Trithrinax campestris (Burmeist.) Drude & Griseb.

Cano, Ángela, Perret, Mathieu & Stauffer, Fred W., 2013, A revision of the genus Trithrinax (Cryosophileae, Coryphoideae, Arecaceae), Phytotaxa 136 (1), pp. 1-53 : 28-45

2. Trithrinax campestris (Burmeist.) Drude & Griseb. in Grisebach (1879: 283), ( Fig. 12 ).

Copernicia campestris Burmeister (1861: 48) . Type:— Burmeister, 1881, pl. 9, fig. 20; Epitype (designated by Krapovickas 2007): ARGENTINA. Córdoba: 3 km al E de Ballesteros, 12 March 1985, Krapovickas & Vanni 39776 (epitype CTES! ( Figs. 13–14 ), isoepitypes F!, MO).

Palms solitary or clustered with up to 6 stems. Stem 2–5 m tall × 20–30 cm diameter, greyish and often with marks of bush fires, leaf scars separated 5–8 cm; aerial roots forming a basal cone reaching up to 30 cm from the ground, the roots rather smooth, light brown, 10 cm long × 0.3 cm of diameter. Leaves 20–30, rigid; leaf sheath up to 74 cm long, dark reddish and loosely covered with a white-waxy indumentum at the base, greyish and glabrous towards the apex, leaf sheath spines up to 22, light brown, 3.3–21 cm long × 3–6 mm wide; petiole 17–43 cm long × 1.5–5.2 cm wide; hastula 1.2–2 cm long × 2.2–3.5 cm wide, rigid adaxially and ca. 1 cm long × 3 cm wide abaxially; leaf blade rigid, pale green, loosely covered with a white-waxy indumentum on both sides, glossy, leaf blade with symmetric or asymmetric base, regularly segmented, palman 0.4–3.0 cm (at the basal segments level) to 12–33 cm (median-apical segments level); leaf segments 20–34, with a bifid, woody and spiny asymmetric apex, basal segments 11–32.5 cm long × 0.5–1.8 cm wide, with an apical splitting of 0.5–7.8 cm, median segments 31–65 cm long × 1.8–4.2 cm wide, with an apical splitting of 2.6–13 cm, apical segments 45–76 cm long × 2–5.3 cm wide, with an apical splitting of 3.6–9.2 cm. Inflorescences 2– 5, about 30–50 cm long; peduncle up to 30 cm long × 1.5–2 cm wide, light brown; peduncular bracts 4, 27– 35 cm long × 5.5–8 cm wide, apically acute and rigid, at fruit stage apart from each other 9.5–10 cm; rachis 50– 53 cm × 0.4–2.5 cm wide at fruit stage, basal portion similar to the peduncle but pale green, medial and apical portions, pale green, the apical portion bearing up to 20 rachillae; rachis-bracts basal rachis bracts 16–21 cm long × 4.5–7 cm wide, medial and apical rachis bracts about 7 cm long × 6–7 cm wide, pale yellow, smooth; branches 5–6 per inflorescence; branches apart from each other 6–10.5 cm apart at fruiting stage, inserted at 10–50° with respect to the rachis axis, basal branches up to 40 cm long × 23 cm wide, medial branches up to 38 cm long × 20 cm wide, apical branches up to 26 cm long × 23 cm wide; rachillae 28–39 per branch, inserted every 0.1–2 cm, subtended by bracts 0.5–1cm long × 0.1–0.2 cm wide, basal rachillae 9–15 cm long × 0.2 cm wide at fruiting stage, inserted at 70–90° with respect to the branches axis, median-apical rachillae 4.5–12 cm long × 0.2 cm wide at fruiting stage, inserted at 30–70° with respect to the branches axis. Flowers inserted every 0.1–0.5 cm, each subtended by a 0.1–0.2 cm long × 0.1 cm wide floral bract; sepals 3 (rarely 5, Krapovickas & Vanni 33754, CTES), 2 mm long × 1–2 mm wide; petals 3 (rarely 5, Krapovickas & Vanni 33754, CTES), 3 mm long × 3 mm wide, not imbricate; stamens 6 (rarely 10, Krapovickas & Vanni 33754, CTES); filaments 6 mm long; anthers 2–3 mm long × ca. 1 mm wide; gynoecium 3-carpellate (rarely 5- carpellate, Krapovickas & Vanni 33754, CTES), carpels up to 3 mm long, ovary 1 mm long × ca. 0.5 mm wide, oblong, style 2 mm long × ca. 0.5 mm wide, stigma hook-shaped. Fruits 2.3–3 cm of diameter, pale yellow, mesocarp 5 mm wide, bitter ( Fig. 12 ). Beccari (1907) reported fruits 1.8 cm in diameter, with slightly lignified, fragile endocarp. Beccari (1931) published a drawing of the fruits based on Lorentz 28 (B) showing that in some cases all three carpels developed; seeds collected by us had a diameter of 1.7 cm and an intrusion seed coat about 7 mm long, whereas Beccari (1907) reported a diameter of 1.2–1.4 cm. We judge these differences to be taxonomically insignificant.

