm in diameter and sometimes branching. There are also plants with a single stem. Ribs at the apex 25, obtuse, 1 cm wide, with rounded areoles, 3 mm high and 2 cm wide, white when young or somewhat grayish; central spines 4, acicular, flexible and weak (bristly), 5 cm long, 1 mm diameter, yellow brown, arranged cross-wise; radial spines 18, acicular, 2 cm long, 1 mm in diameter, weak and flexible, yellow brown, overlap-ping distally with radial spines of surrounding are-oles. Ribs at the base 24–40, obtuse, 5 cm wide and 2 cm high, with circular areoles 3 mm high and 2 cm wide, completely gray; central spines 7, 9 cm long, 3 mm in diameter, acicular, yellow brown; radial spines more than 15, 3 cm long, acicular, and yellow brown. Flowers generally subapical, rarely apical, 15 cm long, with abundant brown hairs 1−2 cm long on the ovary and floral tube; perianth 18 cm in di-ameter; ovary 2–3 cm wide, with floral ovary scales1 cm long, green with brown apex, floral tube scales1–2 cm long, green or yellowish green with brown apex; tepals 4–6 cm long, usually greenish yellow or yellowish white with brown longitudinal lines, rare-ly intense pink; style green, 8 cm long; stigma yellow, 1 cm long. Fruit lateral, 2–3 cm long, 3–4 cm in di-ameter, covered with subulate scales, with axils pre-senting abundant whitish hairs. Seeds 1.4 mm long, 0.8 mm wide.

Figure 1. Trichocereus atacamensis. Photo by Luis Faúndez.


Distribution and habitat: T. ata-camensis is distributed in the arid and semi-arid Chilean NE, Argentine NW and Bolivian South. In Chile it occurs in Region I of Tarapacá and Region II of Antofagasta, in the Ata-cama Desert, and in high Andean communities of the phytogeographic region known as the Prepuna (Ca-brera & Willink 1980), at altitudes of 2400−3700 m. In Argentina it grows in the provinces of Jujuy, Salta, Tucumán and Catamarca, mostly in the Prepuna or protected slopes of the Puna, at altitudes of 2000–3500 m, on rocky slopes, plains and semi-plains. Because of the height of the plant, it is a conspicuous element of the landscape. In Bolivia it occurs in the departments of Oruro and Tarija, at 3500–4000 m altitude.

Taxonomic and nomenclatural comments: Comparing the original descriptions and herbarium material of Cereus atacamensis, Cereus eriocar-pus and Pilocereus pasacanus, there is a high similarity in the habit (erect, 3–15 m high), stem thickness (ca. 40 cm), number of ribs (24–40) and nature of the apical spines (thin and flexible). It is therefore suggested they be grouped under T. atacamensis. The variation in the number of branches is due to the variation in habitat moisture over the area in which this species grows (R. Kiesling, pers. comm.). Britton and Rose (1920) noted the simi-larity between T. pasacana and Cereus atacamensisand proposed they be grouped; Kiesling and Ferrari (2005) and Kiesling et al. (2008) accepted this view.

In a communication sent by Luc Willemse, cu-rator of the herbarium U (Netherlands), he reports that the type specimen of Trichocereus eremophi-lus: Ritter 49a, does not exist in that institution, as quoted in Ritter (:iii. 1979, 1980a) and Eggli et al. (1996).

Herbarium material studied: Argentina, Prov. Catamarca, Dpto. de Belén, Hualfín, 2000 m, 20 Feb. 1947, D. Droghetii s.n. (LIL); Dpto. Belén, Yacutula, 26 Oct. 1974, R. Kiesling 802 (SI); Dpto. Santa María, Cerrillos–Sierra Aconquija, 3100 m, 17 Dic. 1933, Peirano 9797 (LIL); Prov. Jujuy, Tilcara, 27 Oct. 1964, A.L. Cabrera 16340 (LP); Tilcara to Maimará, 29 Oct. 1981, M.M. Costa 2547 (SI); Prov. Salta, Tintín, 3004 m, 3 Dic. 1960, A. Ruíz-L. 21616 (MERL), Dpto. Cachi, Palermo, 20 Oct. 1948, A. Burkart 17616 (SI); Prov. Tucumán, Los Cardones a Santa Rosa (Qda. Amaicha), 2500–3000 m, Dic. 1931, Schreiter 7204 (LIL). Herbario 66736 (LIL). Bolivia, Dpto. Tarija, Prov. Eustaquio Mén-dez, Iscayachi, 19 km hacia Curqui, 3500 m, 10 Nov. 1993, St. G. Beck et al. 22082 (LPB); Dpto. Oruro, Prov. L. Cabrera, Mpio. Salinas de García Mendoza,

19°42'44.7"S; 67°38'43.3"W, 3738 m, Jul. 2006, M. Arakaki and N. Quispe 1723 (LPB). Chile, II Región de Antofagasta, Prov. Antofagasta, Dpto. Cementerio, camino cañería Antofagasta al N cementerio de To-conce, 23 Nov. 1969, H.C. Martin 516 (SI); idem, H.C. Martin 517 (SI).

Other material studied: Chile, I Región de Tara-pacá, Prov. Iquique, city of Iquique, Playa Brava, 12 Nov. 2008 (observed in cultivation).

2. Trichocereus bolligerianus (Mächler & Helmut Walter) S. Albesiano, Haseltonia 17: 18. 2012. (Fig. 2).

Echinopsis bolligeriana Mächler & Helmut Walter. Kakteen And. Sukk. 54 (10): 269, ill. 2003. TYPE: Chile, VI Región del Libertador General Bernardo O’Higgins, Prov. Cardenal Caro, Punta de Lobos, Ene. 2003. Walter 213. (Holotype: SGO!.)

Trichocereus chiloensis var. conjungens F. Ritter, Kakteen Südamerika 3: 1109, ill. 1980. TYPE: V Región de Valparaíso, Küstengebiet. Häufigauch bei Limache, Olmué und Granizo, östlich von Valparai-so FR 228c. (Holotype: U 160273!.) (Name of holo-type is written on herbarium label as “T. chilensis var. conjungens”.)

Iconography: Ritter (1980b), photograph 1062: 1221. Hoffmann and Walter (2004), plate 15: 89. Hunt et al. (2006), photograph 232.4: 232.

Figure 2. Trichocereus bolligerianus.



Extended description: Plants erect or pendulous, 2−6 m long, without trunk. Branches 18 cm in di-ameter. Ribs at the apex 17, obtuse, 0.6–2 cm wide, 3 mm high; with areoles ovate, 2 mm high, 6–8 mm wide, apex yellow, base gray or black; with 1–4 central spines, 0.3 to 2 cm long, acicular, base light green, middle part and apex brown; and with 16 ra-dial spines, 0.5 to 2 cm long, acicular, base light grey or green, middle part green and apex brown. Ribs at the base up to 20, obtuse, 2 cm wide, 10 mm high; with areoles ovate, 2 mm high, 6 mm wide, gray or brown; with 4–6 central spines, 0.2–6 cm long, small-er than 0.7 mm in diameter, acicular, olive green or dark brown; 15 radial spines, 1–2 cm long, acicular, flexible, diameter less than 0.7 mm, olive green or light grey. Flowers subapical, 1 or 3, infundibuliform, 14 cm long, with abundant brown pilosity on ovary and floral tube, perianth 7 cm in diameter, ovary2–3 cm wide, floral scales of the pericarpel 2 cm long, green with apex brown, floral scales of tube 3 cm long, green or yellowish green with apex brown, te-pals 4–6 cm long, greenish yellow or yellowish white with longitudinal lines brown, style green, 8 cm long, stigmas yellow, 1.5 cm long. Fruit subapical, 2–3 cm long and 3–4 cm in diameter, covered with triangu-lar scales, from whose axils emerge abundant brown and grey hairs; pulp white. Seeds 1.8 mm long, 1.0 mm wide.

Distribution and habitat: T. bolligerianus grows in Regions V and VI, on hills of the Coastal Cordil-lera, next to the sea, and on steep slopes (15−20o), where part of the vegetation has been altered by tourist activities.

Taxonomic and nomenclatural comments:Analyzing the original descriptions of Trichocereus chiloensis var. conjungens and Echinopsis bolligeriana,high similarity is observed in the number of ribs and in the number and morphology of spines. Accord-

ingly, we propose to unite these two taxa. Trichocer-eus chiloensis var. conjungens has been considered syn-onymous with Echinopsis chiloensis (Anderson 2005), and lately with T. chiloensis subsp. chiloensis (Kies-ling et al. 2008), but on analyzing the phylogeny of the genus Trichocereus (Fig. 3, Albesiano & Terrazas 2012), morphological (ribs low and sharp) and mo-lecular characters were found that allow its differen-tiation. The individuals in populations of Trichocer-eus bolligerianus observed in Region VI are smaller in size (2 m tall) because of the drastic environmental conditions (higher influence of the cold Humboldt current), compared to the Valparaiso region where populations develop under benign conditions and in more sheltered places.

Herbarium material studied: Chile, VI Región del Libertador General Bernardo O’Higgins, Prov. Cardenal Caro, Punta Lobos, 34o25'33"S; 72o02'36"W; 3 m, 2 Nov. 2008, S. Albesiano, et al. 2039 (AGUCH, MERL); Costa de Pichilemú, 34o27'19"S; 72o01'05"W; 7 m, 2 Nov. 2008, S. Al-besiano et al. 2040 (AGUCH, MERL).

3. Trichocereus chiloensis (Colla) Britton & Rose, The Cactaceae 2: 137, ill. 1920.

Cactus chiloensis Colla, Hortus Ripul. App. 2: 342. 1826. Cereus chiloensis (Colla) DC., Prodr. 3: 465. 1828. Echinocereus chiloensis (Colla) Console & Lem., Rev. Hort. 35: 173. 1864, nom. nud. Echino-cereus chiloensis (Colla) Console & Lem., Illustr. Hor-tic. xi. Misc. 15. 1864. Echinopsis chiloensis (Colla) H. Friedrich & G.D. Rowley, I.O.S. Bull. 3(3): 94. 1974. TYPE: Chile, Región Metropolitana, Santiago, Río San Francisco, 1800 m, 11 Dic. 1924, Wer-dermann 478. (Neotype: K not seen, designated by Hunt & Taylor: 7. 2006; isoneotypes: SI 079802! and MO).



Extended description: Plants erect, 1–6 m high. Branches 10–25 cm in diameter. Ribs at the apex11–17, obtuse, 0.6–3 cm wide, 3−7 mm high; with obovate or oblong areoles 1–4 mm high, 0.6–1.3 mm wide, yellow hairs on the apex and gray or black on the base, or entirely yellow; central spines 1–6, acic-ular, 0.3–15 cm long, brown at the tips and light green at the base, subapical spines gray at the base with brown apex or white with some gray or black spots; radial spines 8–19, 0.4–4 cm long, acicular, rigid, straight, dark brown, green, yellow, gray or white with brown tips. Ribs at the base 11–21, ob-tuse, 1.5 to 4 cm wide, 0.2–2 cm high, with round, oblong, elliptic areoles, 1–5 mm high, 0.5–1.3 cm wide, the oldest sunken, light yellow, brown, light or dark gray or black pilosity; central spines 1–7, gener-ally 4, acicular and straight, 0.2–20 cm long, up to 2 mm in diameter, brown or reddish apex, gray or black base or entirely olive green or dark brown; ra-dial spines 9–24, acicular, flexible or rigid, 1–2.7 cm long, less than 1.5 mm wide, yellow, olive green or light gray with brown, dark gray or black tips. Flow-ers 1–8, 11–21 cm long, subapical with abundant brown hairs on the ovary and floral tube; perianth7–11 cm in diameter; ovary 2–3 cm in diameter, with floral scales 1−3 cm long, with brown apex and dark green center; floral tube scales 1–4.5 cm long, with brown tips, green in the middle and yellowish green base; tepals spatulate, 7 cm long, pink apex and yellowish white base; style green, 10 cm long; stigma yellow, 1–2 cm long. Fruits subapical, 2–4 cm long, 3–5 cm diameter, covered with subulate scales, with abundant gray and brown axillary hairs. Seeds 2 mm long, 1 mm wide.

