New Boraginaceae from Tropical America 4: Three New Species of Cordia from South America James S. Miller Missouri Botanical Garden, P. 0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT. Review of specimens newly collected have been covered in regional accounts (Johnston, for one of the International Cooperative Biodiversity 1930, 1935; Miller, 1988, 2001; Taroda & Gibbs, Groups led to the recognition of three new species 1987). In particular, the South American taxa re¬ closely related to Cordia trachyphylla Marti us, with main poorly known, and their morphological simi¬ which some specimens had previously been con¬ larity makes it difficult to recognize coherent, man¬ fused. All three are members of a group of species ageable species groups. Many species in the section within Cordia sect. Myxa that are characterized by remain undescribed, particularly where the highest having a costate calyx. Three new species within diversity occurs in northwestern South America. this group are described, Cordia hatschbachii, C. Recent novelties include the following three new kingstoniana, and C. koemarae, and a key is pro¬ species. vided to assist in separation of the neotropical spe¬ The three species described below all share a cies of Cordia sect. Myxa with costate calyces. costate calyx and a decidedly acuminate leaf apex Key words: Boraginaceae, Cordia, South Amer¬ and are probably related to Cordia trachyphylla ica. Martius. Although this name has been associated with specimens over a wide geographic range, re¬ cent review of available specimens indicates that Cordia L. is the largest genus in the Boragina¬ Johnston’s (1930) circumscription of a plant with ceae with approximately 325 species that occur ovate to elliptic-ovate leaves less than 16 cm long widely in tropical and subtropical parts of the and 7 cm wide and with an acumen less than 1 cm world; the greatest number of species are restricted long and restricted to Bahia was probably correct, to the Neotropics. Most modern authors have rec¬ and collections from other regions are best assigned ognized six sections in the genus Cordia (Taroda & to the following species. Gibbs, 1986; Nowicke & Miller, 1990). Despite nu¬ merous suggestions that several of the morpholog¬ Cordia koemarae J. S. Miller, sp. nov. T\ PE: Su¬ ically diverse sections of the genus be recognized riname. Sipaliwini: vicinity of the Ulemari as separate genera (Nowicke & Ridgway, 1975), River, 99 km upstream from its confluence most modern authors have continued to treat the with Litani River, low primary forest near river, genus in a broad sense. elev. 150 m, 2°58'18"N, 54°33'14"W, 14 Apr. All six currently recognized sections of Cordia 1998, R. Hammel, S. Koemar & U. Raghoen- have all or most of their species in the New World. andan 21493 (holotype, MO 04974007; iso¬ Sections Gerascanthus (Browne) G. Don, Rhabdo- types. BBS, US), f igure 1. calyx A. DC., Superbiflorae Taroda, and Varronia (Browne) G. Don are all restricted to the New Arbor ad 8 m alta, ramunculis sparse pilosis, trichom- atis 2—2.5 mm longis. Folia persistentia; laminae aniso- World. Species of Cordia sect. Cordia are mostly phyllae, foliis majoribus ellipticis, 12—23.5 cm longis, 6— Caribbean, with an additional species in Somalia 13 cm latis, minoribus orbicularis, ca. 8.5 cm longis, 7.5 and one more widespread on beaches of the Indian cm latis, apice acuminato et caudato, acuminibus 1.5—2 and Pacific Oceans. By far the largest of the sec¬ cm longis, basi rotundata ad obtusam, margine integra, supra serieea, trichomatis 2.5-4 mm longis, appressis, pa- tions is Cordia sect. Myxa (Endlicher) DC., with gina inferiore serieea ad pilosam, trichomatis 2—3 mm lon¬ close to 200 species, and it is the only section well gis; petioli 4—7 mm longi. Infforescentia terminalis vel