Abstract: Counting two new endemic species described in the present article, Bolivia has six Bromelia species; the non-endemic species are B. balansae, B. hieronymi, B. serra, B. villosa. The genus, in Bolivia, is typical of the understory especially of the dry to semihumid forests and of savannas, between 150 and 1,400m. It does not occur in the very humid rain forests.
Bromelia arubaiensis sp. nov. is similar to Bromelia hieronymi, that is present in drier ecoregions in the south of Bolivia, but differs, among others, by wider leaves, a bipinnate, lepidote inflorescence, longer floral bracts and longer violet petals. It differs from B. binotii, found in the Atlantic rain forest of Brazil, by the wider leaves, narrowed between the sheaths and the blades, the bipinnate inflorescence, longer floral bracts and shorter petals.
B. ignaciana sp. nov. is similar to B. balansae but differs by the rose and suberect bracts of the scape and the inflorescence, a more elongate and narrower inflorescence, oblong-lanceolate, acuminate sepals and smaller and rose petals.
Key words: Bromeliaceae, Bromelia arubaiensis sp. nov., B. ignaciana sp. nov., Santa Cruz, Bolivia.
The Bolivian Bromelioideae are less diverse and more widely distributed than the Pitcairnioideae: about 32% are Bolivian endemics (vs. 67% of Pitcairnoideae; IBISCH & VASQUEZ 2000, IBISCH et al. 2001). In spite of a generally very high rate of new discoveries in Bolivia, only a few of the new species of bromeliads that have been described in the last years, belonged to the Bromelioideae. All Bromelia species known until recently (KROMER et al. 1999) were rather `old taxa' (descriptions: 1879-1901). However, as a consequence of our intensified bromeliad collecting activities and studies, in this article, two new species of this subfamily can be described: Bromelia arubaiensis and B. ignaciana. Thus, six Bromelia species are known from Bolivia.
Characteristics of the genus: Plants terrestrial, rarely epiphytic, propagating by rhizomes or lateral shoots; leaves rosulate, with curved spines on the margin; inflorescence composed; sepals free or united, obtuse or attenuate, rarely mucronate; petals usually fleshy, united in a tube with free margins, without appendages; stamens included, the filaments forming a more or less long tube; fruit a succulent berry, relatively large; seeds few to many, complanate, naked.
Key for the species of Bromelia in Bolivia
1. Inflorescence scape absent; rachis densely corymbose. Cerrado, Chiquitano dry forest (100-500m); Santa Cruz => B. villosa
1. Inflorescence scape present; rachis globose or cylindric
- 2. Rachis globose, 6-8cm in diameter, 8-10cm long. Dry Chaco and Chiquitano forests, Cerrado, humid savannas, Tucuman-Bolivian forest, Dry inter-Andean valleys (150m to 1,400m); Beni, Chuquisaca, Cochabamba, La Paz, Santa Cruz, Tarija => B. serra
- 2. Rachis cylindric, 5-7cm in diameter, to 30cm long
- - 3. Inflorescence densely particulate, cylindric, branches to 5cm long
- - - 4. Inflorescence to 22cm long, to 9cm in diameter. Scape bracts red, linear-subelliptic, spreading. Flowers violet with a white margin, petals to 25mm. Cerrado, Chiquitano dry forest (150-800m); Beni, Santa Cruz => B. balansae
- - - 4. Inflorescence to 30cm long, to 6cm in diameter. Scape bracts rose, linear-lanceolate, suberect. Flowers rose with a white margin, petals to 18mm. Cerrado (200-400m); eastern Santa Cruz => B. ignaciana
- - - 4. Inflorescence laxly particulate, pyramidal, branches to 18cm long
- - - - 5. Inflorescence glabrous. Branches to 18cm long, flowers white or lilac. Fruits rose, glabrous. Dry Chaco vegetation (250-700m); (Chuquisaca?), Santa Cruz, Tarija => B. hieronymi
- - - - 5. Inflorescence lepidote. Branches less than 10cm long. Flowers violet. Fruits clear green,lepidote. Pre-Andean (semi-)humid forests (about 500m), northwestern Santa Cruz => B. arubaiensis
GEOGRAPHICAL RANGE: Endemic to Bolivia. Santa Cruz department, provinces of Andres Ibanez and Ichilo.
