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«Andean roots and tubers: Ahipa, arracacha, maca and yacon M. Hermann and J. Heller, editors Promoting the conservation and use of underutilized and ...»

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Promoting the conservation and use of underutilized and neglected crops. 21. 129 The shoots of South American Arracacia species grow from perennial roots that are either woody, wide-ranging and rarely surpassing l-2 cm in diameter, or tuberous, tapering and often more than 4 cm thick at their base. The shoot systems of species with woody roots, such as those of A. elata and A. moschata, are of indeterminate growth. Consequently, their shoots are often over 3 m long and can be seen sprawling over (disturbed) roadside vegetation. The habitats of these species are mostly in temporarily moist or perhumid montane forests (3000 m altitude;

Fig. 15). The inflorescences are borne laterally on distal shoot sections. The plants resprout from basal nodes and they will display synchronously vegetative, flowering and fruiting shoots throughout the year (except for extremely dry periods).

The plant habits and phenologies of tuberous species (A. xanthorrhiza, A. andina, A. equatorialis) are quite distinct. They result from adaptation to warmer and seasonally arid habitats at lower altitudes (below 3000 m). A rootstock consisting of the greatly enlarged and starchy root and compressed stem structures attached to it at soil surface serves as a storage organ allowing the plant to survive rainless periods of up to 8 months during which all aerial plant parts perish. During the rainy season ‘rosettes’ of large and petioled leaves emerge from the rootstock. In

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plants beyond a certain size, also one to several generative shoots of determinate growth can develop. Each of them bears several inflorescences (umbels). Leaves on the generative shoot are much smaller and have reduced petioles. The development of vegetative leaf rosettes and generative shoots is synchronous in A. andina and A.

xanthorrhiza. In greenhouse material of A. equatorialis, the formation of the generative shoot often precedes vegetative growth. This species and A. andina display pronounced dormancy of the rootstock, a trait which is absent in cultivated arracacha. Arracacia colombiana Constance & Affolter, Brittonia 47: 322-323 (1995) Rather slender, caulescent, branching, herbaceous, the foliage and inflorescence sparsely scaberulous to hispidulous, to 1 m tall, from woody roots; leaves ovate, 10cm diameter, biternate or ternate-pinnate, the leaflets ovate, 2-5 cm long, 1-3 cm broad, acute to obtuse, rounded or cuneate at base, mucronulate-serrate, scaberulous to hispidulous on the veins beneath; petiole 10-15 cm long, the petiole and petiolules without definite callous thickenings but usually papillose at the main junctures;

cauline leaves reduced upward with somewhat dilated oblong to oval scarious sheaths, the upper leaves wholly sheathing; umbels pedunculate, the peduncles 5cm long, scaberulous-hispidulous at apex; involucre absent or of 1 or 2 foliaceous bracts; rays 10-25, the 3-5 fertile rays 3.5-7 cm long, the staminate filiform, much shorter, all spreading-ascending; umbellets about 20-flowered, only 2 or 3 flowers perfect, the mature pedicels (3) 8-20 mm long, filiform, spreading; involucel of about 5 linear, entire, unequal bractlets 3-8 mm long, shorter than flowers and fruit; flowers yellow; calyx absent; petals oblong to obovate, l-veined, vein sometimes branched below apex, with a narrow inflexed apex; stylopodium low conical, the styles about 1 mm long, spreading or reflexed; carpophore bipartite, the halves slender, erect;

fruit ovoid-cordate, 3-5 mm long, 4-5 mm broad; little narrowed at apex, cordate at base, the mericarps subterete, glabrous, the ribs low, filiform; vittae rather large, 2 or 3 in the intervals, 4-6 on commissure; seed face deeply sulcate.

List of exsiccatae: COLOMBIA. Cundinamarca: En areas abiertas y en monte muy denso, suelos ricos, arenosos, Cordillera Oriental, entre Bogotá y La Calera, 2650m, 27 Nov 1947, Barkley, García-Barriga & Vanegas 17C752 (Holotype: COL!;

Isotypes: UC!, US!). — Sibaté (2-4 mi S), 13-15 Oct 1917, Pennell 2452 (paratype, F, K, MO, NY, UC, US and US photo # 230). — Meta: Río Arroz well above confluence of Quebrada Pedregal, 29 Aug 1943, Fosberg 20915 (US).