ax¬ represented in all regions of the tropics. The ma¬ illaris, cyrnosa, 8-10 cm latis. Flores bisexuales; calyx jority of the species occur in the Neotropics, but tubularis, ca. 6 mm longus, 10-costatus; corolla alba, tub- about 50 species occur in Africa and Madagascar, ularis, 5-lobata; stamina 5, filis 8—10 mm longis, pubes- centia ad insertionem. and perhaps 20 are known from tropical Asia. The section is difficult taxonomically and has not been Tree 8 m tall, the twigs sparsely but evenly pi¬ treated in a comprehensive manner for the New lose, the hairs 2-2.5 mm long, yellow-brown. World since de Candolle (1845). although species Leaves persistent, dimorphic, larger leaf blades el- Novon 1 1: 421-428. 2001. 422 Novon 3 m m figure 1. Cordia koemarae J. S. Miller. —a. Flowering branch. —b. Inflorescence. —c. Gynoecium. —<1. Corolla opened to show insertion of stamens. —e. External surface of opened calyx. (All from Hummel el ni 21493, MO, holotype.) Volume 11, Number 4 Miller 423 2001 South American Cordia Species liptic, 12-23.5 cm long, 6—13 cm wide, the apex Suriname ICBG is a multi-institutional partnership abruptly acuminate, then caudate, the thin, hair¬ with the three main goals of natural products drug like acumen 1.5—2 cm long, the base rounded to discovery, economic development, and conserva¬ obtuse, sometimes slightly unevenly so, the margin tion. Fieldwork is conducted by members of the entire, the adaxial surface densely sericeous on Missouri Botanical Garden and Conservation Inter¬ young leaves, sparsely sericeous on mature leaves, national in collaboration with staff of the National the hairs tightly appressed, 2.5-4 mm long, the ab- Herbarium of Suriname to collect bulk plant ma¬ axial surface evenly sericeous to pilose, the hairs terial for pharmaceutical evaluation and support bi¬ spreading, 2—3 mm long, the venation brochidod- ological inventory that contributes to conservation romous, the midrib slightly elevated, the secondary initiatives in Suriname. Cordia koemerae is named veins 6 or 7. connected by a series of nearly par¬ in honor of Sylva Koemar, who has helped to or¬ allel tertiary veins, the dimorphic leaves borne op¬ ganize and conduct many of the field expeditions posed to larger leaves in branch axils, smaller, or¬ for the ICBG project and whose tireless efforts in bicular, ca. 8.5 cm long, 7.5 cm wide, petioles 4— the field have helped with the collection of many 7 mm long, canaliculate on the adaxial surface, specimens that have added to our knowledge of the pilose, the hairs ca. 3 mm long. Inflorescences ter¬ flora of Suriname, including the type of this new minal or borne in the axils of branches, cymose, 8— species. 10 cm broad, the branches pilose, the hairs 2-3 mm long. Flowers bisexual; calyx narrowly tubular, Cordia kingstoniana J. S. Miller, sp. nov. TYPE: ca. 6 mm long, 3 mm wide at the mouth, 10-costate, Peru. Loreto: Prov. Requena, Jenaro Herrera, pilose, the hairs spreading, 1.5-2 mm long; corolla elev. ca. 170 m, 4°50'S, 73°45'W. 3 July 1981. white, tubular with spreading lobes, ca. 8 mm long, R. Vdsquez, K. Young & N. Jaramillo 2143 the 5 lobes ovate, ca. 2 mm long, 1.5 mm wide; (holotype, MO 3314002). Figure 2. stamens 5, strongly exserted, the filaments 8—10 mm long, the upper 4—5 mm free, pubescent at the Frutex vel arbor parva ad 3(—12) m alta, ramunculis hirsutis ad strigillosos. Folia persistentia, homomorpha; point of insertion, the anthers bilobed, a round lamina anguste elliptica ad leviter oblanceolatam vel lan- gland present apically between the anther sacs; ceolatam, (9-)10.5-18(-23) cm longa, (2.5-)3-7(- 10.5) ovary ovoid, ca. 1 mm long, 0.5 mm wide, glabrous, cm lata, apice acuminato, acuminibus (0.8—) 1.5— 3 cm lon- the style ca. 3 mm long, the stigmas capitate. Fruits gis, basi rotundata ad