Etymology: — campestris : that is found in the countryside, referring to its usual habitat.

Distribution and ecology: —Central and northern Argentina, in the provinces of Chaco, Córdoba, Corrientes, Entre Ríos, San Luis, Santa Fe, Santiago del Estero and Tucumán ( Fig. 15 ). In Uruguay it is present in the departments of Paysandú, Río Negro ( Gaiero et al. 2012), San José and Soriano, and based on ancient collections the species could be also present in Colonia, although fieldwork efforts should confirm whether these populations are still alive. These palms are present in open hilly areas and in xerophyte plane Espinal forests of thorny trees and shrubs ( Lewis et al. 2009), often affected by anthropogenic fires ( Lewis et al. 2009, Lugo et al. 2011).

Specimens examined: — ARGENTINA. Córdoba: La Unión 3 km al este de Ballesteros, 5 April 1985, Ferruchi 381 ( G, CTES) ; La Unión, 3 km al E de Ballesteros , 153 m, 32° 32' 60''S, 62° 58' 60''W, 12 March 1985, Krapovickas & Vanni 39776, ( CTES, F) ; Entre Chilibroste y Noetinger al lado de la carrilera del tren, ruta 2, km 173, 117 m, 32°21'08.2''S, 062°24'32.7''W, 9 March 2011, Cano 277 ( G, SI) ; Río Segundo, Manfredi, Estación experimental INTA, 22 March 1978, Krapovickas & Vanni 33754 ( CTES) ; Serrezuela , 20 March 1978, Kiesling 1210 ( MO, SI) ; Cerca de Orcosu, Yerba buena etc., 17–18 February 1876, Hieronymus 398 ( P) ; Río Seco , 17 January 1972, Hunziker et al. 21737 ( CORD) ; Sobremonte , 30 January 1987, Hunziker & Subilis 24958 and 24960 ( CORD) ; Departamento de las Minas , February 1876, Hieronymus 412 ( CORD) ; Entre Ríos: Federal , em beira de rodovia, lado esquerdo ca. 200 km de Uruguaiana, BR, 6 March 2008, Tsuji 2481 ( HPL) ; La Paz , 14 km al sureste del acceso a Col. Avigdor, ruta12, 14 September 1986, Krapovickas & Cristóbal 40637 ( CTES, G) ; Villaguay, Mojones Norte , 15 March 1985, Pedersen 14081 ( MO) ; Concepción del Uruguay , 25 March 1875, Lorentz 879 ( CORD) ; San Luis: Ayacucho , 28 December 1885, Kurtz s.n. ( CORD) ; Santiago del Estero: Pozo Hondo, a la entrada de la Estancia La Donosa, km 790, 7 km antes de Pozo Hondo por la ruta 34, 270 m, 27°13'24.8''S, 064°27'51.0''W, 7 March 2011, Cano 276 ( G, SI) ; Tucumán: Leales, Los Puestos , ruta N °9, 1 January 1971, Krapovickas & Cristóbal 17399 ( CTES) ; Leales, Los Puestos, Ruta Nac N °9, 3 May 1971, Krapovickas et al. 18551 ( P) .

URUGUAY. San José: Camino de entrada a Kiyú por Libertad , antes del arroyo Mauricio , 12 September 1969, Del Puerto et al. 8671 ( MVFA) ; Soriano: Palmitas, 1938, Herter 615 ( G) ; Vera, March 1898, Berro 371 ( MVFA) ; Vera, March 1913, Berro 6661 ( MVFA) ; Unknown department: Banda oriental del Uruguay , 1816– 1821, Saint-Hilaire 2464 ( P) .

