BONPLANDIA 16(1-2): 143-180 2007

STERCULIACEAE OF PARAGUAY II WALTHERIA

JANICE G SAUNDERS1

Summary: Saunders, J G 2007 Sterculiaceae of Paraguay II Waltheria Bonplandia 16(1- 2): 143-180 ISSN: 0524-0476

The biodiversity and geographical distribution for Waltheria species in Paraguay are presented to accompany the floristic treatment for the rest of the family given separately in Part I (Cristóbal, 2007) In this work eight species are accepted, described, illustrated, mapped, and a key is given to facilitate their identification Waltheria operculata belongs to sect Stegowaltheria, and the others are from sect Waltheria: W albicans, W carmensarae, W communis, W indica s l , W macrophylla, W procumbens, and W viscosissima Two species are endemic to Paraguay: W procumbens, restricted in distribution, and W macrophylla, collected only once and possibly extinct; both species are in need of local conservation assessment fieldwork

Key words: possibly extinct species, section Stegowaltheria, section Waltheria, homostyly, heterostyly, America

Resumen: Saunders, J G 2007 Sterculiaceae de Paraguay II Waltheria Bonplandia 16(1- 2): 143-180 ISSN: 0524-0476

Se presentan la bíodiversidad y distribución geográfica de Waltheria en Paraguay, que complementa el tratamiento floristico de la familia Parte I (Cristóbal, 2007) En este trabajo se aceptan 8 especies que son descriptas e ilustradas y se acompañan mapas de distribución y una clave para su identificación Waltheria operculata pertenece a la secc Stegowaltheria, y las otras a la secc Waltheria: W albicans, W carmensarae, W communis, W indica s l , W macrophylla, W procumbens, and W viscosissima Puede señalarse que existen dos espe- cies endémicas en Paraguay: W procumbens de distribucion restringida y W macrophylla, coleccionada sólo una vez y posiblemente extinguida Para ambas especies se sugieren tareas locales tendientes a su localización y conservación

Palabras clave: especies posiblemente extinguida, sección Stegowaltheria, sección Waltheria, heterostilia, homostilia, América

Waltheria L Sitella L H Bailey, Gentes Herb 4(9): 349 figs 222-223 1940 Linné, C, Sp Pl: 673 1753 Lophanthus J R Forst & G Forst, Char Gen Shrubs, shrublets, suffrutescent herbs, Pl: 27, t 14 1776 vines, herbs, unarmed, erect to prostrate Astropus Spreng, Neue Entdeck 3: 64 1822 Vestiture of stellate, simple, and glandular

1 Instituto de Botánica Darwinion, Casilla de Correo 22, B1642HYD San Isidro, Argentina E-mail: jsaunders@darwineduar

143 BONPLANDIA 16(1-2) 2007 hairs; glands sessile or capitate Leaves 35, 40) mm x 10-22(-27) mm; dehiscence stipulate, petiolate; laminas serrate, crenate, usually partially loculicidal, rarely bivalvate or dentate Inflorescences distal or disperse, or operculate Seed 14-25(-27, 30) mm x pedunculate or sessile, glomerulate, 10-15(-20) mm, more or less obovoid, composite or secund, compound cymose or obconical, at times obpyramidal and laterally with compound cincinni; primary cymes often 3-faced with sides curved, but not trigonal consisting of 4 bracts2 (these unequal or Embryo straight; cotyledons flat, equal) per floral pair, or 5-bracteate, or 3- suborbicular, cordiform, oblate; endosperm flowered Flowers sessile, rarely short- present pedicellate, perfect, distylous or homostylous Sepals 5, campanulate or tubular, 5-dentate, 5- A tropical and subtropical genus with ca lobed or 5-parted Petals 5, yellow, flat, 60 species, 53 of them American, and one basally clawed; claw adherent basally to species distributed worldwide Six species are staminal tube very slightly, essentially free, extra-American from either NW Australia, or up to one third of petal length (for 02-3 mm) Asia (both Hainan and Malyasia), or Androgynophore absent or reduced, minute Madagascar, or W Africa, or Maui (Hawaii), (07 mm) Stamens 5, opposite petals; or from both the Islas Marquesas and the filaments connate variously for basal third, Society Islands Phylogenetically these extra- halfway, mostly or throughout forming a American species appear to be scattered staminal tube, rarely essentially free; anthers throughout the genus (Saunders, 1993, 1995) 5, fertile, extrorse; thecae 2, parallel, oblong, Waltheria possesses two centers of diversity longitudinally dehiscent Staminodes absent and endemism: the largest in Brazil, a second or rarely present, vestigial, minute, and in Mexico In Paraguay, eight species occur denticulate Pistil 1 Stigma 1, elongate- One of the two endemics, W macrophylla, plumose (column elongate with branches may possibly be extinct Seven of the short), penicillate, (column null, branches Paraguayan species belong to sect Waltheria long, to 15 mm long), or capitate-penicillate One, W operculata Rose, belongs to sect (branches very short, 05 mm long) Style 1, Stegowaltheria K Schum (Schumann, 1886), lateral or eccentric Ovary 1; locule 1; ovules a group with the following synapomorphic 2 (1 abortive) Capsule usually shed together characters: operculate capsule dehiscence, with calyx, with 1 carpel, with 1 seed Capsule subcordiform foliose stipules, and verrucose, with calyx surrounding, with apex free, 2-32(- tuberculate seed testa (Saunders, 1995)

Key to the species of Waltheria in Paraguay

1 Stipules filiform, flexuose, rarely firm and linear-triangular; capsule bivalvate, the 2 valves separating completely 1 W communis A St-Hil 1' Stipules narrowly triangular, triangular, linear-triangular or subovate, never flexuose; capsule not bivalvate 2 Petioles 26-5 mm wide; larger leaves 10-18 cm wide 2 W macrophylla Hassl 2' Petioles < 20 mm wide, or rarely 25 mm wide; larger leaves 1-75(10) cm wide 3 Stems without stellate trichomes, strigose with simple trichomes to 25 mm long; capsule dehiscence operculate 3 W operculata Rose

2 Subsequent bracts described belonging to the primary cyme of floral pair

144 J G Saunders, Sterculiaceae de Paraguay II

3' Stems predominantly with stellate trichomes or glandular-capitate trichomes; capsule dehiscence partially loculicidal, open at apex and upper sides only

4 Leaves acuminate or rarely acute at apices 4 W viscosissima ASt Hil 4' Leaves rounded, obtuse or acute at apices 5 Plants prostrate or procumbent 6 Stems to 04 m long, stiff; flowers distylous; leaves glabrescent, often with black spots or black maculae 5 W procumbens JG Saunders & Soria 6' Stems 06-1 m long, very flexuose; flowers homostylous; adaxial surface of leaves tomentulose, red-spotted below trichomes 6 W carmensarae JG Saunders 5' Plants erect or mostly decumbent, or long lower stems decumbent 7 Capsule interior with endocarp thinly horny (corneous) apically only, for only 04-1 mm in length from apex; petals showy, 48-9 mm x (1)15-38(45) mm; petal lamina obdeltate or obtriangular, subglabrous; petal apical margin truncate with a few simple or V-shaped hairs; claw adherent to basal staminal tube for 07-12(16) mm 7 W albicans Turcz 7' Capsule interior with endocarp thinly to thickly horny (corneous) for half to most of the capsule length from the apex, for ca 08-25 mm; petals small, 3-4(-47) mm x 05-12 mm; petal lamina obovate, elliptic, oblong, rarely obtriangular; petal apical margin usually rounded or obtuse, rarely truncate, vestiture variable, glabrous to densely stellate; lamina adaxial surface vestiture variable, glabrous throughout, or hirsute on distal portion; lamina abaxial surface glabrous to densely stellate on nerves; claw adherent to stamen tube base for 02-08 mm 8 W indica L sl

1 Waltheria communis ASt-Hil Waltheria douradinha A St-Hil, Pl Us Bras 8, Figs 1, 2 and 3 pl 36 1825, syn nov Waltheria boliviensis Turcz, Bull Soc Imp Naturalistes Moscou 36(1): 571 1863, syn nov Saint Hilaire, A, Fl Bras merid 1: 155 1825 Waltheria communis var tomentella K Schum, Waltheria communis var vulgaris (St Hil sic, in Mart, Fl bras 12(3): 59 1886, syn nov sensu K Schum) K Schum, nom illeg Waltheria communis var platyphylla K Waltheria lanata A St-Hil, Fl Bras merid 1: Schum, in Mart, Fl bras 12(3): 59 1886, syn nov 154 1825 Waltheria communis var henningsii K Schum, Waltheria communis var lanata (A St-Hil) K in Mart, Fl bras 12(3): 59 1886, syn nov Schum, in Mart, Fl bras 12(3): 58 1886, syn nov Waltheria communis var dietrichii K Schum, Waltheria gracilis ASt-Hil, Fl Bras merid 1: in Mart, Fl bras 12(3): 59 1886, syn nov 154 1825 Waltheria prostrata K Schum, in Mart, Fl Waltheria communis var gracilis (A St-Hil) K bras 12(3): 56 1886, syn nov Schum, in Mart, Fl bras 12(3): 59 1886, syn nov Waltheria americana L form foliis elliptico Waltheria glabriuscula A St-Hil, Fl Bras lanceolatis K Schum & Hassl, in K merid 1: 155 1825 Schum, Bull Herb Boissier ser 2, 4: 72-73 Waltheria communis var glabriuscula (A St-Hil) 1903, syn nov K Schum, in Mart, Fl bras 12(3): 59 1886, syn nov Waltheria communis var hirta K Schum &

145 BONPLANDIA 16(1-2) 2007

Fig 1 Waltheria communis sl A: branch B: base of leaf with 2 stipules C: primary cyme showing 2 bracteoles and 4 bracts around flower pair D: calyx, opened in adaxial view E: thrum flower pistil F: thrum stamens, tube opened, flat G: petal, adaxial view H: seed, apical view (A-H, Cristóbal 2449) Drawn by Liliana Gómez

146 J G Saunders, Sterculiaceae de Paraguay II

Fig 2 Waltheria communis sl leaf variation in Amambay (A, C-H) and Caaguazú (B), Paraguay A-H: morphological variation from entirely hirsute or lanate to entirely cinereus-tomentellous; A: sparsely hirsute, surfaces red-brown cf var vulgaris, glabriuscula (Hatschbach & Kummrow 48524, CTES) B: densely hirsute, surfaces red- brown var vulgaris, (Schinini 5761, CTES) C: lanate, but lacking distal albo-tomentose leaves, surfaces red-brown cf var lanata (Ferrucci & al$ 710, CTES) D: tomentellous to base, proximal leaves darker above, var tomentella (Soria 5733, CTES) E: tomentellous, tomentous abaxially, concolorous, yellow-olive, var henningsii (Schinini 8994, CTES) F: distally floccose cinereus-tomentellous, basally glabrescent with surfaces red-brown cf var henningsii, cf var platyphylla (Cristóbal & al$ 2457, CTES) G: cinereus-tomentellous, darker adaxial surfaces proximally cf W$ boliviensis (Hatschbach & Callejas 47259, NY) H: cinereus-tomentellous, concolorous pale olive W$ americana var ellipticus lanceolatis (Hassler 7824, SI) Drawn by Francisco Rojas

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Fig 3 Geographical distribution in Paraguay of Waltheria communis sl (l)

Hassl, in K Schum, Bull Herb Boissier ser 2, 4: Xylopodium interior soft, orange or red Stems 73 1903, syn nov mostly 2-3 mm wide, rarely or at base 5-7 mm, Waltheria communis var velutina K Schum & often simple and decumbent, rarely suberect Hassl, in K Schum, Bull Herb Boissier ser 2, 4: or prostrate, at times decurrent from two 73 1903, syn nov stems adjoined; vestiture variable among Waltheria communis var paraibana Monteiro, populations and often within plant, hirsute, Rev Agronomica (Rio de Janeiro) 12(3-4): 197 lanate, tomentose, tomentulose, velutinous, 1953, syn nov glabrescent, successively or mixed, rarely predominantly one type; trichomes mostly Hemicryptophytes, suffrutescent herbs, stellate, rarely mostly simple below stem subshrubs to 6-90 cm tall or long when fertile apices Stipules 5-14 mm long, < 1 mm wide,

148 J G Saunders, Sterculiaceae de Paraguay II

(L:W ratio 40:1 to 10:1), filiform, flexuose, boliviensis) Pistil in thrums 35-57 mm long; rarely both linear-triangular and firm near in pins 57-95 mm long Stigma hemi- inflorescences or stipules all firm (Hassler globose, penicillate in thrums, penicillate or 7824) Leaves spiral, rarely distichous capitate-penicillate (var platyphylla, W Petioles short, 2-9 (15-30) mm long, 02-15 boliviensis) in pins Capsule (21)25-3 x (25) mm wide; abaxial veins prominent La- (17)21-22(24) mm, relatively broad among minas 06-14 cm x 06-92 cm, variable, species, obconical, enclosed within tubular almost always with some that are widest calyx but for apex; dehiscence bivalvate, above midlamina with their margins curved, separating completely along locule suture into then angled, then straight, and with their two separate valves; interior valve surface distalmost basal vein ascending and with yellow horny (corneous) endocarp terminating marginally at the distal third of present completely from apex to base Seed leaf; shape ranging from obovate to widely (18)22-25(3) x 14-16(18) mm wide, obovate, elliptic to widely elliptic, ovate to relatively broad among species, very widely ovate or widely depressed ovate, subobconical-obovoid; apex truncate or ovate-rhombic, ovate-obtrullate, obtrullate, obtuse with a narrow low ridge across the obrhombic; apices obtuse to rounded, rarely apex Chromosomes: Cristóbal (1967: 140- acute, at times apiculate; bases obtuse to 142) reports 2n=12 for A Cristóbal 18 (LIL!), rounded, rarely subcordate or acute; margins with the chromosomes illustrated, reported as serrate or doubly serrate; surface green fresh, W communis var tomentella Pollen: red-brown dried throughout; vestiture often reported as W communis and W douradinha variable between leaf faces or from baseplant in Köhler (1976) to shoot apex, pilose, hirsute, lanate, tomentous, velutinous, tomentellous or Phenology: Flowering year round, peaking glabrescent, commonly tomentellous- in October through January I have observed cinereous adaxially near inflorescences, less in D F, Brazil for the short hirsute plants, often more constant and tomentellous- var vulgaris, that flowers wilt around cinereus on both sides or throughout plant or midday, by curling under at distal lateral hirsute, lanate or glabrescent Inflorescences apices but not at mid-apices of petals leaving often few and distal, rarely numerous and dis- a pointed triangular apex showing, all petals perse; bracts linear, linear-triangular, lacking making a 5-pointed star pattern, unique within towards the inflorescence center Flowers the genus Also observed in São Paulo, and distylous; pin flowers with stigma exserted for illustrated by Ferri (1969: 218) 2-5 mm, with stamens not exserted; thrum flowers with stamens exserted only to 2 mm Vernacular names: Given as Waltheria Calyx lobes 3-ribbed by veins to apex douradinha: “Guaikuru” and “Typycha abaxially, with the veins between ribs obscure guaikuru” in Guaraní, the latter signifying lacking prominent areoles or non-areolate “broom (typycha) made with the guaikuru Petals ca 4-9 x 1-3 mm, bright yellow-gold, plant” on Estancia Sta Teresa in the province of star-like when wilted; lamina obovate, Corrientes, Argentina (Pedersen 82 C; obtriangular, opaque, venose; lamina major orthographic correction for “Tibicha-Guaycuru” veins 5, flat, adjacent basally and divergent at that was found on the original label and its two levels below midlength; apical lamina significance was given by pers comm, C margin rounded and densely stellate; adaxial Brizuela & V Vázquez Rodríguez, 2007); “ore- lamina base densely pilose; abaxial distal la- ja de carnero” that translates as “lamb´s ear”, in mina surface stellate; claw 10-26 mm long, Argentina (Bertoni 616 LIL, U); “Douradinha” slightly broad; claw vein strap-like, flat, in Sta Catarina, Brazil (Smith & al 9332 US); slightly broad Stamens in thrums 46-8 mm “douradinha do campo” in Rio Grande do Sul, long, in pins 3-63 mm long; staminal tube in Brazil (Reitz 515 CTES), in Uruguay as “yerba both 25-53 mm long; anthers 1-17 mm long, de las monjas” signifying in English “nun’s rarely with distal hairs (var platyphylla, W herb” in Herter (1957: 566)

