Phytotaxa 20: 33–46 (2011) www.mapress.com/ phytotaxa / Copyright © 2011 Magnolia Press Article ISSN 1179-3155 (print edition) PHYTOTAXA ISSN 1179-3163 (online edition) Taxonomic and nomenclatural notes on Guatteria australis (Annonaceae) ADRIANA Q. LOBÃO1, RENATO DE MELLO-SILVA2, PAUL J.M. MAAS3 & RAFAELA C. FORZZA4 1 Universidade Federal Fluminense, Departamento de Biologia Geral/Instituto de Biologia, Rua Outeiro de São João Batista, s/n. Campus do Valonguinho. 24020-150, Niteroi, Rio de Janeiro, Brazil; email: alobao@hotmail.com 2 Universidade de São Paulo, Departamento de Botânica, Rua do Matão, 277, 05508-090, São Paulo, SP, Brazil 3 National Herbarium Nederland, Wageningen University branch, Generaal Foulkesweg 37, Wageningen, The Netherlands 4 Jardim Botânico do Rio de Janeiro, Rua Pacheco Leão, 915, Rio de Janeiro, 22460-030, Rio de Janeiro, RJ, Brazil Abstract Guatteria is the largest genus of Annonaceae, comprising ca. 300 species. The genus presents taxonomic problems, and the number of species has been overestimated. Taxonomic revision, description, comments and illustration of G. australis are presented here. As a result, 41 names have been placed in synonymy under G. australis, and three lectotypes are newly designated. Resumo Guatteria é o maior gênero da família Annonaceae compreendendo ca. 300 espécies. O gênero apresenta problemas taxonômicos e o número de espécies tem sido superestimado. Revisão taxonômica, descrição, comentários e ilustração de G. australis é apresentada aqui. Como resultado 41 nomes foram sinonimizados com G. australis e três lectótipos designados. Key words: Atlantic Forest systematics, Annonaceae, Brazilian floristics, Guatteria, Mata Atlantica Introduction Guatteria Ruiz & Pavón (1794: 85) is the largest genus of the family Annonaceae, comprising 307 species (Erkens et al. 2008). It is distributed from southeastern Mexico south to Bolivia and southern Brazil (Barringer 1984). The genus is characterized by the impressed primary vein on the upper side of the leaves, a pedicel with a distinct suprabasal articulation, more or less equal whorls of imbricate petals, and numerous single-seeded stalked monocarps (Erkens et al. 2006, 2007a). Fries (1939) authored the last taxonomic revision of the genus, in which 151 species were recognized from Brazil (106 of which he newly described) and the species classified into 30 sections. Fries (1949, 1953, 1959) changed Guatteria infrageneric classification and created two subgenera, Guatteria subg. Anomalantha (monotypic, comprising only G. anomala R.E.Fr.) and Guatteria subg. Guatteria, with 22 sections, including all Brazilian species. In Guatteria subg. Guatteria there are some difficult species complexes such as the G. trichoclonia complex (distributed throughout the full range of the genus), the G. schomburgkiana complex (occurring in the Guyana Shield and Amazonian regions), and the G. amplifolia complex (Central America; Erkens 2007b). The close affinities among the species in these complexes are generally confirmed by phylogenetic analyses, although some species might be grouped on the basis of homoplastic characters (e.g., the hairs of the G. trichoclonia complex; Erkens 2007b). Accepted by M. Chase: 15 Jan. 2011; published: 1 Mar. 2011 33 The majority of the southeastern Brazilian species of Guatteria (60) are classified in only 10 of the 22 sections of Guatteria subg. Guatteria; only five taxa are not included in these sections (Table 1). Almost all extra-Amazonian taxa occur in the same general geographic area and have morphological similarities. Among these, it is possible to recognize one large complex, G. australis. The similarities among the species of the G. australis complex have been noted previously. Saint-Hilaire (1825) commented on the morphological affinity among G. australis (section Austroguatteria), G. gomeziana, and G. lutea (both section Asterantha) and the possibility that they might be variants of a single species. Martius (1841) observed that G. klotzschiana (Microcalyx) is taxonomically close to G. nigrescens (Asterantha), and Fries (1939) commented that G. australis, G. polycarpa, and G. neglecta (all three Austroguatteria) are impossible to distinguish without the flowers. However, the flowers of the types of all these species overlap in characteristics, and therefore they do not warrant distinction on the specific level. Moreover, Fries (1939) perceived similarities between G. silvatica and G. gomeziana (Asterantha) and also between G. klotzschiana and G. reflexa (both Microcalyx), describing the last from a specimen identified as G. klotzschiana. Lobão & Mello-Silva (2007) suggested maintaining G. glabrescens , G. nigrescens, and G. reflexa on the basis of characters of the leaf blade, but these characters have since proved to be variable after examination of more collections from the Atlantic Forest. Almost all species from southeastern Brazil are known only from their types, and most of these species were described from the same area of the southeastern states of Brazil. For example, G. curvinervia , G. clavigera , G. fruticosa , G. mosenii , G. polycarpa subs. drupacea, G. sordida var. stenopetala, G. sordida var. lancifolia, and G. sordida var. ovalis are all from São Paulo state, the majority from the same locality. Similarly, G. paranensis and G. salicifolia var. erosa are from Paraná state, where only one morphological form of Guatteria is known (A. Lobão, pers. obs.). The presence of G. australis in Paraná and its possible occurrence in Rio de Janeiro and Minas Gerais have been mentioned previously (Fries 1939). TABLE 1: Sections of Guatteria that include extra-Amazonian species (with the exception of section Tylodiscus, see Lobão & Mello-Silva 2007). The taxa and geographic occurrence are sensu Fries (1939). * Amazonian species not included in this study. Section Taxa Occurrence of section Austroguatteria G. australis Saint-Hilaire