PHYTOTAXAISSN 1179-3155 (print edition)

ISSN 1179-3163 (online edition)Copyright © 2013 Magnolia Press

Phytotaxa 94 (1): 13–22 (2013) www.mapress.com/phytotaxa/ Article

http://dx.doi.org/10.11646/phytotaxa.94.1.2

Taxonomic notes on South American Miconia (Melastomataceae). III

RENATO GOLDENBERG1 & MAYARA K. CADDAH2

1 Universidade Federal do Paraná, Departamento de Botânica, Centro Politécnico, Caixa Postal 19031, Curitiba, PR, 81531-970, Brazil; email: renato.goldenberg@gmail.com.2 Universidade Estadual de Campinas, Pós-Graduação em Biologia Vegetal, Cidade Universitária Zeferino Vaz, Campinas, SP, 13083-970, Brazil.

Abstract

We propose nine synonyms and five lectotypifications for South American Miconia (Melastomataceae). Miconia acreana is synonymized under M. erioclada; Miconia depauperata and M. doriana under M. paniculata; Miconia inconspicua var. glabrata under M. petropolitana; Miconia japuraensis under M. juruensis; Miconia martiusiana under M. cinnamomifolia; Miconia maximiliana under M. ibaguensis, Miconia pileata under M. ciliata and Miconia pseudonervosa under M. nervosa. We propose lectotypes for M. acreana, M. doriana, M. juruensis, M. longicuspis and M. pileata.

Key words: Brazil, lectotypes, Miconieae, nomenclature, synonyms, taxonomy, types

Introduction

Miconia Ruiz & Pavon (1794: 60) is a genus with more than 1050 species native to the Neotropics (Goldenberg et al. 2008). As it should be expected for a large polyphyletic group occurring in a region with a recent history of collections and taxonomic revisions (i.e., outside Europe and North America), it is still badly in need of much basic taxonomic work. Paraphrasing a metaphor from J.J. Wurdack (1921–1998), one of the most prolific taxonomists of Melastomataceae, the rug is too big and a lot of sweeping is needed before laying something on it (H. Robinson, pers. comm.). This means that a complete monographic and systematic work on this group is still far from being achieved. There are still many undescribed species in the genus, mostly

from Andean countries, and also a backlog of 18th to early 20th century taxonomy in need of evaluation and revision (Howard & Kellog 1986, Martin et al. 2007, Goldenberg & Martin 2008, Goldenberg et al. 2010, Michelangeli & Goldenberg 2012). This is the case for some of the species studied here, which were known only from the types (some of which are very poor specimens) collected almost two centuries ago. Other species descriptions were based on a sampling that was too poor to adequately understand morphological variation over wide geographic areas. In this paper we tried to solve some of these taxonomic problems regarding poorly circumscribed species, proposing synoymyzations and lectotypifications of some Miconiafrom South America, mostly from Brazil.

Taxonomy and Nomenclature

1. Miconia ciliata (Rich.) De Candolle (1828: 179). Melastoma ciliatum Richard (1792: 109). Acinodendron ciliatum (Rich.) Kuntze (1891: 949). Type:—FRENCH GUIANA. Leblond s.n. in Herb. Richard (Holotype: P!).

Accepted by Eve Lucas: 4 Apr. 2013; published online in PDF: 16 Apr. 2013 13

Synonym:—Miconia pileata De Candolle (1828: 180). Acinodendron pileatum (DC.) Kuntze (1891: 952). Type:—BRAZIL. “Provinciae Minas Geraes limites versus Paranáan”, C.F.P. von Martius s.n. (Lectotype: M-0165639!, here designated). Syntype:—BRAZIL. “Provinciae flum. Nigri, habitat in marginibus sylvarum prope Ega et Coari”, C.F.P. von Martius s.n. (M-0165568!).

Miconia ciliata is a widespread species that ranges from Mexico to Bolivia and southeastern Brazil (Wurdack 1973, Wurdack et al. 1993, Almeda 2009), and one of the most collected species in the family. It shows a lot of variation in density and location of its unbranched, slender trichomes (see Williams 1934 and Wurdack 1973). Some specimens lack trichomes altogether, except for the leaf margins. Several specimens have those trichomes only on the petioles, while some may have a few of them very sparsely scattered along the branches or concentrated on nodal regions. Trichomes on the actual abaxial surface of the leaf or more densely and uniformly distributed along the young branches are less common (see discussion on M. pileata var. latifoliabelow). This variation has misled much of the taxonomic work on the complex, which can be attested by the amount of synonyms already listed for M. ciliata (Cogniaux 1891, Goldenberg 2000, Goldenberg et al. in prep.). In this context, differences attributed to M. ciliata and M. pileata by the taxonomists working on the group after De Candolle (1828) actually do not reflect the character states found in the types. Another issue that contributed to this confusion was the description of several varieties under both species, in an attempt to catalog variation that apparently has a very diffuse pattern and does not follow a geographical pattern. Finally, the circumscription of M. ciliata is also unclear with respect to other closely related species like M. racemosa, (Aublet 1775: 406) De Candolle (1828: 179). We agree with Wurdack (1973: 488) when he stated that the whole Hartigia complex needs an evaluation.