Taxonomic notes: — Trithrinax campestris was originally described as Copernicia campestris by Hemann Burmeister in 1861. He provided a short description of the palm and its habitat, but did not designate a type specimen. After the species was placed in the genus Trithrinax by Drude & Grisebach ( Grisebach 1879), Burmeister (1881) published a drawing of the palm habit in a complementary work, in which details of the leaves and reproductive organs were missing. Burmeister’s drawing was taken as the type of the species. In Beccari’s revision of Trithrinax ( Beccari 1931) , several herbarium specimens of T. campestris collected by Lorentz and Niederlein were cited, some of them deposited in CORD and FI. Krapovickas (2007) designated an epitype in order to complement Burmeister’s drawing; in doing so, he chose a collection gathered in the original locality specified by Burmeister (1861). The epitype collected by Krapovickas and Vanni presents all the vegetative and reproductive organs necessary to accurately identify the taxon.

Local names: —Caranday, carandá ( Báez 1933, del Vitto et al. 1994), carandá de Entre Ríos ( Glassman 1972), palma (del Vitto et al. 1994), saro, caranda-i, palma carandilla ( Cabral & Castro 2007).

Uses:— A mixture of sugarcane fibres and the cellulose extracted from this palm is used to fabricate cardboard boxes ( Luna 1977). The stems are used as posts and shingles due to the fact that plane boards can be produced from them ( Cabral & Castro 2007). The leaves are used to make brooms and brushes, and extracted fibres are used to make ropes or baskets; these fibres were formerly used to fabricate typical Argentinian sandals named alpargatas ( Cabral & Castro 2007). The reticulate leaf sheaths are used to filter the water and to make artisanal dolls and other handcrafts. The fruits are used to feed livestock ( Báez 1933, Cabral & Castro 2007) and also used to prepare an alcoholic beverage and oil. This species is also cultivated for ornamental purposes.

3. Trithrinax schizophylla Drude (1882: 552) . Lectotype (designated by Beccari 1931):— BOLIVIA. Santa Cruz: Chiquitos , September and October 1845, Weddell 3498. (lectotype P! ( Figs. 18–21 ), fragment ex P at FI!) ; Syntype: Amérique Méridionale, d’Orbigny s.n. ( P!) .

Palms solitary or clustered, up to 6 stems. Stem up to 5 m tall × 5.5–15 cm diameter, greyish to pale brown, leaf scars separated 2–3 cm; aerial roots forming a cone reaching up to 10 cm from the ground, the roots smooth to rugose, 1–10 cm long × 0.3–1 cm of diameter. Leaves 12–25, lax to rigid; leaf sheath 18–28 cm long, dark reddish, greyish and smooth towards the apex, leaf sheath spines light brown, 6–31 cm long × 1–4 mm wide; petiole 22–91 cm long × 0.5–3.2 cm wide; hastula 0.7–1 cm long × 1.3–1.8 cm wide adaxially and 0.5–0.7 cm long × 1.3–1.5 cm wide, bifid, abaxially; leaf blade, thin to thick, rigid to flexible, green, loosely covered with a white-waxy indumentum abaxially and adaxially, leaf blade with symmetric or asymmetric base, regularly segmented to biflabellate with a deep split at the middle of the blade forming two identical portions, palman 0.1–2.0 cm (at the basal segments level) to 2–32 cm (median-apical segments level); leaf segments 15–36, pendant to erect, with a bifid, acute or spiny apex, basal segments 22.5–57 cm long × 0.4–2 cm wide, entire or with an apical splitting of 0.8–49 cm, median segments 31.3–82 cm long × 0.8–2.8 cm wide, with an apical splitting of 1.2–49 cm, apical segments 34–84 cm long × 0.6–3.4 cm wide, with an apical splitting of 3.5–59 cm. Inflorescences 1–3, 35– 74 cm long; peduncle 10–36 cm long × 1.0– 3.3 cm wide, white to light brown; prophyll up to 9.5–23 cm long, fibrous, pale-brown towards the apex; peduncular bracts 4–5, 13.5–20 cm long, pale-brown, smooth towards the base and fibrous towards the apex, the peduncular bracts at fruiting stage apart from each other 4.5–7 cm; rachis 30–45 cm × 0.4–2 cm wide at fruit stage; basal rachisbracts 13–15 cm long, medial and apical rachis bracts 5.5–11 cm long; branches 5–6 per inflorescence; branches apart from each other 3.5–10 cm, inserted 10–70° with respect to the rachis axis, basal branches 20– 47 cm long × 11–22 cm wide, medial branches 19–32 cm long × 8–25 cm wide, apical branches 11–28 cm long × 8–21 cm wide; rachillae 37–75 per branch, inserted every 0.1–2.3 cm, bracts up to 0.5 cm long × 0.1– 0.3 cm wide, basal rachillae 8.5–18 cm long × 0.1–0.3 cm wide, inserted at 45–90° with respect to the branches axis, median-apical rachillae 4–14 cm long × 0.1–0.3 cm wide, inserted at 30–90° with respect to the branches axis. Flowers inserted every 0.1–0.8 cm; sepals ca. 2.5 mm long × ca. 1.5 mm wide, basally connate up to ca. 2 mm forming a cupule with three cuspidate tips; petals ca. 3.5 mm long × 4 mm wide, strongly imbricate; filaments 6 mm long; anthers 2 mm long × ca. 0.5 mm wide; gynoecium up to ca. 3.5 mm long, ovary 1 mm long × ca. 0.5 mm wide, ovate, style ca. 2.5 mm long × ca. 0.5 mm wide, erect or slightly curved. Fruits 0.6–1.2 cm of diameter, fleshy, pale yellow; seed 0.5–1 cm of diameter.