G. curvinervia Fries G. densicoma Martius G. dusenii Fries G. dusenii var. subglabra Fries G. fruticosa Fries G. glabrescens Fries G. neglecta Fries G. paranensis Fries G. parvifolia Fries G. parvifolia var. vestita Fries G. polycarpa Fries G. polycarpa subsp. drupacea Fries G. salicifolia Fries G. salicifolia var. erosa Fries Brazil (Paraná, Rio de Janeiro, São Paulo) Asterantha G. acutipetala Fries G. asterantha Fries G. gomeziana Saint-Hilaire G. latifolia (Martius) Fries G. lutea Saint-Hilaire G. mosenii Fries G. nigrescens Martius G. reticulata Fries Brazil (Minas Gerais, Rio de Janeiro, São Paulo) 34 • Phytotaxa 20 © 2011 Magnolia Press LOBÃO ET AL. Section Taxa Occurrence of section Brachystemon G. myriocarpa Fries * G. peckoltiana Fries G. pubens (Martius) Fries G. sellowiana Schlechtendal G. umbrosa Fries G. xylopioides Fries Brazil (Minas Gerais, Pará, Rio de Janeiro) Cordylocarpus G. clavigera Fries Brazil (São Paulo) Dimorphopetalum G. dimorphopetala Fries Brazil (Bahia) Macroguatteria G. boliviana H. Winkler * G. burchellii Fries G. campestris Fries G. eximia Fries * G. ferruginea Saint-Hilaire G. hilariana Schlechtendal G. hilariana var. angustifolia Schlechtendal G. hilariana var. cuneata Fries G. hilariana var. latifolia Schlechtendal G. hilariana var. verrucosa Fries G. lawrancei Fries * G. pogonopus Martius G. saffordiana Pittier * G. speciosa Fries * Brazil (Amazônia, Bahia, Minas Gerais, Rio de Janeiro, São Paulo) Venezuela Bolívia Colômbia Microcalyx G. blanchetiana Fries G. klotzschiana Martius G. reflexa Fries G. silvatica Fries Brazil (Bahia, Rio de Janeiro) Microphyllum G. pohliana Schlechtendal Brazil (Minas Gerais) Oligocarpus G. bahiensis Fries G. odontopetala Martius G. oligocarpa Martius G. sabuletorum Fries * G. schlechtendaliana Martius G. acutiflora Martius G. candolleana Schlechtendal G. eriopoda De Candolle ex Dunal * G. hookeri Saint-Hilaire G. macropus Martius G. mexiae Fries G. microcarpa Ruiz et Pavon * G. minarum Fries G. ouregou (Aublet) Dun. * G. penduliflora Fries G. psilopus Martius G. riedeliana Fries G. sodiroi Diels * G. tomentosa Rusby * G. trichoclonia Diels G. villosissima Saint-Hilaire G. villosissima var. longepedunculata Fries Brazil (Bahia, Minas Gerais, Pará) Trichoclonia TAXONOMIC/NOMENCLATURAL NOTES ON GUATTERIA AUSTRALIS Brazil (Bahia, Espírito Santo, Minas Gerais, Rio de Janeiro) Bolivia Ecuador French Guiana Peru Phytotaxa 20 © 2011 Magnolia Press • 35 Section Taxa Occurrence of section With no section G. sordida Fries G. sordida var. stenopetala Fries G. sordida var. lancifolia Fries G. sordida var. ovalis Fries G. tenuis Fries Minas Gerais São Paulo The morphological boundaries between many of these species have thus become uncertain. Species delimitation elsewhere in Guatteria has been problematic (Erkens et al. 2007) because of the uniformity of reproductive characters among species (van Heusden 1992), the variability of the vegetative characters within species, and the significant changes in petal size and orientation as the flowers mature. Variability in these characters has in a number of cases caused the number of species to be overestimated. Moreover, phylogenetic reconstructions for Guatteria (Erkens et al. 2007) have shown poor resolution in the portion of the trees containing members of the G. australis-complex. The purpose of this study was to review the taxonomy of Guatteria species occurring in southern and eastern Brazil using collections assembled since the time of the last revision (Fries 1939). A taxonomic treatment, description, illustrations, nomenclatural notes, geographic distributions, and phenology of G. australis are presented. Material and methods Collections from the herbaria ALCB, B, BHCB, BM, BR, C, CEPEC, ESA, F, FCAB, FLOR, G, GUA, GFJP, HAMAB, HB, HRB, HRCB, HUEFS, IAN, K, LZ, M, MBM, MBML, MG, MO, NY, OUPR, OXF, P, R, RB, RBR, RFA, RUSU, S, SP, SPF, SPSF, U (incorporated in L), UB, UEC, UPCB, VIC, W, and WU (following Holmgren et al. 1990) were analyzed. Also, field observations were made in several of the main Brazilian vegetation types. The synonyms are listed in chronological order. Names from the same publication are listed in pagenumber order. Material examined by! Taxonomic treatment Guatteria australis Saint-Hilaire (1825: 37). Figs. 1, 2. Guatteria australis var. glabrata Martius (1841: 26), nom. inval. Type:—BRAZIL. São Paulo: Itararé (“Bords du Tareré”), January 1817, Saint-Hilaire 1452 (holotype P!, isotypes P!, 2 sheets). Guatteria gomeziana Saint-Hilaire (1825: 36), syn. nov. Type:—BRAZIL. Minas Gerais: Rio Piracicaba (“Itajuru de São Miguel de Mato Dentro”), Janusary-February 1817, Saint-Hilaire 560 (holotype P!, 2 sheets, isotype K!). Guatteria lutea Saint-Hilaire (1825: 37), syn. nov. Guatteria flava Saint-Hilaire ex Martius (1841: 27), nom. superfl. Type:—BRAZIL. Minas Gerais: Ouro Preto (“entre Vila Rica et Antônio Pereira”), January 1817, Saint-Hilaire 304 (holotype P!). Uvaria hirsuta Vellozo (1827: 238), nom. illeg. non Uvaria hirsuta Jack (1820: 46). Type (not designated): Fl. Flum. tab. 124. Uvaria monosperma Vellozo (1827: 238), nom. illeg. non Uvaria monosperma Lamark (1785: 596). Type (not designated): Fl. Flum. tab. 123. Guatteria hilariana Schlechtendal (1834: 324). Guatteria hilariana f. latifolia Schlechtendal (1834: 324), nom. inval. Guatteria hilariana var. latifolia (Schltdl.) Martius (1841: 35), nom. inval. Type:—BRAZIL. Without locality and date, Sellow s.n. (holotype B!, probable isotype HAL). Guatteria hilariana f. angustifolia Schlechtendal (1834: 324), syn. nov. Guatteria hilariana var. angustifolia (Schltdl.) Martius (1841: 35). Type:—BRAZIL. São Paulo: Santos, April-July 1829, Sellow 5943 p.p. (holotype B! 2 sheets, isotype HAL). 