Plant terrestrial, stemless, flowering to approximately 100cm high, propagating by elongate scaly stolons.
Leaves numerous, rosulate, to 100 to 130cm long.
Sheaths to 80-100mm long, to 45mm wide, glabrous above, white adpressed scales beneath.
Blades linear, attenuate, not pungent, 40 to 50mm wide, brownish towards the base, shiny green towards the apex, the apex recurved, margins serrulate with antrorse castanaceous spines in dense succession, to 5mm long, glabrous above, white adpressed scales beneath.
Scape about 40cm long, to 17mm thick.
Scape bracts subfoliaceous, serrulate, bright red, attenuate. Inflorescence about 25cm long, to 15cm wide, laxly bipinnate, white tomentose-lepidote.
Primary bracts 7-10cm long, 2cm wide, bright red, oblong-elliptic in the lower half, becoming long acuminate from the middle to the apex; the margins serrate, lower ones longer than branches, becoming dry papyraceous, the apex recurved.
Branches distinct, to 10-15cm long, (sub)erect, rachis of lower branches flattened especially toward base, to 12mm wide, to 3-4 mm thick, red, white lepidote.
Floral bracts to 10mm long, 3-4mm wide, longer or equaling pedicels, triangular, dry papyraceous.
Flowers erect, to 40mm long. Pedicel 5-15mm long, slender.
Sepals to 10mm long, to 4mm wide, oblong, triangular, obtuse, white becoming dry papyraceous.
Petals to 17mm long, oblong, obtuse, (rose-lilac) violet with whitish tips.
Ovary inferior, to 10mm long, to 3mm wide. Hypanthium green, white lepidote.
Fruits to 45mm long, about 10mm wide, ellipsoid or clavate.
PARATYPES: BOLIVIA. SANTA CRUZ: Prov. A. Ibanez: Canton Terevinto, Hacienda Arubai, 480m,17°41' 19"S, 63°25'24"W 1 Dec. 1999, flowered in cultivation 7 Aug. 2002, R. Vasquez & J. Coimbra 3468 (LPB, US7.); Prov. Ichilo: El Carmen (SSW of Buena Vista), 17°32', 63°43', 450m, 5 Oct. 1996, I. G. Vargas & S Hurtado 5407 (USZ).
HABITAT: Terrestrial forest species in an pre-Andean area nearby the Andean knee where the Chiquitano Dry Forest and Southwest Amazon ecoregions meet. The forest is mainly evergreen but less diverse than the more humid Amazon forests further northwards.
OBSERVATIONS: The species is similar to Bromelia hieronymi, that is present in drier ecoregions in the south of Bolivia, but differs, among other characters, by wider leaves, a bipinnate, lepidote inflorescence, longer floral bracts and longer violet petals. It is also similar to B. binotii, found in the Atlantic rain forest of Brazil, from which it is distinguished by the wider leaves, narrowed between the sheaths and the blades, the bipinnate inflorescence, longer floral bracts and shorter petals.
The new species is called after the private Arubai reserve of the Coimbra family where it is was discovered. It was illustrated by colored photographs in IBISCH & VASQUEZ (2000) as Bromelia sp. 1.
Table l: Comparison of selected characters of Bromelia arubaiensis sp. nov., B. binotii and B. hieronymi.