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Medicinal uses: Morel & al (1999) report name “oreja do carnero” from Misiones, Ar- moderate antibacterial activity from the gentina; Bertoni 616 (U), reports “pasa por isolated cyclopeptide alkaloids waltherine-A tener propiedades medicinales” and waltherine-B, extracted from the bark covering the roots (xylopodia) of Distribution, habitat, and ecology: See note “douradinha” in Brazil Medicinal uses for below In Paraguay, common in eastern Para- the plants of W communis, formerly known as guay, scarce in western Paraguay; 50-740 m “W douradinha” have been reported by elevation; tropical and subtropical This Schumann (1886), and summarized and distylous species ranges from Northeastern expanded by Cristóbal (1983) Schumann Brazil to Uruguay and northern Argentina, (1886: 3, 12, 110) reports a decoction is used and westward to Bolivia (Santa Cruz) and Pa- as an antisyphilitic Cristóbal says Hoehne raguay Native to, restricted to and refers to Waltheria as being a popular therapy characteristic of “cerrado” sl where against bronchitis, coughs, to weaken tumors, common, except cerradão, occurring in higher to clean old sores, being emollients and level savanna “cerrado” usually not “maturativas”, and is high in steric oils and inundated, with or without low shrubs, or also mucilage Cristóbal further reports that the small trees; labels indicating “cerrado forest”, infusion is made from 20 grams of leaves for “cerrado-scrub”, “cerrado”, “tabuleiros”, each 1 liter of water, and a teaspoon is taken “campos”, “campo limpio”, “campo sujo”, 4-5 times a day Cristóbal cites St Hilaire that occasionally noted to be from “sabanas de the species is used to cure wounds, to treat espartillares”, or tall and low forest especially venereal diseases and respiratory ailments, after disturbance or fires (Zardini & Zavala and because of the mucilage, as an anti- 46740); usually on sandy or clay-sandy soils, inflammatory Cristóbal cites from the thesis red, white or black soils, very rarely from of Dr Manuel Cipriano D´Avila, “Thesis of inundated zones (Zardini & Tilleria 34521), Flora meridional do Rio-Grande-do-Sul” (p at times on serpentine rocks (Zardini & Gue- 65) that it is used medicinally as an infusion rrero 38629), or limestone (Zardini & Gue- for wounds externally, and internally for rrero 41261), or sandstone (Zardini & Aquino bronchitis, laryngitis, and “blenorrhagias” 34813) Plants often occur in disperse Corrêa (1984a: 541) lists under the common colonies, in groups of about 20-30 name “douradinha-do-campo” after a individuals Not usually rhizomatous One Scrophulariaceae (Lindernia crustacea collection in Parque Nacional Cerro Corá no- Benth) the second species W communis and tes unequal pin and thrum flowered plant the third species, W douradinha Waltheria ratios, with more pin-flowered plants communis is listed there as a stimulant, anti- (Ginzbarg & al 554), indicating self- dysenteric, sudorific, emetic, diuretic, highly incompatibility that is not strong recommended for bronchial cattarh and pulmonary disorders, cystitus and Material examined3: Alto Paraguay: Fuerte blenorrhagias Waltheria douradinha is used Olimpo, 14-X-1895, Anisits 2089 (S) Alto as an infusion internally for catarrh, and Paraná: Rva Tatí Yupí, 12-II-1980, Binacional externally to wash wounds, especially 595 (MO); Rva Tatí Yupí, 24-IX-1980, syphilitic irritations distributed in Binacional 563 (MO); Hernandarias, 12-X-1981, homeopathic remedies under the name Buttura 715 (MBM), Rva Biológ Tatí Yupí, 6-X- Stemodia arenaria, the name of another 1987, Caballero Marmori 1301 (CTES); 10 km different plant in the Scrophulariaceae NW de ruta Ciudad del Este-Salto del Guairá, ca- Corrêa notes that Saint Hilaire (1825) listed mino a Itaquyry, 25°01’S 54°59’W, 27-X-1994, these same properties as well as the high Cristobal & al 2470 (CTES); Tatí Yupí, 1901, amount of mucilage Cristóbal (1983) reports Riemel 52 (SI); 20 km N de Hernandarias, 10-I- that Bertoni 616 (LIL), cites the common 1974, Schinini 8116 (CTES, TEX); 4-5 km N de

3 Discrepant spellings of localities given as in Nicora & al 1988

150 J G Saunders, Sterculiaceae de Paraguay II

Hernandarias, Reserva Biológica Tatí Yupí, 24°22’S de Ruta 5, IX-1988, Mereles 1461 (G); ex-campa- 54°35’W, 8-X-1990, Schinini & Caballero mento, proximidades del arroyo Aquidaban Niqui, Marmori 26923 (CTES); Ea Sta Elena, Pira Pyta, albardón del arroyo, 240 m, 22°38’54’’S 25°17’S 54°35’W, 11-X-1990, Schinini & Caballe- 56°01’12’’W, 12-XII-2003, Múlgura de Romero & ro Marmori 27196 (CTES); 4-5 km N de al 3680 (SI); camino entre Ruta 5 y Bella Vista, 42 Hernandarias, Reserva Biologíca Tatí Yupí, km S de Bella Vista,15-V-1974, Schinini 8994 24°22’S 54°35’W, 13-X-1990, Schinini & Caba- (CTES); 32 km SE de Bella Vista, Ayo Negla, 200 llero Marmori 27290 (CTES); 24-26 km E de m, 21-X-1981, Schinini 21486 (CTES, TEX); Ce- Itaquyry, 250 m, 9-X-1995, Schinini & Caballero rro Corá, 16-VIII-1980, Schinini & Bordas 20172 Marmori 29975 (CTES); Réserva Tatí Yupí, (CTES); Colonia Yvypyté [Ybypyte?], 23°S Hernandarias, 6 km, 7-I-1985, Stutz 2141 (G) 56°W, 20-VIII-1980, Schinini & Bordas 20454 Amambay: pascuis ca Pedro Juan Caballero, 8- (CTES); Chiriguelo, 553 m, 22°40S 55°50W, 8- XII-1978, Bernardi 18950 (CTES, G, NY); Par- XII-1997, Schinini & Dematteis 33479 (CTES); que Nacional Cerro Corá, camino al cerro Mura- Colonia Estrella, 45 km NW de P J Caballero, 50 lla, 21-II-1994, Cristóbal & Krapovickas 2363 m, 22°18’S 55°50’W, 8-XII-1997, Schinini & (CTES); Ruta 5, alrededores de P J Caballero, Dematteis 33524 (CTES); camino a Colonia Estre- 23-II-1994, Cristóbal & Krapovickas 2372 lla, 1 km W de Hito 1/44, 50 m, 22°22S 55°45W, (CTES); alrededores de Pedro Juan Caballero, 10-XII-1997, Schinini & Dematteis 33635 ruta 5, 23-X-1994, Cristóbal & al 2457 (CTES); (CTES); Pedro Juan Caballero, 740 m, 8-II-1951 Parque Nacional Cerro Corá, cerca la Casa Fores- Schwartz 11805 (CTES); Parque Nacional Cerro tal, 8-II-1982, Fernández Casas & Molero 6102 Corá, camino a la Administración, XI-1992, Soria (MO, NY); 7 km N de Ruta 5, límite W del Parque 5733 (CTES); Pedro Juan Caballero, 740 m, 8-II- Nac Cerro Corá, 15-IX-1988, Ferrucci & al 710 1951, Schwartz 11805 (CTES); Parque Nacional (CTES, G, TEX); alrededores de Pedro J Caballe- Cerro Corá, 20-I-1981, Vavrek 122 (PY); National ro, 22°34’S 55°44’W, 14-III-1996, Ferrucci & al Park Cerro Corá, 350 m, 22°39’51’’S 917 (CTES); 27 km N de ruta 5, camino a Bella 55[56?]°19’18’’W, 10-XI-1999, Zardini & Báez Vista, 14-III-1996, Ferrucci & al 927-A and 927- 52279 (SI); National Park Cerro Corá, 22°41’10’’S B (CTES); Pq Nac Cerro Corá, 1 km al O de la 56°00’38’’W, 26-II-1997, Zardini & Guerrero rotunda y ca 2 km al SE de Ruta 5, entrada a 46444 ( SI) Caaguazú: cerca de Yhú, unos 4 km Aceite’i, Cancha Cue, 22°40’S 56°02’W, 16-II- al N, 13-XII-1982, Fernández Casas & al 7545 1987, Ginzbarg & al 554 (CTES, PY, TEX); Par- (MO); 5 km E de Caaguazú, Gregory 9602 (LIL); que Cerro Corá, 1 km al SE de ruta 5, entrada a in viciniis Caaguazú, II-1905, Hassler 8966 (G, K, Aceite’i Cancha Cue, 16-II-1987, Ginzbarg 555 NY, S, W); prope Caaguazú, III-1905, Hassler (PY, TEX); Ñuverá, Hassler 1384 (G); in regione 9097 (G, GH, K, MICH, MO, MPU, NY, P, S, W); cursus superior fluminis Apa, ad Paraguarium 5 km E de Caaguazú, Ruta 2, 8-II-1966, Septentrionalem, XI-1902, Hassler 7824 Krapovickas & al 12517 (CTES); Col Oviedo, (isotypes: NY!, SI! of W americana form follius Ruta 2, Km A140, 28-IX-1967, Krapovickas & elliptico lanceolatis K Schum & Hassl); in Cristóbal 13434 (CTES); 115 Km de Asunción, altoplanitie et declivibus Sierra da Amambay, Ayo Itay [Ita?], 15-XII-1986, Perez & al 1429 Lomadas Ponta Pora, XII-1907-1908, Hassler (PY); 22 km al S de Caaguazú, Estancia Itá Carú, 9750 (G, NY, W); Ponta Pora, X-1907-1908, 29-XII-1972, Schinini 5761 (CTES); 4 km N de Hassler 9779 (G); in altoplanitie Sierra de Yhú, 320 m, 14-XII-1982, Schinini 23040 Amambay, X-1912-1913, Hassler4 11384 (BAF, (CTES); 12 km N de Caaguazú, camino a Ihú, 200 G, MO, MPU, NY, S, W); Faz El Buracon, 30 km m, 25°20’S 56°W, 27-III-1993, Schinini & al O de Pedro Juan Caballero, 15-XII-1983, 28142 (CTES); Palomares, camino de Itaquyry Hatschbach & Callejas 47259 (C, CTES, MBM, [Alto Paraná] a Curuguaty [Canindeyú], ca 275 m, NY); Pedro Juan Caballero, 30 km O, 18-X-1984, 24°55’S 55°25’W, 10-X-1995, Schinini & Caba- Hatschbach & Kummrow 48524 (CTES); límite llero Marmori 30035 (CTES); East of Caaguazú, del Parque Nacional de Cerro Corá, al NE, 5 km 390 m, 25°23’47’’S 55°48’11’’W, 28-XII-1994,

4 BAF label collector listed as Hassler, not Rojas; see Schinini (2005)

151 BONPLANDIA 16(1-2) 2007

Zardini & Guerrero 42089 (MO, SI); collecting Cai, 120 m, 23°59’40’’S 55°28’45’’W, 11-VI- area ca 20 sq m J E Estigarriba (=Pastoreo), 1998, Zardini & Chaparro 46679 (MO, SI); 25°23’45’’S 55°48’07’’W, 29-XI-1995, Zardini & Mbaracayú Natural Reserve, Ñanduro Cai, 120 Guerrero 44071 (SI); Arroyo Cambay, 32 km S of m, 23°59’40’’S 55°28’45’’W, 11-VI-1998, Arroyo Yhú, 25°25’S 55°55’W,1- XII-1990, Zardini & Chaparro 48679 (SI); Mbaracayú Zardini & R Velázquez 24720 (CTES, MO); Natural Reserve, Ñanduro Cai, 23°59’39’’S Arroyo Cambay, northern side, 25°25’S 55°55’W, 55°28’44’’W, 31-X-1998, Zardini & Chaparro 12-I-1991, Zardini & U Velázquez 25983 49517 (MO, SI); Mbaracayú Natural Reserve, (CTES); Arroyo Cambay, northern side, 25°25’S vic Aguará-ñú, 24°11’01’’S 55°16’48’’W, 26- 55°55’W, 12-I-1991, Zardini & U Velázquez IV-1999, Zardini & Chaparro 50617 (MO, SI); 26020 (CTES, TEX) Caazapá: Distr Yuty, 20 Mbaracayú Natural Reserve, vic Ñanduro Cai, km E de San Miguel, desvió a 3 de Mayo y 23°59’39’’S 55°28’44’’W, 27-IV-1999, Zardini Capitindy, 11-IX-1987, Arbo & al 2883 (CTES); & Chaparro 50798 (SI); Mbaracayú Natural Plantae Paraguariensis 1885-1895, Ad marginum Reserve, Aguará-ñú, 24°11’01’’S 55°16’48’’W, silvae, pr Yuty, IX-1885-1895 Hassler 950 22-IX-1999, Zardini & Ramírez Benítez 51151 (NY); Santa Ursula, 55 km NE de Yuty, 180 m, (MO, SI); Mbaracayú Natural Reserve, 23-III-1993, Schinini & al 27809 (CTES); Tavaí, 24°11’16’’S 55°16’45’’W, 14-I-1998, Zardini & 250 m, 1-XI-1988, Soria 2542 (CTES, MO); Vera 47927 (SI) Central: 2 km SW del río Sala- Tavaí, 250 m, 1-XI-1988, Soria 2505 (CTES); do, camino de Limpío a Emboscada, 13-XI-1978, Capíí barý, 16-III-1951, Schwarz 12171 (CTES, Arbo & al 1619 (CTES, TEX); San Lorenzo, 24- LIL); between National Park Caaguazú and Tavaí, XII-1936, Archer 4768 (A, US); Lago Ypoa, 1 km N of Tavaí, 26°10’S 55°28’W, 25-XI-1997, 25°57’S 57°24’W, 26-XI-1989, Basualdo & Ortiz Zardini & Benítez 47571 (SI); Estancia Tapytá of 2571 (FCQ); Urbanización Acosta Ñú (entre S Shell Forestry Ltd, on trail to Yuquerí Creek, 150 Antonio e Ytoroó), ca rio Paraguay, 8-V-1994, m, 26°16’26’’S 55°45’47’’W, 14-XII-1999, Zardini Cristóbal & al 2422 (CTES); San Lorenzo, & Brítez 52875 (SI) Canendiyú: cerca Ypé Jhú, 1- Campus Universitario, 17-V-1981, Mereles 1018 XI-1978, Bernardi 18304 (G, MO, NY); ca 20 km (FCQ, G); Asunción, X-1888, Morong 215a (NY); post Ypé Jhú versus Capitan Bado, 12-I-1979 San Lorenzo, Campus Universitario, 17-V-1982, Bernardi 19492 (G); Ñanduro Cai,1-XI-1978, Ortiz 118 (FCQ); Jardín Botánico y Zoológico, [Bernardi] 18337 (G 253804); ca Curuguaty junto Trinidad, Asunción, Reserva Natural, 25°20S al Ayo Schiní, al borde de la carretera, 3-XI-1982, 57°28W, I-1991, Pérez 420 (MO); Jardín Botánico Fernández Casas & Molero 5886 (MO); Apépu, y Zoológico, Trinidad, Asunción, Reserva Natu- iter ad yerbales montium Sierra de Mbaracayú, in ral, 25°20’S 57°28’W, I-1991, Perez 513 (MO); regione fluminis Tapiraguay, VIII-1900, Hassler Jardín Botánico y Zoológico, Trinidad, Asun- 4340 (G, GH, K, MO, MPU, NY, S, W); iter ad ción, Reserva Natural, 25°20’S 57°28’W, III-1991, yerbales montium Sierra de Mbaracayú, in Perez 808 (MO); Paso Ñandeyara, camino del Botá- regione vicine Igatimí, IX-1900, Hassler 4736 (G, nico a Limpio, 1-V-1972, Schinini 4711 (CTES, G, GH, NY, W); iter ad yerbales montium Serra de TEX); Tarumandy-Camino Luque a San Mbaracayú, in altoplanitie et decliviis Sierra de Bernardino, 1-V-1972, Schinini 4733 (CTES); Mbaracayú in camis Ypé Jhú (Serra de Cerro de Areguá, borde el monte, 23-VII-1972, Mbaracayú) X-1900 Hassler 5026 (GH, NY, S, Schinini 5027 (CTES); Ruta 9, 2 km SE de Lim- W); Ypé Jhú, altoplanicie et decliviis Sierra de pio, Paso Ñandeyara, 9-V-1974, Schinini 8794 Mbaracayú, X-1898-99, Hassler 5026 (G) (CTES, TEX); 21 km NW de San Bernadino, 9- [syntype, ann by Schumann W communis var V-1974, Schinini 9194 (CTES); [as Dep Capi- platyphylla K Schum] (G); Ñanduro Cai, rumbo tal] San Lorenzo, Ciudad Universitaria, 6-IX- 54506/45015, 11-X-1996, Jiménez & Marín 1611 1976, Schinini 13564 (CTES, G); Tavarory, Ayo (CTES); Reserva Natural del Bosque Mbaracayú, Abaí near confluence with Río Paraguay, 25°30S 13 km S de Ypé Jhú, Ñanduro Cai, 3-XII-1997, 57°30W, 30-XI-1990, Zardini 24435 (MO); Schinini & Dematteis 33267 (CTES) ; 2 km de Tavarory, 07 km from administration toward Ñanduro Cai, 3-VIII-1987, Soria 1673 (CTES, Arroyo Abai, 25°30’S 57°30’W, 9-XII-1991, FCQ, G); Mbaracayú Natural Reserve, Ñanduro Zardini & Aquino 29332 (CTES, MO, SI);