Etymology: — schizo: splitted, divided; phylla: leaf, in reference to the divided leaf blade.

Distribution and ecology: —From southern Bolivia and northern Argentina, to central and western Paraguay and southwestern Brazil ( Fig. 16 ).

Taxonomic notes: — Trithrinax schizophylla was described by Drude (1882) in Martius’ Flora Brasiliensis, where he designated two syntypes: a collection of Weddell 3498 (prov. Mato Grosso finibus et prov. Chiquitos), and d’Orbigny s.n. (inter prov. Boliviae orientalis Chiquitos et montes pr. S. Cruz de la Sierra). In his revision of Trithrinax, Beccari (1931) designated as lectotype of T. schizophylla the specimen of Weddell 3498 from P. His decision was probably based on the fact that the specimen Weddell 3498 perfectly corresponds to the definition of the species, whereas the specimen of d’Orbigny s.n. corresponds to a mixed collection of a leaf-sheath of Trithrinax sp. and a leaf of Copernicia alba Morong (1893: 246) .

Trithrinax biflabellata was described by Barbosa Rodrigues in his work Palmae Novae Paraguayenses (1899) , based on several unnumbered J. D. Anisits’ collections and one picture studied by him; in the same publication a detailed line drawing was proposed. Anisits’ main palm specimens were part of the Barbosa Rodrigues palm collections, which were destroyed by fire ( Stafleu & Cowan 1976). Other material collected by Anisits has been reported in BP and S ( Kümmerle 1933, Lanjouw & Stafleu 1954); unfortunately no palm material from this collector could be found in these herbaria (Z. Barina and M. Ehn pers. comm.). Consequently, the only original material available is the drawing presented in the original publication ( Fig. 23 ). The latter accurately represents most of the diagnostic characters of the taxon and hence designated here as the lectotype of the species, following the ICBN articles 9.11 and 9.12.

Trithrinax biflabellata was placed under synonymy of T. schizophylla by Henderson et al. (1995). We observed natural populations of both taxa ( Trithrinax biflabellata in the Paraguayan and Argentinian Chaco (type locality) and T. schizophylla in North-western Argentina), and we analysed a total of 41 herbarium specimens from a broad range of localities. As a result, we noticed several morphological and biogeographical differences that lead us to propose two varieties: T. schizophylla var. schizophylla and T. schizophylla var. biflabellata . The main differences are summarized in Tab. 2 and in the key to the varieties of T. schizophylla . A clear geographical differentiation for the two taxa is displayed in the distribution map including the two varieties ( Fig. 16 ).

Barbosa Rodrigues (1899), in his description of T. biflabellata , listed some differences between this taxon and T. schizophylla . However, these do not allow an accurate differentiation of the taxa. Barbosa Rodrigues cited as a distinguishing character the biflabellate leaves for T. biflabellata ; however, this leaf-segmentation type is present in both taxa, although more evident for T. biflabellata . Another character proposed by him was the presence of a single carpel in T. biflabellata versus a 3-carpellate gynoecium in T. schizophylla . Our study shows that all flowers dissected for both taxa consistently show 3 carpels. Barbosa Rodrigues was clearly aware of the existence of two independent taxonomic entities but, probably due to a lack of material and fieldwork, he could not sort out the variation in his description of T. biflabellata . The characters presented in Tab. 2 allow a better differentiation of the two varieties.