Miconia pileata was described by De Candolle (1828) based on Brazilian material seen in the Martius herbarium (now at M), as “Melast. pileatum et ciliatum Schr. et Mart. mss.”. There are three collections at M named as M. pileata that were collected by Martius in three different places of Brazil. One of them, from Pará, has no Schrank handwriting on it but was determined as M. pileata by Cogniaux: it is not the type, but is the only one among the three that has trichomes on the branches, despite the fact that these trichomes are very sparse and not very conspicuous. The other two can be regarded as syntypes, since each one of them has Schrank’s descriptions and determinations, one as Melastoma pileatum (from Minas Gerais) and the other as M. ciliatum (from Rio Negro, Amazonas). Both have no trichomes on the branches, but have a few trichomes on the petiole and a few scattered trichomes on the abaxial foliar nerves, whereas some leaves in the specimen from Amazonas have some trichomes on the leaf surface as well as the nerves. The same features are exactly as found in the holotype of Miconia ciliata (Leblond s.n., herb. Richard, now at P), except for the trichomes on the surface of the leaves: no trichomes on the branches or leaf surfaces, but some on the petioles and on the nerves on the abaxial leaf surface. Since there are almost no differences between the types, the whole discussion by subsequent authors on the differences between these species based on the presence or absence of trichomes on the branches or petioles has no nomenclatural significance, being a reflection of the variability found in this species (see Cogniaux 1886–1888, Cogniaux 1891).

There are some extremes regarding the density and location of the trichomes that remain to be evaluated, like the populations in Central America and Colombia with dense indument (Duke 11284, Mori 7985). These have been treated as M. pileata var. latifolia Cogniaux (1888: 407; see Almeda 2009), but bear no closer resemblance to the typical variety of M. pileata than to M. ciliata.

For a complete list of synonyms, see Cogniaux (1891), Wurdack et al. (1993), Goldenberg (2000), Goldenberg et al. (in press).

Additional material examined:—BELIZE. Cayo, 20 November 1987, M. Balick 1840 (US). BOLIVIA, Beni: Ballivián, 24 May 1988, S.G. Beck 16366 (US). BRAZIL. Amapá: Calçoene, 8 December 1984, S. Mori 17275 (US). Amazonas: Rio Negro, Moreira, 10 November 1971, G.T. Prance 16237 (US). Bahia: Santa Cruz de Cabrália, 17 March 1974, R.M. Harley 17095 (US). Ceará: Guariba, 25 July 1964, A. Castellanos 528(US). Goiás: Niquelândia, 8 May 1998, M.L. Fonseca 1831 (US). Maranhão: São Luís, 1939, R Froes 11635 (US). Minas Gerais: Resplendor, 26 March 1997, E. Tameirão Neto 2358 (UPCB). Mato Grosso: Serra

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Ricardo Franco, 25 September 1978, P. Windisch 2166 (US). Pará: Marapanim, 20 March 1960, W.A. Egler 1392 (US). Pernambuco: Recife, 28 December 1960, S. Tavares 563 (US). Roraima: Serra Tepequem, 16 February 1967, G.T. Prance 4418 (US). COLOMBIA. Antioquia: Planta Providencia, 29 July 1974, J. Denslow 2359 (US). Chocó: Boca Curiche, 20 May 1967, J.A. Duke 11284 (US). ECUADOR. Esmeraldas: San Lorenzo, 13 July 1964, C. Jativa 756 (US). FRENCH GUIANA. Monts Bakra, 15 June 2002, J.J. de Granville 14744 (US). GUATEMALA. Petén, 24 July 1959, E. Contreras 4 (US). GUYANA. Warashema Mnt. Range, 28 February 2007, K.M. Redden 5389 (US). NICARAGUA. Cabo, 11 July 1970, G. Davidse 2310A (US). PANAMA. Cerro Azul, 28 November 1974, R. Hernandez 40 (US); Cerro Jefe, 29 August 1975, S. Mori 7985 (US). PERU. Huanuco: Pachitea, 2 July 1988, B. Wallnöfer 16-2788 (US). SURINAM. Para, 1 December 1996, R. Evans 2609 (US). TRINIDAD. Aripo, 23 March 1959, R.S. Cowan 1158 (US). VENEZUELA. Bolívar: Auyan Tepui, 26 August 1983, G.T. Prance 28201 (US).

2. Miconia cinnamomifolia (DC.) Naudin (1850: 68). Cremanium cinnamomifolium De Candolle (1828: 194). Type:—BRAZIL. No precise locality, C.F.P. von Martius s.n. (Holotype: G-DC-00311256!).

Synonym:—Miconia martiusiana De Candolle (1828: 186). Type:—BRAZIL. No precise locality, C.F.P. von Martius s.n. (Holotype: G-DC-00310942!).

The placement of Miconia martiusiana as a synonym of M. cinnamomifolia has already been suggested by J.J. Wurdack (1970). Nevertheless, the synonymyzation has never been formally proposed. Both types clearly belong to the same species, which is easily recognized by the interpetiolar projections that resemble stipules (Goldenberg 2004, 2009). For more nomenclatural details regarding this species, see Wurdack (1962, 1970).