Distribution and habitat: T. chiloensis is the spe-cies of the genus Trichocereus with the most extensive distribution in Chile, occupying the regions: II, IV, V, VI, VII, VIII, RME, from the pacific coast inland, in river valleys and slopes, from sea level to 1500 m.

Taxonomic and nomenclatural comments: Ac-cording to Article 51.1 of the International Code of Botanical Nomenclature (McNeill et al. 2006), the

name Cactus chiloensis cannot be rejected or replaced even though Colla confused Chile with Chiloé island.

Console and Lemaire (Rev. Hort. 35: 173. 1864) proposed the name Echinocereus chiloensis, without providing a description of the species. Subsequently, Lemaire (Cact. 61: 1868) declared that the name was based on Cereus chiloensis.

The following taxa were excluded from the taxo-nomic treatment, due to the fact that the informa-tion provided in the original description regarding the color and number of central and radial spines is not sufficient for taxonomic identification, and data are missing concerning the characteristics of the stem: habit, height and diameter, in addition to the number of ribs. Furthermore, varieties whose stem diameter is below that observed in the field were not taken into account.

Cereus chiloensis var. brevispinula Salm-Dyck., Cact. Hort. Dyck. (1849): 199. 1850. (Name appears as “C. chilensis var. brevispinula”.)

Cereus chiloensis var. flavescens Salm-Dyck., Cact. Hort. Dyck. (1849): 199. 1850. (Name appears as “C. chilensis var. flavescens”.)

Cereus chiloensis var. heteromorpha K. Schum., Gesamtbeschr. Kakt. 63. 1899. (Name appears as “C. chilensis var. heteromorpha”.)

Cereus chiloensis var. polygona Salm-Dyck., Cact. Hort. Dyck. (1849): 199. 1850. (Sub “C. chilensis var. polygona”.)

Cereus chiloensis var. poselgeri K. Schum., Gesamt-beschr. Kakt. 63. 1899. (Name appears as “C. chilen-sis var. poselgeri”.)

Cereus chiloensis var. pycnacantha K. Schum., Gesamtbeschr. Kakt. 63. 1899. (Name appears as “C.chilensis var. pycnacantha”.)

Cereus chiloensis var. zizkaana K. Schum., Gesam-tbeschr. Kakt. 63. 1899. (Name appears as “C. chilen-sis var. zizkaana”.)

T. chiloensis has high intraspecific variation with respect to growth habit, stem diameter, as well as the number and shape of the spines; as a result, six subspecies are proposed following Schumann (1899)

Subspecies key1. Short plants, 1–1.5 m, forming dense cespitose shrubs ................................subsp. panhoplites1 . Tall plants, greater than 2 m, shrubby or arboreal ............................................................... 22 (1 ). Adult radial spines, flexible, very thin and numerous (14–30) .................... subsp. litoralis2 . Adult radial spines, strong, rigid and few (less than 14) ...................................................... 33 (2 ). Stem diameter 9–10 cm.......................................................................... subsp. chiloensis3 . Stem diameter between 13–25 cm...................................................................................... 44 (3 ). Presence of white spines in subapical areoles,

radial spines in adult areoles 11–13. .............................................................. subsp. eburneus4. Presence of gray spines with brown apices in subapical areoles,

radial spines in adult areoles 7–10. ................................................................ subsp. australis


and Ritter’s (1980b) suggestion.Ritter (1980b) erroneously corrected the spelling

of the name T. chiloensis to T. chilensis.

4. Trichocereus chiloensis (Colla) Britton & Rose subsp. chiloensis

Cereus quisco Remy, Fl. Chil. 3: 19. 1847. TYPE: Chile, Región IV (de Coquimbo), Prov. Elqui, Mpio. Vicuña, 29°51.37'S; 70°22.33'W, 1280 m, 27 Oct. 1997, U. Eggli & B.E. Leuenberger 3052. (Neo-type, designated here: SGO 144120. Isoneotypes: B, CONC, ZSS.)

Trichocereus chiloensis var. borealis F. Ritter, Kak-teen Südamerika 3: 1109. 1980. TYPE: IV Región de Coquimbo, Prov. Elqui, valley of Elqui, Huanta, FR 228d. (Holotype: U!.) (Name appears as “T. chilensisvar. borealis”.)

Iconography: Ahumada and Faúndez (2007): 144.

Extended description: Plants up to 6 m high. Branches 10 cm in diameter. Ribs 11–18, obtuse, 10–15 mm high. Areoles at the apex 5–12 mm wide, one central spine; areoles at the base with 4 central spines, 3–20 cm long, longer and thicker than radials. Radial spinesup to 13, 2 cm long, dark brown when young, then light gray, brown at the tips or completely dark gray. Flowers 13 cm long; te-pals white, spatulate, apiculate at the tips.

Distribution and habitat:Includes Regions IV and V and the Metropolitan area, from the coast inland, and the environs of Limanche, Olmué, Santiago, etc. It is an important element of the landscape of Chile’s Mediterranean area.

Herbarium material studied: IV Región de Co-quimbo, Prov. Limarí, Mpio. Ovalle, 36 km from as-phalt road Ovalle–La Serena, 30°22.42'S; 71°0.49'W, 700 m, 3 Dic. 1994, U. Eggli and B.E. Leuenberger 2570 (SGO); Prov. Elqui, 2 km S of main road La Serena–Vicuña, 620 m, 30°2.79'S; 70°49.17'W, 26 Oct. 1997, U. Eggli and B.E. Leuenberger 3042 (SGO).

5. Trichocereus chiloensis (Colla) Britton & Rose subsp. australis (F. Ritter) S. Albesia-no stat. nov. TYPE: V Región de Valparaíso, Cuesta Las Chilcas, 32°51'05"S; 70°52'39"W, 400-440 m, 5 Nov. 2008, S. Albesiano et al. 2042. (Neotype, designated here: MERL. Iso-neotype: AGUCH.) (Fig. 3).

Trichocereus chiloensis var. australis F. Ritter, Kak-teen Südamerika 3: 1108, ill. 1980. TYPE: Chile, VII Región del Maule, Prov. Talca, Dpto. Lontué, Villa Prat, Abril de 1954, F. Ritter 228a. (Holotype: U, not seen.) (Name appears as “T. chilensis var. austra-lis”.)

Iconography: Ahumada and Faúndez (2007): 141. Hoffmann and Walter (2004), plate 16: 91.


Extended description: Plants 2–4 m high. Stemserect and cylindrical, 13–25 cm diameter. Ribs at the apex obtuse, 13–15, 1–3 cm wide, 3–5 mm high; with obovate areoles, 1–2 mm high, 7–8 mm wide, hairs yellow on the apex and gray on the base, or en-tirely yellow; central spines 1–3, acicular, 1 to 8 cm long, dark yellow to brown at the tips and green or yellow at the base, some subapical spines are gray at the base with brown apex; radial spines 9, from 0.4 to 4 cm long, acicular, yellow, some green or gray with brown tips. Ribs at the base 12–16, from 2 to 4 cm wide, 0.2–2 cm high, with round areoles, some sunken, 1–2 mm high, 6–10 mm wide, light or dark gray or black pilosity; central spines 1–3, acicular, 2–6 cm long, brown or reddish apex, gray or black base, the longest directed perpendicularly to the stem; radial spines 11, acicular, 2 cm long, yellow, green with brown or black tips. Flowers subapical,with cream or brown hairs on the ovary and floral tube, 11–21 cm long, perianth 7–11 cm in diameter; ovary 2–3 cm in diameter, with floral scales ca. 3 cm long, with brown apex and dark green center; floraltube scales subulate, 3–5 cm long, brown tips, green in the middle, and yellow basally; tepals spatulate, 4–7 cm long, with pink apex and yellow base; style

Figure 3. Trichocereus chiloensis subsp. australis.


green, 6–10 cm long; stigma light yellow, up to 2 cm long.

Distribution and habitat: Located in Regions V, VI and VII, on the slopes of the coastal moun-tain range, dominated by the herbaceous plants and shrubs.

Taxonomic and nomenclatural comments: The type specimen (FR 228a) is mentioned (Ritter: iii., 1979, 1980b) as having been deposited in the Na-tional Herbarium of the Netherlands. According to Eggli et al. (1996), the holotype is lost, and there-fore we designate a neotype for the Valparaiso region, which possesses the morphological characteristics of the original description.

In addition, Eggli et al. (1996) list seeds under number FR 228a, at the Zurich Herbarium, but there is no assurance that they correspond to the same type collection of Trichocereus chiloensis var. australis F. Ritter 228a, as the original description does not state that they have been deposited at this herbarium.

Herbarium material studied: Chile, Región Metropolitana, Prov. Cordillera, 8.5 km above pu-ente El Yeso, in the valley of the río Volcán, 1420-1500 m, 1/2/1993, U. Eggli and B. Leuenberger 2299 (SGO). VI Región de O’ Higgins, Prov. Col-chagua, comuna de San Fernando, Manantiales, 34°37'10"S; 71°02'47"W; 338 m, 2 Nov. 2008, S. Albesiano et al. 2038 (MERL, AGUCH); Prov. Cardenal Caro, Comuna de Litueche, a 7 km al NE, 34°05'28"S; 71°41'07"W, 305 m, 2 Nov. 2008, S. Albesiano et al. 2041 (MERL, AGUCH). V Región de Valparaíso, Cuesta Las Chilcas, 32°51'05"S; 70°52'39"W, 400–440 m, 5 Nov. 2008, S. Albesiano et al. 2043 (MERL, AGUCH).

6. Trichocereus chiloensis (Colla) Britton & Rose subsp. eburneus (Phil. ex K. Schum.) S. Albesiano, stat. nov. (Fig. 4).

Cereus chiloensis var. eburnea Phil. ex K. Schum., Gesamtbeschr. Kakt.: 63. 1899. (Name appears as

“Cereus chilensis var. eburnea”). Eulychnia eburnea Phil, nom. nud. (according to Schumann, loc. cit.). Cereus chiloensis f. eburneus (Phil.) Schelle, Handb. Kakteen-kult.: 63. 1907. (Sub “Cereus chilensis f. eburneus”). Trichocereus chiloensis var. eburneus (K. Schum.) F. Ritter, Kakteen Südamerika 3: 1109. 1980, nom. inval. (Art. 33.3). (Name appears as “Trichocereus chilensis var. eburneus”.) TYPE: Chile, IV Región de Coquimbo, 31°28'00"S; 71°09'52"W, 870 m, 6 Nov. 2008, S. Albesiano et al. 2049. (Neotype, designated here: MERL. Isoneotype: AGUCH.)