152 J G Saunders, Sterculiaceae de Paraguay II between Ytoro and Tavarory, 2 km from Tavarory, (SI); Parque Nacional San Luis de la Sierra, Arroyo 25°30’S 57°35’W, 16-X-1991, Zardini & Aquino Tagatiyá-Guazú, 270 m, 22°40’20’’S 57°20’59’’W, 14- 29566 (CTES); Tavarory, 2 km from admin, toward X-1994, Zardini & Ramirez 41430 (SI); Estancia Arroyo Abai, 25°30’S 57°30’W, 9-XII-1991, Zardini Santa Maria, 22°43’50’’S 57°27’05’’W, 16-IX-1994, & Garcete 29251 (CTES); Tavarory, 2 km from Zardini & Vera 41235 (SI) Cordillera: Colonia admin, toward Arroyo Abai, 25°30’S 57°30’W, 9- Ojopoi, al E de Piribebuy, 3-VI-1985, Arenas 2931 XII-1991, Zardini & Garcete 29252 (SI); Estero del (CTES); Emboscada, Compañía, Ltd Trompo, 19-I- Ypoá, Villeta-Puerto Guyrati, 75 km S of Villeta, 1987, Bordas 4007 (CTES, FCQ); 7 km al NW de 25°34’S 57°33’W, 27-XI-1992, Zardini & Guerrero Altos camino a Emboscada, Cnia Bernal Cué, 33862 (MO, SI); Estero del Ypoá, Villeta – Puerto 25°15’S 57°16’W, 6-II-1987, Ginzbarg & al 420 Guyrati Between 7 km S of Villeta, 25°34’S 57°33’W, (TEX); in regione lacus Ypacarai, I-1913, Hassler 7-I-1993, Zardini & Guerrero 34383 (SI); Estero de 11460 (BAF, G, NY); Ruta 2, 10 km E de E Ayala, Ypoá, Villeta – Puerto Guyrati, 7 km S of Villeta, 26-IX-1967, Krapovickas & Cristóbal 13266 25°34’S 57°33’W, 7-I-1993, Zardini & Guerrero (CTES); [illegible, Tobatí ?], I-1969, Schinini 2294 34391 (SI); Tavarory, 1 km E of Río Paraguay, (CTES); 5 km E de Itacurubí, 8-V-1974, Schinini 25°30’S 57°30’W, 30-XI-1990, Zardini & Salinas 9172 (CTES, TEX); camino Emboscada -Nueva Co- 24444 (CTES, MO); Tavarory, 1 km E of Río Para- lombia, 25°10’S 55°15’W, 20-III-1989, Soria 3454 guay, 25°30’S 57°30’W, 30-XI-1990, Zardini & Sali- (CTES, SI); Valenzuela, “ubocaya guazim ty” nas 24509 (MO); Tavarory, 25 km from [mbokaja guasu ty: place where the larger cocote- administration toward Arroyo Abai, 25°30’S ros abound (pers comm, C Brizuela & V Vázquez 57°30’W, 16-XII-1991, Zardini & Tilleria 29477 Rodríguez, 2007)], 26-XII-1950, Sparre & Vervoorst (MO, SI); Estero del Ypoá, Villeta-Puerto Guyrati, 7 7284 (NY); Cerro Tobatí, 12-XII-1993, Tressens & km S of Villeta, 25°34’S 57°33’W, 18-XI-1992, al 4783 (CTES); 10 km E of Rio Salado, 25°09’S Zardini & Tilleria 33699 (SI); Estero del Ypoá, 57°25’W, 11-I-1991, Zardini 25801 (CTES);12 km E Villeta-Puerto Guyrati, between 115 km S of Villeta of Río Salado basin on road to Emboscada, 25°09’S and Puerto Guyrati, 25°36’S 57°36’W, 7-I-1993, 57° 25’W, 18-XI-1991, Zardini & Garcete 28804 Zardini & Tilleria 34521 (SI); border Tavarory- (SI); Eastern side of Río Piribebuy basin, 17 km W Acosta ñu, Paraguay River, 25°27’S 57°33’W, 10-VI- of Arroyos & Esteros, 25°08’S 57°15’W, 18-III- 1993, Zardini & Telleria [Tilleria?] 36086, (SI); 1989, Zardini & Velázques 16453 (CTES, MO); 10 Tavarory, Río Paraguay, 25°30’S 57°30’W, 17-XI- km E of Río Salado, 25°09’S 57°25’W,11-I-1991, 1990, Zardini & Velázques 24161 (MO); Tavarory, Zardini 25801 (CTES) Guairá: Villarrica, 6-X- near Administration, 25°30’S 57°30’W, 17-XI-1990, 1874, Balanza 2659 (G, K); Villarrica, 19-I-1919, Zardini & Velázques 24287 (CTES, MO); Jörgensen 3862 (DS, F, MO); Villarrica, 10-I-1928, Tavarory, Arroyo Abaí near confluence with Río Jörgensen sn (INPA); Villarrica, 10-I-1929, Paraguay, 25°30’S 57°30’W, 30-XI-1990, Zardini & Jörgensen 3862 (MICH, MO, US); Villarrica, IX-1932, Velázquez 24435 (LPB) Concepción: Estancia Sta Jörgensen 3862 (F); Villarrica, Jörgensen 3862 (SI); Maria, 29-I-1896, Anisits 2535 (S); Ruta 5, 35 km E Jörgensen 3862 (LIL); Azucarera de Tebicuary, Ayo de Concepción, 26-II-1994, Cristóbal & Ihacá, [ca 25°37’S 56°40’W], 12-I-1973, Schinini Krapovickas 2394 (CTES); San Rafael, in 5858 (CTES) Itapúa: Trinidad, 27°07’S 55°42’W 30- regione cursus superioris fluminis Apa, iter ad X-1994, Cristóbal & al 2471 (CTES); Capitán Miran- Paraguariam septentionalem, X-1901-1902, da, 42 km N del Hotel Tirol, 27°09’S 55°45’W, 30-X- Hassler 7861 (G, NY); Estancia Aguerito, 1994, Cristóbal & al 2473 (CTES); Capitán Miranda, Potrero Laurel, [not mapped]12-XII-1991, 47 km N del Hotel Tirol, 6-X-1993, Cristóbal & Mereles 4279 (CTES); Ruta 3, 45 km S de Yby- Krapovickas 2294 (CTES); as Dep Campichuelo, yau, 17-V-1974, Schinini 9072 (CTES); 62 km Capitán [Meza?], 18-XI-1949, Montes 7187 (SI); Isla al N del río Ypané, ruta 3, 220 m, 14-XII-1983, Yacyretá, 56°47’W 27°27’S, 25-XI-1988, Tressens & Vanni & al 225 (CTES, PY, TEX); Estancia Be- al 3386 (C, CTES, G); Yacyretá Dam Island Reserve: llo Horizonte, 22°47’50’’S 57°25’29’’W, 16-IX- Western area, 27°26’55’’S 56°49’23’’W, 22-X-1999, 1994, Zardini & Guerrero 41261 (SI); Estancia Zardini & Rodriguez 51648 (SI); Yacyretá Bello Horizonte, 90 m, 22°45’58’’S 57°26’00’’W, Dam Island Reserve: Western area, 27°26’55’’S 13-X-1994, Zardini & Guerrero 41327 56°49’23’’W, 22-X-1999, Zardini &

153 BONPLANDIA 16(1-2) 2007

Rodriguez 51716 (MO, SI) Misiones: several km Zardini 8718 and Zardini 8719 (CTES, G, MO); NW of San Ignacio overlooking Río Alto Paraná, 6- Estero del Ypoá, Paraguari-Carapeguá, 25°41’S II-1982, Anderson 12377 (MICH); 12 km W de San 57°10’W, 27-I-1993, Zardini & Aquino 34813 (SI); Ignacio, camino a Pilar, 15-XI-1978, Arbo & al National Park Ybycu´i, 6 km S of NW corner of 1798 (CTES); 7 km E de San Ignacio, camino a park, 26°04’S 56°46’W, 25-XI-1991, Zardini & Santa Rosa, 26-XI-1993, Arbo & al 6120 (CTES); Garcete 29140 (MO, SI); Est Lago Ypoá: Cerro San Ignacio, 22-X-1951, Burkart 18688 (SI); San Lima, 26°02’S 57°24’W, 22-XII-1993, Zardini & Pedro, 4-XI-1886, Niederlein 1198 (SI); Ruta 12, 3 Guerrero 37627 (SI); Estero del Ypoá, Lago Ypoá: km W de Ayo San Juan, 25-IX-1972, Schinini 5923 Taruma Fondo, along Lago, 25°56’S 57°26’W, 4-I- (CTES); Ea La Soledad, 3 km S de Santiago, 1994, Zardini & Guerrero 37762 (SI); road to Yeré, 56°46’W 27°10’S, 3-4-II-1988, Schinini & Vanni near Río Tebicuary, 26°22’S 57°09’W, 24-II-1994, 26080 (G) Ñeembucú: Paso de Patria, 17-XII-1978, Zardini & Guerrero 38629 (SI); National Park Eskuche 2533-11 (CTES, SI) Paraguarí: Salto de Ybycu´í, NW corner of park along Arroyo Mina, Piraretá, márgenes del arroyo Yhaguy Guazú, 14- 26°01’S 56°46’W, 25-XI-1991, Zardini & Tilleria XI-1978, Arbo & al 1704-Arbo & al 1705 (CTES, 28947 (CTES); National Park Ybycu´í, NW of park, TEX); Ruta 1, río Tebicuary, 15-XI-1978, Arbo & al 26°1’S 56°46’W, 11-IX-1989, Zardini & Velásquez 1790 (CTES); Lago Ypoá, 25°57’S 57°24’W, 15899 (MO); National Park Ybycu´í, 6 km S of NW 26-XI-1989, Basualdo & Ortiz 2571 (CTES); corner of park, along tributary of Arroyo Mina, 2 km N del puente sobre río Tebicuary, frente 26°04’S 56°46’W, 22-VI-1991, Zardini & Velázquez a Villa Florida, 10-X-1993, Cristóbal & 27837 (CTES) Presidente Hayes: Estancia Krapovickas 2299 (CTES); Cordillera de Al- Armonia, 18-I-1900, Anisits 1899 (S); Cerrito, a 8 km tos [ca 2 km N de Paraguari], XII-1900, de Benjamin Acebal, 20-XII-1973, Arenas 1254 Hassler 3621 (GH, K, NY, S, W); in campo (CTES); Estancia Salazar (Retiro primero) 22°59’S prov Chololo, in valle fluminis Y-acá 59°9’W, 12-II-1993, Pérez & al 2872 (CTES); Es- [AyoYhaguy?], X-1900, Hassler 6747a tancia Salazar (Retiro primero) 22° 59S 59° 9W, 14-II- (holotype: G!, isotypes: G!, MO!, NY!, S! of 1993, Pérez & al 2896 (CTES); Estancia Salazar W communis var hirta K Schum & Hassl); (Retiro primero) 22°59’S 59°9’W, 14-II-1993, Pérez Pirebebuy, in valle fluminis Y-acá & al 2898 (CTES); Estancia Salazar (Retiro prime- [AyoYhaguy?], XII-1900, Hassler 6747b ro) 22°59’S 59°9’W, 14-II-1993, Pérez & al (holotype: G!; isotype: G! of W communis 2906 (CTES) San Pedro: 36 km N de Tacuara, var velutina K Schum & Hassl); Frente a Ea La Manina, 24°22’S 56°24’W, 21-X-1994, Florida, orilla río Tebicuary, 4-II-1966, Cristóbal & al 2451 (CTES); 5 km W de San Krapovickas & al 12384 (CTES, TEX); Estanislao, camino a Rosario, 24°38’S Piraretá, 25-XI-1980, Lurvey 257 (MO); 56°28’W, 22-X-1994, Cristóbal & al 2453 Piraretá, 24-X-1981, Lurvey 354 (PY); (CTES); Ruta 5, 37 km de Tacuara, 1-III-1994, Piraretá, 29-X-1981, Lurvey 366 (CTES); Cristóbal & Krapovickas 2407 (CTES); iter Piraretá, 15-IV-1982, Lurvey 517 (PY); Lago ad “yerbales” Montium “Sierra de Ypoá, propiedad “Monitos SA” XII-1988, Mbaracayú”, in regione fluminis Corrientes, Mereles 1902 (CTES, G); Parque Nacional X-1900, Hassler 5834 (G 3 sheets, GH, MO, Yby- cu´í, 4-I-1986, Molas & al 674 (PY); ca MPU [pollen voucher in Kohler 1976], NY, S, 7 km before Piraretá from turnoff at W); 36 km N de San Estanislao, 18-II-1968, Piribebuy, 17-IV-1990, Saunders & al 3208 Krapovickas & al 13967 (LIL, CTES); Colo- (TEX); Piraretá-Cpia de Piribebuy, Salto nia Nueva Germania, 1902 or 1916, Rojas Piraretá, date not indicated, Schinini 4315 10151 (SI); Colonia Nueva Germania, XII- (CTES); cercano a Salto Piraretá, III-1972, 1916, Rojas 13544 (S); 4 km E de San Schinini 8773 (CTES); La Colmena Parque Estanislao, 14-V-1974, Schinini 8869 (CTES); Nacional Ybycui, 27-V-1979, Schmeda sn, Ruta 3, 18 km SE de San Estanislao, 14-V-1974, (FCQ); Col Piraretá, 15-XI-1950, Sparre & Schinini 8877 (CTES); Ruta 3, 10 km S del Río Vervoorst 207 (CTES); Parque Nacional Ybycu´í, Aguaray-Guazú, 14-V-1974, Schinini 8898 5 km N of administration building on road to César (CTES); Ruta 3 y río Tapiracuay, 19-V-1974, Barrientos, 26°03’S 56°50’W, 14-X-1988, Schinini 9086 (CTES); Yaguareté Forest