Key to the varieties of Trithrinax schizophylla

1. Robust palms; leaf blade segments (22–) 28 (–36), mostly lignified-spiny at the apex; plants distributed at 250–1400 m of elevation in SE Bolivia and NW Argentina (Salta and Jujuy) ................................................... var. schizophylla .

- Slender palms; leaf blade segments (15–) 20 (–26), mostly non-spiny at the apex; plants distributed at 100–150 m of elevation in Paraguay and NE Argentina (Chaco and Formosa) .......................................................... var. biflabellata .

3a. Trithrinax schizophylla var. schizophylla ( Fig. 17 ).

Palms robust; Stem up to 5 m tall; leaves rigid; leaf sheath spines 10–31 cm long; petiole 22–51 cm long × 0.7–3.2 cm wide; leaf blade thick, palman 0.5–2.0 cm (at the basal segments level) to 8.5–26 cm (medianapical segments level); leaf segments 22–36, rigid, with a bifid, acute, slightly spiny to lignified-spiny apex, basal segments 22.5–56 cm long × 0.4–2 cm wide, entire or with an apical splitting of 2.7–49 cm, median segments 36–82 cm long × 1.0– 2.8 cm wide, with an apical splitting of 15.5–49 cm, apical segments 39.5–84 cm long × 0.8–3.4 cm wide, with an apical splitting of 15.5–59 cm. Inflorescences 1–3. Flowers 4–5 mm long; sepals 1–2 mm long × 1 mm wide; petals 2 mm long × 1–2 mm wide; filaments 4 mm long; anthers 1–2 mm long × ca. 1 mm wide; gynoecium 2 mm long, ovary ca. 1 mm long × ca. 0.5 mm wide, style ca. 1–2 mm long × ca. 0.5 mm wide. Fruits were not observed at mature stage during the present study but Moraes (2004) described them as being yellow-green to black, 6–8 mm of diameter.

Distribution and ecology: —Distributed from southeastern Bolivia in the departments of Chuquisaca, Santa Cruz and Tarija, to northwestern Argentina in the provinces of Jujuy and Salta ( Fig. 16 ); it grows in xerophytic open areas and in mid-height thorny forests of the Chaco region, from 250 to 1400 m of elevation.

Specimens examined: — BOLIVIA. Chuquisaca: Sud Cinti, ca. de la Comunidad de Las Abras, cañón Piura , 1100 m, 21°05'34''S, 64°16'22''W, 23 October 2005, Lozano & Cardoso 1519 ( MO) ; Luis Calvo, municipio Villa Vaca Guzmán, Cantón Iguembe , Itaco , Trayecto Iguembe-Muyupampa , 1148 m, 20°41'S, 063°47'W, 19 October 2007, Jimenez et al. 552 ( MO) ; Santa Cruz: Chiquitos, September and October 1845, Weddell 3498 ( FI ex P, P); Chiquitos, Pozo del Tigre , 135 km al este de Santa Cruz, propiedad de la empresa ABAFA, alrededores del campamento principal, 250 m, 17°34'S, 061°57'W, 29 September 1994, Vargas & Ortiz 3200 ( F) ; Prov. Andres Ibanez , 5 km SE of Comunidad San Lorenzo, 320 m, 17°49'30''S, 062°50'W, 17 November 1990, Nee & Coimbra 39931 ( MO) ; west Velasco , July 1892, Kuntze s.n. ( NY) ; Provincia Cercado, Cantón-Lugar: Cotoca , Don Lorenzo , Bola , 400 m, 13 April 1990, Moreno 30 ( NY) ; 900 m, 20°07.550'S, 063°28.914'W, Gibbons & Spanner 19963 ( M) ; Tarija: Burdet O'Connor, Berety, 7.9 km (by road) from Entre Ríos to Palos Blancos , 1040 m, 21°26'01''S, 064°00'03''W, 26 March 2007, Nee & Flores 54963 ( MO) ; Burdet O'Connor, Rancho Saururo, 5 km S of highway to Villamontes, 30 km E of Entre Rios , 1200 m, 21°30'S, 064°00'W, 6 September 1987, Killeen 2721 ( NY) ; límite Prov. Chaco / Burdet O'Connor. Puerto Margarita, sobre el río Pilcomayo , 450 m, 25 October 1983, Beck & Liberman 9758 ( NY) ; Burdet O'Connor, Tacuarenda, 35 km al E de Entre Ríos , 1 May 1983, Krapovickas & Schinini 39081 ( CTES, G) .