acutam, margine integra, supra gla¬ bra. pagina inferiore strigosa; petioli 1 —5(— L0) mm longi. unknown. Inflorescentia terminalis et axillaris, 5—10(—15) cm latis. Flores bisexuales; calyx tubularis, 5—6.5 mm longus, lev- Distribution. Cordia koemarae is known only iler 10-costatus, sparse strigillosus ad hirsutum; corolla from the type collection made in the southeastern alba, tubularis, 5—7 mm longa, 5-lobata; stamina 5, filis corner of Suriname near the border with French 10-12 mm longis, pubescentia ad insertionem. Fructus Guiana. drupaceus, albus, putamine inequilateral iter ellipsoideo, 1.3—1.8 cm longo, 0.8—1.1 cm lato. Cordia koemarae is a distinctive species in its pilose indument, caudate leaf apex with a hair-like Shrub or small tree to 3(— 12) m tall, the stems acumen about 2 cm long, and its 10-costate calyx. hirsute or less commonly strigillose, the hairs 0.5— Many species of Cordia sect. Myxa have two op¬ 1 mm long. Leaves persistent, homomorphic; leal posing leaves borne from each branch axil, one of blades narrowly elliptic to slightly oblanceolate or which is decidedly smaller and different in shape lanceolate, (9-)10.5-18(-23) cm long, (2.5-)3-7(- (frequently orbicular) from the other larger leaves. 10.5) cm wide, the apex acuminate, the acumen This appears to be the case with C. koemarae, al¬ (0.8—) I ..5—3 cm long, the base rounded to acute, the though the duplicates of the single known collec¬ margin entire, the adaxial surface glabrous, strigose tion show only a single smaller orbicular leal. The on the midrib, the abaxial surface strigose, the ve¬ flowers of the single known collection do have both nation brochidodromous, the midrib distinct and el¬ stamens and an apparently normal gynoecium in¬ evated on both surfaces, the secondary veins 6 to dicating that this species most probably has bisex¬ 8(—9); petioles I—5(—10) mm long, flattened, round¬ ual flowers, as opposed to the dioecy that charac¬ ed, or shallowly canaliculate on the adaxial surface, terizes the Central American species with costate stiffly pubescent to hirsute, the hairs silaceous. In¬ calyces. florescences terminal and in the axils of leaves and Cordia koemarae was identified from material branches, 5— 10(— 15) cm broad, the branches hir¬ collected in southeastern Suriname during review sute, the hairs 0.5—1 mm long. Flowers bisexual; of specimens collected for the Suriname Interna¬ calyx tubular, 5—6.5 mm long, 2—3 mm wide at the tional Cooperative Biodiversity Group (ICBG). The mouth, lightly 10-costate, sparsely strigillose to hir- 424 No von Volume 11, Number 4 Miller 425 2001 South American Cordia Species sute, the hairs ca. 0.5 mm long, unevenly 3- to 5- (MO). ECUADOR. Napo: Orellana. Parque Nacional Ya- sunf, earretera y oleoductuo de Maxus en construceibn. lobed, the lobes widely triangular to ovate, 1—1.3 Km 54—58, elev. 250 m, 0°48'S, 76°30' W, Aulestia & Andi mm long; corolla white, tubular with reflexed lobes, 714 (MO); Estacion Cientffica Yasunf, Parcela de 50 ha, 5—7 mm long, 5-lobed, the lobes obovate, 2-3 mm elev. 200-300 m, 0°38'S, 76°30'W, Romoleroux A Grefa long, 1.5-1.8 mm wide; stamens 5, strongly exsert- 1835 (MO), 2015 (MO). PERU. Amazonas: Monte Virgin. ed, the filaments 10-12 mm long, the upper 5-6 1 km de bajo de La Poza, Rio Santiago, elev. 180 m, Asuncion 118 (MO); Monte Virgin, 2 km abajo de la coin- mm free, pubescent at the point of insertion, the munidad de Caterpiza, trocha de Mitayar, banda oeste de anthers 1—1.5 mm long; ovary ovoid, 1—1.5 mm la Quebrada Caterpiza, Rio Santiago, Huashikat 610 long, 0.6—0.8 mm wide, glabrous, the style 6—9 mm (MO). Loreto: Prov. Maynas, Caserio de Huanta, Rio Oro- long, glabrous or with a lew appressed hairs along sa, a 5 minutos de la boca, margen izquierda una hora de camino, monte alto, elev. 120 