RECORDS IN BOLIVIA: BENI: Rio Guapore, San Antonio de Manuelito, 25 Jun. 1952, Black & Cordeiro52-15327 (IAN); Prov. Ballivian: road Riberalta-Santa Rosa, 36 km before arriving at Puerto Yata, 21 Oct. 1991, S. G. Beck 20689 (LPB, SEL); Prov. Mamore: "Barranquita"-farm, ca. 18 km S of San Joaquin, 200m,18 Aug. 83,m.J. Balick et al. 1440 (LPB); ibid.: 7 km S of San Joaquin¬Trinidad, 150m, 7 Nov. 1993, M. Moraes 1631 (LPB). SANTA CRUZ: Prov. Chiquitos: Serrania Santiago, 9 km SW of Santiago, 550m, 27 Jul. 1982, W. Till 133 (WU); Prov. Guarayos: rock outcrop between rio Negro and sawmill La Chonta 330m, 15°00'S, 62°37'W terrestrial in Amazon lowland forest, 6 May 2002, R. Vasquez, L.R. Moreno, A. Moreno, O. Moreno, D. Ric & I. Munoz 4488 (VASQ); Prov. Nuflo de Chavez: Lomerio, 12 km N of Las Trancas community, 16031'13"S, 61050'47"W, 450m, 3 Jul. 1995, F Mamani 926 (USZ); Prov. Velasco: Flor de Oro, Noel Kempff Mercado National Park, in Cerrado forest, 200m, 8 June 1993, R. Vasquez, L.R. Moreno & H. Justiniano 1812 (VASQ); ibid.: between Santa Rosa and San Martin, towards Florida, 310m,15°21'35"S, 61°29'32"W on outcrops, 22 Nov. 2001, R. Vasquez, G. Gerlach & L.R. Moreno 4333 (VASQ); ibid.: Cerro Pelado, 35 km from Florida to La Mechita, 320m, 14°36'26"S, 61°29'30"W terrestrial on outcrops, 26 Nov. 2001, R. Vasquez, G. Gerlach & L.R. Moreno 4360 (VASQ); ibid.: 170 km from Santa Rosa de la Roca toward Piso Firme, about 350 m, 14'39'S, 61°35'W terrestrial on granitic rock inselberg, 28 June 1993, P. & C. Ibisch, G. Rauer, D. Rudolph 93.0850 (LPB, FR).
HABITAT: Bromelia balansae is a species of northeastern Bolivia. It is found in open savanna vegetation, especially of the Cerrado type. 150-800m.
OBSERVATIONS: It is similar to B. serra but differs principally by its elongate and cylindric inflorescence, the violet flower color and the more northern distribution.
RECORDS IN BOLIVIA: SANTA CRUZ: Prov. Cordillera: Izozog, May 1934, Cardenas 2755 (GH); Prov. A. Ibanez: 5 km SE of Don Lorenzo communiry, 17'49'S, 62'50'W, 320m,17 Nov. 1990, M. Nee & G. Coimbra 39933 (LPB); 30 km E of Cotoca, between La Bola and Rio Grande, 280m, dry Chaco forest, Mar. 1993, R. Vasquez 2428 (VASQ); ibid., 6 Sept. 1998, R. Vasquez & D. Nash 2971(VASQ); ibid.: 26 Oct. 1999, R. Vasquez, P. Ibisch, M. Nee et al. 3415 (VASQ); ibid.: M. Nee & G. Coimbra S 39933 (LPB); Prov. Nuflo de Chavez: Estancia San Miguelito, 5 km SE of `Puesto Montana', 260m, 29 Mar. 1995, A. Fuentes 584 (LPB). TARIJA: Prov. Gran Chaco: Villamontes, 20 Nov. 1917, Pflanz 942 (M); Prov. O'Connor: Puerto Margarita, Rio Pilcomayo, 21°10'S, 63°50'W 4 May 1983,A. Krapovickas & Schinini 39087 (LPB).
HABITAT: This is a typical Gran Chaco species. It grows in dry forests on poorly drained and temporarily inundated soils, between 250 and 700m, together with Stetsonia coryne (Cactaceae), Copernicia alba, Trithrinax schizophylla (Arecaceae), Aspidosperma quebracho-blanco (Apocynaceae), and others.
OBSERVATIONS: The apical part of the stem and the sheaths of the leaves contain starch consumed by the indigenous peoples of the Chaco (especially Ayoreo). Collected plants are held into the fire until the leaves are toasted. The young stolons are consumed without cooking (ARENAS & ARROYO 1988). The leaves contain fibers that are extracted beating the leaves. Then they are dried, woven and converted into ropes, bags and other articles (ARENAS 1981).