ARGENTINA. Jujuy: Ledesma, Chalicán , ruta 34, 10 May 1971, Krapovickas et al. 18645 ( CTES, P, SI) ; San Pedro de Jujuy abt. 20 km N of San Pedro , 500 m, 24°02.250'S, 064°47.905'W, Gibbons & Spanner 19962 ( M) ; San Pedro de Jujuy, Depto. De Santa Bárbara, about 40 km south of San Pedro del Jujuy on Santa Bárbara road, all along the roadside for about 10 km, 1400 m, 8 October 1938, Eyerdam & Beetle 22848 ( G, K) ; San Ramón de la Nueva Orán, ruta La Estrella-Siete aguas, 123 km al S de San Ramón de la Nueva Orán, 730 m, 23°54'00''S, 64°20'18''W, 13 December 2001, Marcato et al. 328 ( SI, SPF) ; Depto. de Santa Bárbara, Palmeras , a 20 km de Vinalito, 22 October 1964, Cabrera & Fabris 16236 ( K) .

UNKNOWN COUNTRY: Amérique Méridionale [most probably Brazil], Orbigny s.n. (P)

Local names: —In Bolivia these palms are known as palma de sao or saó, palmerita, palma chica, saro, palma chuco , caryay-tí, caryay-mí ( Moraes 2004), utsaho ( Glassman 1972). In Argentina the common names are saro or saho ( Pingitore 1978).

Uses:— The leaves are used to thatch; the segments of young leaves are used to weave hats (sombrero de saó in Bolivia), small bags and fans. These palms are also cultivated for ornamental purposes ( Moraes 2004).

3b. Trithrinax schizophylla var. biflabellata (Barb. Rodr.) An. Cano & F.W. Stauffer comb. nov. ( Fig. 22 ).

Trithrinax biflabellata Barbosa Rodrigues (1899: 2) . Type:— Palm. Paraguay. Tab. 1; PARAGUAY. San Salvador, ad Arroyo Porongo prope Togatiyá et in Chaco inter flumina Pilcomayo et Negro, Anisits s.n. (missing). Lectotype (designated here):―Palm. Paraguay. Tab. 1 ( Barbosa Rodrigues 1899; Fig. 23 ).

Palm slender. Stem up to 5 m tall; leaves flexible; leaf sheath spines 6–21 cm long; petiole 31–91 cm long × 0.5–1.8 cm wide; leaf blade coriaceous-thin, palman 0.1–2.0 cm (at the basal segments level) to 2–32 cm (median-apical segments level); leaf segments 15–26, flexible, with a bifid, acute, non-spiny to slightly spiny apex, basal segments 26.5–57 cm long × 0.5–1.3 cm wide, entire or with an apical splitting of 0.8–43.5 cm, median segments 31.3–74 cm long × 0.8–2 cm wide, with an apical splitting of 1.2–43 cm, apical segments 34–80 cm long × 0.6–2 cm wide, with an apical splitting of 3.5–46.5 cm. Inflorescences 1–3. Flowers 7–8 mm longs; sepals 2–3 mm long × 1 mm wide; petals 3–4 mm long × 4 mm wide; filaments 6 mm long; anthers 2 mm long × ca. 1 mm wide; gynoecium 3–4 mm long, ovary 1 mm long × ca. 0.5 mm wide, style ca. 2–3 mm long × ca. 0.5 mm wide. Fruits were not observed in a mature stage during this work but Beccari (1931) described them of 1 cm in diameter and Lorenzi et al. (2010) reported 0.8–1.2 cm., the latter also measured seeds of 6 mm in diameter.

Etymology: — bi: two; flabellata: fan-shaped. Making reference to the divided leaf blade.

Distribution and ecology: —This palm is distributed in Paraguay and Argentina. In Paraguay it is present in the departments of Alto Paraguay, Boquerón, Presidente Hayes and known in the Department of Concepción only from the protologue. In Argentina it is present in the North-eastern provinces of Chaco and Formosa ( Fig. 16 ). A recently discovered population that may correspond to T. schizophylla var. biflabellata was reported from Mato Grosso do Sul, Brazil, near the frontier of Brazil and Paraguay (Soares pers. comm.), but an accurate identification of this palm is still needed. This variety grows in the lowland Chaco region, in xeromorphic open areas or thorny forests, in terra firme or periodically flooded areas, from 100 to 150 m of elevation.