36 • Phytotaxa 20 © 2011 Magnolia Press LOBÃO ET AL. Guatteria psilopus Martius (1841: 27), syn. nov. Type:—BRAZIL. Rio de Janeiro: Teresópolis: Serra dos Órgãos, June 1834, Luschnath s.n. (holotype BR!). Guatteria nigrescens Martius (1841: 31). Guatteria nigrescens var. oblongifolia Martius (1841: 31), nom. inval. Type:— BRAZIL. São Paulo: Guaratinguetá, December 1817, Martius s.n. (holotype M!, 4 sheets, probable isotype B!). Guatteria densicoma Martius (1841: 32), syn. nov. Type:—BRAZIL. Bahia: Ilhéus, fazenda Almada, December 1818, Martius 711 (holotype M!, isotypes B!, BR!, F!, HAL, K!, NY!, P!). Guatteria klotzschiana Martius (1841: 32), syn. nov. Cananga klotzschiana (Mart.) Warming (1874: 144). Type:— BRAZIL. Rio de Janeiro: “near Tocaja”, without date, Schott s.n. (holotype BR!, probable isotypes B!, F!, US!). Guatteria odontopetala Martius (1841: 33), syn. nov. Lectotype (designated by Fries, 1939):—BRAZIL. Minas Gerais: Manoel Pereira, without date, Pohl 2964 (M!, duplicates B!, BR!, F, G). Guatteria hookeri Saint-Hilaire & Tulasne (1842: 132), syn. nov. Type:—BRAZIL. Rio de Janeiro: Teresópolis, Organ Mountains, banks of the river Paquequer, March 1837, Gardner 306 p.p. (holotype P!, 3 sheets, isotypes BM!, E, F!, G!, K! 3 sheets, LZ, OXF!, NY!, S!). Guatteria neglecta Fries (1939: 308). Guatteria paranensis Fries (1939: 314), nom. illeg. Type:—BRAZIL. Paraná: Morretes, Jacareí (“Jacarehý”), 30 September 1908, Dusén 6780 (holotype S!, 2 sheets, isotypes F!, K!, L, MO!, NY! 2 sheets, P!, U!). Guatteria polycarpa Fries (1939: 310). Type:—BRAZIL. Paraná: Piraquara Roça Nova-Banhado, 30 December 1908, Dusén 7414 (holotype S!, isotypes F!, K!, L, MO!, NY!, P!, U!). Guatteria polycarpa subsp. drupacea Fries (1939: 311), syn. nov. Type:—BRAZIL. São Paulo: Without locality, 26 September 1922, Kuhlmann s.n. (RB 19677) (holotype S!, isotype RB!). Guatteria glabrescens Fries (1939: 311), syn. nov. Type:—BRAZIL. Rio de Janeiro: reserva florestal da fábrica Alliança, 11 January 1922, Kuhlmann s.n. (RB 4483) (holotype S!, isotype F!, RB! 2 sheets). Guatteria fruticosa Fries (1939: 312), syn. nov. Type:—BRAZIL. São Paulo: São José dos Campos, 14 December 1909, Löfgren 531 (holotype S!). Guatteria dusenii Fries (1939: 313), syn. nov. Type:—BRAZIL. Paraná: Guaratuba, Jul 1911, Dusén 13752 (holotype S!, isotypes L!, MO!). Guatteria dusenii var. subglabra Fries (1939: 313), syn. nov. Type:—BRAZIL. Paraná: Paranaguá (“Porto D. Pedro II”), 19 April 1904, Dusén 4497 (holotype S!). Guatteria parvifolia Fries (1939: 314). Type:—BRAZIL. São Paulo: São Paulo, Parque do Estado, 28 October 1931, Hoehne s.n. (SP 28405) (holotype S!, isotypes B!, F, HB!, K!, MO!, SP!, SPF!, US!). Guatteria parvifolia var. vestita Fries (1939: 315), syn. nov. Type:—BRAZIL. São Paulo: Without locality or date, Sellow 211 (holotype B!, probable isotype K). Guatteria curvinervia Fries (1939: 317), syn. nov. Type:—BRAZIL. São Paulo: Barretos, Dec 1917, Frazão 8667 (holotype S!, isotype RB!). Guatteria salicifolia Fries (1939: 316), syn. nov. Type:—BRAZIL. Rio de Janeiro: Rio de Janeiro, Tijuca, 21 December 1874, Glaziou 7508 (holotype B!, isotypes C, K!, MG, P!). Guatteria salicifolia var. erosa Fries (1939: 316), syn. nov. Type:—BRAZIL. Paraná: Morretes, Jacareí (“Jacareý”), 1914, Dusén s.n. (holotype S!, isotype F!). Guatteria mosenii Fries (1939: 321), syn. nov. Type:—BRAZIL. São Paulo: (“Serra do Caracol”), 15 December 1875, Mosén 4002 (holotype S!). Guatteria asterantha Fries (1939: 323). Type:—BRAZIL. Minas Gerais: Viçosa (“road to Cajurý, Canella”), 7 January 1931, Mexia 5485 (holotype S!, isotypes BM, F! 2 sheets, K!, NY!, U!, US!). Guatteria acutipetala Fries (1939: 324), syn. nov. Type:—BRAZIL. Rio de Janeiro Teresópolis, January 1897, Ule 3962 (holotype: B!). Guatteria reflexa Fries (1939: 329), syn. nov. Type:—BRAZIL. Rio de Janeiro: Niterói, Morro da Viração, 28 July 1870, Glaziou 5725 (holotype S!, isotypes B!, C, F!, K!, P!, R!, RB!). Guatteria silvatica Fries (1939: 330), syn. nov. Type:—BRAZIL. Rio de Janeiro: Cantagalo (“Canta Gallo”), 1857, Peckolt 151 (holotype BR!). Guatteria blanchetiana Fries (1939: 331), syn. nov. Type:—BRAZIL. Bahia: Ilhéus, fazenda Almada, without date, Blanchet 2114 (holotype G!, 6 sheets, isotypes B! BM!, F!, K! 2 sheets, MO!, NY!, P! 2 sheets, W!). Guatteria dimorphopetala Fries (1939: 333), syn. nov. Type:—BRAZIL. Between Vittoria (Espírito Santo) und Bahia, without date, Sellow 175 (holotype B!). Guatteria clavigera Fries (1939: 334), syn. nov. Type:—BRAZIL. São Paulo: São Paulo, Serra da Cantareira, 15 December 1933, Koscinsky 214 (holotype S, isotype SP!). Guatteria riedeliana Fries (1939: 339), syn. nov. Type:—BRAZIL. Bahia: Ilhéus, fazenda Almada, November 1821, Riedel 424 (holotype S, isotype NY!, U!). Guatteria penduliflora Fries (1939: 341), syn. nov. Type:—BRAZIL. Rio de Janeiro: Without locality, 1860, Allemão & Cysneiro s.n. (holotype G!, 2 sheets). Guatteria minarum Fries (1939: 346), syn. nov. Type:—BRAZIL. Minas Gerais: Viçosa (“Fazenda Paraiso”), 1 October TAXONOMIC/NOMENCLATURAL NOTES ON GUATTERIA AUSTRALIS Phytotaxa 20 © 2011 Magnolia Press • 37 1930, Mexia 5130 (holotype S!, isotype C, BM!, F!, 2 sheets, K!, NY!, U!). Guatteria hilariana var. cuneata Fries (1939: 397), syn. nov. Type:—BRAZIL. São Paulo: Santos, on Montserrat, northeast part of island São Vicente, 3 November 1826, Burchell 3366-2 (holotype K!, isotype K!, P!). Guatteria hilariana var. verruculosa Fries (1939: 397), syn. nov. Type:—BRAZIL. São Paulo: Santos, Baturoca, 25 January 1875, Mosén 3337 (holotype S!, isotypes B, P!, US). Guatteria sordida Fries (1939: 532). Type:—BRAZIL. São Paulo: Without locality and date, Lund s.n. (holotype C!). Guatteria sordida var. lancifolia Fries (1939: 533), syn. nov. Type:—BRAZIL. São Paulo: “prope Muji”, November 1833, Riedel 1689 p.p. (holotype S, isotype K!). Guatteria sordida var. ovalis Fries (1939: 533), syn. nov. Type:—BRAZIL. São Paulo: “prope Muji”, November 1833, Riedel 1689 p.p. (holotype S!, isotype B!, K!, P!). Guatteria sordida var. stenopetala Fries (1939: 533). Type:—BRAZIL. São Paulo: Mogi (“Mugy”), November 1833, Lund 959 (holotype C!). Guatteria tenuis Fries (1939: 535). Type:—BRAZIL. Minas Gerais: Viçosa, Fazenda do Deserto, 5 January 1931, Mexia 5482 (holotype S!, isotypes BM!, F!, K! U!). Trees or shrubs 1–13(–26) m tall, DBH 6–20 cm. Young twigs glabrous to densely covered with appressed, golden hairs. Leaves: petiole canaliculate, 2 – 8 mm long, blackish, glabrous to densely covered with appressed, golden hairs; blade 5.0–20.0 × 1.5–7.0 (–8.5) cm, chartaceous, narrowly elliptic to elliptic, brown to blackish or bluish when dry, glabrous to sparsely covered with appressed, golden hairs above, glabrous to densely covered with appressed, golden hairs below, primary vein flat on upper side, glabrous, secondary veins distinct, slightly impressed above, raised below, angles with primary vein 70–90?, tertiary veins reticulate, raised on both sides, base acute to attenuate, apex acute to acuminate, acumen 5–15 mm long. Flowers: solitary, axillary; bracts 2, caducous or persistent; pedicels 15–60 mm long, blackish, glabrous to densely covered with appressed, golden hairs; flower buds 4–8 × 5–10 mm, broadly ovoid, with free, valvate sepals, rarely connate when young; sepals 5–10 × 5–7 mm, broadly triangular, valvate, free, rarely connate at base, reflexed, glabrous on both sides; petals on both cycles, 10–38 × 5–20 mm, narrowly elliptic, imbricate, free, greenish, maturing yellowish to pale red, glabrous externally, sparsely covered with curly hairs internally; stamens ca. 2 mm long, connective shield papillate; ovary ca. 2.5 mm long, sparsely covered with erect, golden-brown hairs, stigma papillate. Fruit: torus 4–7 × 5–11 mm, broadly obovoid; monocarps 10–60, ellipsoid, 6–15 × 5–7 mm, atro-vinaceous in vivo, blackish when dry, apex shortly apiculate, stipes 10–30 mm long, green, red to purple in vivo, blackish when dry, glabrous. Seed ellipsoid, 6–10 × 5–7 mm, adhering to pericarp, longitudinally ridged, orange in vivo, brownish to dark brown when dry. Guatteria australis shows enormous morphological variation. In general, it is characterized by leaves of moderate dimensions (5.5–15.0 × 2.0–5.0 cm) and a long flowering pedicel reaching 60 mm in length. The species varies considerably in the size of the leaf and the presence of indumentum (Fig. 1). The leaves are normally glabrous, but the abaxial surface of the leaves, young twigs, and pedicels may be glabrous to densely covered by appressed, golden hairs. In Rio de Janeiro and Paraná, this variation can be found in individuals from the same area (Mello-Silva 1993). For example, in the same area individuals are found with young twigs and leaves with both surfaces glabrous, and others with young twigs and abaxial surface of the leaves densely hairy. In Minas Gerais, however, where G. australis is common, the leaves are normally glabrous. Nomenclatural notes:—Guatteria australis was described from Saint-Hilaire 1452 (Saint-Hilaire 1825). Martius (1841) described G. australis var. glabrata instead of using G. australis var. australis based on the same type. According to Article 26.2 of the ICBN, G. australis var. glabrata is an invalidly published name. Martius (1841) wrongly substituted the name G. lutea for G. flava, G. flava being superfluous according to Article 52.2 of the ICBN. Velloso (1827) described Uvaria hirsuta Vell., but this name is illegitimate, being a later homonym of U. hirsuta Jack (1820) (Article 53.1 of the ICBN). Martius (1841) synonymized U. hirsuta Vell. under G. hilariana Schltdl. Lamarck (1785) described Uvaria monosperma Lam. Velloso (1827) described U. monosperma Vell., but this is illegitimate, being a later homonym (Article 53.1 of the ICBN). 38 • Phytotaxa 20 © 2011 Magnolia Press LOBÃO ET AL. FIGURE 1. Guatteria australis. A. Branch with flower. B–D. Leaves. E. Apex acute, glabrous. F–G. Leaves. H. Base decurrent, glabrous. I. Leaf. J. Base acute, densely covered with trichomes. K–L. Leaves. M. Flower. N. Fruit. (A, M–N. Seele 107 (RB). B. Grande 242 (SP). C. Reginato 223 (RB). D–E. Landrum 2215 (RB). F. Lobão 704 (RB). G–H. Pessoal do Horto Florestal 80 (RB). I–J. Lima 3561 (RB). K. Nadruz 463 (RB). L. Custodio Filho 1571 (RB)). TAXONOMIC/NOMENCLATURAL NOTES ON GUATTERIA AUSTRALIS Phytotaxa 20 © 2011 Magnolia Press • 39 Schlechtendal (1834) described G. hilariana based on Sellow s.n. together with two forms, G. hilariana f. latifolia and G. hilariana f. angustifolia, both with no type cited in the protologue. In the Berlin herbarium there is one specimen identified by Schlechtendal as the type of G. hilariana f. latifolia and another identified as the type of G. hilariana f. angustifolia. The type of G. hilariana f. latifolia is the same type as that of G. hilariana, and should be named G. hilariana f. hilariana. Thus G. hilariana f. latifolia is not a validly published name (Article 26.2 of the ICBN). Martius (1841) indicated himself as the collector of the type of G. psilopus, with the abbreviation “M” in the protologue of the species. Fries (1939) could not trace this collection in either BR (Brussels) or M (Munich) and suggested that Luschnath s.n., Brazil, Rio de Janeiro, Serra dos Orgãos, June 1834, from the “Herbarium Martii” and deposited in BR, and annotated by Martius himself, would be the type instead. Guatteria nigrescens var. oblongifolia should be named G. nigrescens var. nigrescens, thus G. nigrescens var. oblongifolia is an invalid name (Article 26.2, ICBN). Martius s.n. was chosen as the lectotype for G. nigrescens by Fries (1939). Fries (1939) designated the same type, Dusén 6780, for both G. neglecta and G. paranensis, both being legitimate names of equal priority. Thus, the first choice effectively published (Fries 1959: 81) established the priority of G. neglecta, and G. paranensis became a superfluous name (Art. 11.5, ICBN). Taxonomic notes:—Hairy forms of G. australis may resemble G. candolleana. Guatteria candolleana, however, is