GEOGRAPHICAL RANGE: Endemic to Bolivia. Santa Cruz department, San Ignacio municipality, Prov. Velasco, Noel Kempff National Park (it may occur in adjacent Brazil).
Plant terrestrial, stemless, flowering to more than 100cm high.
Leaves numerous, rosulate, to 1m long.
Blades linear, attenuate, pungent, to 35mm wide, serrulate with uncinate spines, to 3mm long, glabrous above, white adpressed scales beneath.
Scape erect, stout, about 30cm long, densely white tomentose-lanate, to 15mm thick.
Scape bracts subfoliaceous, serrulate, rose, attenuate, much longer than the internodes, covering the scape completely, basal ones to 35cm long, upper ones to 18cm.
Inflorescence about 30cm long, less than 15cm wide, paniculate, cylindric, white tomentose-lanate.
Primary bracts like scape bracts, exceeding branches, to 11cm long, lanceolate with a long acuminate apex or short cuspidate apex.
Branches short, hidden by primary bracts, to 3-4cm long, (sub)erect, few-flowered.
Floral bracts linear-lanceolate, acute, 25mm long, 10mm wide, exceeding the ovary, semilate, rose.
Flowers subsessile, 40-50mm long.
Sepals free, oblongate-lanceolate, acuminate, to 18mm long, to 6mm wide, conduplicate-carinate, diminutely serrulate at base and clearly serrulate toward the apex.
Petals cuneate, apically cucullate, erect, not very opened during anthesis, 18mm long, connate for 4mm, 7mm wide, rose with white tips.
Stamens included, filaments more than half-connate.
Ovary inferior, to 25mm long, to 6mm wide.
Hypanthium to 7mm long, green, white lanate.
The new species is illustrated on the cover of the present journal (plants at Las Gamas, locality of the paratype R. Vasquez et al. 2300; photograph Roberto Vasquez).
Table 2: Comparison of selected characters of Bromelia ignaciana sp. nov. and B. balansae
PARATYPES: SANTA CRUZ: Prov. Velasco: PNNKM, Las Gamas, 13 Jun. 1994, R. Vasquez, L.R. Moreno, O. Moreno, H. Justiniano, D. Ric & A. Moreno 2300 (VASQ).
HABITAT: Terrestrial between rocks at the Caparu table mountain and nearby the Las Torres camp of the Noel Kempff Mercado National Park.
OBSERVATIONS: B. ignaciana is similar to B. balansae in the general habit and the cylindric inflorescence with short floral branches. However, it differs by its rose and suberect bracts of the scape and the inflorescence (not red and spreading), the more elongate and narrower inflorescence, the oblong-lanceolate and acuminate (not elliptic and obtuse) sepals, and the smaller and rose (not violet) petals.
The new species is dedicated to the San Ignacio municipality, one of the greenest and most forest-rich municipalities of Bolivia, thanks to the Noel Kempff Mercado National Park, with a high coverage of protected areas.
TYPE: Argentina. Lorentz & Hieronymus s. n. (LT: GOET).
GEOGRAPHICAL RANGE: Bolivia, Paraguay y Argentina. In Bolivia: departments of Beni, Cochabamba, La Paz and Santa Cruz.
Plant terrestrial, to 50cm high.
Leaves 1-1.5m long, 3-5cm wide, with a serrate margin, with antrorse and retrorse, unciform spines; apical ones red at maturity.
Scape short. Bracts subfoliaceous, red.
Flowers in short fascicles, 5-9 flowered.
Sepals free, 15 x 5mm, white.
Petals white with a violet band towards the apex, connate at base.
Fruits yellow when ripe, edible. Fig. 7.