Extended description: Plants erect, shrubby, 2 m high. Lateral branches arising at ground level, light green, 13 cm wide, the apex with abundant spines, acicular, brown, 1 mm diameter. Ribs at the apex 12, obtuse, 2 cm wide, 4 mm high, with obovate areoles,1 mm high, 8 mm wide, yellow hairs on the apex and gray on the base; central spines 4, acicular, 4–15 cm long, 1.5 mm diameter, white with some black spots; radial spines 10, acicular, 3 cm long, less than 1 mm diameter, white and rigid. Ribs at the base ob-tuse, 12, 3 cm wide, 1 cm high, with obovate areoles,1 mm high, 1.3 cm wide, with entirely dark gray to black pilosity; central spines 4, acicular 6–12 cm long, 1.5 mm diameter, dark brown, radial spines 12, acic-ular, 2.4 cm long, black. Flowers subapical, 5–8, 13 cm long, perianth 8 cm in diameter; ovary and floral tube with scales 1.2 cm long, with brown apex, green base; from whose axils emerge abundant brown hairs, 2 mm long; tepals 3–6 cm long, brown apex, white base.

Distribution and habitat: IV Region of Coqui-mbo, Province of Choapa, on hillside that do not ex-ceed 900 m, and where the dominant physiognomic type is xerophytic scrub less than 2 m high, featuring the columnar cacti Eulychnia and Trichocereus.

Taxonomic and nomenclatural comments: This taxon morphologically resembles T. chiloensis subsp. australis, but differs in having young spines that are white, and a greater number of radial spines in adult areoles.

Figure 4. Trichocereus chiloensis subsp. eburneus


Philippi proposed the name of Eulychnia eburnea(nom. nud.), without providing a diagnosis or mor-phological description for the plant. Subsequently, Schumann (1899, pp. 59–60) considered that this name corresponded to a variety of Cereus chiloensis,since Philippi described only three species of Eulych-nia: E. acida, E. breviflora and E. castanea.

According to Article 33.3 of the International Code of Botanical Nomenclature (McNeill et al. 2006), the combination T. chiloensis var. eburneus (K. Schum.) F. Ritter is invalid, as Ritter did not provide basionym information: name, place and date of pub-lication.

Herbarium material studied: Chile, IV Región de Coquimbo, Prov. Elqui, Mpio. Vicuña, 32 km S of Vicuña, 30°11.53'S; 70°39.61'W, 1650 m, 28 Oct. 1997, U. Eggli and B. E. Leuenberger 3058 (SGO); Prov. Limarí, Mpio. Río Hurtado, 30°16.03'S; 70°39.64'W, 1300 m, 28 Oct. 1997, U. Eggli and B. E. Leuenberger 3068 (SGO); Prov. de Choapa, 31°28'00"S; 71°09'52"W, 870 m, 6 Nov. 2008, S. Albesiano et al. 2049 (AGUCH, MERL).

7. Trichocereus chiloensis (Colla) Britton & Rose subsp. litoralis (Johow) Faúndez, Darwin-iana 45(2): 237. 2007. (Fig. 5).

Cereus litoralis Johow, Revista Chilena Hist. Nat.25: 157. 1921. Trichocereus litoralis (Johow) Looser, Revista Chilena Hist. Nat. 33. 598, ill. 1929. Echi-nopsis litoralis (Johow) Friedrich & G. D. Rowley, I.O.S. Bull. 3 (3): 96. 1974. Echinopsis chiloensissubsp. litoralis (Johow) M. Lowry, Cactaceae Syst. Init. 16: 8. 2003. TYPE: Chile, V Región de Valpara-iso, San Antonio, 5 Nov. 1997, U. Eggli, B. Leuen-berger and S. Arroyo 3110. (Neotype: SGO 146136! designated by Faúndez and Kiesling: 237. 2007. Iso-neotypes: B, CONC, ZSS.)

Iconography: Looser (1929), plate. 114: 599. Hoffmann (1989), plate 10: 89. Hunt et al. (2006), photograph 233.4: 233.


Extended description: Plants 3 m high. Stems:trunk 50 cm high; branches 12–18 cm in diameter; in some individuals, the branches are initially pro-cumbent and then become erect, in other plants all branches are erect. Ribs at the apex 17, obtuse, 1 cm wide, 3 mm high; with obovate areoles 1 mm high, 6 mm wide, with yellow hairs on the apex and gray on the base; central spines 2–6, acicular, 1–6 cm long, shorter ones with brown apex, green base, longest ones gray or dark yellow; radial spines 15–19, acic-ular, 2 cm long, some spines are green with brown apex or entirely gray or light brown. Ribs at the base17–21, obtuse, 2–4 cm wide, 0.3 to 1 cm high. Are-oles circular, 2–5 mm high, 1 cm wide, gray or black. At the upper edge of the areola is a transverse fur-row which results in depression of that part of the areole. Central spines 4–7, 2–4 cm long, acicular, 1.5 mm in diameter, olive green or light to dark brown. Radial spines 15–24, 1–2 cm long, acicular, 0.7 mm in diameter, green with brown tips. Flowers 1 or 2, subapical, 12–16 cm long, 11 cm in diameter when open (4 cm in closed flowers), ovary 1.5 to 2.2 cm wide, floral tube scales 1–2 cm long; floral upper scales 2–3 cm; tepals 5 cm long, style 8 cm long; stig-mata 1 cm long. Fruits 5 cm diameter, 4 cm long, covered by abundant brown and gray hairs.

Distribution and habitat: T. chiloensis subsp.litoralis occurs in Regions IV and V, extending along the coast of the Aconcagua, from Papudo south, pos-sibly in Valparaiso and Santiago and also Zapallar and Cachagua, on plains and hillsides near the coast, at 33−100 m altitude, habitat dominated by herba-ceous and shrub layers including Fuchsia lycioideswith some scattered saplings.

Taxonomic and nomenclatural comments: Be-cause of the resemblance in appearance (tall plants over 2 m, and shrubs), stem diameter (greater than

Figure 5. Trichocereus chiloensis subsp. litoralis


10 cm), rib number (ca. 21) and rib shape (obtuse, separated by acute and undulating grooves) T. litora-lis has been considered a subspecies of T. chiloensis.

The subspecies litoralis, in turn, differs from the subspecies chiloensis, by the number and morphology of the radial spines: flexible, thin and numerous (14–30) vs. strong, rigid and few (less than 14).

The stems of some individuals of T. chiloensissubsp. litoralis are prostrate and cespitose, similar to those of T. spinibarbis, but differ from the latter by the larger diameter of the stems (12–18 cm vs. 4–10 cm), and rib number (17–21 vs. 8–15), in addition to the smaller size of the central spines in mature ar-eoles (4 cm vs. 10 cm).

Looser (1929) found that the differentiating characteristics (number of stigmas and flower orien-tation), between T. litoralis and T. chiloensis, indicat-ed in the original description of C. litoralis (Johow 1921), are of little differential value, since some indi-viduals of this species have the same number of stig-mas (18) and their flowers have the same orientation (north).

Herbarium material studied: Chile, IV Región of Coquimbo, Prov. Choapa, 14.5 km on Panameri-cana N of turnoff from Panamericana to Los Vilos, in a steep valley between to road and the sea, 100 m, 10 Nov. 1991, U. Eggli and B. Leuenberger 1664, 1668 (SGO), Mpio. Mincha, Caleta Oscuro, 3 km W of Puerto Oscuro, 31o25.27'S; 71o35.68'W, 20-100 m, U. Eggli, B.E. Leuenberger and S. Arroyo 3087 (SGO) p.p.; Prov. Elqui, Mpio. Coquimbo, 22 km W Tongoy towards Puerto Aldea, 30o18.51'S; 71o35.52'W, 31 Oct. 1997, U. Eggli, B.E. Leuen-berger and S. Arroyo 3086 (SGO); V Región de Valparaíso, Prov. Petorca, La Ligua, Los Molles, 32°13'59"S; 71°30'46"W; 21 m, 5 Nov. 2008, S. Albesiano et al. 2044 (MERL, AGUCH); Quili-marí, Petorca, La Ligua, Los Molles, 32°07'17"S; 71°27'47"W; 33 m, 5 Nov. 2008, S. Albesiano et al. 2047 (MERL, AGUCH).

8. Trichocereus chiloensis (Colla) Britton & Rosesubsp. panhoplites (K. Schum.) S. Albesiano, stat.nov. (Fig. 6).

Cereus chiloensis var. panhoplites K. Schum., Gesa-mtbeschr. Kakt. 63: 1899. (Sub “Cereus chilensis var. panhoplites”). Trichocereus chiloensis var. panhoplites(K. Schum.) F. Ritter, Kakteen Südamerika 3: 1109, ill. 1980, nom. inval. (Art. 33.3). (Sub “Trichocereus chilensis var. panhoplites”). TYPE: Chile, IV Región de Coquimbo, Provincia Limarí, Ovalle, Higueritas Unidas, 30°22'16"S; 71°14'00"W, 477 m, 6 Nov. 2008, S. Albesiano et al. 2052. (Neotype: MERL, designated here. Isoneotype: AGUCH.)

Iconography: Ritter (1980b), photograph 1059: 1221.

Original description.

Extended description: Plants short, stems 1.0–1.5 m tall, cespitose. Branches originate at ground level, 16 cm in diameter. Ribs at the apex 11 obtuse, 3 cm wide, 7 mm high with obovate areoles 4 mm high, 1.3 cm wide, apex yellow and the rest gray. Central spines at the apex 1–3, 8 cm long, 2 mm di-ameter, acicular, absolutely straight and gray; radialsat the apex 6–10, 2.0−2.4 cm long, acicular, three of them with green base and brown tip, the oth-ers gray. Ribs at the base 12, obtuse, 3 cm wide, 7 mm high, with elliptical areoles 2 mm high, 1.3 cm wide, spaced 10–15 mm apart and completely gray, except in the area where new spines arise, where the hairiness is yellow. Central spines 3–4, 6−9 cm long, 2 mm wide, straight, acicular, gray with black apex; radials 9–11, 3 cm long, 1.5 mm in diameter, acicu-lar, gray with black tip. Fruit 3.5 cm long, 4 cm in diameter and covered with brown hairs.

Distribution and habitat: Near Ovalle, in the coastal plains, near the coastal range, at 500 m alti-tude; landscape dominated by herbaceous and low shrub layers.

Taxonomic and nomenclatural comments:Britton and Rose (1920) and Anderson (2005) con-sidered that this taxon is a synonym of T. chiloensis.Based on the original description and the material collected, we found evidence to consider it a subspe-cies of T. chiloensis, for example, the low stature and cespitose stems branching at ground level, whereas other subspecies may exceed 2 m and form shrubs or exhibit an arborescent habit, branching a few meters above the ground.

According to Article 33.3 of the International Code of Botanical Nomenclature (McNeill et al. 2006), the combination T. chiloensis var. panhoplites(K. Schum.) F. Ritter is invalid, as Ritter establishes no basionym or place or date of publication.

Herbarium material studied: Chile, III Región of Atacama, Prov. Huasco, Mpio. Vallenar, 31 km SW of the Panamericana on the gravel road to Mina Algarrobo, 28o46.74'S; 70o58.02'W, 1100−1250 m,

Figure 6. Trichocereus chiloensis subsp. panhoplites


7 Dic. 1994, U. Eggli and B.E. Leuenberger 2591 (SGO); IV Región de Coquimbo, Talinay, close to Ovalle, 24 Sep. 1957, C. Muñoz P. 4306 (SGO).