a distinct species, characterized by leaves with an obtuse or cordate base and a short pedicel. Guatteria penduliflora, G. hookeri, and G. psilopus have been treated as synonymous with G. candolleana (Lobão & Mello-Silva 2007). However, these three species are distinct from G. candolleana in their leaves with an acute base and by the long pedicels, and they fit within the range of morphological variation of G. australis. Therefore, they are synonymized with this species. Lobão and Mello-Silva (2007) treated G. klotzschiana, G. acutipetala, and G. silvatica as synonymous with G. nigrescens. Some authors (e.g., Martius 1841, Fries 1939, Lobão et al. 2005, 2006, Lobão & MelloSilva 2007) have considered G. nigrescens to be a species distinct from G. australis. Guatteria nigrescens has larger leaves than those of the type of G. australis, but our studies have shown that the leaves of G. australis are variable in length. For this reason, G. nigrescens is here treated as synonymous with G. australis, as previously proposed by Maas et al. (2002). If G. nigrescens is synonymous with G. australis, then consequently G. klotzschiana, G. acutipetala, and G. silvatica become new synonyms of G. australis. Comparison of the types of G. australis, G. gomeziana, and G. lutea proved that they are extremely similar to each other, and therefore G. gomeziana and G. lutea are now considered as synonyms of G. australis. These three names have equal priority, but the name G. australis has been adopted because it is a widely used name and an appropriate specific epithet. The morphological similarities among these and the possibility that they might be variants of a single species was stressed by their author, Saint-Hilaire (1825). Martius (1841) described G. odontopetala based on Blanchet 2114 and Pohl 2964. According to Fries (1939) these collections represent two species. He selected Pohl 2964 (M) as the lectotype of G. odontopetala and described G. blanchetiana based on Blanchet 2114. All types of Guatteria hilariana and its infraspecific taxa occur within the same geographical area (Fries 1939). They share many features of the type of G. australis, differing mainly in leaf size and indumentum. These features are highly variable within the populations of G. australis, however, and fall entirely within the range of variation of G. australis. The type of G. dusenii is similar to that of G. australis, only differing from the latter in its dense indument on young twigs and pedicels. However, since the density of hair cover is notoriously variable, this does not support distinction at the specific level. Guatteria glabrescens shares all features with G. australis, except for the decurrent leaf base. Distinguishing G. glabrescens based only on this highly variable feature is not justified. Maintaining G. glabrescens, G. nigrescens, and G. reflexa based on characters of the leaf blade, as done by Lobão & Mello-Silva (2007), in our opinion no longer seems appropriate because these characters have proven to be variable upon examination of more collections from the Atlantic Forest. Therefore they are now treated as synonyms of G. australis. 40 • Phytotaxa 20 © 2011 Magnolia Press LOBÃO ET AL. The types of both G. salicifolia and G. minarum have slender elliptic leaves, resembling the leaves of some Salix species. However, this extreme leaf morphology occurs throughout the range of G. australis and is not by itself sufficient to sustain the two species as distinct entities; G. salicifolia and G. minarum are treated as synonyms of G. australis. Additional Guatteria species described from Bahia are included in the synonymy of G. australis because their types were found to fall within the range of variation now known for G. australis. The types of G. blanchetiana and G. riedeliana, for example, have larger leaves than the type of G. australis, but they all share so many other features, such as the long pedicel and chartaceous leaves with cuspidate apex, that these types clearly fall within the same species concept. Guatteria densicoma and G. dimorphopetala, on the other hand, share with G. australis the size of the lamina, long pedicels, and elliptic, chartaceous leaves. All other names, viz. G. clavigera, G. curvinervia, G. fruticosa, G. gomeziana, G. lutea, G. mosenii, G. paranensis, G. polycarpa subsp. drupacea, G. reflexa, G. sordida var. stenopetala, G. sordida var. lancifolia and G. sordida var. ovalis are treated as synonymous with G. australis because their types fit within the range of morphological variation of G. australis. Geographical distribution and habitats:—Guatteria australis is distributed from Bahia to Rio Grande do Sul, reaching eastern Goiás to the west (Fig. 2). It occurs in all kinds of forests in southeastern Brazil, reaching elevations of 2,000 m at Campos do Jordão, São Paulo, and is common in disturbed areas. It is frequent in the Zona da Mata of Minas Gerais and in São Paulo, where the complete range of variation of the species can be found. It has also been found in gallery forests in Minas Gerais and São Paulo and hydrophilic forests in Bahia. FIGURE 2. Distribution of Guatteria australis ( ). Phenology:—Guatteria australis has been collected with flowers and fruits during the entire year. Many populations show intense flowering and fruiting, frequently on the same individual. Conservation status:—Low risk. Guatteria australis is widely distributed in the Atlantic Forest and generally represented by large and widespread populations and therefore is at low risk of extinction. Nevertheless, it may become locally threatened where its populations have only a few individuals, such as in the states of Goiás and Rio Grande do Sul. TAXONOMIC/NOMENCLATURAL NOTES ON GUATTERIA AUSTRALIS Phytotaxa 20 © 2011 Magnolia Press • 41 FIGURE 3. Guatteria australis. A–B. Branch with flower. C. Immature flower. D. Mature flower. E–F. Immature fruits. (Photographs: A–B, D–F Lobão, C. Paula, E. Matos). Acknowledgements The authors thank Gea Zijlstra, David Johnson (reviewer), Roy Erkens (reviewer), Mariana Saavedra, and Cassia Sakuragui. We also thank all the curators of the herbaria visited, especially M, C, G and P. This research was supported by the FAPERJ, CAPES, and IAPT Foundations. Renato de Mello-Silva thanks the 42 • Phytotaxa 20 © 2011 Magnolia Press LOBÃO ET AL. Margaret Mee Amazonian Trust for supporting a stay at RBG-Kew, and visits to BM, OXF, P and U herbaria (incorporated in L), for studying Annonaceae. Renato de Mello-Silva and Rafaela C. Forzza are CNPq research fellows. References Barringer, K. (1984) A new species of Guatteria (Annonaceae) from Panama. Annals of the Missouri Botanical Garden 71: 1186–1187. Erkens, R.H.J., Maas, P.J.M., Chatrou, L.W., Schatz, G.E. & Zamora, N. (2006) Seven taxonomic discoveries in Annonaceae from southeastern Central America. Blumea 51: 199–220. Erkens, R.H.J., Koek-Noorman, J., Maas, J.W. & Maas, P.J.M. (2007A) Classification of a large and widespread genus of Neotropical trees, Guatteria (Annonaceae) and its three satellite genera Guatteriella, Guatteriopsis and Heteropetalum. Taxon 56: 757–774. Erkens, R.H.J., Chatrou, L.W., Maas, J.W., Ginkel, M.V. & Maas, P.J.M. (2007B). Taxonomic problems in the Central American Guatteria amplifolia complex (Annonaceae) cannot be elucidated by AFLP analyses and sequence markers. Pp. 101–115. In: Erkens, R.H.J. From morphological nightmare to molecular conundrum. Phylogenetic, evolutionary and taxonomic studies on Guatteria (Annonaceae). Ph.D. thesis. Erkens, R.H.J., Westra, L.Y.T. & Maas, P.J.M. (2008) Increasing diversity in the species-rich genus Guatteria (Annonaceae). Blumea 53: 467–514. Fries, R.E. (1900). Beiträge zur Kenntnis der Süd-Amerikanischen Anonaceen. Kongliga Svenska VetenskapsAkademiens Handlingar 34: 9–19. Fries, R.E. (1939) Revision der Arten einiger Anonaceen-Gattungen V. Acta Horti Bergiani 12: 289–540. Fries, R.E. (1959) Annonaceae. 171 p. In: Engler, H.G.A. & Prantl, K.A.E. (eds.). Die natürlichen Pflanzenfamilien 2 ed., 17a. Duncker & Humblot, Berlin. Heusden, E.C.H. van. (1992) Flowers of Annonaceae: morphology, classification, and evolution. Blumea Supplement 7: 1–218. Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. (1990) Index herbariorum: Part I. The herbaria of the world. 8th ed. The New York Botanical Garden, New York. 693pp. Jack, W. (1820). Malayan Miscellanies 1: 46. Lamarck, J.B.A.P.M. (1785) Encyclopédie méthodique. Botanique 1(2). Panckoucke. Paris. Lobão, A.Q., Araujo, D.S.D. & Kurtz, B.C. (2005) Annonaceae das restingas do Estado do Rio de Janeiro. Rodriguésia 56: 85–96. Lobão, A.Q., Forzza, R.C. & Mello-Silva, R. (2006) Annonaceae da Reserva Biológica da Represa do Grama, Descoberto, Minas Gerais, Brasil, com uma nova espécie, Unonopsis bauxitae. Rodriguésia 57: 137–147. Lobão, A.Q. & Mello-Silva, R. (2007) Guatteria (Annonaceae) do estado do Rio de Janeiro, Brasil. Rodriguésia 58: 859–884. Maas, P.J.M., Maas-van de Kamer, H., Junnika, L., Mello-Silva, R. & Rainer, H. (2002, "2001") Annonaceae from central-eastern Brazil. Rodriguésia 52 (80): 65–98. Martius, C.F.P. (1841) Annonaceae. Pp. 1–63 In: Martius, C.F.P. & Eichler, A.G. (eds.). Flora brasiliensis 13. Frid. Fleischer, Leipzig. Mello-Silva, R. (1993) Annonaceae. Pp. 43–51. In: Mello, M.M.R.F., Barros, F., Chiea, S.A.C., Wanderley, M.G.L., Jung-Mendaçolli, S.L. & Kirizawa, M. (eds.). Flora Fanerogâmica da Ilha do Cardoso 3. Instituto de Botânica, São Paulo. Ruiz, H.L. & Pavón, J.A. (1794) Guatteria P. 85. In: Ruiz, H.L. & Pavón, J.A. (eds.). Florae peruvianae, et chilensis. Prodromus. La Imprenta de Sancha, Madrid. Saint-Hilaire, A.F.C.P. (1825) Flora Brasiliae meridionalis 1. A. Belin, Paris. Saint-Hilaire, A.F.C.P. & Tulasne, L.R. (1842) Revue de la flore du Brésil méridional. Annales des Sciences Naturelles, Botanique, série 2. 17. Schlechtendal, D.F.L. (1834) De Anonaceis brasiliensibus herbarii regii berolinensis. Linnaea 9: 315–331. Velloso, J.M.C. (1827) Florae fluminensis icones. Typographia National, Rio de Janeiro. Warming, J.E.B. (1874) Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1873. TAXONOMIC/NOMENCLATURAL NOTES ON GUATTERIA AUSTRALIS Phytotaxa 20 © 2011 Magnolia Press • 43 Index of exsiccatae Abreu N. L. 132; Aguiar O. T. 433; Albuquerque J. E. 1835 , 1860; Allemão s/n (G); Almeida A. L. 30; Alves L. J. 321; Amorim A. M. 4516, 6887; Anderson W. R. 35703, 35909; Andrade A. P. M. 775; Anunciação E. A. 11, 12; Aparicio A. 56; Aragaki S. 53, 466, 467, Araujo D. S. D. 1365, 4762, 6490, 6549, 6580, 7293, 10407, 10496; Araújo F. S. 122; Araujo s/n (OUPR 225), s/n (OUPR 252); Arbo M. M. 7714, 7811; Árbocz G. F. 2338, 4205, 32705, 32706; Arzolla F. A. R. D. P. 525; Assis L. C. S. 441, 606, 883, 887, 939, 944, 1001, 1014; Assis M. A. 33, 400; Auler L. 1; Ávila N. S. 5, 466; Badini J. s/n (OUPR 222), s/n (OUPR 23341), s/n (OUPR 23342); Baitello J. B. s/n (SP 258840); Barbiellini A. A. s/n (SP 40584); Barbosa E. 37, 265, 556, 631; Barreto H. L. M. 925, 3206; Barreto K. D. 814, 1100, 1469, 3152; Barros A. M. M. 2423; Barros F. 691, 709, 1571, 1928, 2037, 2375, 2965; Barros W. D. 31, 1191; Batista E. R. 16, 64; Belém R. P. 583, 3062, 3328; Benko-Iseppon A. M. 16; Bernacci L. C. 931, 1008; Bianchini R. S. 1444; Blanchet J. S. 2114 ; Boone W. 484; Borges C. M. O. s/n (VIC 14981); Borges R. A. X. 789; Bovini M. G. 507, 552, 639, 650, 1085, 1123, 1941; Brade A. C. 7238, 7239, 16780, 18733, 20949; Braga J. M. A. 937, 2730, 4706; Braidotti J. C. 134; Brina A. E. s/n (BHCB 39314); Britez R. M. 398, 1455, 24790; Burchell W. J. 1839, 2014, 3429, 3054 pp, 3054 pp, 3134-2, 3366-2, 3737-2, 4078, 4612; Busquetti 96; Buzato S. 22500; Campos D. P. 4; Carneiro J. 1299, 1481; Carvalho A. M. V. 4123, 5148; Carvalho J. P. M. s/n (SPSF 8716); Carvalho-Sobrinho J. G. 80, 104; Castro A. G. 195; Castro R. M. 465, 515; Catharino