This recently described species from a limited geographic range in Colombia is similar to Neonelsonia acuminata and the Mexican Arracacia filipis. Whether this is a ‘good’ species will be seen when more material and especially fully mature specimens become available. Arracacia tillettii Constance & Affolter, Brittonia 47: 324-327 (1995) Stout, caulescent, branching, parsley-scented, the foliage and inflorescence sparsely tomentulous, to 1 m tall, from a thick taproot; leaves thick membranous, triangularPromoting the conservation and use of underutilized and neglected crops. 21.

ovate, 10-25 cm diameter, 2-3 ternate or ternate-pinnate, the leaflets lanceolate to ovate-lanceolate, 5-8 cm long, l-3.5 cm broad, acute or acuminate, cuneate, rather coarsely serrate and occasionally lobed or incised, paler and prominently reticulateveined and scaberulous beneath; petiole 20-35 cm long, both petioles and petiolules papillose at the main junctures, sheathing, the lower third of sheath lustrous nearwhite; cauline leaves reduced upward with somewhat dilated oblong scarious sheaths, the upper wholly sheathing; umbels pedunculate (or umbels sometimes sessile), the peduncles 5-15 (20) cm long, tomentulous at apex; involucre of 3-5 linear to ovate-lanceolate entire to 3-lobed bracts 5-20 mm long; rays 15-20, the 8-10 fertile rays 4-10 cm long, the staminate filiform and much shorter, all spreading-ascending;

umbellets 15-25-flowered, the mature pedicels to 10 mm long, spreading; involucel of about 5 linear to lanceolate, entire or few-lobed bractlets 2-5 mm long, shorter than flowers and fruit; flowers light yellowish green, the petals oblong-oval; stylopodium low conical, the styles spreading, about 1 mm long; carpophore bipartite; immature fruit ovoid-elliptical, 4-5 mm long, 3-4 mm broad, obtuse at apex, rounded at base, glabrous, the ribs narrowly winged (?); vittae and seed face not seen.

List of exsiccatae: COLOMBIA. Guajira: Sierra de Perijá, Cerro Pintada, 3200 m, 26

Apr 1987, Cuadros & Gentry 3543 (paratype, JBGP, MO, UC). — VENEZUELA. Zulia:

Distr. Maracaibo, Campamento ‘Monte Viruela’ (10°25’ ca. 13” N, 72° 52’ ca. 42” W), on tepuí-like limestone massif 5 x 2.5 km on international boundary, Serranía de

Valledupar, Sierra de Perijá, ca. 3100 m, 21-28 Jul 1974, Tillett 747-1194 (Holotype:

VEN!; Isotypes: COL!, MO!, NY!, UC!).

This species has recently been described on the basis of two collections from an inaccessible area in the Serrania de Perijá near the border between Venezuela and Colombia. According to Constance and Affolter (1995b), it is difficult to assign generically, partly because of a lack of mature fruit. Arracacia tillettii has affinities with Myrrhidendron and its isodiametric fruit recalls Neonelsonia. Carpologically, A.

tillettii is very similar to A. colombiana. Arracacia moschata (Kunth) DC., Prodr. 4: 244 (1830) Conium moschatum Kunth in H.B.K., Nov. Gen & Sp. 5: 12, pl. 430 (1821).

Plants stout, caulescent, branching, 0.5-2 m tall, the foliage glabrous, the inflorescence scaberulous, from tuberous roots. Leaves ovate, 10-30 cm long, 8-15 cm broad, bipinnate or ternate-pinnate, the leaflets ovate-oblong to ovate, acute, 2-5 cm long, 1-3 cm broad, spinulose-serrate and incised to pinnatifid toward base, paler beneath, squamulose-tufted on the rachis above. Petioles 10-20 cm long. Cauline leaves with oblong inflated sheaths. Inflorescence branching, the peduncles axillary. Involucre usually 0. Rays 10-25, rather stout, spreading-ascending or spreading, scaberulous, 5-13 cm long. Involucel of 3-8 linear to lanceolate, entire or few-toothed bractlets 6mm long. Petals maroon (rarely yellow), oval; styles slender, the stylopodium conical. Pedicels 10-30 mm long, scaberulous. Carpophore bifid ca. half its length.