9. Trichocereus deserticola (Werderm.) Looser,

Chil. Hist. Nat. 33: 614. 1929. (Fig. 7).Cereus deserticolus Werderm., in Notizbl. Bot. Gart.

Berlin-Dahlem 10: 764. 1929. Trichocereus deserti-colus (Werderm.) Backeb., Kaktus-ABC 202. 1936, nom. inval. (Art. 53.1). Echinopsis deserticola (Wer-derm.) H. Friedrich & G.D. Rowley, I.O.S. Bull.3(3): 95. 1974. TYPE: Chile, Prov. Antofagasta, Dpto. Taltal, Taltal, 500 m, Oct. 1925, Werdermann 869, p.p. (Lectotype: B, designated by Leuenberger & Eggli: 690. 1996. Isolectotypes: U!, F, G, K, LIL, MO!, S, US.)

Trichocereus fulvilanus F. Ritter, Kakt. And. Sukku-lent. 13: 165-167, ill. 1962. TYPE: Chile, Taltal, Jun. 1954, F. Ritter 263. (Holotype: U 117827!)

Trichocereus serenanus F. Ritter, Kakt. And. Sukku-lent. 16 (11): 212. 1965. TYPE: Chile, 15 km to the east of Serena, F. Ritter 533. (Holotype: U 116996!)

Iconography: Backeberg (1959), photographs 1109-1111: 1143-1144. Hoffmann (1989), plate 9a: 87.


Extended description: Plants up to 1.5 m tall, forming dense, low, shrubs. Stems slightly arched and cylindrical, 4 to 10 cm in diameter, the base procumbent. Epidermis opaque and finely velvety. Ribs at apex 8–12, 1 cm wide, 1 cm high, obtuse, with acute grooves; areoles obovate, 4–6 mm apart, the young 4 mm high, 1.0–1.4 cm wide, with hairs light brown on the upper half and dark brown or reddish orange on the bottom half. Spines mostly straight, some slightly undulating; centrals 4, up to 8 cm long, 1.5 mm in diameter, acicular, arranged crosswise, some growing upward, black with a yel-low base or dark gray with black spots, some cov-ered by lichens (inhabiting the damper areas); radials

9–16, acicular, 0.6−3 cm long, five of them yellow-ish brown, the others gray with black apex. Ribs at the base 10–15, 1 cm wide, 0.7 to 1 cm high; areoles2–5 mm high, 0.5−1.4 cm wide, with brown, gray or black pilosity; central spines 4–6, 2–10 cm long (two are the longest) 2 mm in diameter, white or olive green with brown spots, acicular, growing hori-zontally; radials 14–20, 1.1 cm long, acicular, brown to greenish brown. Flowers generally apical, 1 or 2, 8−13 cm long, with abundant brown hairs on the

ovary and floral tube; perianth 6 cm in diameter; ovary 2.0–2.5 cm wide; floral scales at middle of floral tube 2 cm long with brown edges and green center; floral scales at upper (distal) portion of floral tube 4.5 cm long, with apex red to pink with yellow edges and green center; tepals 4.3–5.3 cm long, 1.3−2.0 cm wide, white to yellow with pink longitudinal lines; style green, 5 cm long, 3 mm wide; stigma light yel-low, up to 2 cm long and with over 10 lobes. Fruitsmostly apical 4–6 cm long, covered with subulate scales, with abundant brownish axillary hairs 4−6 mm long. Seeds 1 mm long.

Distribution and habitat: Trichocereus deserti-cola grows in Regions II, III and IV (Kiesling et al.

Cereus deserticolus Werd. nov. spec. – Caulis erectus, simplex vel prope basim parce ramosus, altitudine 1–1,5 m, 7–10

cm diam., apicem versus angustatus; costae 8–10, ad 1,5-2,5 cm altae, supra areolas valde incisae; sinus acutí; areolae ad 1,2-

1,5 cm approxi-matae, orbiculares vel ellipticae, ca. 1 cm diam., tomento densuspullo dein griseo obtectae; aculei numerosi,

15-25, juventutedide pulli dein grisei, inaequales, peripherici aciculares vel subulata,1–1,5 cm longi, centrales ca. 1–3, ad 12

cm longi, rectiarcuati; flores ex areolis junioribus erumpentes, infundibuliformes,7–8 cm longi; tubus atque ovarium squamis

lanceolatis, acuminatis, sursum gradatim ad 1 cm accrescentibus lanaque pulla ex squamorum axillis oriunda onsite; perigonii

phylla exterior lanceolata, ad 3,5 cm longa, interior oblongo–lanceolata, ca. 4 cm longa,ca. 1,2–1,3 cm lata, basi attenuata,

albida; stamina numerosa in-sequalia in tubi pariete ascendentia, quam petala multa breviora; stylus staminibus longioribus

subaequilongus, ca. 4 cm longus; stigmata ca. 16 ad 1 cm longa; fructus ignotus.

Chile: Prov. Antofagasta, Dept. Taltal, bei Taltal, in ca. 500 m

Höhe (blühend im Oktober 1925 – Werdermann n. 869).

Die Pflanzen wuchsen in einer vom Küstennebel reichlichbefeuchteten Schlucht oberhalb der Hafenstadt Taltal auf

felsigemBoden oder zwischen grobem Gesteinsschutt. Am Grunde nieder? egend und häufiger verzweigt waren die Säu-

len gegen die Spitzem ca. 1 m bis 1,5 m lang aufgerichtet und 7–10 cm dick. DiePlanzen standen einzeln oder zu klein-

en, aber dichten Heckenmsammengeschlossen. Nur wenige Exemplare hatten schon ihretrichterförmigen, reinweißen

und relative kurzen Blüten entwickelt.

Die neue Art ist zweifellos nahe verwandt mit Cereus nigripilis Phil., welchen Britton et Rose, Cactaceae II, p. 139,

wieder als Synonym zu Cereus (Trichocereus) coquimbanus (Mol.) K. Sch. stellen. Sieweicht jedoch von diesem durch die

geringere Zahl der durch scharfeFurchen voneinander getrennten Rippen, die bis auf die langen und Stark entwickelten

1–3 Mittelstacheln viel schwächere Bestachelungund die relativ kurzen Blüten ab.

Figure 7. Trichocereus deserticola.


2008), in the coastal mountain range on rock bodies or among large stone rubble, with slopes that exceed 20 degrees inclination, between 250 and 600 m al-titude. Accompanying species include: Heliotropium stenophyllum (Boraginaceaae), Encelia canescens (As-teraceae), Adesmia argentea (Fabaceae), among others (Hoffmann 1989; Gajardo 1995).

Taxonomic and nomenclatural comments: After comparing the original descriptions to herbarium material of Cereus deserticolus, Trichocereus fulvila-nus and Trichocereus serenanus, it is proposed they be grouped as synonyms of C. deserticolus, because of their great similarity in appearance (small shrubs 1.5 m high), stem thickness (less than 10 cm), num-ber of ribs (fewer than 13), number of central spines (ca. 4) and spine shape (acicular, straight, and some slightly undulating).

The holotype of Cereus deserticolus (Werdermann 869) deposited in the Berlin herbarium, was de-stroyed in 1943 in World War II. Fortunately, the isotypes, which were kept separately from the gen-eral herbarium, were preserved. Unfortunately, some were severely damaged by water, as mentioned by Werdermann (1944), in his list of duplicates, refer-encing series No. 837 to 875, which were replaced by another set of duplicates, including C. deserticola,and Werdermann 869, which were distributed to other herbaria. Leuenberger and Eggli (1996) evalu-ated the Werdermann 869 specimen duplicates and concluded that they were a mix, namely that the flowers were from a Trichocereus species, but that the stem fragments presented the anatomical characters of Eulychnia.

According to Leuenberger and Eggli (1996), in the original description of C. deserticolus the color of the spines (dirty to blackish gray) and the color of the hairs of the areole (blackish to gray), correspond to Eulychnia iquiquensis, and not to C. deserticolus.We do not agree with that assessment, because at the Taltal commune we find individuals of C. deserticoluswith central spines at the apex black or dark brown, with clear green base, but the ones on the central part of the stem are gray, and the hairs are also gray, as mentioned in the original description of C. deser-

ticolus.Herbarium material studied: Chile, II Región

of Antofagasta, Prov. Antofagasta, Mpio. Taltal, 6 km E of (above) Paposo on road connecting to the Panamericana, 25°0.97'S; 70°26.21'W, 550-600 m, 15 Dic. 1994, U. Eggli and B. Leuenberger 2653 (SGO); 21 km N of Taltal towards Paposo, c. 1 km inland into Qda. Hueso Parado, somewhat to the N of Punta Agua Dulce, 25°14.1'S; 70°25.4'W, 220 m, 17 Dic. 1994, U. Eggli and B. Leuenberger 2664 (SGO); Cerro Perales, immediately E of Taltal, 25°25.5'S; 70°25.24'W, 600-800 m, 10 Oct. 1997, U. Eggli 2915 (SGO); Qda. Huanillos, Comuna Tal-tal, 25°57'35"S; 70°38'39"W, 256 m, 7 Nov. 2008, S. Albesiano et al. 2055 (MERL, AGUCH); Parque Nacional Pan de Azúcar, Comuna Taltal, Las Lomi-tas, 25°59'0.5"S; 70°36'43"W, 693 m, 8 Nov. 2008, S. Albesiano et al. 2058 (MERL, AGUCH). III Región de Atacama, Prov. Chañaral, Falda Verde, 26°17.41´S; 70°38.27’W, 400-450 m, 5 Oct. 1997, U. Eggli 2888 (SGO); Qda. León, 26°56'01"S, 70°44'30"W, 329 m, 9 Nov. 2008, S. Albesiano et al. 2060 (MERL, AGUCH); Prov. Copiapó, Co-muna Copiapó, Qda. Matamoros, 27°53'48"S; 71°05'10"W, 193 m, 10 Nov. 2008, S. Albesiano et al. 2061 (MERL, AGUCH); Prov. Huasco, Mun. Huasco, 11 km N of Huasco Bajo, 28°23.38'S; 71°11.18'W, 20-70 m, 20 Oct. 1997, U. Eggli and B.E. Leuenberger, 2999 (SGO); Comuna Vallenar, 28°94'41"S; 68°72'11"W, 88 m, 11 Oct. 2010, S. Albesiano et al. 2089 (MERL).

10. Trichocereus faundezii S. Albesiano sp. nov. TYPE: Chile, IV Región de Coquimbo, Prov. Choapa, 31°40'25"S; 71°17'11"W, 197 m, 5 Nov. 2008, S. Albesiano, L. Faúndez, R. Flores and P. Saldivia 2048. (Holotype: MERL. Isotype: AGUCH 64088). (Figs. 8 & 9).

Plantae caespitosae 80 cm altae. Rami decumben-tes, aliqui arcuti, 14 cm diametri, viridulae. Costae 14−18, tuberculatae, obtusae, 2 cm latae et 3 mm altae. Areoles oblongae, 1 mm altae, 8 mm latae, ci-nereae. Spinae rigidae, aciculae et cinereae, centrales

Figures 8 & 9. Trichocereus faundezii.