154 J G Saunders, Sterculiaceae de Paraguay II

(Sustainable Forest Systems site): around dietrichii by ovate leaves with erose margins asserradero, 210 m, 23°47’46’’S 56°12’41’’W, 12-III-1998, Zardini & Guerrero 48297 (SI); Obs 2: Hassler 7842, described as W Yaguareté Forest (SFS site): around asserradero, americana form folliis elliptico lanceolatis 210 m, 23°47’46’’S 56°12’41’’W, 12-III-1998, Zardini (Fig 2 H), possesses numerous inflorescences & Guerrero 48298 (SI); Yaguareté Forest (SFS along the stem as W indica, and firm but site): trail to Céspedes Cué, 182 m, 23°48’37’’S curved stipules linear-triangular throughout, 56°06’47’’W, 12-III-1998, Zardini & Vera 48209 both features otherwise found as variable (SI);Yaguareté Forest (SFS site): northern states within plants of W communis sl Its boundary, 23°46’49’’S 56°11’49’’W, 27-V-1997, ribbed petioles, its tomentulose leaves with Zardini & Zavala 46740 (SI);Yaguareté Forest uppermost basal veins extending to leaf tips, its (SFS site): around Aserradero, 23°47’46’’S distylous flowers, its calyx 3-ribbed, tubular 56°12’41’’W, 27-V-1997, Zardini & Zavala 46880 and tomentulose, and lacking areoles or (SI); Primavera, 21-IX-1954, Woolston 330 (K, sericeous hairs, its penicillate stigma and its NY) Department not indicated: [Guairá or bivalvate capsule indicate closer affinity to the Paraguarí?] Isla Alata, Tebicuary Mi, 17-XI-1978, broad variation in W communis sl than to Bernardi 18766 (G, MO); Sued Paraguay, IX- that of W indica sl 1892, Kuntze s/n (NY); Central Paraguay, 1888- 1889, Morong 201 (WIS, US); sin loc, X-1985, Obs 3: In Paraguay, floristic studies Ortiz 547 (G ex PY); collector unknown, loc indicate it occurs only in dry “campo” unknown, herb 77 (G); Cerro Mbatavi, 28-XI- (Chodat, 1901: 69) or “campo” (Jiménez & 1981, Soria & Zardini 1871 (MO) al, 2000; Basualdo & al, 2005: 93-94) Collections in Brazil indicate occurrence in Obs 1: Eleven varieties have been “campo limpio”, “campo sujo”, “cerrado”, recognized in Brazil, six previously noted for but never in “cerradão” because it is a Paraguay: Waltheria communis vars heliophyte (Saunders, 1995) In Brazil, W tomentella, platyphylla, lanata, vulgaris and communis is floristically listed for cerrado hirta and velutina (Schumann, 1886, 1903) (“cerrado” sensu strictu or “ cerrado”, or Schumann did not accept or reject W “campo limpio”) in central zones (Filgueiras boliviensis or W douradinha, but included & Pereira, 1990; Ferri, 1996; Batalha & original material of W boliviensis Mantovani, 2001), and further south, noted to unknowingly as var tomentella, and be “characteristic of and exclusive to” “cam- specimens like W douradinha with var pos” of central Santa Catarina according to gracilis Geographically isolated and Klein in Cristóbal (1983: 28) To the southwest, peripheral forms in its large geographical Cardoso & al (2000) indicate it comprises 20% range stablize for long or very short stem frequency of transect groundcover of native length and greater or lesser amounts of Elyonurus muticus grassland “campo” in an tomentose or hirsute vestiture The central unburned state in the Pantanal, Mato Grosso zone plants of its entire range show varieties The entire mapped distribution for W communis introgressing in São Paulo, Minas Gerais, shown in Saunders (1995: 45, 445) appears Goias, and in Amambay, Paraguay (Saunders, congruent with distribution of cerrado on the 1995) Amambay representatives (Fig 2) Planalto, except that it extends further south demonstrate the entire range of morphological beyond the distribution of cerrado portrayed in variation seen elsewhere, supplemented in Cavalcanti & Joly (2002: 354), or portrayed for Ferri (1969: 218), Correa (1984a: 541), Cris- Paraguay in Basualdo & Soria (2002: 12) If tóbal (1983: 27), Tressens & al (2002: 371) restricted and exclusive of campo as suggested and Cristóbal (2005: 350) Another variety above, then the distribution of W communis described not placed in synonymy here, W given here (Fig 3) suggests that distribution of communis var erosa Buxb, Oesterr Bot Z the cerrado biome (sensu Oliveira & Marquis, 73:122 1924 needs review of original mate- 2002) is more extensive than previously shown rial By description, it differs from var in Paraguay

155 BONPLANDIA 16(1-2) 2007

2 Waltheria macrophylla Hassl (bi-lobed) or truncate, with base tapering and Figs 4 and 5 attenuate to claw, membranous, glabrous or subglabrous, adaxial surface with glandular areas at base but midvein not raised; claw a Hassler, E, Bull Herb Boissier ser 2, 7: 725-726 little broad, flat, vein strap-like (dried), 1907 adherent to staminal tube for 05-07 mm Stamens in thrums 64-75 mm long and Shrubs, erect or spreading, 03-08 m tall exserted for 3-5 mm, in pins 42-53 mm long, Xylopodia gnarled; bark 2 mm thick with exserted for 2 mm; staminal tube short in transverse slits; interior tan, ligneous Branch thrums, 09-15(2) mm long, in pins stems 18-38 cm long, 3-10 mm diam, exceptionally short among other species, only to compressed from stems adjoined, slightly 05-06 mm Pistil in thrums 51-62 mm long, curved above or straight, golden woolly to exserted 15 mm, in pins 72-89 mm long and glabrescent at base with stellate trichome rays exserted 4 mm Stigmas very dimorphic, in pins 2-5, to 15 mm long; cortex surface dull obovoid, penicillate with branches spreading; in orange-brown Leaves distichously arranged thrums clavate-elongate-plumose, obtrulloid, Petioles 12-35(40) mm long, wide and compressed, complanar or flattened; basal compressed, (16)26-40(50) mm wide La- branches of thrums upwardly directed, minas orbicular-reniform, orbicular, widely decurrent, continuous with column; stigmatic elliptic, widely depressed ovate, widely branches in both pins and thrums short Capsule depressed cordiform, as wide or wider than and seed unknown Pollen: unknown long, (2)45-163 cm x (18)47-18 cm, at midstem abruptly large and very wide; apices Phenology: Flowering but not with fruit in obtuse-rounded, rounded, or sometimes 2- March Presumed to be matinal lobed or 2-parted at plant bases; bases cordate up to 25 cm, or rounded; margins slightly Distribution and habitat: Subtropical uneven, crenate-dentate; adaxial vestiture Endemic to Caaguazú, known only from the golden woolly and sessile glandular, type collection made in 1905 “Campis siccis, becoming moderately densely tomentose; campis combustis”, dry “cerrado” in pink and adaxial stellate trichomes septate with bases white pebbly, sandy soil; probably recurrent connate into a short stalk, scraping away after fires Fieldwork is needed for its easily; abaxial vestiture when young golden- conservation assessment, as it is potentially woolly on veins and paler, velutinous extinct or rare Hassler collections from the between veins, remaining densely stellate, same type locality habitat of W macrophylla soft; adaxial surface discolorous by for other Paraguayan endemics or species glabrescence, green, yellowish olive dried, limited in distribution offer additional slightly coriaceous but wrinkled localities to search for it: (Arbo, 1987: 16, 22, Inflorescences golden-woolly, secund Bracts 40, 61) unequal, obovate, unequally lobed, lanceolate, linear, abaxial trichomes V-shaped or simple, Material examined: Caaguazú: prope adaxial trichomes simple Flowers showy, ca Caaguazú in campis siccis or campis combustis, 5-11 mm diam, distylous, with stamens and III-1905 Hassler 9151 (holotype G!; isotypes G!, pistil exserted in both pins and thrums, with a GH!, LIL!, MICH!, MO!, NY!, P!, S!, W!) strongly melliferous odor if rehydrated Calyx 6-75 mm long, sericeous, tomentose; tube translucent; lobe veins prominent, with all 3 3 Waltheria operculata Rose major veins continuing from base to apex, Figs 5 and 6 smaller veins non-areolate Petals 5-10 mm x 15-3 mm, long-clawed (for more than midlength), pale lemon yellow; lamina apex Rose, J N, Contr US Natl Herb 5: 183 obtriangular with apical margin emarginate 1899

156 J G Saunders, Sterculiaceae de Paraguay II

Fig 4 Waltheria macrophylla$ A: branch B: thrum pistil C: pin pistil D: thrum flower E: 2 of 5 thrum stamens, petal bases F-G: bracts H: calyx opened, adaxial surface I: cut petiole, abaxial surface Drawn by Liliana Gómez

157 BONPLANDIA 16(1-2) 2007

Fig 5 Geographical distribution in Paraguay of Waltheria macrophylla (o), W$ operculata (n), W$ viscosissima (p), W$ procumbens (*), W$ carmensarae (u) and W$ albicans (l)

Turnera valleana Standl & L Williams, Ceiba 1: all resulting in a corymbiform or umbelliform 88 1950 shaped plant; strigose; trichomes simple 1-25 Waltheria excelsa Turcz var hirsuta Hassl, mm long, never stellate Stipules dimidiate- Addenda Pl Hassl 12 1917 Nomen nudum cordate (half heart-shaped) or rarely dimidiate-ovate (half-ovate), 22-11 mm x Annual herbs, upright to 60 cm high, (1)2-4 mm, broad among species Leaf lami- yellow-green Sterile parts with vestiture sim- nas: 1-5 cm long, 02-2 cm wide, oblong to ple, stellate hairs absent or rare Stems from narrowly ovate, obtuse at bases, acute at erect central axis, later with up to 8 basal apices, serrate at margins, setose or hirsute laterals bowed, with laterals curved and on surfaces; trichomes all or mostly simple, spreading from base, with apices dichotomous, rarely V-shaped or stellate; palmate basal

158 J G Saunders, Sterculiaceae de Paraguay II

Fig 6 Waltheria operculata$ A: plant B: stipule, abaxial face C: leaf, adaxial surface D: detail of leaf trichomes, adaxial surface E: compound cincinni, flower pairs removed F: calyx G: detail of stem H: corolla and stamens, together tubular I: pistil from homostylous (monomorphic pin?) flower J: seed, lateral view K: capsule showing strigose operculum at apex, membranous portion below, seed contour visible within (A, Saunders 1626; E, F, H-K, Saunders 1616, both from México; B-D, G, Fiebrig 1382) B-D, G: drawn by Liliana Gómez, A, E, F, H-K: drawn by Branton Whetstone

159 BONPLANDIA 16(1-2) 2007 veins not always obvious Flowers (Bolívar), Honduras (Choluteca), and homostylous or monomorphic pins with pistil México (Oaxaca) In Brazil from “caatinga”, extended above stamens Calyx lobes in fruit “chaco”, or thorn-scrub on shallow sandy, stage with veins multi-areolate and prominent pebbled soils, or old lake bed deposits, or river Petals small for section Stegowaltheria, ca 3- floodplains with well-drained sparsely covered 8 x 1-2 mm, pale yellow; lamina oblong, damp ground glabrous, membranous; claw short, thin Stamens 17-48 mm long; staminal tube Material examined: Alto Paraguay: Chaco (08)15-18(48) mm long Pistil 27-44 mm septentrionalis, 21°S, Olimpo Berg [= Fuerte Olim- long; stigma lanceolate, distended-plumose, to po], X-1907, Fiebrig 1382 (holotype of W excelsa 24 mm long; branches disperse to 02 mm Turcz var hirsuta Hassl: G!) Concepción: long Capsule 28-35 x 14-19 mm; Around Sargento José E López (“Puentecinho”), dehiscence operculate; operculum tardily shed 22°22’58’’S 56°53’50’’W, 16-I-2000, Zardini & by circumferally tearing away from the Guerrero 53927 (CTES) membranous transparent portion extending for 2/3 of capsule length from below the operculum nearly to base; operculum interior 4 Waltheria viscosissima A St-Hil with yellow hard thick bony (osseous) Figs 5 and 7 endocarp; endocarp lacking in rest of capsule; operculum locular suture tardily dehiscent or not after separation from the rest of the Saint Hilaire, A, Fl Bras merid 1(4): 150 capsule Seed 22-25 x 12-15 mm, 1825 obconical-obpyramidal shape, somewhat 3- Waltheria hirsuta Presl, Reliq haenk 2: 152 faced as a pyramid but not trigonal due to the 1835 sides being curved rather than flat; seed apex Waltheria tubiflora Klotzsch, Linnaea 14: 300 with an obtuse broad mucronate ridge with 1840 Nomen nudum two sulci on either side of ridge; apical view Waltheria mollis Willd ex K Schum, in Mart, of seed with contour oblong-rectangular; testa Fl bras 12(3): 53 1886 Nomen nudum verrucose, tuberculate Pollen: reported for Waltheria machrisiana L B Smith, Contr Sci Conzatti 3673 (US) in Köhler (1971) Nat Hist Mus Los Angeles County 23: 10 1958 figs 5-6 Phenology: Flowering in October, January in Paraguay; in Brazil flowering December- Shrubs, subshrubs, 07-2 m tall, either April, peaking Feb-April Flowers close in the erect, firm-stemmed, or weak, thin-stemmed middle of the morning and reclining or clambering Stems reddish, densely pilose-glandular or hirsute-glandular; Uses: None known in Paraguay In Piauí glandular-capitate hairs long and short, 05 – (Brazil) it is locally “grazed principally by 12 mm long; glandular-capitulae often red- horses, followed by burros, cattle, goats and violet, at times viscid with sticky exudates; sheep, that eat the stems, leaves, flowers and underlayer sparse, whitish, with stellate fruits” (Bona Nascimento 115, CTES) trichome rays sinuous and soft Leaf laminas: ca 2-11 x 1-7 cm, cordiform; apices Distribution and habitat: In tropical Pa- acuminate, less often acute; bases cordate to raguay from Alto Paraguay and Concep- rounded; margins serrate or dentate; tooth ción in cerrado forest and scrub on red apex single-setose; surfaces discolorous; sandy soil Common in Brazil (Bahía, adaxial surface green, appressed pubescent, Ceará, Goiás, Mato Grosso, Minas Gerais, pilose or glabrescent; abaxial surface Paraiba, Pernambuco, Piauí, Rio Grande tomentose, whitish, soft Flowers distylous, do Norte, Río de Janeiro), with sporadic with one sessile and one pedicellate in floral collections in Bolivia (Santa Cruz), pair; pedicels to 12 mm Calyx very glandu- Colombia (Santa Marta), Venezuela lar, basally rounded especially in fruit; lobes

160 J G Saunders, Sterculiaceae de Paraguay II

Fig 7 Waltheria viscosissima$ A: branch B: stem C: leaf tooth D: petal lamina without claw showing raised adaxial midvein and dark glandular areas at base E: pin flower, calyx base rounded, hairs not drawn, distal petals cut away, stigma curvation artifactual F: seed, lateral view showing circumferally sulcate apex (A-C, Krapovickas & Schinini 32432, CTES, from Bolivia; D-E, Saunders 3234 FAA material, CEPEC, from Brazil; F, Gentry 5077 from Venezuela) Drawn by Liliana Gómez

161 BONPLANDIA 16(1-2) 2007

much longer than the short tube; smaller lobe rich in “óleo etéro” and mucilage The infusion veins obscure between marginal veins and preparation and administration is summarized midveins; midvein and marginal veins in Cristóbal (1983) Pharmacological testing of extending to the apex Petals showy, 55-10 x the hydrobutanol phase of the ethanolic extract 15-5 mm, yellow-orange; laminas obdeltate, from aerial plant parts concluded that the membranous, glabrous, with adaxial surface cardiac and peripheral actions induced from with midvein raised at base and glandular their administration to rats were probably areas present, with base narrowly rounded or mediated by stimulation of cardiac and auriculate; claws long, thin, adherent to endothelial muscarinic receptors (Vasques & staminal tube for 2-3 mm Stamens in thrums al, 1999) 6-75 mm long, in pins 27-45 mm; staminal tube in pins and thrums 20-32 mm Pistil in Distribution, habitat and ecology: A thrums 38-52 mm long Stigma erect, slightly subtropical and tropical species, tropical in Pa- dimorphic, cylindrical (pins), ovoid (thrums), raguay from Amambay and Concepción, elongate-plumose, in thrums with branches probably below 500 m, occurring in scrub on longer, denser Capsule 3-32(4) x (18)22- cliffs of a table mountain (Zardini & Cardozo 27 mm, relatively broad among species, 44956) at 270 m Disjunct between South obtrulloid; exterior chartaceous; interior America and Mexico Common in Brazil thinly corneous only at apex for ca14 mm (Alagoás, Amazonas, Bahía, Ceará Distrito long and not below; dehiscence partially Federal, Espírito Santo, Goiás, Maranhão, loculicidal along locule suture at apex and Mato Grosso, Mato Grosso do Sul, Minas continuing on both sides a little, disintegrating Gerais, Pará, Paraiba, Pernambuco, Rondônia, secondarily and breaking into pieces below Tocantins) and Mexico (Jalisco, Nayarit, portion with hardened endocarp where wall is Oaxaca, and Sinaloa); sporadic in Guyana weak and thin Seed 21-27 x 14-2 mm, (Roraima), Bolivia (Santa Cruz), Colombia broad among species, subobovoid but (Magdalena, Santa Marta, Vichada), Vene- subobconical below upper third; apex zuela (Amazonas, Anzoátegui, Apure, Bolí- exceptional among species with an obtuse var, Guárico, Portuguesa, Sucré, Zulía) In mucronate ridge and circumferally sulcate Brazil, found bordering gallery forests in ce- below; cuticle with mosaic of brown and rrado, burned forest roadsides, in sandy, rocky yellow rectangular bands; testa smooth or clay soils Distylous forms are from South Pollen: reported in Köhler (1973, 1976); in America Homostylous forms are restricted to Melhem & al (1983) as W rotundifolia (see Mexico and a few collections in Colombia at Obs below) Santa Marta from the upper elevations, although distylous forms are found at the Phenology: Flowering December and June in lower elevations in Santa Marta (Saunders, Paraguay Flowers close shortly after midday 1995; Cristóbal & al, 2005)