Additional material examined:—BRAZIL. Bahia: Nova Viçosa, 10 December 1984, G. Hatschbach 48781 (US). Espírito Santo: Cariacica, 6 May 2008, R. Goldenberg 1117 (RB, UPCB). Minas Gerais: Alpinópolis, 12 February 1998, R. Goldenberg 461 (UEC, UPCB). Paraná: Adrianópolis, 18 October 2005, R. Goldenberg 754 (UPCB). Rio de Janeiro: Rio de Janeiro, 24 November 1983, C. Angeli 554 (US). Santa Catarina: Itajaí, 7 January 1955, R. Klein 1013 (US). São Paulo: Cruzeiro, 5 April 1995, R. Goldenberg 64(UEC).

3. Miconia erioclada Triana (1871: 111). Acinodendron eriocladum (Triana) Kuntze (1981: 951). Type:—PERU. Province? Tarapoto, s.d., R. Spruce s.n. in Herb. Benth. (Holotype: K-000006171!).

Synonym:—Miconia acreana Ule (1915: 360). Type: —BRAZIL. Acre: Rio Acre, Seringal São Francisco, September 1911, E. Ule 9676 (Holotype: B, destroyed; Lectotype: K-000536155!, here designated).

Ule (1915) described M. acreana without comparing it to any other species of Miconia, but indicated that it would key near M. carassana Cogniaux (1891: 346). Cogniaux (1891) distinguished M. erioclada from M. carassana by the dense or sparse unbranched trichomes on the abaxial surface of the leaf in the former, and the dense stellate trichomes on the leaf of the latter. The type of M. acreana has a sparser indument than most plants of M. erioclada, but the trichomes do not differ between them, whereas they are strikingly different from the ones found in M. carassana. It is impossible to understand why Ule compared his species to M. carassana instead of M. erioclada, from which it does not differ.

Trichome density and also leaf width are variable within M. erioclada. The specimens from Acre, Brazil, and a few from neighboring localities in Peru (Foster 3530, 11737, 11884, Gentry 18753) usually have narrower leaves with sparser indument on both leaf surfaces than plants from other sites, but these differences are not discrete enough to be worthy of formal infraspecific status. Other features, such as stamen morphology and the indument on young branches, inflorescences and hypanthium are identical.

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Additional specimens examined:—BOLIVIA. Beni: Ballivián, 31 October 1989, D.N. Smith 13807 (US). Cochabamba: Chapare, 20 November 1980, T.B. Croat 51291 (US). La Paz: Sud Yungas, 24 December 1987, R. Seidel 2215 (US). Santa Cruz: Ichilo, 12 December 1989, M. Nee 38079 (US). BRAZIL. Acre: Jordão, 6 February 2009, R. Goldenberg 1294 (RB, UPCB); Marechal Thaumaturgo, 3 May 2001, L.G. Lohmann 464(NY); Santa Rosa, 15 March 2002, D.C. Daly 11357 (NY, UPCB). COLOMBIA. Putumayo: Puerto Asis (Rio San Miguel), 13 December 1940, J. Cuatrecasas 11049 (US). Santander: San Vicente de Chucurí, 27 Feb 1964, L. Uribe-Uribe 4693 (US). ECUADOR. Esmeraldas: Esmeraldas, 8-9 April 1967, B. Sparre 15329(US). Morona-Santiago: Limón, 28 March 1974, G. Harling 12925 (US). Napo: Anangu, 6 July 1983, J.E. Lawesson 39765 (US). Pastaza: Montalvo, 18 July 1980, B. Øllgard 34643 (US). PERU. Amazonas: Bagua, 3 November 1978, P. Barbour 4398 (US). Cuzco: Paucartambo, 6-7 August 1974, R.B. Foster 3004 (US). Huanuco: Pachitea, 25 October 1967, J. Schunke 2252 (US). Junín: Tarma, 9 October 1976, R. Ferreyra 18607 (US). Loreto: Pucallpa–Tingo María Road, 27 March 1977, A. Gentry 18753 (US); Maynas, 8 October 1973, M. Rimachi 610 (US). Madre de Dios: Manu, 23 August 1976, R.B. Foster 3530 (US); 11 October 1986, R.B. Foster 11737 (US); 16 October 1986, R.N. Foster 11884 (US). San Martin: Lamas, 12 September 1937, C.M. Belshaw 3406 (US).

4. Miconia ibaguensis (Bonpl.) Triana (1871: 110). Melastoma ibaguense Bonpland (1815: 105). Tschudyaibaguensis (Bonpl.) Grisebach (1860: 47). Acinodendron ibaguense (Bonpl.) Kuntze (1891: 951). Type:—VENEZUELA. Sucre: Sta. Maria, A. Bonpland 371 (Holotype: P-00136503!).

Synonym:—Miconia maximiliana De Candolle (1828: 186). Type:—BRAZIL. No precise locality, C.F.P. von Martius s.n. (Holotype: G-DC-00310943!).