m, Diaz et al, 476 (MO); the base, the stigmas capitate. Fruits drupaceous, Prov. Maynas. Yanamono, Explorama Tourist Campon Rfo white, subtended by the slightly expanded, cupu- Amazonas between Indiana and mouth of Rfo Napo, elev. late calyx, the exocarp evidently strigillose to near¬ 120 m, 3°28'S, 72°48'W, Gentry et al. 29129 (MO-2); ly glabrous, endocarp asymmetrically ellipsoid, Prov. Alto Amazonas, Andoas, Rfo Pastaza near Ecuador border, elev. 210 m, 2°48'S, 76°28'W, Gentry et al. 29783 1.3—1.8 cm long, 0.8—1.1 cm broad. (MO); Prov. Maynas, Iquitos, Estacion Biologica Allpa- Distribution. Cordia kingstoniana is wide¬ huayo, elev. 130 m, 3°53'S, 73°25'W, Jaramillo & Marcos 709 (MO); Prov. Maynas, Distrito Pebas, Rfo Ampiyacu, spread in western Amazonia in Peru, Ecuador, and cerca a estacion IMARPE, 1 km antes de Quistocha, Re- adjacent Brazil, and it should be expected in south¬ villa 936 (MO); Prov. Maynas, Distrito Iquitos, Rfo Nanay, eastern Colombia. earretera del Caserio de Lupuna to Santa Clotilde, in front This is a relatively widespread species known of Bella Vista. Rimachi 1786 (MO); Prov. Maynas, Distrito from a significant number of collections. This dis¬ Iquitos, Rfo Nanay near mouth, earretera de Picumyacu, trocha del fundo “Monte Azul", elev. ca. 160 m, Rimachi tinctive species is remarkably constant in appear¬ 6633 (MO); Prov. Maynas, Distrito Iquitos, earretera del ance and can easily be recognized by its slender caserio de Puerto Almendras, monte alto, arcilloso, cerca twigs and short-petioled narrowly elliptic leaves al terreno de la UNAP, elev. ca. 140 m, Rimachi 6948 that are prominently acuminate. Romoleroux & Fos¬ (MO); Prov. Maynas, Puerto Almendras, elev. 122 m, 3°48'S, 73°25'W, Vdsquez & Jaramillo 4238 (MO); Prov. ter 2015 and Asuncion 118 are unusual among col¬ Maynas, Indiana, Explorama Reserve, elev. 108 m, lections of this new species in their large leaves. 3°30'S, 73°05'W, Vdsquez et al. 13449 (MO); Prov. May¬ The name Cordia trachyphylla has been applied to nas, Iquitos, Allpahuayo, Estacibn Experimental del In¬ specimens of C. kingstoniana, but leaves of the two stitute de Investigaciones de la Amazonia Peruana (IIAP), species are quite distinct. Cordia kingstoniana has Vdsquez et al. 13869 (MO), Vdsquez & Jaramillo 14066 (MO), Vdsquez et al. 16239 (MO), Vdsquez 16713 (MO), narrowly elliptic to slightly oblanceolate or lance¬ Vdsquez et al. 23544 (MO). olate leaves that are close to parallel sided and have an acumen usually more than 1.5 cm long, Cordia hatschlmchii J. S. Miller, sp. nov. TYPE: while C. trachyphylla has more ovate leaves, defi¬ Brazil. Espfrito Santo: Mun. Linhares, Reserva nitely not parallel sided, and with an acumen less Elora da Sooretama, Rod. BR-101, 8 Apr. than 1 cm long. 1984, G. Hatschbach 47742 (holotype, MO This species is named in honor of David G. I. 3648229; isotypes, MBM, MO). Figure 3. Kingston of Virginia Polytechnic Institute and State University who has contributed greatly to our Frutex 2 m alta, ramunculis pubescentibus. Folia per- knowledge of the medicinal value of plant-derived sistentia; lamina oblanceolata ad anguste ellipticam, compounds, particularly those used to treat cancer. 14.5—30 cm longa, 3.8—6.3 cm lata, apice acuminato, basi Prof. Kingston is the group leader and coordinates attenuata ad acutarn, margine integra et leviter revoluta, supra scabra, pagina inferiore pubescenti; petioli 4—6 mm the ICBG project. longi. Inflorescentia terminalis, cymosa, 6—9.5 cm lata. Flores bisexuales; calyx tubularis, 5-5.5 mm longus, 10- Additional specimens examined. BRAZIL. Amazon¬ costatus; corolla alba, tubularis, 6.5-8 mm longa, 5-lob- as: Sao Paulo de Olivenya, Mori et al. 9182 (MO). Para: ata; stamina 5, fibs ca. 8.5 mm longis, pubescentia ad Mun. de Itaituba, estrada Santarem—Cuiaba. BR 