Vernacular names: “malva-branca” in Material examined: Amambay: in altoplanitie Brazil (Corrêa 1984b: 43-45), “malva” near Sierra de Amambay, XII-1912/1913, Hassler 11160 Belem in Pará, Brazil (Archer 7816 NY), (G); Estancia Don Juancito, Cerro Ysau (grupo ce- “malva preta” (Silva 79 NY) in Pará, Brazil rros AQUA), 270 m, 22°40'12’’S 56°09'35’’W, 12-VI- 1996, Zardini & Cardozo 44956 (CTES) Concepción: Uses: None known in Paraguay A zwischen Río Apa und Río Aquidabán, 1908-1909, commercial bast fiber source in northern Fiebrig 4772 (E, G, GH, K) Brazil (Archer 7816, NY), used for forage and floristic improvement in Minas Gerais Obs: The inflorescence structure for (Pott 3398, CTES) Corrêa (1974, 1984b) Waltheria viscosissima was studied, described, states it has medicinal properties, and is used and illustrated by Bayer (1994: 32), reported as to treat bronchitis, coughs, soften tumours W rotundifolia based on Irwin & Soderstrom and clean old ulcers, or as an emollient being 6845 (SPF) from Brazil That voucher identity

162 J G Saunders, Sterculiaceae de Paraguay II is corrected here It also serves as the voucher apical suture; apex interior with thin white for pollen descriptions reported as W horny (corneous) endocarp or yellowish thick rotundifolia in Melhem & al (1976) bony (osseous) endocarp only at apex for 1 mm in length; wall below apex lacking hardened endocarp interiorly, rather 5 Waltheria procumbens JG Saunders & membranous transparent from outside to Soria within and for about 2/3 the length of entire Figs 5 and 8 capsule Seed 18-20 x 11-13 mm, subobovoid, with a conical projection at base; testa essentially smooth (only obscurely and Saunders, J G & N Soria, Darwiniana 43: 205- minutely verrucose) Pollen: reported for 209, fig 2 2005 Hassler 7842 (BM, MPU!, S!) as W prostrata in Köhler (1976) Shrubs, diminuitive, prostrate or procumbent, to 40 cm long when flowering Phenology: Flowering October through Branches with up to 4 orders of branching; March Flowers presumed to be matinal plantbase stems to 1 cm diam, very hard- wooded, with wood camel brown; non-basal Distribution and habitat: Endemic, stems virgate, thin, mostly 1 mm diam, stiff, apparently with restricted distribution, to the subapically terete, dark brown Vestiture district of Bellavista in the department of minutely stellate-puberulent and minutely Amambay in Paraguay, ocurring there along glandular; glandular hairs with capitulae often the upper rapids of the Apa river and in the red Stipules near inflorescences triangular region of the upper Aquidaban River Found Leaf laminas: to 5 x 3 cm, heteromorphic with in sand deposits in “cerrado” contiguous with margins entire or often unequally 5-7 lobulate, Brazilian “cerrado”, in “campo”, or “prados narrowly ovate to widely ovate, widely con rodales de bosquete, abundante ganado” elliptic; apices obtuse to rounded; bases (with livestock), up to 200 m Tropical rounded, subcordate; margins dentate, serrate, or biserrate, prominently glandular with costal Material examined: Amambay: a pocos km de vein thick abaxially; adaxial surface Bellavista en dirección a San Carlos [22°7’60’’S subglabrous, appressed setulose, glabrescent, 56°31’W], 12-II-1982, Fernandez Casas & heterotrichous simple or stellate; abaxial Molero (JC#) 6238 (NY); in arenosis in surface stellate-pubescent; either surface regione cursus superioris fluminis Apa, iter ad often with black stains or black spots Bracts Paraguariam septentrionalem, XI-1901-1902, unequal, the widest one broadly ovate or Hassler 7842 (holotype: G!; isotypes: A!, BM, C!, elliptical Flowers distylous, showy due to G!, LIL!, MICH!, MO!, MPU!, NY!, S!, W!); petals exceeding calyx length by 2-5 mm Potrero Guaraní, 12 km SE de Administración, Calyx exceptionally short among species, 3- Estancia Santa Teresa [ca 16 km SW by road of 35 mm long Petals ca 5-9 x 1-4 mm, pale Ea Zapallo, 22°37’26’’S 56°38’5’’W], 20-III-1991, yellow, membranous, subglabrous; claw thin Soria 4458 (CTES); Estancia San Victor [ca 16 km Stamens in thrums 43-52 mm long, in pins W along road 3 km S of Charagua from road Bella 22 mm; staminal tube short, in thrums and Vista-Yby Yaú, 22°37’60’’S 56°28’W], Potrero Toro, pins 12-13 mm long Pistil in thrums 3-34 25-X-1991, Soria 4930 (CTES) mm long, in pins 54 mm Stigma oblong, elongate-plumose Calyx lobes in fruiting stage with very prominent veins; veins in 6 Waltheria carmensarae J G Saunders basal third of lobe with the laterals arched Figs 5 and 9 inward to the midvein, with a vertical vein spur at the arch apex; lobe midvein alone extending to apex Capsule 2-25 x 12-18 Saunders, J G, Darwiniana 43: 202-205, fig 1 mm, dehiscence partially loculicidal across 2005

163 BONPLANDIA 16(1-2) 2007

Fig 8 Waltheria procumbens$ A: bracts of primary cyme B: petal, adaxial face C: capsule, lateral view, with darker seed visible through membranous base below chartaceous apex D: seed, lateral view E: seed, apical view F: style and stigma of pin flower, a few stylar branches removed in front G: upper calyx tube and lobe of one segment, without hairs drawn, adaxial surface H: calyx with capsule, hairs not drawn I: stipule, adaxial view J: plant portion, prostrate, flowers closed (A-J, Hassler 7842) Drawn by Francisco Rojas Figures reproduced from Saunders (2005)

164 J G Saunders, Sterculiaceae de Paraguay II

Fig 9 Waltheria carmensarae$ A: branch, prostrate B: homostylous flower pistil C: homostylous flower D: portion of upper calyx, adaxial view E: stamens, pistil apex, and one petal, others removed above adherent base F: petal, adaxial face G: capsule cut in half, interior view, showing shaded corneous portion at apex and clear membranous portion below apex H: seed, lateral view I: seed, apical view J: stipule, adaxial view K: 4 bracts of primary cyme, abaxial view (A-K, Cristóbal & Saunders 3213) Drawn by Francisco Rojas Figures reproduced from Saunders (2005)

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Subshrubs, procumbent vines, 06-10 m sandy soil (Schulz 7700) In Argentina where long when flowering Xylopodium present, more known, occurring in “matorrales”, “pal- soft-wooded, interior tan Vestiture on stems, mares”, palm savannas, savannas, or “campos petioles, leaves, minutely tomentulose, altos arenosos” in deep riverine sand dunes, trichomes stellate, and abundantly and more médanos, loose riverbank sand, or very sandy prominently minutely glandular-capitate, to 0 loam, usually from 45-70 m 5 mm long, capitulae often red Stems very flexuose, yellow-green, or reddish beneath, dull Material examined: Itapúa: Yacyretá Dam Island brown; below apices compressed and Reserve, western area, 27°26’55’’S 56°49’23’’W, 22- decurrent for 6-10 nodes Leaf laminas ca 1-7 X-1999, Zardini & Rodríguez 51651 (CTES, SI) cm x 1-6 cm, ovate to widely ovate, ovate- Ñeembucú: Paso de Patria, 10-XII-1950, Schulz elliptic; apices widely obtuse to rounded; ba- 7700 (CTES, LIL, TEX) ses subcordate to rounded; margins shallowly dentate and glandular-prominent; teeth with costal vein thickened beneath; adaxial 7 Waltheria albicans Turcz surfaces stellate-pubescent or puberulent, red- Figs 5 and 10 veined and red below stellate trichomes; abaxial surfaces stellate-tomentulose Primary cyme bracts unequal, the widest one narrowly Turczaninow, P K N S, Bull Soc Imp ovate or narrowly elliptical Flowers Naturalistes Moscou 31(2): 214 1858 homostylous Calyx lobes multi-areolate veined Petals ca 5-8 x 1-2 mm, yellow, Suffrutescent herbs 02-12 m tall when obtriangular, subglabrous, membranous; claw flowering, parts yellow-green, whitish from thin, adherent for 15-2 mm Stamens 25-43 trichomes Stems all decumbent, or with a sin- mm long; filament tube 2-36 mm Pistil 34- gle mainstem erect and with the laterals 47 mm long Stigma oblong, elongate- decumbent and long from base and shorter plumose Capsule 25-32 x 14-2 mm, toward the apices; compressed for ultimate 4- partially loculicidally dehiscent at apex along 6 nodes; indumentum very soft along most of suture and along upper sides slightly; stemlength, sericeous, velutinous, puberulent, hardened endocarp white slightly horny densely glandular-capitate; glandular hairs (corneous), present only shortly at apex inte- short and long, to 12 mm long, with their rior for ca 04 mm in length, absent below; capitulae more often pale yellow than dark subapical wall mostly membranous yellow or orange Leaf laminas: ca 1-10 cm x transparent below Seed 20-23 x 12-15 mm, 1-9 cm, variable in shape and vestiture among oblong-obovoid, base obliquely obtuse populations, cordiform, widely ovate, ovate, Pollen: reported in Saunders (2005) widely depressed ovate, narrowly ovate, rarely elliptical-oblong; apices obtuse, Phenology: Flowering October and rounded, less often acute; bases cordate, December in Paraguay, but in adjacent Argen- rounded; margins dentate; adaxial surface tina, with more collections, during September densely to sparsely appressed sericeous, through June, peaking from March-April pubescent; adaxial trichomes stellate and sim- Flowers close by midday ple (rarely all simple), some glandular- capitate; rays of those stellate hairs often less Distribution and habitat: Subtropical and than 7-8 in number, often unequal in length; tropical Restricted to Mesopotamia along the abaxial surface vestiture denser, paler, Parana and Uruguay Rivers in northern Ar- stellate; adaxial surface glaucous, with folds gentina and adjacent Paraguay, except for one or not and flat, only plicate at edges disjunct collection from the middle Orinoco Inflorescences very sericeous with the River in Venezuela In Paraguay found in trichomes in white tufts among bracts, densely dunes with palm savannas and forest islands glandular-capitate Bracts of the primary (Zardini & Rodríguez 51651) or on hills with cyme unequal with the widest bract ovate,

166 J G Saunders, Sterculiaceae de Paraguay II

Fig 10 Waltheria albicans$ A: plant B: stem C: thrum flower D: 2 thrum stamens, dissected, showing adherence of two petal claw bases (upper portion cut off) with base of stamen tube E: capsule, exterior view showing the clear membranous portion below the upper shaded chartaceous portion exteriorly matching where corneous inside F: seed, lateral view G: bracts (three bracts and a wider prophyll) of flower pair, adaxial view (A-B, Cristóbal & al$ 2434; C- G, Cristóbal & al$ 2444) Drawn by Liliana Gómez

167 BONPLANDIA 16(1-2) 2007 ovate-elliptical Flowers distylous or 2444, Mereles & Degen 5706 and 6191), in homostylous, showy with distally broad petals undegraded (Cristóbal & al 2433 and 2434) 1-3 mm longer than calyx Calyx lobe veins in or degraded chaqueña open-forest-savanna fruiting stage very prominent, with the margi- with Schinopsis heterophylla and nal veins arched and anastomose to midvein in Aspidosperma quebracho-blanco, with the lower third of lobe, with the veins at Bumelia excelsa and Prosopis spp (Vanni & lobebase between marginals lacking areoles al 2192, in cultivated zone; Schinini & Bor- or with only one pair, with midvein above arch das 15050 and 15088), xerophytic forest compressed with the midvein edges uneven up borders (Mereles 6547 and 6587), open to the apex by its decurrent anastomosed forest with Bromeliaceae (Cristóbal & smaller veinlets Petals 48-65(90) x (11)15- Krapovickas 2387), in transitional zones 30(45) mm, pale yellow; laminas obdeltate, between chaqueña-chiquitania forests altered obtriangular, subglabrous, membranous, with pastizal or forest borders, sometimes with apical margin truncate with only 2-3 hairs that Lonchocarpus nudiflorens and Schinopsis are V-shaped or simple; adaxial midvein cornuta (Pérez de Molas & al 8608), or prominent from lamina base to midlength; claw disturbed forests borders at roadside (Cristó- thin, adherent for 07-12(16) mm to staminal bal & al 2455) Common in Brazil from dry tube base Stamens in thrums 45-72 mm long, “caatinga”, naturally occurring on rocky in pins 3-4 mm long, in homostyles 25-39 mm slopes with pebbly, sandy, soil from 300-500 long Pistil in thrums 25-41 mm long, in pins (1000) m Sporadically found in (with some 42-68 mm long, in homostyles 29-4 mm long citations for new country records included in Stigma cylindrical, elongate-plumose Capsule parentheses) in Bolivia (Chuquisaca, Santa 2-3 x 1-16(2) mm; dehiscence partially Cruz: Nee 47723 (NY); La Paz: Beck 4739 loculicidal; interior with thin white horny (CTES)), Argentina (Salta), Guyana (corneous) endocarp only at apex for less than (Rupununi: Goodland 931 (NY), Schomburgk 1 mm of length (04 mm in distylous plants), 67 and 500), Venezuela (Bolivar: Holst & van with the hardened endocarp lacking below; der Werff 2537, TEX), Colombia (Cali: Ra- subapical wall membranous transparent mos 689), and México (Oaxaca) Flowers are comprising 2/3 of entire capsule length, when distylous in Brazil, Paraguay, and Guyana and touched with dissecting needle without are homostylous in Paraguay and Argentina consistency or resistence; base more consistent, chartaceous, green or tan Seed Material examined: Alto Paraguay: Línea 19, 18-23 x 1-14 mm, tilted oblong-subrhomic 32 km N del cruce Lagerenza-Agua Dulce, with a conical projection at base Pollen: 19°59'04’’S 60°03'168’’W, 14-IV-1997, Mereles reported in Köhler (1976) 6587 (CTES); Parque Cué, 20°07' 217’’ 61°41'14’’W, 31-III-1995, Mereles & Degen 5978 Phenology: Flowering September and (CTES pin p-s=14); Tyto Lagerenza-Agua Dulce, December through May in Paraguay, peaking 19°59'00’’S 60°32'08’’W, 18-V-1996, Mereles & in May For Brazil, Paraguay and Bolivia Degen 6351 (CTES); Cercanias de Base Aerea combined flowering all year, peaking in Feb- Adrián Jara, camino a Ea Karenina,14-V-2003, Mar and July Flowers are open until midday Pérez de Molas & Navarro 8608 (CTES); Mayor Pedro Lagerenza, 20°S 60°45'W, 6-IV-1978, Distribution and habitat: In Paraguay Schinini & Bordas 15050-A and 15050-C (CTES, occurring in the drier, western part of the Cha- ex parte con W indica); loc cit Schinini & Bordas co Boreal in consolidated dunes or sandy soil 15050 (BAB); Mayor Pedro Lagerenza, [182 m], below 500 m Found in virgin “campo” 20°S 60°45'W, 16-IV-1978, Schinini & Bordas arborescent sandy high savanna (“espartillar”) 15088 (pin CTES) Amambay: 5 km N de ruta 5, with Tabebuia aurea, Shinopsis balansae, camino a Bella Vista, 26-II-1994, Cristóbal & Astronium sp , Elyonurus sp , Aristida sp Krapovickas 2387 (CTES); 45 km E de Yby Yaú, and Eragrostis sp , (Pérez & al 2630), 23-X-1994, Cristóbal & al 2455 (CTES) Boque- “espartillares” (Cristóbal & al 2441, and rón: Fortín Nueva Asunción, N del aeropista mili-