Specimens examined: — PARAGUAY: Alto Parana: Camino a Fuerte Olimpo, entre Fn. C. A. Lopez y Fn. Torres, 15 May 1981, Krapovickas et al. 37435 ( CTES) ; Estancia Laguna Ganso , 22°34'3''S, 59°36''51'' W, 25 September 2002, Mereles et al. 8824 ( CTES) ; Boquerón: Campo Loro , 4 July 2003, Vogt 47 ( FCQ) ; Aproximadamente a 70 km de Filadelfia, terreno perteneciente a la Cooperativa Menonita , 116 m, 22°24'25.5''S, 059°20'38.2''W, 28 February 2011, Cano & Palacios 272 ( FCQ, G) ; Filadelfia, Col. Fernheim ( Río Verde ?) parque de recreo para la salud mental, 15 September 1990, Vanni et al. 2161 and 2162 ( G) ; Filadelfia Col. Fernheim, 10 km E de Campo Grande , 21°45'S, 059°W, 15 September 1990, Vanni et al. 2156 ( G) ; Filadelfia Coletado a cerca de 500 km de Ponta Porã por estrada e de 120 km em linha reta da fronteira com Brasil (região de Porto Murtinho-MS), 20 November 2009, Lorenzi et al. 6780 ( HPL) ; Newland km 412, ruta Carlos A. López, km 2 del cruce Loma Plata , camino a Newland 22°34'41.58"S, 59°49'52.23"W, 20 September 1989, Peña & Molas 221 ( PY) ; Tyto Neuland-Campo Aceval , 23 July 1992, Mereles & Degen 4553 ( FCQ) ; Entrada a la Estancia Pitiantuta , 21°16'13''S, 59°36'19''W, 8 February 2005, Vogt & Mereles 232 ( FCQ) ; Colonia Mennonita, Parcialidad Sanapana , 22°49’S, 59°49’W, Mereles 4162 ( CTES, FCQ, G) . Presidente hayes: Distrito Teniente Irala Fernández, Estancia Salazar, potrero 5 (aproximadamente 1 km después del peaje interno de la Estancia ), yendo hacia el norte, costado derecho de la carretera, 113 m, 23°04'42.3''S, 059°13'21.3''W, 27 February 2011, Cano & Palacios 271 ( FCQ) ; Ruta Fortín General Bruguez, entre km 50–55, 25 July 1995, Pietrobom da Silva & Zardini 2155 ( SPF) ; Monte Lindo , ruta trans-Chaco, km 210, 6 December 1995, Krapovickas & Cristóbal 44186 ( CTES, K, MO) ; Monte Lindo , 28 November 1988, Caballero-Marmori 1643 ( ITAIPU) ; Pozo Colorado, Estancia Victoria, ca. 40 km SE of Pozo Colorado, collection site ca. 3.5 km E of the headquarters for Estancia Victoria , 131 m, 23°43.763'S, 058°33.782'W, 23 November 2002, Noblick & Rios-Otero 5273 ( FTG) ; Ruta Transchaco , 21 km NW Tte. Irala Fernández, 10 May 1994, Krapovickas et al. 45257 ( CTES, G) ; De pozo Colorado , 93 km ante civitatem Concepción, 4 March 1980, Bernardi 20166 ( BM, G) ; Estancia La Perla , 23°26'S, 059°34'W, 14 October 1986, Pedersen 14637 ( G) ; Estancia Ñ, 24°16'21.0''S, 58°33'27.0''W, 1 February 2005, de Egea & Peña-Chocarro 768 ( BM) ; Estancia Santa Elisa , 17 September 1994, Zardini & Guerrero 41281 A ( NY) ; Estancia Vanguardia , 24°17'2.00''S, 58°39'11.0''W, 13 September 2004, de Egea et al. 498 ( BM) ; Pilcomayo river basin, trail to Fortín General Bruguez , 100 m, 24°55'25''S, 058°11'07''W, 4 June 2000, Zardini & Guerrero 54705 ( FTG) . Unknown department: Chaco, Hassler 2707 ( FI ex B); Paraguay, S. E. en montes (orillas), Hassler 12701 ( G) . ARGENTINA. Chaco: Castelli, General Güemes, 20 km al N de Asustado, Ruta 95 (camino de tierra), 24 April 1997, Castro 14 ( CTES) ; Depto. de Río Teuco, puente Saladillo, Ruta 95, a 115 km de Saenz Peña , 19 March 1952, Schulz 8302 ( K) ; Formosa: Las Mercedes , ruta 81, 28 km NW de Pirané, 8 June 1971, Krapovickas et al. 19570 ( F) ; 200 m, 24°36.635'S, 060°44.178'W, Gibbons & Spanner 19961 ( M) ; Dep. Patiño, Colonia Muñiz , a 8 km de Las Lomitas, 3 June 1985, Dell’Arciprete 27 ( CTES) ; Dep. Pirane, Loc. Palo Santo , 6 December 1972, Maruñak et al. 419 ( CTES) .