E. L. M. 475, 542, 665, 768, 1240; Cavalcanti D. C. 294; Ceccantini G. C. 129; Cervi A. C. 2636, 3506; Chagas F. s/n (FUEL 2021); Chiea S. C. 414, 700; Coelho J. C. R. 700; Colares J. E. R. 85; Cordeiro I. 1264, 1649, 1935, 1979, 2383, 2798, 2906, s/n (SPF 43643); Correia C. M. B. s/n (RB 295531); Costa L. V. 345, s/n (BHCB 11936), s/n (BHCB 17666), s/n (BHCB 22504), s/n (BHCB 52468); Cruz A. M. R. s/n (247061), s/n (SP 247069), s/n (SP 247081); Cruz J. M. 18; Custodio Filho A. 612, 1470, 1505, 1571, 1624, 1628, 1749, 1873, 1928, 2029, 2144, 2241, 2524, 4584; Damasceno Junior G. A. 29298; Damazio L. s/n (OUPR 219), s/n (OUPR 220); Davis P. H. D 60645, D 60774; Dedeca D. s/n (SP 69618); Demuner V. 34, 223, 432, 449, 616, 631, 736, 917; Destefani A. C. C. 224; Dombroushi P. K. 1631, 7122, 24160; Dombrowski L. T. 179, 1072; Duarte A. P. 2297, 3204, 3115, 4307, 4975, 6801, 8455, s/n (RB 4614); Ducke A. s/n (RB 16584); Dusén P. K. H. 4497, 6780, 4144, 7414, 13746, 13752, 15364, s/n (S); Echternacht L. 215; Edwall 4189; Ehrendorfer F. 73825; Elias S. I. 72, 302; Emmecrich M. 6112; Equipe seção Botânica s/n (SP 268361); Esteves G. L. CFCR 15457; Esteves R. 4, 24; Fadelli L. 258, 322; Farág P. R. 478; Farah F. T. 2178, 2181, 2312; Faria P. C. L. s/n (CESJ 34484); Farias D. S. 164; Farinaccio M. A. 450, 585, 664, 713; Fernandes D. 636, 677; Ferreira F. M. 463; Ferreira G. M. P. 28; Ferreira R. M. 49; Ferreti A. R. 66, 95, 133, 132, 156; Fiaschi P. 34, 434, 488, 514, 662, 731, 1298, 1095; Ficher D. C. H. 8; Figueiredo N. 14411, 15607; Filho A. M. s/n (RB 78224); Folli D. A. 2258, 2522, 3436; Fontana A. P. 984; Forero E. 8144, 8720; Forzza R. C. 1717, 2030; Fraga C. N. 1842, 1970; França F. 2589; França G. S. 144, 505; Francisco E. M. s/n (MBM 257088), s/n (MBM 257089); Franco G. 2967; Franco G. A. D. C. 438; Frazão 8667; Furlan A. 681, 912, 1101; Galetti M. s/n, col. Saibidela 1116, col. Saibidela 120, col. Saibidela 146; Gandolfi S. 11924, s/n (ESA 33500), s/n (UEC 79124); Garcia F. C. P. 77, 290; Garcia R. J. F. 263, 309, 331, 546, 716, 759, 954, 986, 996, 1003, 1963; Gardner G. 306 p.p., 4403; Gehrt A. s/n (SP 3703), s/n (SP 35285), s/n (SPF 71929); Gentry A. H. 49358, 49813, 58789; Geraldino H. C. L. 121; Géza Árborez 181, 553, 574; Gianotti E. 14924; Giordano L. C. 1782, 2024; Glaziou A. F. M. 1596, 2481, 2677, 2981, 3858, 3878, 5725, 7504, 7505, 7506, 7508, 8253, 8254, 10223, 10224, 13513, 15824, 17462; Godoy J. R. L. 59, 124, 143, 168, 169, 170, 171, 172, 173; Godoy S. A. P. 599; Goes M. s/n (SPF 105301); Goés O. C. 46, 63, 432, 793, 1069; Gomes B. Z. 93, 95; Gomes J. 65; Gomes V. 104, 2486; Gonçalves S. B. 257; Gorenstein M. R. 14, 115; Gorgatti L. 2, 4, 5, 6; Gottsberger G. 11-3468, 11-12268A, 11-12268B, 11-15168, 1128969, 11-91179, 12-12468, 12-27168, 12-29969, 13-1268, 13-111066, 21-7175, 21-241186, 21-28872, 21-9274B, 22141174, 23-211275, 26-51172, 32-71090; Grande D. A. 242; Grombone-Guaratini M. T. 11, 12, 307; Guedes L. M. 3119, 5480; Guedes R. R. 242, 2159, 2173; Hage J. L. 30, 448, 1044, 1391, 1232, 1280, 1446, 1475, 1768; Halfeld s/n (OUPR 234); Handro O. s/n (SP 50441); Harley R. M. 17870, 21888; Hatschbach G. G. 968, 1860, 5300, 5302, 5521, 7505, 13636, 14620, 30375, 32115, 35777, 37005, 40207, 40693, 41821, 42600, 50840, 51735, 52556, 53497, 53736, 54915, 56162, 57916, 58179, 59840, 61472, 68859, 69291, 71521, 72736; Heringer E. P. 472, 18566; Hoehne F. C. s/ n (S), s/n (SP 1145), s/n (SP 1472), s/n (SP 1518), s/n (SP 1869), s/n (SP 3002), s/n (SP 5066), s/n (SP 5725), s/n (SP 17873), s/n (SP 26558), s/n (SP 28405), s/n (SP 29355), s/n (SP 39251), s/n (SP 73147), s/n (SPF 77748), s/n (SPF 77750); Hoehne W. s/n (SPF 67487); Honda S. 801, s/n (PMSP 494); Horta M. 5030; Irwin H. S. 22031, 22483; Ivanauskas N. M. 869; Jardim J. G. 491, 1163, 4163, 4422, 4474, 4890; Jesus N. G. 1160; Jung S. L. 434, 450; JungMendaçolli S. L. 570; Junior A. J. G. s/n (VIC 29537); Kawall M. 362; Kinoshita L. S. 95-67; Kirizawa M. 866, 1148, 1271, 1426, 1454, 1980, 2022, 2405, 2411, 2605, 2620, 2819, 2882, 3061, 3195, 3209, 3410; Kitakawa A. Y. s/n (SPF 124837), s/n (SPF 124838); Klein R. 178; Klein V. L. G. 923 962, 1202; Kollmann L. 527, 819, 858, 892, 1501, 44 • Phytotaxa 20 © 2011 Magnolia Press LOBÃO ET AL. 1765, 1948, 2142, 2655, 3774, 4480, 6570, 7989; Konno T. 111; Koscinsky M. 214, 235; Kozera C. 802; Krieger L. 11145, 11516, 13351, 19308, 21127, 25039; Kuhlmann J. G. s/n (RB 4483), s/n (RB 19677), s/n (RB 19677), s/n (RB 81370), s/n (VIC 2126); Kuhlmann M. 219, 531, 575, 1541, 2261, 2598, 2676, 4580, s/n (SP 36278), s/n (S), s/n (S); Kummrow R. 1442, 1656, 3211; Kurtz B. C. 117, s/n (RB 290853); Landrum L. R. 2215, 4067; Leitão Filho H. F. 278, 1305, 1351, 2211, 10237, 10392, 10758, 10774, 10784, 15358, 22836, 27766, 32663, 34722, 34734, 34761; Leite E. C. 276, 829; Leitman M. 383; Leoni L. S. 2077, 2098, 4081, 5378, 5546, 6260; Lima G. P. s/n (RB 429710); Lima H. C. 3405, 3561, 3695, s/n (RB 290843), s/n (RB 290852); Lima L. R. 388, 419; Lima M. P. M. 169; Lima S. 13245; Limões J. A. 1, 53; Lindeman J. C. 1977, 2330, 3058, 4069, 5646, 5764, 5864, 3058A, 5299B; Lobão A. Q. 444, 446, 447, 466, 490, 494, 497, 499, 526, 530, 534, 565, 625, 645, 646, 647, 648, 675, 677, 684, 693, 702, 714, 720, 727A, 738, 743, 770, 1288, 1289, 1302, 1303, 1315, 1355, 1358; Lobb W. 37, s/n (K); Löfgren 531, 1455, 3339, 4189, 4190, 4577; Lombardi J. A. 529, 5538, 6232, s/n (BHCB 7286); Lopes M. A. 775; Lopes M. M. M. 558; Lopes W. P. 518, s/n (VIC 17067); Los M. M. 101; Loureiro D. M. 222; Luchiari C. 696; Lund 959, s/n (C); Luschnath s/n (BR); Lutz B. 733; Maas P. J. 3222, 6969, 7035, 7086, 7087, 8816, 8837; Macedo I. C. C. 60; Magalhães M. 307; Major Novaes s/ n (R 60771), s/n (R 60774), s/n (R 60777); Mamede M. C. H. 94, 106; Mannu M. s/n (FUEL 3741); Manu M. s/n (FUEL 3648); Marassi R. D. 24; Markgraf F. 10121, 13059; Marques M. C. 304; Marquete N. 474; Marquete R. 162, 