1−3, diametri 1 mm, 3.5 cm longae; radiales 9−12, longae 2 cm, diametri 0.5 mm, divaricatae. Flores 2−6, subapicales, 11 cm longae, 9 cm diametri, ovar-ius 2 cm diametri, squamae florales cum copiosis pilis brunneis, tepali albi cum apice roseo.

Morphological description: Plants cespitose, 80 cm tall. Branches decumbent, some arched, 14 cm diameter, grayish green. Ribs at the apex 18, tuber-culate, obtuse, 2 cm wide, 3 mm high; with obovate areoles, 1 mm high and 8 mm wide, gray. Spines acicular and gray; central spines 1–3, 1 mm diam-eter, 2.5 cm long; radials 9, 2 cm long and 0.5 mm in diameter. Ribs at the base 14, tuberculate, obtuse, 2.5 cm wide and 3 mm high; with obovate areoles 1 mm high and 7 mm wide and gray; spines gray and acicular, central 3, of 3.5 cm long, radials 12 and 1.5 cm length. Flowers 2–6, subapical, 11 cm long, 9 cm in diameter, ovary 2 cm diameter; floral scales of the base 6 mm long, upper 2 cm long, green base, brownish pink apex and axils where abundant dark brown hairs originate; tepals 5 cm long; deep pink in flower buds later becoming white with a brownish pink apex as adults.

Etymology: Species dedicated to Luis Faúndez, the Chilean botanist, enthusiast, who kindly accom-panied us and guided the explorations in the field in arid and semi–arid Chile.

Taxonomic and nomenclatural comments: The habit of this species (cespitose plants, less than 1.5 m) is similar to that of T. spinibarbis, but their differ-ences are set forth in Table 1.

11. Trichocereus macrogonus (Salm-Dyck) Riccob., Boll. Reale Orto Bot. Palermo 8: 236. 1909.

Cereus macrogonus Salm-Dyck, Cact. Hort. Dyck.:203. 1849 [1850]. Echinopsis macrogona (Salm-Dyck) H. Friedrich & G.D. Rowley, I.O.S. Bull. 3(3): 96. 1974. TYPE: Peru, Dpto. Junin, Prov. Tarma, be-tween Acobamba and Palca, 2890 m, 1 Oct. 1982, D. Smith 2456. (Neotype: SI!, designated by Albe-siano & Kiesling: 32. 2012; isoneotype: MO!)

Iconography: Backeberg (1959), photograph 1075: 1120. Ritter (1980a), photographs 448−449: 764. Hunt et al. (2006), photograph: 235.3: 235.


Extended description: Stems erect, 2.5−5 m tall. Branches 6−15(−20) cm diameter. Ribs at the base 7−8, obtuse, 2.5 cm wide, 2 cm tall. Areolescircular, 6 mm diameter, all grey or dark brown; a transverse dermal furrow present just above each ar-eole. Spines acicular; centrals 1−3, ca. 3.3 cm long, brown with black points; radial spines ca. 8, 2−8 cm long. Flowers mostly near the stem apex, rarely at the apex, funnelform, 21 cm long, 2 cm in diameter at the level of the ovary; the latter densely covered with clear brown hairs; scales on the ovary 1 cm long, green with brown apex, scales on the tube 2.5 cm long, green or yellowish green with brown apex; te-pals 9 cm long, mostly yellowish green or pale yellow with brown longitudinal lines, rarely deep pink; stylegreen, 8 cm long; stigma yellow, 1 cm long. Seedsbroadly ovoid (ratio of length to width = 1.3 to 1), small (0.9−1.1 mm long), dull, without keel, anticli-nal walls of the testa straight.

Distribution and habitat: Frequently cultivated in Bolivia, Colombia, Chile, Ecuador and Peru, as an ornamental, for fences, and for medicinal or magi-cal purposes. Its most plausible wild origin is the medium to high valleys of the Peruvian Andes, at al-titudes of 2000−3000 m (Fernández and Estupiñan 2005; Albesiano and Kiesling 2012).

Taxonomic and nomenclatural comments: This species is characterized by the stems with ribs that are few (up to 10), high (2 cm) and wide (2.5 cm), with widely set areoles (2.5 cm), and the presence of a transverse dermal furrow just above each areole. It also presents a high infraspecific variation in the number and shape of the spines (Albesiano and Kies-ling 2012).

11a. Trichocereus macrogonus (Salm-Dyck) Riccob. var. macrogonus.

Trichocereus peruvianus Britton & Rose, The Cac-taceae 2: 136. 1920. Echinopsis peruviana (Britton & Rose) H. Friedrich & G.D. Rowley, IOS Bull. 3 (3): 97. 1974. Trichocereus pachanoi f. peruvianus (Britton & Rose) F. Ritter, Kakteen Südamerika 4: 1324. 1981. TYPE: Peru, Dept. Matucana, vicinity of Matucana, 9 Jul. 1914, J.N. Rose & Mrs. J.M. Rose 18658

T. faundezii T. spinibarbis

Branch diameter

14 cm 10 cm

Number of ribs at the apex

18 12

Tuberculate ribs

Present Absent

Width of ribs 2 cm 1 cm

Length of central spines

2.5 cm 10 cm

Table 1. Differences between T. faundezii and T. spinibarbis in terms of morphological characters.


(Neotype, US761324! Isotype, NY 386193! desig-nated by Albesiano & Kiesling: 32. 2012)

Cereus rosei Werd., in Backeberg, Neue Kakteen73: 101. 1931. As a new name for T. peruvianus Britton & Rose. TIPO: Bolivia, Dpto. La Paz, Prov. Larecaja, Collabamba, 5 Nov. 2002, R. Kiesling et al. 10041. (Neotype: LPB, designated here.)

Trichocereus puquiensis Rauh & Backeb. in Backeb. Descr. Cact. Nov. 20. 1956. Echinopsis puquien-sis (Rauh & Backeb.) H. Friedrich & G.D. Rowley,I.O.S. Bull. 3(3): 97. 1974. Echinopsis peruviana(Britton & Rose) H. Friedrich & G.D. Rowley subsp. puquiensis (Rauh & Backeb.) Ostolaza, Cactaceae Consensus Init. 6: 8. 1998. TYPE: Bolivia, Prov. B. Saavedra, Puente Camota, highway to Charazani, 15°13'39"S; 68°45'36"W, 1433 m, 12 Sep. 2004, L. Cayola et al. 1533. (Neotype: LPB, designated here.)

Trichocereus tacnaensis F. Ritter, Kakteen Südameri-ka 4: 1326. 1981. TYPE: Perú nordöstlich von Tacna, 3000-3500 m, 1959, F. Ritter 994. (Holotype: U!.)

Iconography: Backeberg (1959), photograph 1075: 1120. Anderson (2005): 235, 240. Hunt et al. (2006), photograph 236.1: 236.

Extended description: Plants erect (sometimes decumbent), 5 m high. Stems bluish green or light green. Branches 16−20 cm in diameter, erect and strong. Ribs at the apex 7, obtuse, 2.5 cm wide and 3 cm high, with circular areoles, spaced 2 cm apart, 1–3 mm high, 6 mm wide, gray or dark brown; cen-tral spines (in apical areoles) 1–3, acicular, 3–10 cm long, brown with black apex; radial spines 7–10, acicular, 1–2 cm long, stiff, brown with black tip. Ribs at the base 8–10, obtuse, 1.2 cm wide, 1.3 cm high; with circular areoles 1–3 mm high, 6–12 mm wide, spaced 2–3 cm apart, gray with black or dark brown hairs; central spines (in basal areoles) 3–4, acicular, straight, 5 cm long, 1 mm diameter, black apex, brown middle, and gray base; radial spines 7–20, 1–3 cm long, acicular, rigid, brown with black apex. Flowers apical and subapical, 21 cm long, with brown hairs on the ovary and floral tube; perianth 7cm in diameter; ovary 2 cm wide, floral ovary scales 1–3 cm long, green with brown apex; floral tube scales 2.5 cm long, green or yellowish green with brown apex; tepals 6–9 cm long, yellowish green or yellowish white with brown longitudinal lines; style green, 10 cm long; stigma yellow, 1–2 cm long. Fruits subapical, dark green, 2–4 cm long, 3–5 cm in diameter, covered with subulate scales, from whose axils emerge brown and gray hairs. Seeds black, 2 mm long and 1 mm wide.

Distribution and habitat: Native of Peru (Borg 1937), in the valleys of the department of Lima, at 2500 m, in the valley of Puquio, at 3300 m, south-east of Tacna at 3000–3500 m, and northeast of Es-tique (Ritter 1981). It is also distributed in Bolivia (1500 m), on nearly bare rocky hillsides.

Taxonomic and nomenclatural comments: In 1920 Britton and Rose described Trichocereus peru-vianus, as a species resembling T. macrogonus by the number (7–10) and characteristics of the ribs (2 cm high and 2.5 cm wide), the color of its stems (green–blue or light green), by the presence of a groove above each areole, and by the distance between the areoles (2.5 cm), and so it is considered synonymous with the type variety.

Werdermann, considering this taxon under Ce-reus, had been obligated to rename it as Cereus rosei,to avoid the homonymy with Cereus peruvianus (L.) Miller (Borg 1937).

In the original description of Trichocereus pu-quiensis, we observe that the morphological char-acters are coincident with those of the type variety (light green branches of 15 cm diameter, ribs 8–10, the presence of a dermal furrow just above each ar-eole, and differentiation between central and radial spines). T. puquiensis is therefore proposed as a syn-onym of T. macrogonus var. macrogonus.

Trichocereus tacnaensis also is considered a syn-onym of this variety, in spite of the 12–15 cm diam-eter of its branches, which may be due to high levels of environmental moisture.

Herbarium material studied: Bolivia, Dpto. La Paz, Prov. Larecaja, Collabamba, 5 Nov. 2002, R. Kiesling et al. 10041 (LPB); Prov. B. Saavedra, Pu-ente Camota, highway to Charazani, 15°13'39"S; 68°45'36"W, 1433 m, L. Cayola et al. 1533, 1534 (LPB). Perú, Dpto. Tacna, above Tacna, 74 km Weg Charaña-Tacna, ca. 3500 m, F. Ritter 994 (121188 SGO).

11b. Trichocereus macrogonus (Salm-Dyck) Riccob. var. pachanoi (Britton & Rose) S. Al-besiano & R. Kiesling, Haseltonia 17: 32, ill. 2012. (Figs. 10 & 11).

Trichocereus pachanoi Britton & Rose, The Cac-taceae 2: 134, ill. 1920. Cereus pachanoi (Britton & Rose) Werderm., in Backeb, Neue Kakteen: 73. 1931. Echinopsis pachanoi (Britton & Rose) H. Friedrich & G.D. Rowley, IOS Bull. 3 (3): 96. 1974. TYPE: Ecuador, Cuenca, 17 to 24 Sep. 1918, J.N. Rose, A. Pachano & G. Rose 22806. (Holotype: US, lost.) (Lectotype: NY 386191!, designated by Albesiano &

Key to the varieties

1. Spines of the older areoles 18−20, 3−4 of them prominent, longer, stronger, and more robust (ca. 5 cm long, 1 mm in diameter). Plants up to 5 m tall, branches erect or ascending; stems stouter, 16−20 cm in diameter. ..........................................T. macrogonus var. macrogonus

1'. Spines of the older areoles 3−7, ca. 0.5−2 cm long, and less than 1 mm in diameter. Plants up to 2−4 m tall; stems erect, becoming parallelstems more slender, 6−11(−15) cm in diameter. ................................T. macrogonus var. pachanoi


Kiesling 17: 32. 2012.)Trichocereus santaensis Rauh & Backb., in Backeb.