Arracacha (Arracacia xanthorrhiza Bancroft) Fruit lance-ovoid, 6-8 mm long, 3-4 mm broad, the ribs prominent, acute; vittae solitary to several in the intervals, usually 4 on the commissure. (Description taken from Mathias and Constance 1976.) List of exsiccatae: ECUADOR. — Azuay: “Crescit in frigidis Provinciae de los Patos,

prope Teindala, 1400 hex. “, Humboldt & Bonpland 2163 (P. holotype). — Bolívar:

Canton Guaranda, parroguia Salinas, en Las Tres Mercedes, a 4.8 km desde Salinas hacia Guanjo, 3200 m, 10 May 1994, Tapia & Cazar 33 (UC). — Carchi: 10 km de El Angel hacia Tulcán en la Panamericana antigua, antes de llegar a los frailejones, 3000-3300 m, 3 Feb 1995, Hermann & Korntheuer 1373 (UC) — El Frailejón on road Tulcán-El Carmelo, 3300 m, 6 Mar 1974, Harling & Andersson 12518 (NY). — Julio Andrade-Playón de San Francisco road, Cochaseca, 6 July 1978, Boeke & Jaramillo 2363 (NY). — Las Peñas, between la Rinconada and San Gabriel, 3150 m, Asplund 7182 (S, UC). — “Hauca & Tusa”, 2700-3000 m, Lehmann 4675 (K). - Ca. 8 km S of Tulcán, 2500 m, Hitchcock 21005 (GH, NY, US). — Páramo del Angel, 3400 m, Sparre 14241 (S). —Road Tulcán-El Pun, 3500 m, Mexía 7580 (UC, US). — Cotopaxi: Road Quevedo-Latacunga, Zumbagua, 3500 m, Harling, Storm & Ström 8906 (GB). — Imbabura: Mojanda, ca. 10 km SSW of Otavalo, 2900-3150 m, Sparre 13462 (S). — Mojanda, on road Otavalo-Minas, 3200 m, Sparre 16823 (S). — Cerro Cotacachi above Lago Cuicocha, ca. 3300 m, Asplund 20247 (S). — Sine loco, Sodiro 80, 80/17 (Q). — Napo: Near Archidona, Jameson 724 (BM). — Pichincha: Quito, Jameson 30 (BM, W).

— Quitensian Andes, Jameson s.n. (K). — Cerro Pichincha, Jameson s.n. (US); Hall 40 (K); Benoist 2423 (P, UC). — Cráter de Pululahua, N of Quito, ca. 2800 m, Barclay, Juajibioy & Tinajero 7901 (UC). — Road Quito-Santo Domingo de los Colorados, km 46, 2000 m, Dodson & Thien 1064 (LA, WIS).— Tungurahua: Road Paso-Ambato, near Río Ambato, 2850 m, Heinrichs 71 (G, M, NY). — San Fernando, AmbatoGuaranda, km 12, 3300 m, Sparre 18410 (S). — Prov. unknown: Sine loco, Spruce 5794 (BM, NY); Jameson 30 (K).

This species is, with Arracacia xanthorrhiza, one of the two properly referred classical species in the genus. Arracacia moschata is a well-defined taxonomical entity and can be easily recognized because of its distinctive (spinulose-serrate) leaf and seed shape (Fig. 16). The seeds, and to a lesser extent the leaves, have a pleasant umbelliferous fragrance, somewhat reminiscent of the resinous odour of certain conifers and Citrus.

It is common in the páramos of central and especially northern Ecuador, on both sides of the Cordillera, between 3000 and 3300 m altitude. However, it has not yet been collected in Loja and it seems to be absent from Peru. Arracacia moschata might also occur in southern Colombia, in habitats similar to the ones in which the species is common in northern Ecuador.

I have seen large populations of this species in Carchi (Hermann & Korntheuer 1373) (Fig. 16), where it occurs in wet habitats regularly exposed to fog and mist. In these situations, A. moschata is associated with Coriaria, Ericaceae and Espeletium.