De Candolle (1828) was aware of Melastoma ibaguense, but treated it as one of his “Melast. incertae”. In the same publication, he also described M. maximiliana. Cogniaux (1888) accepted M. maximiliana and placed it near Miconia ibaguensis, distinguishing both by the young branches and hypanthia lacking simple trichomes in the former, and present in the latter. Nevertheless, there are actually a few scattered simple trichomes on the branches of M. maximiliana’s type, and these are similar to the ones in M. ibaguensis. Miconia ibaguensis is a widespread species that ranges from Mexico to Paraguay and southern Brazil (Wurdack 1973, Wurdack et al. 1993, Almeda 2009), in which this variation in the amount of trichomes on the branches and hypanthium is evident.

Additional material examined:—BELIZE. Cayo, 22 May 1996, M. Nee 46804 (US). BOLIVIA. La Paz: Nor Yungas, 19-20 October 1982, J.C. Solomon 8553 (US). BRAZIL. Acre: 10 October 1968, G.T. Prance 7960 (US). Amazonas: Manaus, 9 October 1982, S.S. Renner 872 (US). Bahia: Rio de Contas, 18 September 1989, G. Hatschbach 53449 (US). Ceará: Crato, 19 January 1983, T. Plowman 12751 (US). Distrito Federal: 16 June 1982, J.H. Kirkbride (US). Espírito Santo: Guarapari, 5 September 1980, B. Weinberg 222 (US). Goiás: Caiapônia, 18 October 1964, G.T. Prance 59453 (US). Mato Grosso: Salto do Céu, 25 October 1995, G. Hatschbach 63856 (US). Minas Gerais: São Roque de Minas, 29 September 1995, J.N. Nakajima 1404 (US). Pará: Itaituba, 14 April 1983, M.N. Silva 3 (US). Paraná: Antonina, 30 September 1977, G. Hatschbach 40200(US). Rio de Janeiro: Resende, 22 October 1981, J.P.P. Carauta 3885 (US). Roraima: 10 December 1954, B. Maguire 40198 (US). São Paulo: Campo Alegre, 24 September 1940, J.F. Toledo s.n., SP 43172 (US). COLOMBIA. Cesar: Rio Oro, 19 July 1974, H. Garcia-Barriga 20575 (US). COSTA RICA. Cartago: 7 May 2002, D.S. Penneys 1531 (US). ECUADOR. Zamora-Chinchipe: Zumba, 12 October 1988, G. Harling 25146(US). EL SALVADOR. Sierra de Osicala, 1929, S. Calderon 2503 (US). GUATEMALA. Petén: 13 May 1970, R. Tun 1070 (US). GUYANA. Potaro-Siparuni: 16 January 1993, T.M. Henkel 791 (US). HONDURAS. Lempira, 8 March 1969, A. Molina 24141 (US). MEXICO. Oaxaca: 28 December 2001, D.S. Penneys 1448(US). NICARAGUA. Cabo, 11 July 1970, G. Davidse 2312 (US). PANAMA. Chiriquí: Boquete, 10 March 1963, W.L. Stern 1947 (US). PARAGUAY. San Pedro: Cororó, 20 April 1995, A. Schinini 29379 (US). PERU.

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Loreto: Gran Pajonal, June 1976, G.A.J. Scott 884 (US). SURINAME. Paramaribo, 15 November 1960, K.U. Kramer 2064 (US). VENEZUELA. Portuguesa: Guanare, 12 March 1982, R. Liesner 12548 (US).

5. Miconia juruensis Pilger (1905:172). Type:—BRAZIL. “Estrema, Jurua Miry, Rio Jurua” (probably State of Acre, near Cruzeiro do Sul), September 1901, E.Ule 5827 (Holotype: B, destroyed; Lectotype: K-000536204!, here designated; Isolectotypes: HBG-518982, MG!, US [fragm.]!).

Synonym:—Miconia japuraensis Cogniaux (1909: 253). 190. Type:—BRAZIL. “Rio Japurá, Jupará (probably “Japurá’”), 19 September 1904, A. Ducke 6794 (Holotype: BR-5202263!; Isotypes: MG!, US-00121024 [fragm.]!).

Miconia japuraensis was published in 1909, in an issue that was intended to be printed in 1907. This means that it was prepared in less than two years after the publication of Miconia juruensis. It is not clear whether Cogniaux (1909) was unaware of M. juruensis or simply misled by a different sectional placement of the species (sect. Miconia by Pilger vs. sect. Tamonea by Cogniaux), but he did not mention it, and both types are very similar. They share the same minute sessile glands and sparse stellate trichomes (these mostly on the nerves) on the abaxial surface of the leaves, whorled inflorescence branches, obtuse-denticulate or undulate-denticulate calyx lobes, glandular trichomes on the filaments and glandular connective appendages. The only apparent difference between them is that the trichomes on the filaments from the type of M. juruensis are less dense than in the type of M. japuraensis. As for the sectional placement, Cogniaux (1909) correctly placed this species in Section Tamonea.