Descr. Cact. Nov. 20. 1957 (1956). Echinopsis san-taensis (Rauh & Backb.) H. Friedrich & G.D. Row-ley, I.O.S. Bull. 3(3): 97. 1974. (Lectotype, designat-ed here: Backeberg. Die Cactaceae 2: 1113. 1959.)

Trichocereus torataensis F. Ritter, Kakteen Sü-damerika 4: 1325, ill. 1981. TYPE: Perú, Torata, 3000 m, 1953, F. Ritter 1467. (Holotype: U!.)

Trichocereus huanucoensis Johnson, nom. nud.Iconography: Backeberg (1959), photographs

1072–1074: 1119-1120. Madsen (1989), plate 7: 29. Ritter (1981), photographs 1186, 1193: 1550. Reyna and Flores (2001), photographs 1–2: 28-29. Anderson (2005): 238. Hunt et al. (2006), photo-graph 235.4: 235.


Extended description: Plants 2–4 m high, branched from the base. Stems erect, straight, ar-

ranged in parallel, cylindrical, dark green, light green when young, 6–11(–15) cm in diameter. Ribs obtuse, generally 6–8(–13), 2.5 cm high, with areoles ob-ovate, round, 3−6 mm wide and spaced 1.5−2.7 cm apart, hairs yellow in the apical (superior) part of the areole, and light gray in the basal (inferior) part of the areole. One central spine; up to 6 radials, acicu-lar, brown, aimed in various directions, up to 2 cm long, and less than 1 mm wide. Flowers 1–4, api-cal and subapical, 24 cm long, with black curly hairs 2 cm long on the ovary and floral tube; perianth 20 cm in diameter; floral ovary scales subulate, ca. 2–5 mm long, dark green; floral tube scales subulate, 20 mm long, with brown tips, green in the middle and yellow at the base; ovary 3 cm in diameter; tepalsspatulate, 11 cm long, white; style white with green base, 14–19 cm long; stigma light yellow, up to 3.5 cm long. Fruits green, 3 cm in diameter, 6 cm long, covered by subulate scales, from whose axils emerge

black hairs. Seeds black, 2 mm high and 1.3 mm wide

Figure 10. Trichocereus macrogonus var. pachanoi (photo by Daniel Schweich.

Figure 11. Trichocereus macrogonus var. pachanoi (Quito, Ecuador, photo by H. Smidlock).


Distribution and habitat: This is the variety of Trichocereus with the northernmost distribution. It is found in Bolivia, Chile, Ecuador, and Peru, on the cliffs along the rivers, at altitudes up to 3000 m. It is also grown as an ornamental plant in gardens and parks, and as living fences (Madsen 1989). It is a possible native of southern Ecuador and northern Peru, where it is known as San Pedro (Borg 1937).

Taxonomic and nomenclatural comments: US type herbarium sheet specimen is not found (they are not on the list of Herbarium types, and a let-ter with curator, Rusty Russell on September 2008, confirms that the sample has not been found since 1941); therefore, the NY sheet has been designated as a lectotype.

It is befitting to designate the variety T. macrogo-nus var. pachanoi because of its proneness to having diminutive spines (less than 1 cm in length, often a small fraction of that length), even if this characteris-tic exhibits phenotypic plasticity reflecting the influ-ence of different environmental conditions.

Trichocereus torataensis is regarded a synonym of the variety pachanoi by the stem diameter (usually 7 cm), the distance between the areoles (up to 2.5 cm), and by having few (less than 7) radial spines and a single central spine. Anderson (2005) and Hunt et al. (2006) consider T. torataensis a synonym of E. pe-ruviana, and a synonym of Trichocereus macrogonus var. macrogonus, but records of the latter show an increased number of spines (20), 3–4 of which are longer and more robust (5 cm long, 1 mm diameter).

There are Trichocereus huanucoensis Johnson (2287797–2287798MO!, 00386189NY!) specimens marked as “isotypes” in the herbarium of the Mis-souri Botanical Garden (MO) and at the New York Botanical Garden (NY), but this species has no de-scription or diagnosis, making its name a nomen nudum. Here it is considered a synonym of T. mac-rogonus var. pachanoi on account of the similarity in the number of ribs (6), distance between the areoles (2–2.5 cm), and low number of spines (5), which are comprised of one central and 4 radials.

Herbarium material studied: Perú, Dpto. Mo-quegua, Cuajones, ca. 3000 m, F. Ritter 1467 (121146 SGO).

12. Trichocereus nigripilis (Phil.) Backeb., Cactus Journ., Great Brit. 5: 49. 1937. (Fig. 12).

Cereus nigripilis Phil., Fl. Atacam. 23 (146): 197. 1860. Trichocereus chiloensis (Colla) Britton & Rose var. nigripilis (Phil.) Espinosa, Revista Chilena Hist. Nat. 41: 292. 1937. Trichocereus coquimbanus sensu Britton & Rose var. nigripilis (Phil.) Borg, Cacti:181. 1951. Echinopsis nigripilis (Phil.) H. Friedrich & G.D. Rowley, I.O.S. Bull. 3 (3): 96. 1974. TYPE: Chile, Coquimbo, Nov. 1853, R. A. Philippi 35. (Lectotype: SGO 52682! designated by Muñoz: 92. 1960.)

Iconography: Backeberg (1959), photograph 1111: 1144.


Extended description: Plants cespitose, up to 1.2 m high, forming dense, low, shrubs. Stems slightly arched, cylindrical, 5–7 cm in diameter. Ribs at the apex 11, obtuse, 1 cm wide, and 1 cm high, the grooves undulating; areoles round, 3 mm high, 7 mm wide, brown in the apical (superior) part of the ar-eole, and gray in the subapical (inferior) part of the areole. Spines straight; centrals 4, acicular, 2 mm in diameter, 5 cm long, both apical and subapical cen-trals black; radials 13, acicular, 0.5 mm in diameter, 0.7−2.0 cm long; radials of areoles in both apical and subapical regions of branches are black. Ribs at the base 10, obtuse, 1 cm wide, 0.8 mm high; with

Figure 12. Trichocereus nigripilis.


areoles 5 mm high, 8 mm wide, round,with dark gray pilosity; central spines 4, 6 cm long, acicular, 1.5 mm in diameter, light gray with black tips; radials12, 1 cm long, acicular, 5 mm in diameter, light gray. Flowers 13 cm long; ovary and floral tube covered with black pilosity; stigma erect, 1–2 cm long. Fruitsapical, 3.5 cm long. Seeds 1.5 mm long, 1 mm wide, with a dark yellow hilum–micropyle region.

Distribution and habitat: Species distributed in Region III of Atacama and Region IV of Coquimbo, from sea level up to altitude of 700 m.

Taxonomic and nomenclatural comments: The following annotation by U. Eggli and B. Leuenberg-er appears in the sheet of the T. nigripilis type speci-men: “flower is of Trichocereus, vegetative parts of Eulychnia.” But on reviewing the vegetative charac-ters of the areoles (dark brown) and spines — shape (bristly, acicular and rigid), number (17), length (2 to 5 cm long) and color (gray) — we found that these characters agree with the original description of T. nigripilis; thus, vegetative parts do not correspond to Eulychnia but to T. nigripilis.

Herbarium material studied: Chile, III Región de Atacama, Prov. Huasco, Comuna Vallenar, 10 km al sur de Vallenar, 28o39'48"S; 70o45'49"W, 614 m, 6 Nov. 2008, S. Albesiano et al. 2054 (MERL, AGUCH); Qda. León, 26o56'01"S; 70o44'30"W, 330 m, 9 Nov. 2008, S. Albesiano et al. 2059 (MERL, AGUCH). IV Región de Coquimbo, al N de la Serena, 27.5 km al N de Ruta 5, al N de Juan Soldado, 29°49'S; 71°16'W, 11 Oct. 2010, S. Albe-siano et al. 2083 (MERL).

13. Trichocereus pectiniferus S. Albesianosp. nov. TYPE: Chile, IV Región de Coquimbo, Juan Soldado, 29°43'26"S; 71°18'45"W, 166 m, 6 Nov. 2008, S. Albesiano, L. Faúndez, R. Flores and P. Saldivia 2053. (Holotype: MERL. Isotype: AGUCH 64080.) (Fig. 13)

Trichocereus coquimbanus sensu Britton & Rose, The Cactaceae 2: 139. 1920, non Cactus coquimba-nus Molina, Sag. Stor. Nat. Chili: 170. 1782, non Cereus coquimbanus sensu K.Schum., Monatsschr. Kakteenk. 6: 166. 1896, non Echinopsis coquimbanasensu Friedrich & G.D. Rowley, I.O.S. Bull. 3 (3): 94. 1974.

Plantae caespitosae altae 80 cm. Rami decum-bentes, aliqui arcuti, 5 cm diametri. Costae 11, obtusae, 2 cm latae. Areoles ovobadae rotundae-que, 3 mm altae, 1.2 cm latae, cinereae. Spinae centricae 4, in cruce, aciculae subulataeque, 2 mm latae, 1.5–6 cm longae, cinereae cum adul-tae sunt. Spinae radialis 15, aciculae, 1.3–2.2 cm longae, cinereae, adultae crescunt adpresas et in pectinis forma. Fructus apices et subapices, virides, 3 cm longi et 4.5 cm lati et cum squamis pilosis, quarum axillarum emergunt copiosos pilos brun-neos cinereosque, pulpam albam, semina nigra.

Morphological description: Plants cespitose, 80 cm tall. Branches up to 15 in number, decumbent,

some arched, 5 cm diameter. Ribs at the apex 11, ob-tuse, 2 cm wide; with obovate areoles 2 mm high, 1.2 cm wide, yellow apex and gray base. Central spines4, 1.5−6 cm long, acicular, 2 mm in diameter, some with dark brown apex, others green at the base and brown at the tip, others light gray with brown apex, some arched; radials 15, most acicular, few setose (one per areole), 2 cm long, light brown. Ribs at the base 10, obtuse, 2 cm wide and 7 mm high; with round areoles 3 mm high and 7 mm in diameter, gray. Central spines 4, acicular, 5 cm long, light gray, ar-ranged crosswise and divaricate (ca. 75°) to the stem, intersecting with the central spines of neighboring areoles; radials in adult areoles 12, acicular, 1.3 cm long, gray, pectinate, appressed to stem (ca. 15°). Flowers: Floral remnants short, with abundant black wool. Fruits apical or subapical, 3 cm long and 4.5 cm in diameter, covered with subulate scales, 3 mm long and abundant, brown, and gray, hairs emerging from axils. Seeds with black testa.

Iconography: Hoffmann (1989), plate 9b: 87. Etymology: Species characterized by the distribu-

tion of radial spines in adult areoles, where the radi-als are appressed to the surface of the stem and pecti-nate (organized like the teeth of a comb).

Distribution and habitat: It grows in Region IV of Coquimbo, at altitudes from sea level to 500 m, associated with Eulychnia sp., Copiapoa sp. and Opuntia sp.