168 J G Saunders, Sterculiaceae de Paraguay II tar, [54 m] 20°43'S 61°56'W, 24-III-1986, Brunner flowers of stable distylous populations, except 1628 (PY, pin); Ruta Transchaco, 29 km SE de Nue- departing from the usual 02-03 mm long free va Asunción, 13-V-1994, Cristóbal & al 2433 filament length (CTES); Ruta Transchaco, 25 km SE de Nueva Asunción, 13-V-1994, Cristóbal & al 2434 Obs 2: Several examples appear to be (CTES); Ruta Transchaco, 25 km SE de Nueva intermediate with W indica Vanni 3494 by Asunción, 15-V-1994, Cristóbal & al 2441 its small petals, only 5 x 1 mm, its short claw (CTES); Parque Valle Natural, 8 km de Colonia only 1 mm long, its leaf trichome ray number Neuland, 22°36'S 60°05'W, 13-III-2005, Fortunato of 7-13, is like W indica, but its greater petal & al 8777 (CTES); 40 km S Linea 10 4ta División adherence to staminal tube for 15 mm is of Tyto a Agropil, 26-V-1994, Mereles & Degen W albicans Secondly, Cristóbal & al 5706 (CTES); Cercanías del Fortín Platanillos, 27- 2455 by its petals hirsute basally on the X-1995, Mereles & Degen 6191 (CTES); Estancia adaxial surface, and a few different capsules San Ramón, 22°41'30’’S 60°30'W, 14-XII-1992, with extended corneous endocarp (to 14 Pérez & al 2630 (CTES); Colonias Mennonitas, mm), is like W indica, but other capsules are proximidades de Isla Poi, 22°33'22’’S 59°45'08’’W, similar to those of W albicans, with short 2-V-95, Vanni 3494 (CTES); Cnía Fernheim, corneous endocarp restricted to the apex Filadelfia, 14-IX-1990, Vanni & al 2133 (pin Thirdly, Cristóbal 3954 has trichomes on CTES, TEX); 9 km S de Loma Plata, Propiedad de adaxial leaves with stellate ray number Ebenfeld, 25-II-1991, Vanni & al 2192 (pin representative of both species, many with 10 CTES); Parque ‘Valle Natural’, 12 km S de rays like W indica as well as 4-8 like W Filadelfia, 128 m, 22°20'S 60°01'60’’W, 28-II-1991, albicans, with very large leaves to 8 cm Vanni & al 2478 (thrum CTES); Cnía wide, almost twice the normal size for W Mennonitas, Campo Loa Nasue, 80 km W de albicans, with adaxial sericeous trichomes Filadelfia, IV-1997, Vanni & López 3954 (CTES); twice the size of normal ones in W albicans between Parque Nacional Teniente Agripino is peculiar for its small very long pedunculate Enciso and Nueva Asunción, 20° 54'21’’S suborbicular or ovate-orbicular leaves at the 61°50'27’’W, 27-I-1995, Zardini & Acosta 42265 plant base and anthers indehiscent with low (SI); (ex Nueva Asunción Department) between amounts of pollen Fourthly, Cristóbal & al Parque Nacional Teniente Agripino Enciso and 2455 has a extensively corneous capsule like Nueva Asunción, 20°54'14’’S 61°50'46’’W, 27-I- W indica, but large apically broad petals like 1995, Zardini & Acosta 42361 (SI) Presidente W albicans Hayes: Isla Poí, 22°28'S 59°40’’W, 16-V-1994, Cris- tóbal & al 2444 (CTES) 8 Waltheria indica L sensu lato Obs 1: Waltheria albicans is distylous in Figs 11 and 12 Brazil, with larger corollas, anthers and a much longer stigmatic column Homostylous populations apparently not stable for organ Linné, C, Sp Pl 673 1753 type species reciprocity length are found in Paraguay, with Waltheria americana L, Sp Pl 673 1753 difference between length of pistil and stamens widely ranged but usually not Subshrubs, erect, 05-15(2) m tall, variable spatially separated, including one collection among populations Taproot 1 cm diam with stamens longer than pistil (Vanni 2133), Stems mostly 2-3 mm diam, to 10 mm at base, see below Only Mereles & Degen 5978 is a terete below apices, tomentose, cannescent- pin, with 02 mm separation between pistil tomentulose; stellate trichomes at times with and stamen Vanni 2133 “thrum-homostyle” purple residue inside; glandular-capitate hairs with anther base touching stigma tip, differs absent or short with their capitulae pale by stamens 38 mm long, stamen tube 24 mm yellow, dark purple, or rose Stipules firm, long, free filaments 10 mm long, linear-triangular, linear, to 2-8 mm long measurements falling within the range of pin Petioles 2-65 cm x 1 mm, oblate in

169 BONPLANDIA 16(1-2) 2007

Fig 11 Waltheria indica sl A: flowering branch B: homostylous flower C: 2 segments of the upper calyx tube opened and two lobes, in adaxial view D: adaxial petal, showing the adherent base of the petal claw to the base of the stamen tube (without anthers) E: pistil F: capsule exterior, showing darkened chartaceous portion G: capsule cut in half, interior in lateral view, showing darkened corneous endocarp H: seed, lateral view (A, E-H, Schinini 1859, CTES; B-E, Saunders 1854 liquid preserved material from Brazil) Drawn by Liliana Gómez

170 J G Saunders, Sterculiaceae de Paraguay II

Fig 12 Distribution in Paraguay of Waltheria indica sl (l) transsection, adaxially canaliculate Leaf lami- olive, abaxially paler, whitish-olive or nas generally to 8 cm x 4 cm, 11-8(-14) cm yellowish-olive; basal palmate veins long, 05-6 (-95) cm wide, narrowly ovate to terminating at margins at or below midlamina very widely ovate, elliptical, obovate, oblong; Inflorescences at numerous leaf nodes, disper- apices acute, obtuse, rounded; bases se glomerules, sessile or subsessile, peduncles subcordate, rounded, obtuse, acute; margins to 35 cm long Bracts uniform in shape and serrate, dupliserrate, large and small serrate; width, linear-lanceolate, at times narrowly adaxial face tomentose, tomentulose, with ovate Flowers homostylous Corolla not stellate trichome rays (5-)8-13(-18); abaxial much exceeding calyx apices Calyces 33- vestiture denser; surfaces concolorous, 50 mm long, tomentose, sericeous entirely or grayish, or discolorous, dark adaxially, gray or on veins; lobe veins in fruiting stage multi-

171 BONPLANDIA 16(1-2) 2007 areolate with 4-6 areoles, at times fewer herbs as an anti-inflammatory for aspirin-like Petals pale yellow, with center darker or not, qualities (Abbott & Schimazu, 1985; Neal, small, 3-4(-47) mm long, 05-12 mm wide; 1965) Morton (1981) lists its internal use for lamina obovate, elliptic, oblong, rarely diarrhea, sore throat, fever, pain, rheumatism, obtriangular; apices usually rounded, obtuse, bladder ailments, blood enrichment, febrifuge, rarely truncate, often with many stellate hairs abortifacient, contraceptive, its external use at margins; adaxial surface glabrous or pilose for edema, ulcerated wounds, and its bark fiber distally; abaxial surface glabrous or stellate- used to make cordage, sugar sacks, and vested; claw adherent to staminal tube base sandals variously in Latin America Pais & al for only 02-08 mm Stamens 2-4 mm long; (1968) have isolated and identified the filament tube 18-3 mm long; anthers small, structures of the cyclopeptide alkaloids, 05-08 (11) mm long Pistil equal to or slightly adouetines X, Y, Y´ and Z found in W indica, longer than stamens, 25-5 mm long Capsule reported as W americana The pharmacologically 2-3 x 1-17 mm; dehiscence partially active flavonol, kaempferol-3-O-galactoside, loculicidal; interior with endocarp thinly horny has been noted from W indica (Petrus 1990) (corneous) for half or more of length, for 08- Reed (1974) found it to be the major alternate 25 mm from apex, consistent and resistant to host plant of an important cotton pest the touch with a probe Seed 14-25 x 07-13 (Lepidoptera: Noctuidae: Earias biplaga) due mm Chromosomes: 2n=40 (Harvey, 1966:160 to higher resistence of Earias in DDT sprayed from Sierre Leone, Kamabai for Harvey 94, areas than the usually more damaging cotton K) as W americana and in Harvey (1967:66) boll weevil pest Heliothis armigera in East revised to W indica; 2n=26 (Krishnappa & Africa, suggesting eradication of W indica in Munirajappa, 1980: 536, as W indica); n=20 Tanzania for pest control as W indica (Gill, 1978: 22 from Tanzania, Dar-es-salaam for Gill 90 DSM); 2n=24 Distribution and habitat: Global (Bahadur & Sidhu, 1975 in Love, A Taxon distribution in subtropical and tropical zones 24: 501-516 1975, as W indica), 2n= 24 Roadside weed, in old pastures, cotton fields, (Bahadur & Srikanth, 1983: 103, 105 from and secondary growth, and disturbed areas, in India, provenant from Chilkur Forest Reser- rock crevices on top of plains, in “matorral”, ve, cultivated in experimental garden of “campos altos”, “orillos de montes”, forest Kakatiya University, Warangal, voucher not borders or slopes, in “picada de la selva”, in indicated) as W indica See also Note 1 acaulous palm cerrados, “campo cerrados”, below Pollen: variable, reported in Köhler cerrado scrub forest, inundated savannas, (1971, 1976) riverbanks, in sandy, clay, disturbed or impoverished soils, on limestone or basalt Phenology: Flowering year round, but rock outcrops, from 150-500 m primarily May-June and again in October- February Flowers are open until late Material examined: Alto Paraguay: 1 km al E afternoon de línea 10, por línea 3, alrededor de una aguada a 100 m del camino, 19°14’S 61°2’W, 27-X-1992, Vernacular names, uses and economic Fortunato, Ramella & Palese 3649 (BAB); importance: None are known for Paraguay Destac 4 de Mayo, Picada Otazú, tyto Cerro Martínez (1979), Morton (1981), and Corrêa Léon, s/f-II-1989, Mereles & Ramella 2620 (1974) list many vernacular names in Latin (CTES); Campo Grande al norte por linea 28 America, but Corrêa (1974) gives the only Chaparrales transicionales del cerrado sobre one, “malva-veludo”, for South America substrato calcareo, assoc Tabebuia (Brazil) Corrêa (1984b: 43-44) lists it as the selachidentata-Terminalia argentata, 132 m, second species (as W americana) for “malva- 19°46'48’’S 58°46'54’’W, 10-V-2003, Pérez de Molas branca” used as an antisyphilitic In Hawaii, & Navarro 8201 (CTES); Ea Campo Grande al W indica is considered to be one of the ten norte por linea 28, Bosques chiquitanos de transi- most recognized and locally used medicinal ción al Chaco sobre suelos biendrenados,

172 J G Saunders, Sterculiaceae de Paraguay II assoc Simira aff Sampaioana-Acosmium camino de Limpio a Emboscada, 13-XI-1978, Arbo cardinaspii, borde de bosque, 151 m, 19°44'22’’S & al 1617 (CTES); E de Limpio, 27-IV-1985, Bordas 58°48'07’’W, 11-V-2003, Pérez de Molas & Navarro 3945 (CTES); Capiatá-Barrio Anahí, 14-V-1985, 8324 (CTES); Picado Otazú (Oeste) [20°19’S Bordas 4042 (CTES); Río Salado, 35 km al E del 60°28’W] entre 4 de Mayo y Cerro León, 27-II- pueblo Ypacaraí por la Ruta 2, 25°24’S 58°46’W, 25- 1989, Ramella & Mereles 2413 (CTES); Mayor IV-1985, Brunner 1072 (CTES); in regione lacus Pedro Lagerenza, 20°S 60°45’W, 6-IV-1978, Ypacarai, IV-1913, Hassler 12196 (BAF, NY); Jar- Schinini & Bordas 15050 (CTES); Cerro Léon, dín Botánico, I-1913, Pavetti & Rojas 10307 20°26’S 60°15’W, en picada de la selva hacia el (CTES); Tarumandy-Camino Luque a San cerro, 30-IX-1979, Schinini & Bordas 17843 Bernardino, 1-V-1972, Schinini 4731 (CTES); Lim- (CTES); Parque Nacional Defensores del Chaco: pio-Piquete-Cué, 25°07’S 57°30’W, 8-VI-1992, Madrejón Area, 20°27’32’’S 59°47’25’’W, 15-II- Zardini & Aquino 31800 (CTES); Estero de Ypoá 1999, Zardini & Godoy 50484 (CTES) Alto Pindoty, top of meseta, 150 m, 25°37’S 58°31’W, 17- Paraná: Reserva Biológia Itabó, sendero VI-1992, Zardini & Aquino 32060 (CTES); Estero Tangará, ca 80 km N of Puerto Stroessner, 252 del Ypoá, Villeta-Puerto Guyrati, 105 km S of m, 24-V-1989, Krapovickas & Cristóbal 43201 Villeta, on Arroyo Pikysyry, 25°36’S, 57°34’W, (CTES); Estancia Bertoni (Rio Paraná), 25°38’S 26-X-1992, Zardini & Guerrero 34196 (CTES); 54°36’W, 22-VII-1994, Zardini & Florentin border Tavarory-Acosta ñu, creek, affluent of 40054 (MO, SI) Amambay: Parq Nac Cerro Paraguay River, 25°27’S 57°32’W, 10-VI-1993, Corá, 1 km de la cada [casa?] del quardaparques, Zardini & Jara 36125 (CTES); Tavarory, 3 km S 22°40’S 56°03’W, 4-V-1989, Fortunato & al from administration, 1 Km E of Rio Paraguay, 1072 (BAB, CTES); Parq Nac Cerro Corá, 1 km 25°30’S 57°30’W, 9-XII-1991, Zardini & Tilleria de la cada [casa?] de guardaparques, camino a 29229 (CTES, 2 sheets); 25 km W from entrance Cerro Muralla, 22°40'S 56°03'W, 4-V-1989, to Tavarory, 25°30'S 57°30'W, 18-V-1992, Fortunato & al 1089 (BAB, CTES); 9 Km SE de Zardini & Tilleria 31779 (CTES); Estero del Bella Vista, 24-VI-1977, Krapovickas & Schinini Ypoá, Villeta-Puerto Guyrati, 7 km S of Villeta, 32614 (CTES); Parque Nac Cerro Corá, camino a 25°34'S, 57°33'W, 18-XI-1992, Zardini & Colonia Naranjaí, próximo naciente arroyo Tilleria 33698 (MO, SI); Itá Ybaté-Pindoty, 150 Aquidabán, 22°40'S 56°5'W, 2-V-1992, Morrone & m, 25°37’S 57°31’W, 27-VI-1991, Zardini & Ru Pensiero 535 (SI); camino entre Ruta 5 y Bella Velázquez 28090 (CTES); Tavarory, 1 km Vista, 42 km S de Bella Vista, 15-V-1974, Schinini southeast from entrance, 120 m, 25°28’S 8986 (CTES); Ea Sta Irene, Arroyo Negla, 22°20'S 57°31’56W, VI-1994, Zardini & Vera 39470 56°30'W, 17- XI-1996, Schinini & Barrail 31693 (CTES) Concepción: prope Concepcion, Iter ad (CTES); 2-3 Km al SE de Colonia Estrella, 500 m, Paraguariam septentrionalem, X-1902, Hassler 22°20'S 55°48'W, 9-XII-1997, Schinini & Dematteis 7603 (NY); de Valle Mi a Concepción, Estancia 33593 (CTES); San Luis, 4-IV-1951, Schwarz San Luis + 3 km saliendo del casco, 194 m, 12268 (CTES); Estancia Carmen de la Sierra, 22°24’S, 57° 25’W, 15-X-2000, Kiesling & al Potrero Lili, 22-X-1991, Soria 4679 (CTES) Boque- 9657 (SI); between Paso Horqueta and Estancia rón: Col Menno, Lolita, 23°S 59°35'W, 10-IX-1990, San Fernando, 22°58’48’’S 57°21’37’’W, 1-VII- Vanni & al 1838 (CTES); Estación Experimental de 1994, Zardini & Guerrero 39842 (CTES) Cor- Col Neuland, 22°45’S 60°W, 13-IX-1990, Vanni & dillera: 13 Km E del camino Pirabebuy- al 2095 (CTES) Canendiyú: Reserva Natural del Paraguari, en dirección del Valenzuela, 2-III- Bosque Mbaracayú, 13 km S of Ypé-jhú, 1994, Cristóbal & Krapovickas 2402 (CTES); Ñanduro Cai, 3-XII-1997, Schinini & Cerro de Tobatí, ca 4,5 km al S de Tobatí, 12,2 Dematteis 33286 (CTES); Mbaracayú Natural km de Rta 2 en Caacupé, entrando 0,5 km al O Reserve, Ñanduro Cai, 23°59’40’’S en camino de tierra y caminando al NNE hasta 55°28’45’’W, 11-VI-1998, Zardini & Chaparro la cumbre de cerro, 9-II-1987, 25°20’S 48698 (CTES); Mbaracayú Natural Reseve, 57°05’W, Ginzbarg & al 439 ( CTES, NY); San around Ñanduro Cai, 23°59’39’’S Bernardino, 26-V-1964, Gregory & al 11493 055°28’44’’W, 27-V-1999, Zardini & Chaparro (CTES); Tobatí, 24-V-1964, Krapovickas & 50764 (CTES) Central: 2 km SW de río Salado, Cristóbal 11449 (CTES); Cerros