RECORDS IN BOLIVIA: BENI: Prov. Ballivian: Espiritu, in the zone of influence of the Rio Yacuma, 200m,13 Apr. 1980, S G. Beck 2533 (LPB); ibid.: EBB, 40 km E of San Borja,l4°50'S, 66'40'W, 200m, 16 Apr. 1991, T. Killeen 2864 (LPB); Prov. Cercado: Trinidad, Misiones Guarayos, Sep. 1926, Werdermann 2300 (LPB, S); Prov. Maraban: 30 km of Trinidad to San Joaquin, 14'10'S, 64'50'W, 2 Nov. 1993, M. Moraes et al. 1473 (LPB, SEL); Prov. Yacuma: Beni Biological Station, Estancia El Porvenir, nearby station, 14'50'S, 66°20W 250m, 12 Jul. 1988, E. Vallanueva et al. 858 (LPB); ibid.: 23 Dec. 1988, M Moraes 944 (LPB); Prov. Mamore: 7 km S of San Joaquin toward Trinidad, 150m, 7 Nov. 1993, M. Moraes et al. 1631(SEL). COCHABAMBA: Prov. Ayopaya: Atispaya,1,340m,16°36' S, 66043'W, terrestrial in dry inter-Andean valley with deciduous forest at the transition towards a humid forest, 22 Jun. 2001, R. Vasquez, G. Navarro, M. Fernandez, F. Miranda & H. Rocha 4159 (VASQ). LA PAZ: Prov. Franz Tamayo: Rio Bilipisa, 10 km NW of Apolo, 14°36' S, 68'27'W, 1,100m, 5 Jul. 1997, M. Kessler, J Gonzales, K Bach, L Jimenez & A. Portugal 11037 (LPB, SEL, GOET); Prov. Iturralde: Luisita, 13 '05'S, 67°15'W 180m, 28 Febr. 1984, SG. Beck & Haase 10102 (LPB). SANTA CRUZ: Prov. A. Ibanez: Canton Terevinto, Hacienda Arubai, 480m, 17Y'41' 19"S, 63Y'25'24"W, terrestrial in transitional habitat between Cerrado savanna to humid forest, 1 Dec. 1999, R. Vasquez & J. Coimbra 3469 (VASQ); ibid.: International Airport of Viru-Viru, Warnes, 17°39'46"S, 63°69'24"W 380m, 3 Febr.1994, M. Menacho & Sea 364 (USZ); Prov. Chiquitos: 1945, Weddel13520 (P); ibid.: 55 km from San Jose a Quimome, 380m, 17°42'S, 61°14'W, terrestrial on rock outcrops within Cerrado, 14 Jul. 2002, R. Vasquez, D. Ric & A. Osinaga 4567 (VASQ); Prov. Cordillera: Izozog, May 1934, Cardenas 2756 (GH); Campo Leon, 150 km southeast of Santa Cruz de la Sierra, 350m, terrestrial in dry Chaco forest, 12 Sept. 1999, R. Vasquez, et al. 3362 (VASQ); ibid.: Curuyuqui, 50 km SE of SC on rio Parapeti, 18°45'S6"S, 62°13'S9"W, 350m, 25 Oct. 91,A. Gentry et al. 75214 (USZ); ibid.: Charagua, 5 km from the town, Rio Ovai, 19'46'S, 63'1 2' W 800m, 13 Apr. 1990, I. G. Vargas 471(USZ); ibid.: Banados del Izozog, Estancia Cachari, 410m,14 Mar. 1991, G. Navarro & I. G. Yargas 394 (USZ); ibid.: Copere, 16 km S of Las Brechas, 19'40'S, 62'08'W, 12 Oct. 1992, M. Hllegas et al. 260 (USZ); ibid.: Boyuibe, 55 km towards the border with Paraguay, via F. Villaz6n, 620m, 4 Oct. 1983, S.G. Beck & M. Liberman 9418 (LPB); Prov. Florida: N of Pena Colorada, ca. 1,300m, terrestrial on slopes with deciduous Chaco Serrano forest, 13 Oct. 1999, R. Vasquez, F. Quispe, et al. 3400 (VASQ); ibid.: 5 km of Mataral on road to Vallegrande, 18°09'S, 64°13'W 1,400m, 15 Dec. 1990, M. Nee 40291 (USZ); ibid.: 2,5 km NE of main highway at Mairana, quebradaYesera,l8°06'30"S, 63°57'W 260 m, 27 Jan. 1994, M. Nee & I. G. Yargas 44673 (USZ); ibid.: Refugio Los Volcanes, 3 Km al NE de Bermejo,l8°06' S, 63'36'W, 1,050m, 5 Oct. 1997, M. Kessler J. Gonzales, K. Bach & E. Rapp 12337 (LPB, SEL, GOET); Prov. A. Ibanez: 30 km east of Cotoca, between La Bola and the Rio Grande, 280m, in dry Chaco forest, Mar. 1990, R. Vasquez 1853 (VASQ); ibid.: 26 Oct. 1999, R. Vasquez, P. Ibisch, M. Nee et al. 3416 (VASQ); Prov. Ichilo: 13 km NE of Buena Vista on highway to Portachuelo, near turnoff to San Javier, 17°24'S, 63°34'W 300m, 21 May 1991, M. Nee 40504 (USZ); ibid.: 13 km NE of Buena Vista on highway to Portachuelo, 17°24'S, 63°34'W 300m, 21 May 1991, M. Nee 40504 (LPB); ibid.: 4 km N of Buena Vista on road to Laguna Madrejon,l7°25'S, 63°40'W 315m, 31 Oct. 1990, M. Nee 39672 (LPB); ibid.: road from Buena Vista to Portachuelo, 10 km NE of Buena Vista, 2 km W of turnoff to San Javier, 17'24'S, 63'35'W, 300m, 6 Dec. 1989, M. Nee 37979 (LPB); Prov. Nuflo de Chavez: Lomerio, 12 km N of Las Trancas community, l6°31'13 "S, 61°50'47"W 450m, 12 Dec. 1995, F. Mamani & A. Jardim (USZ); ibid.: Estancia San Miguelito, Pascana WCS camp, 17-45'S, 61-47'W, 400m, 26 Jul. 1995, A. Fuentes 1148 (USZ); Prov. Velasco: Jul. 1892 Kuntze s. n. (NY); ibid.: Noel Kempff Mercado National Park, Serraniade Huanchaca, 13°59'5,60°43'W 800m, 3 Dec. 1987, W. Thomas et al. 5561 (USZ); ibid.: Bajo Paragua Forest Reserve, 5 km from the sawmill on the road to Empalme,l4°32' S, 61'30'W, 400m, 8 Apr. 1994, M. Saldias et al. 3 758 (SEL).
HABITAT: The Bromelia species with the widest distribution. It is found in lowland dry forests, as well as in dry to semihumid inter-Andean valleys and the montane Chaco and the Tucumano-Bolivian forest, from 150m to 1400m.
OBSERVATIONS: As in B. hieronymi, the young stems are consumed by the indigenous people of the Gran Chaco. In the quechua-speaking areas the plant is known as "chaguar" while in "guarani dominated" areas the species is called “caraguata” or "garabata" (ARENAS & ARROYO 1988).
TYPE: Brazil. Glaziou 22191(holotype: B).
GEOGRAPHICAL RANGE: Brazil and Bolivia. In Bolivia department of Santa Cruz, province of Nuflo de Chavez.
Leaves forming a rosette; blades to 1m long, the apical ones intensely red; large sheaths forming a bulbous structure.
Inflorescence sessile, densely corymbose, with many flowers, densely whitish to brownish lanate.
Floral bracts spathulate, 18mm long, obtuse.
Flowers 5cm long.
Sepals 15-18mm long, free.
Petals to 2cm long, blue-violet, connate at base.
Ovary cylindrical. Fig. 8.
RECORDS IN BOLIVIA: SANTA CRUZ: Prov. Nuflo de Chavez, 75 km northeast of Concepcion, Palmarito community, S. Beck 25440 (LPB); ibid: between San Javier and Concepcion, ca. 400m, terrestrial in dry Chiquitano forest, 31 Jul. 2000, R. Vasquez, J. Rivero, ME. Vasquez & C. Vasquez 3 764 (VASQ).
HABITAT: The species grows in Cerrado and open dry Chiquitano forests, between 300 and 500m