Taxonomic and nomenclatural comments: Brit-ton and Rose combined T. coquimbanus (Molina) Britton & Rose, unaware that the original descrip-tion of Cactus coquimbanus Molina (170. 1782) cor-responds to a species of Eulychnia. Among the veg-etative traits, Molina highlighted growth habit (erect

Figure 13. Trichocereus pectiniferus.


and high) and sizeable spines 20 cm long (“which were once used to sew socks”), while the plant de-scribed by Britton and Rose presents cespitose habit and spines that do not exceed 8 cm in length. For that reason, we propose this entity as a new species of Trichocereus distributed in the Province of Coqui-mbo.

It differs from T. glaucus and T. spinibarbis by the position of external spines in adult areoles. In T. faundezii spines are appressed to the stem (diver-gence angle of ca. 15 degrees) and pectinate, while in T. glaucus and T. spinibarbis they are divaricate (di-vergence angle of ca. 75 degrees), and radial.

Paratypes: IV Región de Coquimbo, Prov. Elqui, Mpio. La Serena, 1 km E of El Peñón, 30°8.68'S; 71°12.93'W, 160–250 m, 4 Dic. 1994. U. Eggli and B.E. Leuenberger 2574 (SGO); Ruta 5, km 514, Comuna Higuera, cuesta de Buenos Aires, 29°31'S; 71°12'W, 270 m, 10 Oct. 2010, S. Albe-siano et al. 2084 (MERL); camino a los Choros, 29°13'S;71°32'W, 270 m, S. Albesiano et al. 2085 (MERL); entrada a los Choros, 29°13'S; 71°32'W, 90 m, 10 Oct. 2010, S. Albesiano et al. 2086 (MERL).

14. Trichocereus skottsbergii Backeb., in Skottsb., Acta Horti Gothob. 18: 146, ill. 1950. (Fig. 14).

Echinopsis skottsbergii (Backeb.) H. Friedrich & G.D. Rowley, I.O.S. Bull. 3 (3): 97. 1974. Echinop-sis chiloensis (Colla) H. Friedrich & G.D. Rowley subsp. skottsbergii (Backeb. ex Skottsb.) G.J. Charles. Cactaceae Syst. Init. 19: 13. 2005. TYPE: Skottsberg 834. (Lectotype: GB, lost, designated by Charles: 13. 2005.)

Trichocereus skottsbergii Backeb. ex Skottsb. var. breviatus Backeb., Descr. Cact. Nov.: 20. 1956. Echi-nopsis skottsbergii (Backeb.) H. Friedrich & G.D. Rowley var. breviata (Backeb.) H. Friedrich & G.D. Rowley. I.O.S. Bull. 3 (3): 98. 1974. TYPE: Chile, IV Región de Coquimbo, Dpto. Ovalle, Fray Jorge, 200 m, Nov. 1925, E. Werdermann 885. (Neotype: SI, designated here.)

Iconography: Backeberg (1959), photographs 1105-1108: 1141-1142. Ritter (1980b), photograph 1070: 1223.


Extended description: Plants erect, mostly about 2 m high but can exceed 3 m (Luis Faundez, pers. comm.). Stems: Lateral branches originating above the base, grayish green, 10–15 cm in diameter with abundant acicular apical spines less than 0.5 mm in diameter, light yellow. Ribs at the apex 16, obtuse, 1.8 cm wide, 5 mm high, with obovate areoles 1 mm high and 1 cm wide, yellow apex and gray base; cen-tral spines 3, acicular, 2–12 cm long, some flexible and weak (bristly), 5 mm diameter, some not sharp, yellow brown, smaller spines have green base and brown apex, the largest spines gray with some black spots and brown tip; radial spines 13−26 (both at the apex and at the base of the stem), 2–6 cm long, acic-ular, less than 1 mm in diameter, flexible and weak (bristly), somewhat pointed (but not sharply), with green base and brown apex. Ribs at the base 12–17, obtuse, 2 cm wide and 1 cm high, with obovate ar-eoles 1 mm high and 0.5–1 cm wide, completely dark gray to black; with 4 central spines, 7 cm long, 3 mm in diameter, acicular, green with brown apices; radialspines 13, to 2.5 cm long, acicular, dark green with brown apices, or totally white. Flowers 1–3, subapi-cal, 8–12 cm long; ovary 2.5 cm wide, covered with subulate scales to 3 mm long with abundant brown and black hairs; floral tube 5 cm in diameter, covered with scales, from whose axils arise substantial black and brown hairs, 6 mm long; outer tepals spatulate, 1.7 cm long, pink apex and green base, with a pink

midline; inner tepals spat-ulate, 3 cm long, white with pink apex; style yel-low, 8 cm long; stigmayellow, 7–13 mm in length. Fruits 3 cm long and 2 cm in diameter.

Distribution and habitat: In Coquimbo, in Fray Jorge National Park, located 30 km west of Ovalle, and in Talinay National Park, south west of Ovalle. Located on the slopes of the coastal range, where the dominant phys-

Figure 14. Trichocereus skottsbergii.


iognomic vegetational types are stumpy and spinose scrub, from sea level to 200 m.

Taxonomic and nomenclatural comments:Backeberg (1950) established that T. skottsbergii and T. deserticola are similar, but characters were found to differentiate them (Table 2).

Within a population, there are individuals that are set apart by the smaller size of their flowers and central spines, these are named the breviatus variety; however, these features are within the morphologi-cal species, and as a result, the variety must pass into the synonymy of the species, as proposed by Ritter (1980b).

Charles (2005) proposed T. skottsbergii as a sub-species of T. chiloensis, but when we studied the mor-phology of the apical radial spines, some differences were observed: In T. skottsbergii those spines are flex-ible, dull, and no longer than 6 cm, while in T. chi-loensis they are rigid, sharp and up to 4 cm in length. Accordingly, it is proposed T. skottsbergii be separated from T. chiloensis at the specific level. In addition, Charles did not realize that this species had been described in one of Scottsberg’s papers (vide supra), where the description was attributed to Backeberg, who must be considered the author of the species.

Claes Persson, curator of the GB herbarium (Sweden) indicates (in litt.) that the type specimen, Skottsberg 834, does not exist in that institution, as cited by Charles (2005).

Herbarium material studied: Chile, IV Región de Coquimbo, Dpto. Ovalle, Fray Jorge, 200 m, Nov. 1925, E. Werdermann 885 (SI); Prov. Elqui, Mpio. La Serena, Punta Teatinos, 29°49.29'S; 71°17.36'W, 10-100 m, 5 Dic. 1994, U. Eggli and B.E. Leuen-berger 2579 (SGO); Puerto Oscuro, al borde de la playa, 31°25'21"S; 71°35'24"W, 46 m, 6 Nov. 2008, S. Albesiano et al. 2050 (AGUCH, MERL).

15. Trichocereus spinibarbis (Otto ex Pfeiff.) F.

Ritter, Kakt. And Sukkulent. 16 (11): 212. 1965.

(Fig. 15).

Cereus spinibarbis Otto ex Pfeiff., Enum. Diagn. Cact.: 86. 1837. Echinocereus spinibarbis (Otto ex Pfeiff.) K. Schum., Monatsschr. Kakteenk. 5: 124. 1895. Echinopsis spinibarbis (Otto ex Pfeiff.) Hoffm.-

Grob. Cactaceas. 1989. TYPE: Chile, III Región de Atacama, cerca de Vallenar, 28°35'S; 70°46'W, 11 Oct. 2010, S. Albesiano et al. 2088. (Neotype: MERL, designated here.)


Extended description: Plants cespitose, up to 80 cm high, forming dense low shrubs. Stems slightly arched, and cylindrical, 6 cm in diameter. Ribs at the apex 9, obtuse, 1 cm wide, 0.8 mm high; with areolesround and obovate, 5 mm high, 1 cm wide, yellow in areoles at the apices of the branches, and gray in areoles below the apices of the branches. Spines: cen-trals 3–4 in areoles below the apices of the branches, arranged crosswise, 3–12 cm long, acicular, bristly, flexible, curved, 2 mm wide, gray with brown tips;radials 10–15, 1.5 cm long, acicular, 0.5 mm in di-ameter. Ribs at the base 9, obtuse, 1 cm wide, 1 cm high; with areoles 8 mm high, 1 cm wide, round, dark gray pilosity; central spines 4, 2–9 cm long, acic-ular, 1.5 mm in diameter, light gray with black tips; radials 22, acicular, 1 cm long, 0.5 mm in diameter,

T. deserticola T. skottsbergii

Branches emerging at

ground level50 to 80 cm

above the ground

Number of ribs at apex

8−12 16

Branch diameter

to 10 cm to 15 cm

Length of radial spines at

apexto 3 cm to 6 cm

Table 2. Differences between T. deserticola and T. skottsbergiiaccording to the morphological characters.

Figure 15. Trichocereus spinibarbis.


light gray with brown tip. Flowers 13 cm long; ovaryand floral tube covered with black pilosity; stigmaerect, 1–2 cm long. Fruits apical, 4 cm long. Seeds 1.5 mm long, 1 mm wide, with a dark yellow hilum-micropyle region.

Distribution and habitat: T. spinibarbis occurs in Region II of Antofagasta and Region III of Ata-cama, at an altitude of 200 m.

Taxonomic and nomenclatural comments:Salm-Dyck (44, 199. 1850), created an infrageneric classification for Cereus taxa, suggesting C. spinibar-bis should be placed in the subsection Velutini, based on its opaque and fine velvety epidermis, and the spacing between the areoles (4–6 mm). The observa-tion made by Salm-Dyck is deemed reliable, as Salm-Dyck was Pfeiffer’s student and was well acquainted with his plants. Labouret (334. 1853) mentioned the size of the spines, ca. 30 cm long. This had not been mentioned in the original description. Schumann (62. 1899) transferred C. spinibarbis to the synonym of Cereus chiloensis, without providing comments, although C. spinibarbis and C. chiloensis differ in general appearance (dense low shrubs usually less than 1.5 m vs. shrubs or arborescent plants greater than 1.5 m), branch diameter (4–10 cm vs. 10–25 cm), and fine velvety epidermis in C. spinibarbis, a feature absent in C. chiloensis. Subsequently, Brit-ton and Rose (1920) transferred Cereus spinibarbisto the genus Eulychnia [Eulychnia spinibarbis (Otto ex Pfeiff.) Britton & Rose, The Cactaceae 2: 82–83, f. 122] without providing details about this change. Additionally, the morphological characters described do not match the original description of Eulychnia breviflora Phil. Later Ritter (1980b) established that Cereus spinibarbis belongs to the genus Trichocer-eus, given that the velvety epidermis in Chilean cacti occurs only in this genus, and passed E. spinibarbisBritton & Rose (non Cereus spinibarbis Otto ex. Pfei-ff.) into the synonymy of Eulychnia breviflora Phil.

Herbarium material studied: Chile, II Región de Antofagasta, Prov. Antofagasta, Comuna Taltal, 25°26'S; 70°31'W, 11 Oct. 2010, S. Albesiano et al. 2091 (MERL). III Región de Atacama, 37 km de Punta de Choro, 210 m, 11 Oct. 2010, S. Albesiano et al. 2087 (MERL); Prov. Huasco, Comuna Vallenar, cerca de Vallenar, 28°35'S; 70°46'W, 11 Oct. 2010, S. Albesiano et al. 2088 (MERL).

16. Trichocereus undulosus S. Albesiano sp. nov.TYPE: Chile, IV Región de Coquimbo, Provincia

Limarí, Peña Blanca, 30°48'32"S; 71°34'29"W,

359 m, 6 Nov. 2008, S. Albesiano, L. Faúndez, R.