Miconia juruensis has been collected in the western Amazon basin, in the eastern portions of Colombia, Peru and Bolivia (not in Ecuador), and in Brazil, in the states of Amazonas and Acre. The collections from Peru, Bolivia and Acre sometimes lack the sessile glands on the abaxial leaf surfaces, as well as the trichomes on the filaments, but this does not seem to be sufficient to assign formal infraspecific rank.

Additional specimens examined:—COLOMBIA. Caquetá: Solano, 7 March 1945, E.L. Little 9649 (NY, US). PERU. Loreto: Lower Rio Huallaga, 1929, L. Williams 4230 (US); Maynas, 24 June 1972, S. McDaniel 16319 (US); Maynas, 24 April 1973, M. Rimachi 243 (US); Maynas, 1 August 1975, M. Rimachi 1828 (NY, US); Maynas, 22 May 1978, A. Gentry 22145 (US); Maynas, 30 December 1982, S. McDaniel 26573 (US); Requena, 24 August 1982, F. Encarnacion 2649 (US); Rio Nanay, 1929, L. Williams 481 (US). Ucayali: Purús, 6 February 2000, J.G. Graham 709 (UPCB); Purús, 25 February 2000, J.G. Graham 1134 (UPCB). BOLIVIA. Beni: Ballivián, 8 November 1985, J.C. Solomon 14627 (US). Pando: Manuripí, 19 September 1981, J.C. Solomon 6329 (NY, US); Manuripí, 8 August 1982, C.R. Sperling 6537 (NY, US). Santa Cruz: Ichilo, 31 October 1990, M. Nee 39648 (NY, US). BRAZIL. Acre: 11 August 1933, B.A. Krukoff 5396 (MO, NY, RB, US); Jordão, 5 February 2009, F.A. Michelangeli 1295 (RB, UPCB); Jordão, 6 February 2009, R. Goldenberg 1296 (RB, UPCB); Mâncio Lima, 11 November 2007, R. Goldenberg 942 (NY, RB, UPCB). Amazonas: Humaitá, September-October 1934, B.A. Krukoff 6093 (MO, US); Humaitá, September-October 1934, B.A. Krukoff 6248 (US); Rio Javari, Miraflor, 30 July 1973, E. Lleras P16932 (INPA, MO, NY, US).

6. Miconia longicuspis Cogniaux (1891: 851). Acinodendron longicuspe (Cogn.) Kuntze (1891: 951). Type:—BRAZIL. Rio de Janeiro: “Alto Macahe”, 21 April 1888, A. Glaziou 16918 (Lectotype: BR-5213115!, here designated; Isolectotypes: BR-5213177!, F-0063794, K!, MO-313816!, NY-00038512!, P!, R-09557!, Z-015130). Syntype:—BRAZIL. Rio de Janeiro: “Alto Macahe”, 1884, R. Mendonça 460 (Syntype: US!).

Miconia longicuspis is very similar to M. paniculata (see below), but differs from it by its persistent subulate bracteoles (which can be seen even in young fruits), campanulate hypanthium and triangular calyx lobes with small external teeth. It occurs in Eastern Brazil from Espírito Santo and Minas Gerais to Paraná. The collections from São Paulo and Paraná listed below have been previously identified as M. paniculata(Goldenberg 2009, Wanderley et al. 2011, Goldenberg 2012, Meyer et al. 2012).

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The syntype mentioned by Cogniaux (1891; Mendonça, from Minas Gerais) has no indication of province, but it is actually from “Alto Macahe”, which is in the state of Rio de Janeiro.

Additional specimens examined:—BRAZIL. Espírito Santo: Itaguaçu, 20 May 1946, A.C. Brade 18313(US); Santa Teresa, 19 July 1989, W. Boone 1333 (US), 13 July 2007, R. Goldenberg 892 (NY, UPCB). Minas Gerais: Delfim Moreira, 21 April 1939, M. Kuhlmann s.n. (US); Ouro Preto, 17 May 2011, A.C. Tonaco 178(UPCB); Santa Bárbara, April-June 1885, E.A. Vainio 33477 (US); Santa Bárbara, 28 May 1983, J.R. Pirani 723 (US). Paraná: Tunas do Paraná, 21 April 2007, E. Camargo & R. Goldenberg 101 (UPCB). Rio de Janeiro: Itatiaia, 25 March 1942, A.C. Brade 17283 (US); Itatiaia, 9 March 1982, S.S Renner 737 (US); Nova Friburgo, 10 July 1990, H.C. Lima s.n. (RB 290298, US). São Paulo: Paraibuna, 26 March 1984, G.J. Shepherd 15830 (UEC).

7. Miconia nervosa (Sm.) Triana (1871: 111). Melastoma nervosum Smith (1812: 31). Acinodendronnervosum (Sm.) Kuntze (1891: 245). Type (fide Wurdack et al. 1993):—“JAMAICA”, coll. unknown(Holotype: LINN-HS782-22).

Synonym:—Miconia pseudonervosa Cogniaux (1887: 337). Acinodendron pseudonervosum (Cogn.) Kuntze (1891: 952). 1891. Type:—BRAZIL. Minas Gerais: near Engenho do Mello Franco, J.E. Pohl 2889 (Lectotype: BR-5211982!, fide Goldenberg et al. 2010 ; Isolectotypes: LE, NY-00229333!, W).