163, Km 1229. Silva 385 (MO). Rondonia: vicinity of Santa Bar¬ insertionem. bara, 15 km E of Km 1 17, Prance & Ramos 7181 (MO); Shrub 2 m tall, the stems stiffly pubescent, the Porto Velho, Represa Samuel, forest along PR-3 road, ca. 10 km SE of dam, 8°49'S, 63°24'W, Thomas et al. 5133 hairs spreading, 0.5-1 mm long. Leaves persistent; <— Figure 2. Cordia kingstoniana J. S. Miller. —a. Flowering branch. —b. Fruit asymmetrical in persistent calyx. —c. Gynoecium. —d. Corolla opened to show insertion of stamens. —e. External surface of opened calyx. —f. Lower surface of leaf base. (All from Vdsquez et al. 2143, MO, holotype.) 426 Novon I*igure 3. Cordia hatschbachii J. S. Miller. —a. Flowering branch. —b. External surface of opened calyx, G\ noecium. —d. Corolla opened to show insertion of stamens. (All from Hatschbach 47742, MO, holotype.) Volume 11, Number 4 Miller 427 2001 South American Cordia Species leaf blade oblanceolate to narrowly elliptic, the dioecy. However, the South American C. trachy- broadest point above the middle, 14.5-30 cm long, phylla, C. trichoclada DC., C. chamissoniana Don, 3.8—6.3 cm wide, the apex acuminate, the base at¬ C. gardneri I. M. Johnston, C. sipapoi Gaviria, C. tenuate to acute, the margin entire and slightly rev¬ decipiens 1. M. Johnston, and the three new species olute, the adaxial surface scabrous, very rough to described here are quite diverse morphologically, the touch, coarsely and stiffly pubescent along the may not be a monophyletic group, and are probably primary and secondary veins, the abaxial surface a different lineage from the species with costate stiffly pubescent, densely so on the primary and calyces in Central America. Yet costate calyces is secondary veins, venation brochidodromous, the an easily observable character useful for recogni¬ midrib slightly elevated above, distinct and terete tion of species, so the key presented below is for below, coarsely pubescent on both surfaces, the purposes of identification and is not meant to con¬ secondary veins 8 to 12; petioles 4-6 mm long, vey any sense of relationship. canaliculate above, coarsely pubescent. Inflores¬ cences terminal, cymose, 6—9.5 cm broad, coarsely Key to the Neotropical Species <>e Cordia sect. Myxa pubescent, the peduncles 0.8—5.5 cm long. Flowers with Costate Calyces bisexual; calyx tubular, 5—5.5 mm long, 1.5—2 mm wide at the mouth, 10-costate, densely pubescent, la. Leaf margins serrate to minutely denticulate; Central America and Mexico. 5-lobed, the lobes triangular, ca. 1 mm long; corolla 2a. Leaf base acute, obtuse, or rounded. white, tubular, 6.5-8 mm long, 5-lobed, the lobes 3a. Leaves at least 3 times as long as spreading to reflexed, ovate, ca. 2.5 mm long, 1.8 wide; stigma lobes clavate; calyx mm wide; stamens 5, the filaments ca. 8.5 mm long, 3.7—6.5 mm long. the upper 4 mm free, pubescent at the point of 4a. Upper leaf surface evenly stri- gose; corolla 7.5—10.5 mm insertion, the anthers ca. 1.5 mm long; ovary ovoid, long; southern Mexico to Pan¬ ca. 1.5 mm long, 1 mm wide, glabrous, the style ama .C. diversifolia ca. 5 mm long, the stigmas capitate. Young fruits 4b. Upper leaf surface glabrous to drupaceous, subtended by the calyx, the exocarp scabrous; corolla 5.8—6.5 mm long; Guatemala .... C. skutchii minutely strigillose, mature fruits unknown. 3b. Leaves usually less than 3 times as Distribution. Cordia hatschbachii is known only long as wide; stigma lobes filiform; calyx 3-4 mm long; Mexico to Nic¬ from the type collection from Atlantic coastal for¬ aragua .C. salvadorensis ests in Espi'rito Santo. 