Local names: —In Brazil locals call this palm carandilla, carandai ( Lorenzi et al. 2010), or buriti ( Cabral & Castro 2007). In Paraguay it is known as karandilla ( Gauto 2009) and in Argentina as carandillo ( Pingitore 1978) or palma carandilla ( Cabral & Castro 2007).

Uses:— The stems are used as torches and in construction ( Hahn 1990, Gauto 2009), the palm heart or palmito is eaten and the leaves and the spines are used to make handcrafts such as baskets, fans and a variety of useful objects (Arena 1981, Gauto 2009). The leaves are also used for thatch or strong fibre ( Hahn 1990), and the fruits are eaten by local people ( Gauto 2009). This species is also cultivated for ornamental purposes ( Lorenzi et al. 2010).

Figures

FIGURE 12. Morphology of Trithrinax campestris. A. Habit (Santiago del Estero, Argentina), scale bar: 50 cm; B. Leaf blade, scale bar: 10 cm; C. Detail of hastula (adaxial view), scale bar: 1 cm; D. Segment apex, scale bar: 2 cm; E. Scanning electronic micrograph of anthetic flower, arrow pointing to the stigma, scale bar: 2 mm; F. Flower at full anthesis, arrow pointing to the stigma, scale bar: 2 mm; G.Infructescence showing fruits at different developmental stages, scale bar: 10 cm; H. Ripe fruit showing perianth remnants, scale bar: 1 cm I. CS of the fruit and the seed, scale bar: 1 cm; J. Seed, scale bar: 1 cm; K. LS of seed showing the homogeneous endosperm, scale bar: 1 cm.

FIGURE 13. Epitype of Copernicia campestris, Krapovickas & Vanni 39776 (CTES); barcode: CTES0000146. Credits: herbarium CTES. (1/2).

FIGURE 14. Epitype of Copernicia campestris, Krapovickas & Vanni 39776 (CTES); barcode: CTES0000147. Credits: herbarium CTES. (2/2).

FIGURE 15. Geographic distribution of Trithrinax campestris (yellow circles).

FIGURE 16. Geographic distribution of Trithrinax schizophylla and its two varieties T. schizophylla var. schizophylla (yellow circles) and T. schizophylla var. biflabellata (red circles).

FIGURE 17. Morphology of Trithrinax schizophylla var. schizophylla. A. Habit (Salta, Argentina), scale bar: 60 cm B. Leaf blade, scale bar: 10 cm; C. Detail of hastula (adaxial view), scale bar: 1 cm; D. Segment apex, scale bar: 2 cm; E. Inflorescence at full anthesis, note the fibrous-reticulate apex of the peduncular and rachis bracts (Krapovickas 18645, CTES), scale bar: 10 cm; F. Scanning electronic micrograph of the flower, arrow pointing to the stigma, scale bar: 1 mm.

FIGURE 18. Lectotype of Trithrinax schizophylla, Weddell 3498 (P), barcode: P00725553. Credits: herbarium P. (1/4).

FIGURE 19. Lectotype of Trithrinax schizophylla, Weddell 3498 (P), barcode: P00725554. Credits: herbarium P. (2/4).

FIGURE 20. Lectotype of Trithrinax schizophylla, Weddell 3498 (P), barcode: P00725555. Credits: herbarium P. (3/4).

FIGURE 21. Lectotype of Trithrinax schizophylla, Weddell 3498 (P), barcode: P00725556. Credits: herbarium P. (4/4).

FIGURE 22. Morphology of Trithrinax schizophylla var. biflabellata. A. Habit (Boquerón, Paraguay), scale bar: 50 cm; B. Leaf blade, scale bar: 10 cm; C. Detail of hastula (adaxial view), scale bar: 1 cm; D. Segment apex, scale bar: 20 cm; E. Inflorescence at late anthesis, scale bar: 10 cm; F. Scanning electronic micrograph of the flower, arrow pointing to the stigma, scale bar: 1 mm.

FIGURE 23. Lectotype of Trithrinax biflabellata reproduced from Palm. Paraguay. (1899) Tab. 1.