177, 272, 413, 829, 1232, 1239, 1759, 1895, 3719; Martinelli G. 550, 3064, 3181, 9841, 10539, 10541, 10560, 11713, 11823, 12258, 12986; Martini A. 3012; Martins D. 103; Martins F. R. 2131, 2180, 15872; Martins S. E. 242; Martius C. F. P. 711, s/n (M s/n), s/n (M s/n); Martuscelli P. 197; Matthes L. A. F. 669, 7748, 7749, 7750, 7751; Mattos J. 9099, 13180, 13926, 13932, 14271; Medeiros D. A. 14; Meireles L. D. 723, 852, 1051; Mello-Silva R. 35, 992, 1239, 1254, 1258, 1616, 1630, 1633, 1728, 1800, 1805, CFCR 5752; Melo E. 2471; Melo M. M. R. F. 425, 557, 579, 579, 634; Mendonça 758; Messias M. C. T. B. 869, s/n (OUPR 1395); Mexia Y. 4945, 5024, 5086, 5130, 5131, 5289, 5482, 5485 ; Miers J. 4018, s/n (OXF); Mizue Kirizawa 2022; Moraes P. L. R. 744, 767, 787, 788, 789, 823, 851, 1085; Morawetz W. 11-241280, 11-261280, 11-281280, 11-4181, 11-101280, 11-211280, 11-71280, 11-91280, 12-4181, 12-101280, 12-281280, 13-281280, 21-7181, 21-11181, 21-12381, 21-201280, 21-261280, 22-4181, 12101280, 31-12181, 31-29181; Mori S. A. 11860, 13858; Mosén 3337, 4002; Mota N. F. O. 513; Mota R. C. 1594; Moura J. 715; Moura L. C. 23; Müller F. 434; Muniz C. F. S. 505; Nadruz M. 463, 499, 506, 572; Negrelle R. A 496, A 637, s/n (R 169520); Neto J. A. A. M. 775, 2084, 21346, 23553; Neto S. J. S. 1460; Nogueira J. E. 230; Novaes C. s/ n (SP 2064); Occhioni P. 113; Ogata H. 792, s/n (PMSP 3855); Okano R. M. C. s/n (VIC 13028); Oliveira C. A. L. 1739; Pabst G. F. J. 5934; Paes L. E. 3; Paiva M. R. C. s/n (SPF 139066); Paixão J. L. 698; Pardo C. S. 725; Pastore J. A. s/n (SPF 112645), s/n (SPSF 8362); Paula C. H. R. 724, 780; Peckolt 151; Pedra E. F. 8; Pedroni F. 223, 235, 30449, 30450; Pedrosa D. S. 1133; Pedroso 26; Peras M. s/n (OUPR 2217); Pereira B. A. S. 3347, 3362; Pereira E. 18, 454, 655, 3831, 3851, 4273, 4868, 5120, 6898; Peron M. 767, 769, 881; Pessoa S. V. A. 124, 144, 438, 454, 463, s/ n (RB 290840); Pessoal do Horto Florestal 80, 81; Pickel C. s/n (SP 53377); Pifano D. S. 196, 207; Pinheiro F. 276, 708; Pinheiro R. S. 443; Pinto C. G. 19; Pioker F. C. s/n (SPF 139096); Pirani J. R. 552, 1367, 4418, 4753, 5102; Pizziolo W. 342; Pohl 2964, s/n (BR s/n); Prance G. T. 6950; Puttemans A. s/n (RBR 2057); Queiroz J. M. 30132; Queiroz L. P. 6496; Ramalho S. R. 935; Reginato M. 223; Regnell A. F. III-252; Reis A. 149, 150; Reitz R. 1408, 2490, 2819, 4680, 5725, 8325, 8787; Ribas O. S. 32, 253, 264, 302, 777, 809, 854, 961, 2118, 3003, 3028, 3388; Ribeiro J. E. L. S. 202, 532, 629; Ribeiro R. 967, 1015, 1061, 1100, 1108, 1136, 1141, 1171, 1219, 1243, 1773, 2056, 2112; Richard A. s/n (B s/n); Riedel L. 424, 1172, 1178, 1689 pp; Rinoli J. s/n (BHCB 16390); Robim M. J. 351, s/n (SPSF 8383); Rodela L. G. T3QA-2; Rodrigues R. R. 196, 14973, s/n (SPF 47165); Romaniuc Neto S. 131, 199, 1071; Rosa N. A. 3710; Rosa P. s/n (RB 81409); Rosquel M. B. 294, 373, 471; Rossi L. 19, 34, 39, 45, 80, 103, 131, 131-A, 557, 818, 2024, 2123, s/n (PMSP 557), s/n (SPF 47164), s/n (SPF 71529), s/n (SPF s/n), s/n (SPF s/n); Rotta E. 91; Sá C. F. C. 1233, 2591; Saavedra M. M. 518; Saint-Hillaire A. F. C. P. 304, 560, 1452, s/n (P s/n), s/n (P s/n), s/n (P s/n); Sakuragui C. M. 368; Salimena F. R. s/n (CEJF 31013), s/n (CESJ 31397); Salino A. 5906; Sambuichi R. H. R. 858A, 1335ª; Sampaio D. 55, 74; Sampaio P. S. P. 153; Sanches C. D. 11, 59 ; Sanchez M. 2 ; Sant’Ana S. C. 183, 861; Santiago S. A. 29; Santos H. 145; Santos T. S. 1347, 1771, 2224, 2850, 3402, 3775; Sattler D. 88; Sazima M. 16935; Scaramuzza C. A. M. 379; Scheel-Ybert R. 335; Schenk H. 4079; Schott H. W. s/n (BR 997337), s/n (997330), s/n (BR); Schwacke C. A. W. 8907; Seele C. 107, 205; Sellow F. 175, 211, 4510, s/n (B), 5943 p.p.; Shepherd G. J. 8201, 10972; Shirasuna R. T. 73; Silva A. F. 126, 1296, 1353, 1362,, 1468, 1479, 1496, 1514, 1542, 1549, 1565, 2573; Silva A. G. s/n (OUPR 3547); Silva F. C. 704; Silva Filho C. A. 64; Silva G. P. 6082; Silva G. s/n (SPSF 13305); Silva J. M. 73, 277, 1288, 2858, 3320, 3927; Silva J. S. 2; Silva S. M. 1324, 24562, s/n (SPF 49419); Silva S. P. C. 654; Silveira A. L. 101; Simao-Bianchini R. 15, 493, 517, 851, 924, 932; Siqueira M. F. 22026; Skvortzov B. s/n (SP 106042); Smith L. B. 14889; Soares A. 109; Sobral M. 5491, 5740, 6603; Solórzano A. 32; Sousa V. C. 4156, 4465, 4760, 8796, 9018, 9289, 9292, 10559, 21617, 21674, 21694, 23211, s/n (SPF 202277); Souza TAXONOMIC/NOMENCLATURAL NOTES ON GUATTERIA AUSTRALIS Phytotaxa 20 © 2011 Magnolia Press • 45 F. M. 187; Souza H. C. 102; Souza H. M. s/n (SP 268302), s/n (UEC 66944); Souza L. C. 175; Souza W. S. 24787, 24788, 24789, 25246, s/n (SPF 49420), s/n (UEC 51701); Stehmann J. R. 4011, s/n (BHCB 20303); Stubblebine W. H. 13193; Sucre D. 2415, 3184, 3476, 6590, 7385, 7445, 7812, 7965, 9516, 10588; Sugiyama M. 304, 853, 854, 864, 874, 1043, 1390, 1398; Sztutman M. 259; Takahasi A. 359; Tamashiro J. Y 649, 1157, 1278, 1642, 10552, 18620, 18711, 21280, 176969; Tameirão Neto E. 63, 758, 2785, 2899, 2900, 3497, 3721, 4252, 4253; Tatagiba F. 47; Tatto L. s/n (RB 45792); Tavares R. A. M. 57; Thomas W. W. 3920, 6010, 9872, 11224, 11240, 11583, 12006, 13790 ; Thomaz L. D. 790, 1291 ; Toledo F. R. N. s/n (BHCB 22766) ; Toledo Filho D. V. 10708; Tomasulo P. L. B. 347; Toniato M. T. 29270; Tweedie J. s/n (K); Torezan J. M. 512, 721; Uhlmann A. 70; Ule E. 969, 3962, 4525, 26X122; Urbano 8885; Valente G. E. 1096, 1097; Valões J. s/n (SPF 105309); Vasconcelos M. F. 78; Vaz A. F. 566, 643; Veloso H. 65C; Versieux L. M. 22; Vervloet R. R. 72, 2189; Vidal M. M. 28 ; Vieira C. M. 63, 301 ; Vieira M. C. W. 1010; Vieira M. F. 486, 649; Vitta F. A. s/n (SPF 75606), s/n (SPF 143379); Walter B. M. T. 88; Webster G. L. 25556 ; Weir J. s/ n (BM), s/n (K s/n); Wendt T. 183; Wesenberg J. 550; Wilms W. 405 A; Záchia R. 722; Zampa P. C. s/n (CESJ 31173); Zborowski M. 6; Zipparro V. B. coleção saibadela 308, coleção saibadela 420. 46 • Phytotaxa 20 © 2011 Magnolia Press LOBÃO ET AL.