2b. Leaf base cordate; Mexico to Guatemala This species is quite distinctive in its long (14.5- .C. cordiformis 30 cm) narrowly oblanceolate leaf blades that are lb. Leaf margins entire; South America. somewhat clustered near the ends of branches. This 5a. Leaves greater than 15 cm wide. 6a. I ^eaves bullate; southeastern Brazil species is named in honor of Gert Hatschbach, from .C. trichoclada the Museu Botanico Municipal in Curitiba, Brazil, 6b. Leaves not bullate; Brazil and Peru whose collections and research have contributed so . C. chamissoniana much to our knowledge of the forests of coastal Bra¬ 5b. Leaves less than 13 cm wide. zil. The type collection is one of many invaluable 7a. Leaves elliptic or elliptic-ovate, ca. twice as long as wide. collections that he has made documenting the di¬ 8a. Stems and leaves pilose, the versity and conservation importance of this highly hairs more than 1 mm long. threatened region so rich in endemic species. 9a. Leaf apex caudate; ovary glabrous; Suriname Cordia sect. Myxa is a large, diverse group, and .C. koemarae recognition of groups of related species within the 9b. Leaf apex acute to atten¬ section has proven difficult. One character that is uate; southeastern Brazil very distinctive in the section, is easily visible on .C. gardneri 8b. Stems and leaves various, the specimens, and occurs in only a limited number of hairs less than 1 mm long. species is the presence of a costate calyx. This 10a. Ovary glabrous. character facilitates species recognition but most 11a. Leaf base acute; probably does not circumscribe a monophyletic southeastern Bra¬ group. The four species with costate calyces in zil . . C. trachyphylla lib. Leaf base obtuse Central America (C. cordiformis I. M. Johnston, C. to rounded; Vene¬ diversifolia A. DC., C. salavadorensis Standley, and zuela .C. sipapoi C. skutchii I. M. Johnston) are probably closely re¬ 10b. Ovary pubescent; north¬ lated, as they also share serrate leaf margins and western Brazil C. decipiens 428 Novon 7b. Leaves lanceolate to narrowly ellip¬ Johnston, I. M. 1930. Studies in the Boraginaceae VIII. tic or oblanceolate, at least 3 times Observations on the species of Cordia and Tournefortia as long as wide. known from Brazil, Paraguay, Uruguay, and Argentina. 12a. Hairs on steins erect; leaf base Contr. (Tray Herb. 92: 3-89. rounded to acute; Brazil, Ec¬ -. 1935. Studies in the Boraginaceae X. The Bor¬ uador and Peru C. kingstoniana aginaceae of northeastern South America. J. Arnold Ar¬ 12b. Hairs on stems spreading; leaf bor. 16: 1-64. base attenuate to acute; south¬ Miller, J. S. 1988. A revised treatment of Boraginaceae eastern Brazil . . C. hatschbachii for Panama. Ann. Missouri Bot. Card. 75: 456-521. -. 2001. Boraginaceae. In: W. I). Stevens, C. Ulloa Acknowledgments. Financial support for the U., A. Pool & 0. M. Montiel (editors). Flora de Nica¬ ICBG project was provided by the National Insti¬ ragua. Monogr. Syst. Bot. Missouri Bot. Card. 85. 1: tutes of Health, the National Science Foundation, 435-455. and the U.S. Agency for International Development, Nowicke, J. W. & J. E. Ridgway. 1975. Pollen studies in under Grant No. UO1 TW/CA-00313. Joshua Ro¬ the genus Cordia (Boraginaceae). Amer. J. Bot. 60: 584-591. senthal, of the Fogarty International Center, NIH. -& J. S. Miller. 1990. Pollen morphology of the and James Rodman, NSF, have provided tremen¬ Cordioideae (Boraginaceae): Auxemma, Cordia, and Pa- dous support and encouragement for the ICBG pro¬ tagonula. PI. Syst. Evol. [Suppl. 5]: 103-121. ject. 1 also thank Barbara Alongi for the illustra¬ Taroda, N. & P. E. Gibbs. 1986. Studies on the genus tions. Cordia L. (Boraginaceae) in Brazil. 1. A new infrage¬ neric classification and conspectus. Revista Brasil. Bot. 9: 31—42. Literature Cited -&-. 1987. Studies in the genus Cordia L. Candolle, A.P. de. 1845. Boraginaceae. In: Prodromus (Boraginaceae) in Brazil. 2. An outline taxonomic re¬ Systematis Naluralis Regni Vegetabilis 9: 466—501. vision of subgenus Myxa Taroda. Hoebnea 14: 31-52.