Cogniaux (1887) distinguished M. pseudonervosa from M. nervosa based on petiole length and leaf bases and apices. He described the leaves of M. nervosa as subsessile, but this is a condition seldom found in its populations (see Wurdack 1973, 1980, Wurdack et al. 1993), even in the ones listed in his treatment. Some collections listed as M. pseudonervosa actually have short-petiolate leaves along with long-petiolate ones on the same plant (Riedel 2700 in P; Pohl 2899, dupl. in NY). As for the leaf bases and apices, there is a whole gradient from long-attenuate to acute-cuneate (M. nervosa) to slightly attenuate to obtuse bases (M. pseudonervosa), and long-acuminate (M. nervosa) to short-acuminate apices (M. pseudonervosa). This gradient agrees with a north to south axis, i.e. the plants collected in Minas Gerais and São Paulo (see Goldenberg 2009) actually have longer petioles and leaves with broader bases, as described for M. pseudonervosa. Nevertheless, the populations collected near its type locality (Goiás) and neighboring states, like Mato Grosso and the Federal District show a lot of variation regarding these characters. As already discussed by Goldenberg et al. (2010), much of this variation, as well as leaf size and hair density, may be explained by the fact that some of the plants collected at southern localities grow under drier conditions, in seasonal forests encroached by cerrado, than the plants collected in the Amazon basin and further north. Despite this morphological pattern associated with a geographical distribution, we see no reason to keep these populations as distinct entities. Miconia nervosa, as defined here, occurs from Belize and Guatemala to Bolivia and Southeastern Brazil.

For the complete list of synonyms under M. nervosa see Goldenberg et al. (in prep.); for information on types and synonymyzations under M. pseudonervosa see Goldenberg et al. (2010) and Melastomataceae.net (2007-2013).

Additional material examined:—BELIZE. Stann Creek, 24 January 1957, P.H. Gentle 9291 (US). BOLIVIA. Santa Cruz: Velasco, 29 July 2000, D.C. Wasshausen 2238 (US). BRAZIL. Acre: Mâncio Lima, 13 November 2007, R. Goldenberg 970 (RB, NY). Amazonas: Maraã, 20 October 1982, C.A. Cid 3406 (US). Amapá: Macapá, 15 March 1962, J. Mattos 10042 (US). Bahia: Camacan, 9 April 2006, R. Goldenberg 848(CEPEC, UPCB). Distrito Federal: 20 July 1966, H.S. Irwin 18297 (US); 24 August 1981, J.F. Kirkbride 4347 (MBM, NY, US). Espírito Santo: Linhares, 31 October 1983, G. Martinelli 9759 (US). Goiás: “Aldea Caretas do Pedro Terceiro”, J.B.E. Pohl 1178 (K); J.B.E. Pohl 1705 (syntype of M. pseudonervosa, NY); Goiás Velho, 9 May 1973, W.R. Anderson 9946 (US); Luziânia, 5 September 1978, E.P. Heringer 17332 (NY); Pirenópolis, 9 June 1995, F.C.A. Oliveira 388 (UEC). Maranhão: Curutapera, 17 September 1985, W.L. Baleé 1035 (US).

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Minas Gerais: Uberlândia, 10 August 1994, R. Romero 1099 (US); “Engenho do Mello Franco”, J.B.E. Pohl 1197 (syntype of M. pseudonervosa, BR, K, NY); no precise locality, J.B.E. Pohl s.n. (BM). Mato Grosso: Aripuanã, 8 October 1973, G.T. Prance 18243 (US); “Expedition Base Camp”, 16 September 1968, R.M. Harley 10059 (NY, US); “Reserva do Cabaçal”, 9 May 1995, G. Hatschbach 62557 (MBM). Mato Grosso do Sul: Rio Verde de Mato Grosso, 28 August 1973, G. Hatschbach 32455 (MBM, US); Sonora, 17 August 2002, V. Pott 5670 (UPCB). Pará: Almeirim, 19 December 1986, M.J. Pires 1540 (US). Pernambuco: Caruaru, 4 April 1995, M. Tscha 51 (US). Rondônia: Porto Velho, 4 October 1979, G. Vieira 232 (US). Roraima: Serra do Tepequém, 20.Feb.1967, G.T. Prance 4572 (US). São Paulo: Cajuru, March 1857, A.F. Regnell III.23 (syntype of M. pseudonervosa, BR-fragm., P, S); Campinas, A.E. Severin 160 (syntype of M. pseudonervosa, S); Franca, June 1834, L. Riedel 2700 (syntype of M. pseudonervosa, P-2 sheets); Itirapina, November 1992, R. Goldenberg 43 (UEC, US). Tocantins: Tocantinópolis, 27 March 1976, G. Hatschbach 38402 (US). COLOMBIA. Antioquia: San Carlos, 30 November 1988, G. McPherson 13309 (US). COSTA RICA. Alajuela: 8 July 1976, J. Utley 5355 (US). ECUADOR. Imbabura: 25 July 1992, A. Freire-Fierro 2350 (US). FRENCH GUIANA. Saul: 13 December 2007, O. Tostain 1184 (US). GUATEMALA. Izabal: 20 March 1972. E. Contreras 11430 (US). GUYANA. Potaro-Siparuni: 23 November 1995, H.D. Clarke 600 (US). HONDURAS. Atlántida: 26 June 1970, G. Davidse 2193 (US). PANAMA. Isla de Barro Colorado, 5 January 2007, R. Perez 1691 (US). PERU. Loreto: Maynas, 18 September 1990, J.J. Pipoly 12111 (US). SURINAM. Brokopondo: 27 November 2003, T.R. van Andel 4599 (US). TRINIDAD. Arima Valley, 24 March 1959, R.S. Cowan 1209 (US). VENEZUELA. Bolívar: Foraneo Aripao, 1 May 1988, G. Aymard 6718 (US).