Promoting the conservation and use of underutilized and neglected crops. 21. 133 Fig. 16. Herbarium specimen of Arracacia moschata collected in Carchi, Ecuador. (Lot.: 10 km de El Angel hacia Tulcán en la Panamericana antigua, 0°40’ N, 77°53’ W, 3300 m asl, 3 February 1995, vern. ‘sacha zanahoria’, Hermann & Korntheuer 1373, UC.) Scale: 10 cm.

134 Arracacha (Arracacia xanthorrhiza Bancroft) Its shoots, which emerge from a lignified and compact rootstock, reach several meters in length and sprawl over roadside thickets. The woody roots can surpass 1 m in length but rarely exceed 2 cm in thickness (Tapia & Cazav 33). Arracacia elata Wolff, Bot. Jahrb. 40: 304 (1908) Arracacia Pennellii Constance, Bull. Torrey Bot. Club 76: 71 (1949).

Arracacia Wigginsii Constance, ibid. 43.

Plants stout, caulescent, branching, vining or scrambling, 1-8 m long, the foliage glabrous, minutely scaberulous in the inflorescence, from tuberous roots. Leaves triangular-ovate to ovate, 8-30 cm long, 10-25 cm broad, ternate or 1- or 2-pinnate, the leaflets lanceolate to ovate-lanceolate, acute or acuminate, 2-6 cm long, 0.8-4 cm broad, spinulose-serrate and usually incised or lobed towards base, paler beneath, squamulose-tufted on the rachis above. Petioles 15-45 cm long. Cauline leaves with oblong inflated sheaths. Inflorescence branching, the peduncles terminal and axillary.

Involucre usually 0. Rays 15-30, slender, spreading-ascending, scaberulous, 3-7.5 cm long. Involucel of 3-10 linear to lanceolate, entire to toothed bractlets 2-15 mm long.

Petals greenish-yellow, oval to obovate; styles slender, the stylopodium conical.

Pedicels 3-8 mm long, scaberulous. Carpophore bifid, ca. l/4 its length. Fruit ovoid to oblong-ovoid, tapering at apex, 4-8 mm long, 3-5 mm broad, the ribs prominent, obtuse; vittae solitary to several in the intervals, usually 2 or 4 on commissure.

List of exsiccatae: COLOMBIA. Cundinamarca: Río San Francisco, above Bogotá, 13 Sep 1917, Pennell 1932 (NY type of A. pennellii Constance, GH, US). - Macizo de Bogotá, Quebrada del Rosal, 3000 m, 29 June 1939, Cuatrecasas 5700 (US). - Norte de Santander: Between Mutiscua and Pamplona, 3400 m, 23 Feb 1927, Killip & Smith 19728 (GH, NY, US). - Santander: Cordillera Oriental, páramo de Santurbán, entre Cuesta Boba y el extremo oeste, 3400 m, 27 July 1940 Cuatrecasas & Barriga 10314 (US). - Páramo de Romeral, 3800-4100 m, 29-30 Jan 1927, Killip & Smith 18541 (GH, NY, US). - Vicinity of Vetas, thickets along stream, 3100-3200 m, 16 Jan 1927, Killip & Smith 17347 (GH, MO, NY, US). — Quebrada de País, north of La Baja, dense forest,

ca. 3200 m, 31 Jan 1927, Killip & Smith 18781 (GH, MO, NY, US). —ECUADOR. Azuay:

Cruz Pampa region above Baños (ca. 15 km SW of Cuenca), 9000-10000 ft. elev., 29June 1945, Camp E-3934 (NY). - Panamerican Highway, 65-70 km S of Cuenca, shrubby páramo and roadbanks, 3300-3400 m, 3 Jan 1981, Balslev 1426 (NY). -Canton Cuenca, Zurucucho, forest edge, 3200 m, 11 Aug 1978, Boeke, Jaramillo & Narvaez 2631 (NY). - Road Cuenca-Angas, roadside, 3050 m, 28 Dec 1976, Boeke 664 (NY). — Cerro Soldados (W of Sayausid) below Quinoas, 3400 m, Harling, Storm & Ström 8762 (GB, UC). — Laguna de Surucuchu, 3100-3200 m, Sparre 18855 (S). — Pan American Highway 40 km S of Cuenca, 3300 m, Wiggins 10769 (UC, holotype of A. wigginsii;

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