173 BONPLANDIA 16(1-2) 2007

de Tobatí, 1900, IX-1902, Hassler 6123 (NY); de montes terreno arenoso”, 27-I-1943, Pavetti Caacupé, Cerro, aprox 53 km de Asunción, 9- & Rojas 10307 (CTES); Paraguay [Guairá:?] II-84, Pérez 228 (SI); Serranía de Tobatí, Mese- Fernando de la Mora, Km 8, I-1968, Schinini ta Ybytú Silla, 297 m, 25°12’S 57°07’W, 25-VIII- 1855 (SI) 1988, Zardini 6747 (CTES); eastern side of Río Piribebuy basin, 17 km W of Arroyos y Esteros, Obs 1: Among W indica sl collections 25°08’S 57°15’W, 19-V-1990, Zardini 20097 there are shrubs 1-15 m tall, with a calyx (CTES); Cerro Tobatí, 27-XI-1987, Zardini & tomentulose, only sericeous on the veins, Degen 3798 (CTES); Tobatí, “Ybytú Silla” bearing stems very shortly cannescent- mesa, Southern area, 297 m, 25°12’S 57°07’W, tomentulose, with short purple glandular- 8-II-1991, Zardini & A Salem 26110 (BAB); capitate hairs Most interesting is the Tobatí, “Ybytú Silla” mesa, northern area, 297 possession of purple residual deposits in the m, 8-II-1991, Zardini & Velázquez 26168 stellate trichomes similar to W ladewii Rusby, (MO) Guairá: Colonia Independencia, endemic to Beni, Bolivia from wet savannas serrania del Ilyturusú, I-1967, Schinini 14 (SI); These Paraguayan collections are found in the Colonia Independencia, II-1968, Schinini 1647 provinces of Cordillera, Paraguarí, Pres (CTES); Mbocyaity-Melgarejo, Cerro Naville, Hayes, Central and Boquerón, some from low 25°43’S 56°21’W, 6-VI-1992, Zardini & Guerre- elevations (150 m), from rock outcrops of ro 32466 (CTES) Itapuá: Yacyretá Dam Island basalt, or limestone in sandy or clay soils, and Reserve: Eastern area, Aña Cua, 27°23’45’’S need futher study: (Cristóbal & Krapovickas 56°39’08’’W, 23-X-1999, Zardini & Gamarra 2402, Ginzbarg & Davalos 439, Hassler 51873 (CTES); Yacyretá Dam Island Reserve: road 6123, Zardini 20097, Arbo 1759, Sparre & to Aña Cua, 27°26’41’’S 56°45’00’’W, 24-X-1999, Vervoorst 930, Mereles 3533, Pavetti & Zardini & Gamarra 51974 (CTES) Misiones: Rio Rojas 10307, Zardini & Velázquez 28090, Tebicuary, 26°23'43’’S 57°07'45’’W, 28-VII-1994, Zardini & Tilleria 29229, Zardini & Zardini & Guerrero 40156 (CTES) Paraguarí: Acquino 32060, Vanni & Schinini 2095) Salto de Piraretá, márgenes del arroyo Yhaguy- Guazú, 14-XI-1978, Arbo & al 1759 (CTES); Obs 2: Some plants appear to be Piraretá above Salta Pirareta, 17-IV-1990, Saunders intermediate with W albicans Vanni 1838 by & al 3211 (CTES 2 sheets, NY); Cerro Sto To- its long capsule (22-24 mm) with slight más, 29-XI-1950, Sparre & Vervoorst 730 (CTES); extension of corneous endocarp (09-10 mm) Cerro Yaguarón, 25°37’S 57°15’W, 24-X-2000, is slightly similar to W albicans, but in its Keisling & al 9824 (SI); Acahay Massif, rough, extensively woody branches, plicate easternmost peak, base of peak, 25°52’S 57°08’W, elliptical leaves that are tomentose, petal 27-I-1992, Zardini & Aquino 30025 (CTES); Pala- shape and size (37-40 x 08 mm and slight cios Mountain, 25°25’S 57°10’W, 24-VI-1989, adherence of the petal (only 05 mm) to the Zardini & Velásquez 13179 (CTES) Presidente stamen tube, and anther size is more similar to Hayes: Puerto Militar, frente a Concepción, 8- W indica Another example from Paraguarí is XII-1989, Mereles 3533 (CTES); Estancia Zardini & Aquino 30025 with some Zalazar, laguna, 22°58'50’’S 59°07'05’’W, 13- inflorescences deltate shaped, others sessile, XII-1996, Zardini & Fernández 45975 (CTES) glomerulate and with petals and capsule more San Pedro: Ruta 3 y Río Tapiracuay, en orillos like W indica del río, 19-V-1974, Schinini 9087 (CTES) Department not indicated: Plantae Para- Obs 3: Other plants appear intermediate guarienses 1885-1895, Hassler 384 (NY, 2 with W communis One is Saunders & al sheets); in dumeto ad ripam lacus Ypacaraí, IV- 3211 (CTES 267041, plant A), locally 1900, Hassler 4063 (NY); Plantas del Paraguay: sympatric with W communis It possesses [illegible], Jörgensen 3861 (SI); [loc illegible], basalmost rounded or suborbicular leaves, a 3-I-1928, Jörgensen 3861 (BAB); cult Jardín style uniformly wide to base, a stigma Botánico, Universidad Nacional de Tucumán, penicillate similar to W communis, and the 2 provenance: “Paraguay, habita orillas examples (NY, CTES) are variable

174 J G Saunders, Sterculiaceae de Paraguay II

Note 1: Citation of 2n=14 for W indica in illustrations, and am grateful to Dr Carmen Fedorov (1974:705), Whetstone (1983: 19) and Cristóbal, Dr Fernando Zuloaga, and Dr Billie in Bahadur & Srikanth (1983:103) is erroneous Turner for financing them I thank Dr Larry J and must be revised to W ovata Cav It is a Dorr for directing my attention to the misinterpretation of Brizicky (1966: 73), erroneous chromosome number citation of W clarified there on p 74 citing W ovata from indica in the literature, Dr Carlos Brizuela Diers (1961: 450) Bates (1967) also reported and Professor Verónica Vázquez Rodríguez for W ovata (as W reticulata) 2n=14 for facilitating or providing Guaraní Although Robyns (1964) listed W ovata as a translations and two anonymous reviewers for synonym of W indica, and Brizicky did not, it helpful comments I thank Dr Massimiliano is usually considered as a separate species, the Dematteis for sending the map template, and various references summarized in Dorr the editor of Darwiniana, Dr Raúl Pozner, for (1999) Other first reports for reissued counts permission to reissue figs 8 & 9 Finally, I 2n=14 or 2n=28 for W indica alluded to in thank Dra Cristóbal for the opportunity to Bahadur & Srikanth (1983:103) have not been revise the Paraguayan Waltheria, and along seen The only other verified chromosome with Lic Sara Tressens, for bibliographic, count reports I have seen for other species in photographic, mapping and herbarium the genus are n=5 for W acuminata (Bates, Waltheria resources shared 1976), the lowest count in the Sterculiaceae, and 2n=12 for W communis (Cristóbal, 1967), neither species ever considered to be Literature Cited synonyms of W indica

Note 2: The name W americana L var ABBOTT, I A & C SHIMAZU 1985 The geographic glandulosa R E Fries (1908), has the type origin of the plants most commonly used for medi- collection cited from Paraguay, Estancia Sta cine by Hawaiians J Ethnopharmacol 14: 213- María, Anisits 2041, herb Regnell that has 222 not been seen By description it differs from ARBO, M M 1987 Turneraceae In R Spichiger (ed) Flora del Paraguay Conservatoire & Jardin W americana by intermixed glandular botaniques de la Ville de Genève & Missouri stipitate and tomentose vestiture, dense on Botanical Garden, Ginebra 65 pp calyx lobes and prophylls BAHADUR, B & SIDHU 1975 In A Löve (ed) Taxon 24: 501-516 1975 [Waltheria indica 2n=24] Acknowledgments —— & R SRIKANTH 1983 Pollination biology and the species problem in Waltheria indica complex Phytomorphology 33: 96-107 [Waltheria indica I thank the directors of the herbaria cited by 2n=24 for 4 plants of heterostyles and homostyles on p 105] acronym where Paraguayan material was BASUALDO, I & N SORIA 2002 100 Especies del studied (AS, BAB, BAF, BM, C, CTES, E, F, cerrado en Paraguay Departamento de Botánica FCQ, G, GH, K, LIL, LL, MBM, MICH, MO, FCQ-UNA, San Lorenzo, Paraguay, & Missouri MPU, NY, P, PY, S, SI, TEX, U, US, W) Botanical Garden, St Louis 220 pp This study was realized with generous support ——, N SORIA, R DEGEN, M ORTIZ & E from Ing Agr Ramón Palacios, the Instituto ZARDINI 2005 Catálogo de la flora vascular de de Botánica Darwinion, The University of Tavaí, Dpto Caazapa, Paraguay Rojasiana 7(1): Texas at Austin, and through funds from the 27-99 UT Graduate School, Billie Turner fellowship BATALHA, M A & W MANTOVANI 2001 Floristic composition of the cerrado in the Pe-de-Gigante award, Alexopolos fellowship award, and the Reserve (Santa Rita do Passa Quatro, Southeastern National Science Foundation dissertation Brazil) Acta Bot Bras 15(3): 289-304 grant BSR-8800899 I gratefully acknowledge BATES, D 1967 Chromosome numbers in the Malvales Lic Liliana Gómez, Prof Francisco Rojas I Gentes Herb 10(1): 39 and Branton Whetstone for the exquisite —— 1976 Chromosome numbers in the Malvales III

175 BONPLANDIA 16(1-2) 2007

Gentes Herb 2(3): 144-147 Sterculiaceae In J A Steyermark, P E Berry & B BAYER, C 1994 Zur Infloreszenzmorphologie der K Holst, (eds) Flora of the Venezuelan Guayana, Malvales Diss bot 212 Cramer: Berlin, Stuttgart P E Berry, B K Holst, K Yatskievych (eds), Vol 280 pp 9 Rutaceae-Zygophyllaceae, pp 248-280, Missouri BRIZICKY, G K 1966 The genera of Sterculiaceae in Botanical Garden Press, St Louis the southeastern United States J Arnold Arbor 47: DIERS, L 1961 Der Anteil an Polyploiden in den 60-67 Vegetationsgürteln der Westkordillere Perus Z CARDOSO, E L, SM ARAÚJO CRISPIM, C A G BOT 49: 437-488 [Waltheria ovata 2n=14, p RODRIGUES & W BARIONI, Jr 2000 Aerial 450] biomass and primary production of herbaceous DORR, L J 1999 Sterculiaceae In P M Jørgensen & communities in a native grassland of Elyonurus S León –Yanez (eds) Catalogue of the Vascular muticus under annual burnings in the Pantanal, Plants of Ecuador pp 919-922 Missouri Botanical Brazil Pesq Agropecu Brasil [online] 2000 vol Garden Press St Louis 35, no 8, [cited 2006-11-01], pp 1501-1507 FEDOROV, A A 1974 Chromosome numbers of Available from:  FERRI, MG 1969 Plantas do Brasil: Espécies do Cerra- CAVALCANTI, R B & C A JOLY 2002 Biodiversity do Editôra Edgard Blücher Ltda, Editôra da and Conservation Priorities in the Cerrado Region In Universidade de São Paulo, São Paulo 240 pp PS Oliveira & R J Marquis (eds) The cerrados of FILGUEIRAS, T S & B A da S PEREIRA 1990 Brazil: Ecology and Natural History of a Neotropical Flora do Distrito Federal En M Novaes Pinto Savanna pp 351-367 Columbia University Press, (org) Cerrado: caracterização, ocupação e perspec- New York tivas, pp 331-388, Brasília, Editora Universidade CHODAT, R 1901 Sterculiaceae In R Chodat (ed) de Brasília Plantae Hasslerianae 1 [:96-98; Waltheria 97], Bull FRIES, R E 1908 Studien über die amerikanische Herb Boissier ser 2, no 1: 402-404 [Waltheria Columniferenflora, Kongl Svenska Vetensk Akad 403] Handl 42(12): 7-67 Taf 1-7 [Waltheria 13-15, CORRÊA, M P 1974 Dicionário das Plantas Úteis do Tafel 3] Brasil Vol V Rio de Janeiro Instituto Brasileiro GILL, L S 1978 Chromosome number reports in de Desenvolvimento Forestal Ministério da Agri- Tanzania II Adansonia 18: 19-24 [n=20 on p 22 as cultura Waltheria indica for Tanzania: University of Dar- —— 1984a Dicionário das Plantas Úteis do Brasil es-salaam campus, Gill 090 DSM] Vol 2: letras Car-E Rio de Janeiro, Ministério da HARVEY, M J 1966 In A Löve (ed) IOPB Agricultura, Imprensa Nacional chromosome number reports VII Taxon 15: 155- —— 1984b Dicionário das Plantas Úteis do Brasil 163 [2n=40 on p 160 for Sierra Leone, Kamabai, Vol 5: letras M-R Rio de Janeiro, Ministério da Harvey 94 K, as Waltheria americana] Agricultura, Instituto Brasileiro de —— 1967 In A Löve (ed) IOPB Chromosome Desenvolvimento Forestal number reports IX Taxon 16: 62-66 [corrects CRISTÓBAL, C L 1967 Cromosomas de Malvales Harvey 1966 listing from W$ americana to W$ indi- Kurtziana 4: 139-142 ca] —— 1983 Esterculiáceas In R Reitz, Flora ilustrada HASSLER, E 1917 Addenda ad Plantas Hasslerianas, catarinense, Parte I As Plantas: Fasc ESTE: 1-57, 1-20 [Sterculiaceae 11-12; Waltheria 12] est 1-10, IOESC, Itajaí, Santa Catarina HERTER, G F 1957 XIV Flora Ilustrada del Uruguay —— 1998 Sterculiaceae In A T Hunziker & A Anton Fasc XIII Entrega XIIIa: Celastraceae II- (eds) Flora Fanerogámica Argentina, Fasc 57: 1- Cactaceae Waltheria douradinha: Plate 2184 on p 32 PROFLORA (CONICET) Museo Botánico, 566 IMBIV, Cordoba JIMÉNEZ, B, S KNAPP, G MARIN & M PEÑA- —— 2005 Sterculiaceae In A Burkart & N M CHOCARRO 2000 Listado preliminar de plantas Bacigalupo (eds) Flora ilustrada de Entre Rios vasculares de la reserva natural del bosque (Argentina), Parte 4 Dicotiledoneas Arquiclamideas Mbaracayú, Paraguay Rojasiana 5(2): 141-271 B: Geraniales a Umbelliflorales, pp 333-351 KÖHLER, E 1971 Zur Pollenmorphologie der Gattung [Waltheria pp 348-350], I N T A, Buenos Aires Waltheria L (Sterculiaceae) Feddes Repert, 82: —— 2007 Sterculiaceae de Paraguay I Ayenia, 125-153 Byttneria, Guazuma, Helicteres, Melochia y —— 1973 Über einen bemerkenswerten Sterculia Bonplandia 16(1-2): 5-142 Pollendimorphismus in der Gattung Waltheria L ——, J G SAUNDERS & P E BERRY 2005 Vorläufige Mitteilung über die heterostyle W$