8. Miconia paniculata (DC.) Naudin (1950: 245). Cremanium paniculatum De Candolle (1828: 194). Acinodendron paniculatum (DC.) Kuntze (1891: 952). 1891. Type:—BRAZIL. “Provinciae Minas Geraes, prope Villam Ricam”, C.F.P. von Martius s.n. (Holotype: M-0165633!).

Synonym:—Miconia depauperata Gardner (1843: 346). Type:—BRAZIL. “Organ Mts.” (state of Rio de Janeiro), January 1838, G. Gardner 394 (Holotype: K-000536150!; Isotypes: BM-000799245!, FI-004737, K-00053619!, NY00229125!, P!, US-00120950!).

Synonym:—Miconia doriana Cogniaux (1887: 376). Type:—BRAZIL. “Serra de Friburgo, Terras do Im. Davide Bacelli” (state of Rio de Janeiro), 3-7. Feb.1883, J. de Saldanha 7072 (Holotype: B, destroyed; Lectotype: BR-5210985!, here designated; Isolectotypes: P!, R!).

As currently circumscribed, M. paniculata can be recognized by the leaves with mite domatia (nerves basally joined by a membrane on the abaxial surface), very early caducous bracts and bracteoles, and flowers with obconical hypanthium and long, subulate external calyx teeth. These plants can have 5-merous, diplostemonous flowers, but most specimens have 6-merous pleiostemonous flowers with up to 22 stamens. This species occurs from Bahia and Minas Gerais to Santa Catarina.

As for the species synonymized here, M. doriana was described by Cogniaux (1887) based on its pleiostemonous flowers (with 15 to 20 stamens), acute calyx lobes, these half of the hypanthium length, and stamens with unappendaged connectives. In the same key where he distinguished M. doriana from the other species in the M. paniculata complex, Cogniaux contradicted himself by separating M. depauperata and M. paniculata (which should have obtuse calyx lobes) from M. tristis, due to their triangular to subulate calyx lobes (that should be supposedly found only in M. doriana). The types of both M. paniculata and M. depauperata actually have distinct subulate calyx teeth that do not differ from the ones found in M. doriana, despite some variation in length. As for the connective appendages, Cogniaux did not use this character in his following treatment (Cogniaux 1891), apparently because the stamens in the type of M. doriana actually have small dorsal appendages.

The remaining issues are the flower merosity and the number of stamens. The type of M. depauperataclearly does not agree with Cogniaux’s description: among its several duplicates, one can find buds with five or six calyx lobes, and stamens varying from 17 (US) to 22 per flower (K). The type of M. paniculata has

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strictly 5-merous buds, but none of them were dissected in order to count the stamens. Nevertheless, there are just a few recent collections of plants with 5-merous, diplostemonous flowers (Santos 176, Pinheiro 1334, both from Bahia) that do not differ at all, in other aspects, from the plants with pleiostemonous flowers.

The collections reported as this species by Goldenberg (2009) and Wanderley et al. (2011) for São Paulo and by Meyer et al. (2012) and Goldenberg (2012) for Paraná actually belong to M. longicuspis Cogn. (see above).

Additional specimens examined (apart from historical collections, only flowering material cited):—BRAZIL. Bahia: Arataca, 14 April 2006, R. Goldenberg 865 (UPCB); Castro Alves, 27 May 1987, L.P. Queiroz 1568 (US); Una, 14 June 1971, T.S. Pinheiro 1334 (US); Una, 8 December 1987, E.B. dos Santos 176(US). Espírito Santo: Santa Maria do Jetibá, 5 January 2008, R. Goldenberg 1028 (NY, UPCB); Santa Teresa, 15 January 1986, H.Q.B. Fernandes 1777 (US). Minas Gerais: Viçosa, 7 April 1930, Y. Mexia 4576 (NY, US). Paraná: Bocaiúva do Sul, 23 January 1963, G. Hatschbach 9699 (US); Bocaiúva do Sul, 20 December 1972, G. Hatschbach 31023 (US); Campina Grande do Sul, 16 December 1962, G. Hatschbach 9558 (US); Piraquara, February 2004, R. Goldenberg 616 (UPCB), Piraquara, 1 March 2008, R. Goldenberg 1157(UPCB); São José dos Pinhais, 27 December 1960, G. Hatschbach 7740 (US). Rio de Janeiro: S.loc., s.d., A. Glaziou 13854 (US); Itatiaia, 11 November 1945, A.C. Brade 17432 (US); Itatiaia, 30 January 1950, A.C. Brade, 20229 (US); Petrópolis, 29 November 1959, G. Pabst 5208 (NY, US); Teresópolis, January 1890, J.T. de Moura 374 (BR; cited in Flora Brasiliensis). Santa Catarina: Blumenau, 15 December 1959, R. Klein 2339(NY, US); Ibirama, 26 January 1957, R. Klein 2206 (US); Palhoça, 18 January 1956, R. Reitz 2440 (US); Vidal Ramos, 29 January 1958, R. Reitz 6348 (US). São Paulo: São Paulo, 4 January 1921, F.C. Hoehne 5986 (NY). No precise locality: F. Sellow s.n. (US, 2 sheets), F. Sellow 109 (US).