Flores and P. Saldivia 2051. (Holotype: MERL.

Isotype: AGUCH). (Fig 16).

Planta arborea, 2.5 m altae, ramificatae ex trunci 11 cm diametri. Ramae atrovirentes, erec-tae, 15 cm diametri. Costae 20, obtusae, 2 cm latae et 1 cm altae. Costae versantur separatae per sulcos acutos et ondulatos. Forma ondulata debe-tur quem costae sunt crassiores altitudinem areoles

et qui areoles costarum vicinarum alternantur. Spinae rigidae, aciculae et atrovirentes; centrales 4, diametri 2 mm et 3.5 cm longae; radiales 12, diametri 0.5-1 mm, 1.5 cm longae. Flores sub-apicalis, infundibuliformes, 13 cm longae, ovarius 2.5 cm diametri et tubus floral cum copiosis pilis (pelos) cinereis, tepali albi cum apice brunneo. Fructus virides, 3 cm latae et 2.5 longae, cum co-piosis pilis cinereis. Semina nigra.

Morphological description: Plants 2.5 m high, stem a trunk 11 cm in diameter. Branches dark green, 15 cm in diameter. Ribs at the apex 20, obtuse and tuberculate; with obovate areoles 2 mm high, 8 mm in diameter. Central spines 3, 4 cm long, white with brown markings, 1 mm in diameter in the areoles at the apex of the branches, and 2 mm in diameter in the more basal areoles. Radial spines 12, acicular, 1 cm long, 8–10 mm wide, white with brown spots. Ribs at the base 20, obtuse, tuberculate, 2 cm thick, 1 cm high; with obovate areoles 2 mm high, 8 mm in diameter, entirely gray. Central spines 4, 2−4 cm long, 2 mm wide, acicular, dark green. Radial spines 12, 1.5 cm long, 0.5–1 mm in diameter, acicular, dark green. Flowers 3–10, subapical, 13 cm long, 6 cm in diameter; ovary 2.5 cm wide, middle floral scales 2 cm long, green with brown midstripe, with abun-dant axillary gray hairs; tepals 6 cm long, white with a brown midline; style 8 cm long; stigma lobes 1 cm

Figure 16. Trichocereus undulosus.


long. Fruits green, 3 cm diameter, 2.5 cm long, cov-ered with subulate scales, with abundant axillary gray hairs, green exocarp, and white mesocarp and endo-carp. Seeds black.

Etymology: The epithet refers to the shape of the rib: markedly undulating, giving a tuberculate ap-pearance.

Distribution and habitat: This species grows in the Region of Coquimbo, at 400 m altitude, in low scrub, on rocks and protruding areas alongside Puyasp. (Bromeliaceae).

Taxonomic and nomenclatural comments: T. undulosus and T. skottsbergii resemble each other in the tuberculate shape of their ribs, but also have characters that differentiate them (Table 3).

17. Trichocereus uyupampensis Backeb., in Backeb. & F.M. Knuth, Kaktus-ABC: 412 (205). 1936.

Trichocereus uyupampensis Backeb., Blätt. Kak-teenf.: 7. 1935, nom. nud. Echinopsis uyupampensis(Backeb.) H. Friedrich & G.D. Rowley, I.O.S. Bull.3(3): 99. 1974. TYPE: photograph: 1194. (Neotype, designated here: Ritter. Kakteen Südamerika 4: 1553).

Trichocereus glaucus F. Ritter, Kakt. And. Sukku-lent. 13: 180, ill. 1962. Echinopsis glauca (F. Ritter) H. Friedrich & G.D. Rowley, I.O.S. Bull. 3 (3): 95. 1974. TYPE: Southern Peru, Tambo river, Julio 1954, F. Ritter 270. (Holotype: U 117004!.)

Trichocereus glaucus F. Ritter f. pendens F. Ritter, Kakt. And. Sukkulent. 13: 181. 1962, nom. inval. (Art. 37.1). Trichocereus glaucus F. Ritter var. pendens(F. Ritter) Backeb., Kakteenlex. 437. 1966. Echinopsis glauca (F. Ritter) H. Friedrich & G.D. Rowley f. pen-dens (F. Ritter) H. Friedrich & G.D. Rowley, I.O.S. Bull. 3 (3): 95. 1974

Iconography: Ritter (1980b), photograph 1078: 1225.


Extended description: Plants cespitose, stemsbranching from the base, shrubs. Branches decum-bent or pendent, 1–2 m long, 4–8 cm diameter, young stems green opaque, turning gray–green with age. Ribs 7−9, crenate, 2.5 cm wide, 7.5–12.5 mm high, with round areoles 1–2 cm apart, 5.0–7.5 mm in diameter, all gray or light brown. Apical spinesblack to brown, spines in the middle and lower parts of stem becoming gray; radial spines 7–10, rigid, strong, acicular, 2–15 mm long; central spines 3–6, outstanding, straight, subulate, 1.5 mm diameter, 2–8 cm long. Flowers 13−19 cm long, open in the evening or at night, aromatic; green receptacle with scales 2–5 mm long, subulate, fleshy, green; dark brown hairs; nectaries 3–4 mm in diameter, tubu-lar, 13–23 mm long; floral tube 4–6.5 cm long, 2–3 cm wide at the top, internally greenish white, exter-nally grayish green, bearing hair (as on the recep-tacle), subulate scales, larger than 1 cm long, dark green; stamens white, green at the bottom, inserted 2–3 cm from the base of the tube, extending to the top where they end; style pale green with apex white, sometimes brown, 10–11 cm long, of which 14–18 mm are the lengths of the stigmata, which vary in number from 13 to 16 and are a very pale yellow; tepals 5–8 cm long, 1–2 cm wide, with apex serrated

or smooth, mucro-nulate, basally nar-rowing to 2/3 to 3/4 of maximum width, the outer te-pals shorter, spatu-late, with the apex mucronulate, the inner tepals white or somewhat pink.Fruits 4 cm long, 3

T. undulosus T. skottsbergii

Color of branches dark green grayish green

Number of ribs 20 12–17

Spines at the apex thick (1 mm) and rigid thin (less than 0.5 mm) and flexible (bristly)

Color of spines at the apex white with brown spots. Gray with little black dots and brown tip.

Thickness and size of adult central

spines2 mm and 3.5 cm long 1.5 mm and 7 cm long

Table 3. Differences between T. undulosus and T. skottsbergii in terms of morphological characters.


cm in diameter, apex truncate. Seeds 1.2 mm long, 1 mm wide, 0.6 mm thick, testa glossy black; (Ritter 1962, 1980b, Backeberg 1976).

Distribution and habitat: In southern Peru, de-partment of Arequipa, in the mountains of the lower Rio Tambo, and the department of Monquegua, in the Ilo region. In northern Chile, Region XV of Arica and Parinacota, province of Arica, south of Arica, on the cliffs near the coast, at 400–600 m al-titude (Backeberg 1976, Anderson 2005, Kiesling et al. 2008).

Taxonomic and nomenclatural comments:Backeberg (1935) mentioned Trichocereus uyupam-pensis as a nomen nudum when he described the veg-etation in the arid region of southern Peru. Months later Backeberg (1936) published formally the diag-nosis and morphological description of this species. Subsequently, Backeberg (1976) proposed T. glaucusas a variety of T. uyupampensis, but sufficient com-mon morphological characters were found (form of growth, stem diameter, number of ribs, number and shape of radial spines, etc.), to suggest grouping them.

Ritter (1962) mentioned the pendulous stem form (“pendens”) which occurs south of the region of Arica in Chile. The growth habit of the stem is thought to be due to the habitat where the spe-cies grows, either on the rocky cliffs near the coast (where it has a pendulous habit) or inland, on steep rocky walls around rivers (where it exhibits an erect habit). It is proposed here that the morphological concept of T. glaucus be extended.

What is more, according to Article 37.1 of the International Code of Botanical Nomenclature (Mc-Neill et al. 2006) Ritter did not validly publish the name T. glaucus f. pendens as he did not designate a type specimen.

Excluded species

Eulychnia coquimbana (Molina) S. Albesiano,

comb. nov.

Cactus coquimbanus Molina, Sag. Stor. Nat. Chili.: 170. 1782. Cereus coquimbanus (Molina) K. Schum., Monatsschr. Kakteenk. 6: 166. 1896. Tricho-cereus coquimbanus (Molina) Britton & Rose, TheCactaceae 2: 139. 1920 (except description and fig-ure). Echinopsis coquimbana (Molina) Friedrich & G.D. Rowley, I.O.S. Bull. 3 (3): 94. 1974. TYPE (Neotype designated here): plate VI, in: C. Skotts-berg. 1950. Acta Horti Gothob. 18.)

Eulychnia spinibarbis (Otto ex Pfeiff.) Britton & Rose, The Cactaceae 2: 82-83, ill. 1920. TYPE (Neo-type, designated here): plate VIIb, in: C. Skottsberg. 1950. Acta Horti Gothob. 18.

In creating the new combination, Trichocereus co-quimbanus, Britton and Rose erroneously identified a kind of Trichocereus considered to be T. pectiniferus.The species described by Molina, that has tall, erect stems and 20 cm long spines (“which then were used to crochet socks”), corresponds to the genus Eulych-nia; while the plant described by Britton and Rose

is cespitose, one meter high, with spines that do not exceed 8 cm in length.

The combination of Echinopsis coquimbanus is included because Friedrich and Rowley conducted the transfer of all species of Trichocereus published by Britton and Rose (1920) to the genus Echinopsis,without performing a nomenclatural and taxonomic study of each species.

Eggli and Walter (2012) propose rejecting the name Cactus coquimbanus Molina, because the char-acters mentioned in the original description (“Cactuserectus, longus, 10-angularis: angulis obtusis, spi-nis longissimus rectis”) are insufficient for knowing whether they correspond to a species of the genus Eulychnia or to one of the genus Trichocereus, both of which occur in the surroundings of the city of Coquimbo. Our observations of the vegetative char-acters permit us to conclude that the description of C. coquimbanus corresponds to a species of the genus Eulychnia, and accordingly the description of T. pec-tiniferus (syn. Trichocereus coquimbanus) corresponds to cespitose plants which do not exceed 80 cm in height and which have short (6 cm) spines, while the number of ribs in Trichocereus chiloensis (11−21) is greater than 10.

ACKNOWLEDGMENTS

I wish to express my acknowledgment to Ayra Faúndez, Luis Faúndez, Omar Ferrari, Rodrigo Flores, Susana Freire, Roberto Kiesling, Iris Peralta, Patricio Saldivia, Cecilia Scoones and Detlev Metz-ing for their valuable help with the field work, in the laboratory and with the editing of the manuscript. To the directors and curators at the consulted herbar-ia (Stephan Beck, Norma Deginani, Laura Iharlegui, Eduardo Méndez, Inés Meza, Mélica Muñoz, Nora Muruaga and Luc Willemse), for their kindness in resolving doubts, sending pictures and loaning the material required. To José Monserrat (Latin transla-tion) and Leopoldo Arrieta (English translation). To the German Cactus Society, the RLB-Andrew W. Mellon Foundation Grant 2010–2011 and to the Organization of American States (OAS) for grant-ing a Doctoral Scholarship to S. Albesiano. Special thanks go also to the reviewers of the manuscript, who contributed in great degree to its completeness.

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