176 J G Saunders, Sterculiaceae de Paraguay II

viscosissima St Hil Grana 13: 57-64 —— 1995 Systematics and evolution of Waltheria —— 1976 Pollen dimorphism and heterostyly in the (Sterculiaceae-Hermannieae) PhD dissertation genus Waltheria L (Sterculiaceae), pp 147-161 In I The University of Texas, Austin Available at K Ferguson & I Muller (eds), The Evolutionary PROQUEST, Ann Arbor, MI Significance of the Exine Linn Soc Symp Series —— 2005 New species and new records of Waltheria No 1 from Paraguay, Argentina and Venezuela KRISHNAPPA, D G & MUNIRAJAPPA 1980 In P Darwiniana 43: 201-211 Goldblatt 1981: 355, 392 Index of chromosome SCHININI, A 2005 Teodoro Rojas, Biografía, Viajes y numbers Ann Missouri Bot Gard 1979-1981 [on especies dedicadas Rojasiana 7: 101-149 p 536 in Original citation in IOPB Chromosome SCHUMANN, K 1886 Sterculiaceae In C Martius number reports LXVIII Taxon 29: 535-536, 1980 (ed) Flora brasiliensis 12(3): 2-114, Figs 1-24 but volume missing at SI to reconfirm] [Waltheria 50-68 tab 11-14] MARTINEZ, M 1979 Catálogo de Nombres Vulgares —— 1903 Sterculiaceae In R Chodat & E Hassler de Planta Vasculares, Fondo de Cultura Económica, (eds) Plantae Hasslerianae 2 [:300-308; Waltheria Mexico 304-305] Bull Herb Boissier ser 2, 4: 68-76 MELHEM, T, M SILVESTRE & N LUCAS 1976 [Waltheria 72-73] Pollen morphological studies in Sterculiaceae TRESSENS, SG, R VANNI & M G LÓPEZ 1999 In Hoehnea 6: 23-32 MM Arbo & S G Tressens (eds) 2002 Flora del MOREL, AF, GEHRKE, ITS, MOSTARDEIRO, Iberá pp 201-338 Eudene, Corrientes 613 pp MA, ETHUR, EM, ZANATTA, N, & MA- VASQUEZ, CA, SF CORTES, MS SILVA, & IA de CHADO, E C S 1999 Cyclopeptide alkaloids MEDEIROS 1999 Phytotherapy Res 13(4): 312- from the bark of Waltheria douradinha 317 Phytochemistry 51(3): 473-477 WHETSTONE, R D 1983 The Sterculiaceae in the MORTON, J F 1981 Atlas of Medicinal Plants of flora of the southeastern United States Sida 10(1): Middle America Charles C Thomas, Springfield 15-23 NEAL, M L 1965 Gardens of Hawaii Special Publication 50 Bernice P Bishop Museum, Honolulu 924 pp Index of the species treated NICORA, E, R KIESLING, D RODRÍGUEZ, M E MULGURA, G GIBERTI & M DONATH 1988 Índice geográfico paraguayo Instituto de Botánica 1 W communis A St-Hil Darwinion, ined 2 W macrophylla Hassl OLIVEIRA, PS & R J MARQUIS (eds) 2002 The 3 W operculata Rose cerrados of Brazil Columbia University Press New 4 W viscosissima ASt-Hil York 5 W procumbens J G Saunders & Soria PAIS, M, J MARCHAND, F X JARREAU, & R GOUTAREL 1968 No 180 Alcalöides 6 W carmensarae J G Saunders peptdiques V Structures des adouétines X, Y, Y´et 7 W albicans Turcz Z, alcalöides du Waltheria americana L 8 W indica L (Sterculiaceae) Bull de la Soc Chimique de France 18 (3): 1145-1148 PETRUS, A J A 1990 Polyphenolic components of Index of collectors Waltheria indica Fitoterapia 61(4): 371 REED, W 1974 Populations and host-plant preferences in Earias spp$ (Lepidoptera: Noctuidae) in East Binaciónal, 563 (1) Africa Bull Entomol Res 64(1): 33-44 ROBYNS, A & J CUATRECASAS 1964 Binaciónal, 595 (1) Sterculiaceae In W H Lewis (ed) Flora of Anderson, W R, 12377 (1) Panama Part VI Family 117 Ann Missouri Bot Anisits, J D, 1899 (1), 2089 (1), 2535 (1) Gard 51: 69-107 Arbo, M M & al, 1705 (1) SAINT HILAIRE, A 1825 Plantes usuelles des Arbo, M M, G I Lavia, MG Pelligrini & R brasiliens 8, pl 36-80 Beron , 6120 (1) SAUNDERS, J G 1993 Four new distylous species of Arbo, M M, A Schinini & I Basualdo, 2883 (1) Waltheria (Sterculiaceae) and a key to the Mexican Arbo, M M, S G Tressens, A Schinini & S and Central American species and species groups Ferrucci, 1617 (8), 1619 (1), 1704 (1), 1759 Syst Bot 18: 356-376 (8), 1790 (1), 1798 (1)

177 BONPLANDIA 16(1-2) 2007

Archer, W A, 4768 (1) Hatschbach, G & R Kummrow, 48524 (1) Arenas, P, 1254 (1), 2931 (1) Jiménez, B & G Marín, 1611 (1) Balansa, B, 2659 (1) Jörgensen, P, s/n (1) Basualdo, I & M Ortiz, 2571 (1) Jörgensen, P, 3861 (8), 3862 (1) Bernardi, L, 18304 (1), 18337 (1), 18766 (1), Kiesling, R, E A Ulibarri & M Quintana, 9657 18950 (1), 19492 (1) (8), 9824 (8) Bordas, E, 3945 (8), 4007 (1), 4042 (8) Krapovickas, A, 13434 (1), 13967 (1) Brunner, D H, 1628 (7), 1072, (8) Krapovickas, A & C L Cristóbal, 11449 (8), Burkart, A 18688 (1) 13266 (1), 13434 (1), 43201 (8) Buttura, 715, (1) Krapovickas, A, C L Cristóbal, & L Z Ahuma- Caballero Marmori, G, 1301 (1) da, 13967 (1) Cristóbal, C L & al, 2433 (7), 2441 (7) Krapovickas, A, C L Cristóbal & R A Palacios, Cristóbal, C L & A Krapovickas, 2294 (1), 2299 (1), 12384 (1), 12517 (1) 2363 (1), 2372 (1), 2387 (7), 2394 (1), 2402 (8), Krapovickas, A & A Schinini, 32614 (8) 2407 (1) Kuntze, O, s/n (1) Cristóbal, C L, A Krapovickas, R M Harley & Lurvey, E, 257 (1), 354 (1), 366 (1), 517 (1) A Schinini, 2451 (1), 2453 (1), 2455 (7-8), 2457 Mereles, F, 1018 (1), 1461 (1), 1902 (1), 3533 (8), 4279 (1), 2470 (1), 2471 (1), 2473 (1) (1), 6587 (7) Cristóbal, C L, A Krapovickas & A Schinini, Mereles, F & R Degen, 5706 (7), 5978 (7), 2422 (1), 2433 (7), 2434 (7), 2441 (7), 2444 Degen, 6191 (7), Degen, 6351 (7) (7) Mereles, F & L Ramella, 2413 (8), 2620 (8) Eskuche, U, 2533-11 (1) Molas, L & al, 674 (1) Fernández Casas, J & al, 7545 (1) Montes, J E, 7187 (1) Fernández Casas, J & J Molero, 5886 (1), 6102 Morong, T, 201 (1), 215a (1) (1), 6238 (5) Morrone, O & J Pensiero, 535 (8) Ferrucci, M S, A M Gonzalez, A Schinini & R Múlgura de Romero, M E, C Calviño, S Beron, 917 (1), 927–A (1), 927-B (1) Martínez & N O´Leary, 3680 (1) Ferrucci, M S, R Vanni & L Ferraro, 710 (1) Niederlein, (1) Fiebrig, K, 1382 (3), 4772 (4) Ortíz, F, 118 (1), 547 (1) Fortunato, R H, N M Bacigalupo & N Soria, Pavetti & T Rojas, 10307 (8) 1072 (8), 1089 (8) Pederson, T M, 3861 (8) Fortunato, R H, R Gómez, M Morales & L Pérez, L, 228 (8), 420 (1), 513 (1), 808 (1) Pérez, 8777 (7) Pérez, L, & al, 1429 (1) Fortunato, R H, L Ramella & R Palese, 3649 (8) Pérez, L, S Bertoni, T Florentin Peña & A Boga- Ginzbarg, S & al, 420 (1) do, 2872 (1), 2896 (1), 2898 (1), 2906 (1) Ginzbarg, S, J Dávalos & I Fox, 439 (8) Pérez, L S Bertoni, M Quintana, B Benitez & A Ginzbarg, S & J Dávalos, I Fox, C Rodriguez, Bogado, 2630 (7) 554 (1) Pérez de Molas, L & G Navarro, 8201 (8), 8324 Gregory, W, 9602 (1), 9690 (1) (8), 8608 (7) Gregory, W C, A Krapovickas & J Pietrarelli, Ramella, L & F Mereles, 2413 (8) 11493 (8) Riemel, 52 (1) Hassler, E, 384 (8), 950 (1), 3621 (1), 4063 (8), Rojas, T, 10151 (1), 13544 (1) 4340 (1), 4736 (1), 5026 (1), 5834 (1), 6026 Sanguis, s/n (1) (1), 6123 (8), 6747 (1), 6747a (1), 6747b (1), Saunders, J, N Soria & I Basualdo, 3208 (1), 7603 (8), 7824 (1), 7842 (5), 7861 (1), 8966 3211 (8) (1), 9097 (1), 9151 (2), [Rojas?] 9750 (1), Schinini, A, 14 (8), 1647 (8), 1855 (8), 2294 (1), 9779 (1), 11160 (4), 11384 [probably =Rojas 4315 (1), 4711 (1), 4731 (8), 4733 (1), 5027 1384](1), 11460 [probably a Rojas coll in (1), 5761 (1), 5858 (1), 5923 (1), 8116 (1), herb Hassler, see Schinini 2005: 128] (1), 8773 (1), 8794 (1), 8869 (1), 8877 (1), 8898 12196 (8) (1), 8986 (8), 8994 (1), 9072 (1), 9086 (1), Hatschbach, G & R Callejas, 47259 (1) 9087 (8), 9172 (1), 9194 (1), 13564 (1), 21486

178 J G Saunders, Sterculiaceae de Paraguay II

(1), 23040 (1) Zardini, E M & G Garcete, 28804 (1), 29140 (1), Schinini, A & O Barrail, 31693 (8) 29251 (1), 29252 (1) Schinini, A & E Bordas, 15050 (8), 15050-A (7), Zardini, E M & J Godoy, 50484 (8) 15050-B (8), 15050-C (7), 15050 (7), 15088 Zardini, E M & L Guerrero, 32466 (8), 33862 (7), 17843 (8), 20172 (1), 20454 (1) (1), 34196 (8), 34343 (1), 34383 (1), 34391 Schinini, A & G Caballero Marmori, 26923 (1), (1), 37627 (1), 37762 (1), 38629 (1), 39842 27196 (1), 27290 (1), 29975 (1), 30035 (1) (8), 40156 (8), 41235 (1), 41261 (1), 41327 Schinini, A & M Dematteis, 33267 (1), 33286 (1), 42089 (1), 44071 (1), 46444 (1), 48297 (8), 33479 (1), 33524 (1), 33593 (8), 33635 (1), 48298 (1), 53927 (3) (1) Zardini, E M & V Jara, 36125 (8) Schinini, A & R Vanni, 26080 (1) Zardini, E M & A Ramirez, 41430 (1) Schinini, A, R Vanni & S Cáceres, 27809 (1), 28142 Zardini, E M & S Ramírez Benítez, 51151 (1) (1) Zardini, E M & J Rodríguez, 51648 (1), 51651 (6), Schmeda, s/n (1) 51716 (1) Schulz, A G, 7700 (6) Zardini, E M & Salema, 26110 (8) Schwarz, G J, 11805 (1), 12171 (1), 12268 (8) Zardini, E M & A Salinas 24444 (1) Soria, N, 1673 (1), 2505 (1), 2542 (1), 3454 (1), 4458 Zardini, E M & T Telleria [Tilleria?], 36086 (5), 4679 (8), 4930 (5), 5733 (1) (1) Soria, N, & E Zardini, 1871 (1) Zardini, E M & T Tilleria, 28947 (1), 29229 (8), Sparre, B & F Vervoorst, 207 (1), 730 (8), 7284 29477 (1), 31779 (8), 33698 (8), 33699 (1), (1) 34521 (1) Stutz, L C, 2141 (1) Zardini, E M & Velázquez, 15899 (1), 16453 (1), Tressens, S, T Ahti, L Ferraro & V Maruñak, 4783 24161 (1), 24444 (1), 24509 (1) (1) Zardini, E M & C Velázquez, 24435 (1) Tressens, S G, V Maruñak, C L Cristóbal & M Zardini, E M & R Velázquez, 13179 (8), de Pompert, 3386 (1) 24720 (1), 26168 (8) Vanni, R, 225 (1), 3494 (7-8) Zardini, E M & Ru Velásquez, 28090 (8) Vanni, R & al, 2478 (7) Zardini, E M & U Velázquez, 24287 (1), Vanni, R & G López, 3954 (8) 25983 (1), 26020 (1), 27837 (1) Vanni, R, S Ferrucci, P Cowan, R Duré & A Zardini, E M & M Vera, 39470 (8), 41235 (1), Schinini, 225 (1) 47927 (1), 48209 (1) Vanni, R, A Radovancich & A Schinini, 1838 [a] Zardini, E M & S Zavala, 46740 (1), 46880 (1) (7-8), 1838 bis [b] (8-7), 2095 (8), 2133 (7), 2192 (7) Woolston, A L, 330 (1) Index of common names Zardini, E M, 6747 (8), 20097 (8), 25801 (1) Zardini, E M & al, 8718 (1), 8719 (1) Zardini, E M & A Acosta, 42265 (7), 42361 (7) “douradinha” (W communis as W douradinha: Zardini, E M & P Aquino, 29332 (1), 29566 (1), Brazil: SC: Smith & al 9332, US) 149 30025 (8-7), 31800 (8), 32060 (8), 34813 (1) “douradinha do campo” (W communis as W Zardini, E M & P Báez, 52279 (1) douradinha: Brazil: RGS: Reitz 515, CTES) Zardini, E M & A Benítez, 47571 (1) 149, 150 Zardini, E M & R Britez, 52875 (1) “guaikuru” (W communis: Argentina: Corrientes: Zardini, E M & W Cardozo, 44956 (4) Pedersen 82, C; orthography corrected from Zardini, E M & I Chaparro, 48679 (1), 48698 Guaycuru and meaning given by pers comm, (8), 49517 (1), 50617 (1), 50764 (8), 50798 C Brizuela & V Vázquez Rodríguez, 2007) (1) 149 Zardini, E M & R Degen, 3798 (8) “malva”(W viscosissima: Brazil: Pará: Archer 7816, Zardini, E M & J Fernandez, 45975 (8) NY) 162 Zardini, E M & A Florentin, 40054 (8) “malva branca” (W viscosissima in Brazil, Corrêa, Zardini, E M & R Gamarra, 51873 (8), 51974 (8) 1974) 162

179 BONPLANDIA 16(1-2) 2007

“malva branca” (W indica as W americana in carmensarae 143, 145, 158, 163, 165 Brazil, Corrêa, 1984) 172 communis 143, 144, 146, 147, 149, 150, 155, 174 “malva preta” (W viscosissima Brazil: Pará: Silva var dietrichii 145, 155 79, NY) 162 var erosa 155 “malva-veludo” (W indica in Brazil, Corrêa, var glabriuscula 145 1974) 172 var gracilis 145 “oreja de carnero” (W communis as W douradinha: var henningsii 145 Brazil: Campo Grande: Bertoni 616, U) 150 var hirta 145, 154, 155 “typycha guaikuru” [W communis as W var lanata 145, 155 douradinha: Argentina: Corrientes: Pedersen var paraibana 148 82, C) corrected (pers comm, C Brizuela, var platyphylla 152, 155 2007) from “Tibicha-Guaycuru” on specimen var tomentella 145, 149, 155 label Pedersen 82, C] 149 var velutina 148, 154, 155 “yerba de las monjas” (W communis: Uruguay: var “vulgaris” 145, 149, 155 Herter, 1957: 566) 149 douradinha 145, 149, 150 douradinha var gracilis 155 excelsa var hirsuta 158 Index of scientific names glabriuscula 145 gracilis 145 hirsuta 160 Astropus 143 indica 143, 145, 155, 169, 170, 171, 172, 174 Elyonurus muticus 155 lanata 145 Lindernia crustacea 150 ladewii 174 Lophanthus 143 machrisiana 160 Scrophulariaceae 150 macrophylla 143, 144, 156, 157, 158 Sitella 143 mollis 160 Stemodia arenaria 150 operculata 143, 144, 156, 158, 159 Turnera valleana 158 ovata 175 Waltheria 143, 144, 150, 175 procumbens 143, 145, 158, 163, 164 sect Stegowaltheria 143, 144, 160 prostrata 145, 163 sect Waltheria 143, 144 reticulata 175 acuminata 175 rotundifolia 162, 163 albicans 143, 145, 158, 166, 167, 168, 174 tubiflora 160 americana 145, 172, 175 viscosissima 143, 145, 158, 160, 161, 162 americana var glandulosa 175 americana form foliis elliptico lanceolatis not seen: 145, 151, 155 Waltheria americana var glandulosa boliviensis 145, 155 Waltheria communis var erosa

Original recibido el 1 de noviembre de 2006; aceptado el 3 de abril de 2007$

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