9. Miconia petropolitana Cogniaux (1888: 613). Acinodendron petropolitanum (Cogn.) Kuntze (1891: 952). Type:—BRAZIL. Rio de Janeiro: Petrópolis, “Itamaraty”, 1886, A. Glaziou 16021 (Lectotype BR-5213450!, fide Slanis & Goldenberg 2011; Isolectotypes: K-000536148!, P!).

Synonym:—Miconia inconspicua var. glabrata Wurdack (1962: 166). Type:—BRAZIL. Santa Catarina: Campo Alegre, “Morro do Iquererim”, 9 May 1957, R. Reitz & R. Klein 4808 (Holotype: US-00121017!; Isotypes: GH-00072821, HBR, K-000536162!, NY-00229189!, S-05-3597, UC-1243702).

Miconia inconspicua var. glabrata was originally compared only with the typical variety of M. inconspicuaMiquel (1849: 542), from which it differs mostly by its sparser and smaller trichomes, but also smaller flowers and incomplete thecal septum (Wurdack 1962). In the same treatment for the Flora of Santa Catarina where this variety was described, there is a description of M. petropolitana, but no comparison was made with it, neither in the protologue of the variety, nor in the comments under this species. Both were kept apart in the key due to their leaf margins (entire or indistinctly undulate in M. petropolitana vs. undulate-serrate in the variety), but this character difference is not consistent outside of Santa Catarina: the type of M. petropolitana , from Rio de Janeiro, has distincty dentate leaf margins. Nevertheless, the diagnostic features mentioned above for M. inconspicua var. glabrata (size and density of trichomes; thecal septum completion) match perfectly with the ones found in M. petropolitana. Although the specimens listed under M. inconspicua var. glabratausually have smaller leaves and also obtuse leaf bases that are broader than in M. petropolitana, they seem to represent only a local variation. Since intermediate forms are found in Paraná and São Paulo, we see no reason to recognize infraspecific status for these populations.

For illustrations and lectotypification of M. petropolitana, see Slanis & Goldenberg (2011).Additional specimens examined: ARGENTINA. Misiones: Depto. San Pedro, 23 September 1978, P.R.

Legname 5556 (LIL). BRAZIL. Espírito Santo: Arituba, 21 August 1982, R. Ribeiro 286 (US). Minas Gerais: Cristina, 22 October 1989, R. Mello-Silva 95 (US); Viçosa, 2 October 1930, Y. Mexia 5136 (US). Paraná: Campina Grande do Sul, 28 August 1948, G. Hatschbach 984 (US); Campo Largo, 22 September 1976, L.T.

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Dombrowski 6365 (US); Piraquara, 23 September 1992, S. Ziller 234 (US); Tijucas do Sul, 17 August 1958, G. Hatschbach 4989 (US). Rio de Janeiro: Itatiaia, 10 October 1977, P.J.M. Maas 3154 (US). Rio Grande do Sul: Caxias do Sul, 3 February 2003, A. Kegler 1450 (US); Farroupilha, 21 October 1984, R. Wasum 470(US). Santa Catarina: Campo Alegre, R. Reitz 4799 (US); Lauro Müller, 22 August 1958, R. Reitz 7032 (US), 23 August 1958, R. Reitz 7051 (US); Mafra, 7 September 1957, R. Reitz 4855 (US); Paulo Lopes, 17 December 1973, A. Bresolin 876 (US); Porto União, 16 September 1962, R. Klein 3068 (US). São Paulo: Jundiaí, August 1995, R. Goldenberg 94 (US). No precise locality: s.d., Sello s.n. (US).

Aknowledgements

The authors thank Fernanda A. Carvalho for the pictures of the specimens from M, the staff at HBG for the type image of M. juruensis, and Fabian Michelangeli and an anonymous reviewer for the comments on the text. R.G. received funding from CNPq (Proc. 302325/2009-3), NSF (DEB-0818399) and CAPES (BEX 9095/11-0), and M.K.C. a grant from CNPq. We also thank Vicki Funk and US Herbarium staff for the support during R.G.’s sabbatical at this institution.

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