Nomenclatural and taxonomic changes in Paepalanthus (Eriocaulaceae) from São Paulo and Minas Gerais, Brazil

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ISSN 0075-5974, Volume 65, Number 2

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Nomenclatural and taxonomic changes in Paepalanthus (Eriocaulaceae) from São Paulo and Minas Gerais, Brazil Marcelo Trovó1 & Paulo Takeo Sano1 Summary. The largest genus of the Eriocaulaceae, Paepalanthus, presents many taxonomic problems. Some of these were identified during studies of Eriocaulaceae from the flora of São Paulo State and Caparaó National Park. Here, we propose changes in nomenclature as a solution to such issues, based on type collections, recent collections and field observations. These changes are in agreement with the taxonomic species concept, and the rules established by the International Code of Botanical Nomenclature. We define six lectotypes: P. gneissicola, P. caparoensis, P. caldensis, P. lundii, P. oerstedianus and P. striatus, and six synonyms: P. gneissicola = P. acantholimon, P. loefgrenianus = P. aequalis, P. multicostatus = P. calvus, P. scopulifer = P. caparoensis, P. neocaldensis = P. flaccidus and P. macrotrichus = P. lundii. We also present comments on morphology, protologue and type collections. Key Words. Caparaó National Park, lectotypes, nomenclature, São Paulo State, synonyms, taxonomy.

Introduction Paepalanthus Mart. nom. cons. is the largest genus of Eriocaulaceae, with c. 550 species, mainly distributed in the Neotropics on rocky outcrops of the Espinhaço Range, in the Brazilian savanna, and in the Tepuis of Venezuela. Several species are endemic, usually being restricted to one locality (Giulietti & Hensold 1990; Stützel 1998). Paepalanthus presents the greatest morphological variation within the family, thereby resulting in several species and supra-specific category descriptions (Bongard 1831; Koernicke 1863; Ruhland 1903; Silveira 1908, 1928). Bongard (1831) described 83 species, mostly from Brazil, mainly based on the Riedel collection deposited in LE. In the taxonomic treatise on Eriocaulaceae for Flora Brasiliensis, and essentially from material deposited in B, Koernicke (1863) described many other Eriocaulaceae species. At the beginning of the 20th century, two authors independently described numerous species. Ruhland (1903), also working in the Berlin herbarium, provided the ultimate revision of the whole family, and Silveira (1908, 1928), a Brazilian geologist, described dozens of new species based on his own collections deposited in R. Bongard, Koernicke, and Ruhland had never undertaken fieldwork in Brazil, and relied on a limited number of specimens to study. Hence, they were unaware of the large infra-specific variability in Eriocaulaceae. Although Silveira had experience in observing plants in their natural environment, he was an amateur botanist, his botanical concepts usually not being in accordance with those of European botanists.

Consequently, they all tended to describe many taxonomic synonyms or taxa with problematic typification. As these studies have been the basis for work on Paepalanthus, the elucidation of such taxonomic inconsistencies is one of the main goals for improving comprehension of the genus as a whole. Current approaches in the study of Paepalanthus are focused on taxonomic revisions. Hensold (1988), TissotSqualli (1997) and Sano (2004) revised some of the infra-generic categories. Nevertheless, there are no recent studies available on a number of important groups. One reason for this is that morphological plasticity within these groups as a whole is as yet to be fully explored. Therefore, floristic studies are important for any taxonomic progress in the group, since they pave the way towards detecting and solving differences as regards both taxonomy itself and nomenclature. Giulietti & Parra (1994) and Parra & Giulietti (1997), for instance, proposed important changes in nomenclature for solving issues regarding the Flora of the Pico das Almas (Giulietti & Parra 1995) and Flora of the Serra do Cipó (Parra 1998). In a similar effort, we herein propose changes in nomenclature to prepare monographs on Paepalanthus for the Flora of São Paulo State as well as that of the Caparaó National Park (Trovó et al. 2007).

Material and methods The nomenclatural changes proposed here were based on intensive fieldwork, type collection analysis and recent material deposited in the following herbaria (acronyms according to Holmgren et al. 1990): B, BM,

Accepted for publication June 2010. 1 Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo, CEP 05508-900, São Paulo, Brazil. e-mail: [email protected]; [email protected]

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BR, BHCB, C, CEPEC, CESJ, CVRD, ESA, ESAL, F, G, GFJP, HB, HBG, HBR, HRB, HRCB, HUEFS, HUFU, IAC, K, L, LE, LL, M, MBM, NY, OUPR, P, R, RB, S, SP, SPF, U, UEC, UPS, US, W, WU. All cited specimens were observed and analysed by the authors. In the present work we adopted the taxonomic species concept proposed by Stuessy (1990). All changes in nomenclature are in accordance with the rules established by the International Code of Botanical Nomenclature (McNeill et al. 2006).

Nomenclatural and taxonomic changes

Paepalanthus acantholimon Ruhland (1903: 163). Type: Brazil, Minas Gerais, Serra de Caparaó, Feb. 1890, W. Schwacke 6711 (holotype B; isotype P). Paepalanthus gneissicola Silveira (1928: 124, Tab. 77). Type: Brazil, Minas Gerais, in Serra do Caparaó, Minas Geraes, altitudine 2700 m, Sept. 1912, A. Silveira 597 in Herbarium Silveira (lectotype R, selected here), synon. nov. NOTES. Paepalanthus acantholimon was described by Ruhland (1903), based on a single specimen collected in the Caparaó Range (W. Schwacke 6711). The diagnostic features of P. acantholimon are a small rosette with a rhizome and, generally, the presence of one scape. This species grows on rocks, forming small cushions. Later, Silveira (1928) described P. gneissicola, which was collected in the same range (Silveira 597). However, the characteristics outlined by the latter (Silveira 1908) did not serve to distinguish the two species, thereby giving rise to the proposition that they are synonymous. There are two labels on the material mentioned in the Paepalanthus gneissicola protologue. One is Silveira’s and the other refers to another collection from the same locality (O. A. D. n. 5, 10/VII/41). As the material consists of un-mounted plants, we were unable to state which is the type collection. We selected a single plant to be mounted on a separate sheet and designated this as the lectotype.

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(1829) distinguished the species by the presence of linear leaves with the same-sized scapes and sheaths with a truncated apex. The same features were used by Ruhland (1903) to describe P. loefgrenianus, which was collected in São Paulo State (Loefgren & Edwall 2227). Consequently, a new synonym is proposed.

Paepalanthus calvus Koern. (Koernicke 1863: 391). Type: Brazil, Minas Gerais, Minas Geraes, alt 7000 ped., G. Langsdorff s.n. (holotype B; isotype LE). Paepalanthus multicostatus Ruhland (1903: 148, Tab. 17). Type: Brazil, Minas Gerais, Serra da Piedade, June 1893, A. Glaziou 20532 (holotype B; isotypes C, K, P), synon. nov. NOTES. Koernicke (1863) described Paepalanthus calvus collected in Minas Gerais (Langsdorff s.n.). The species bears glabrous leaves, large capitula and dark-brown involucral bracts. Ruhland (1903) described P. multicostatus collected on the Piedade Range in Minas Gerais (Glaziou 20532). In the protologue, Ruhland (1903) pointed out the similarity between these two species. According to recent collections and careful analysis, both species are the same and must be considered as synonymous.

Paepalanthus caparoensis Ruhland (1903: 145). Type: Brazil, Minas Gerais, Serra de Caparaó, in rup. humidis frequens, 9 Feb. 1890, W. Schwacke 6712 (lectotype B, selected here). Paepalanthus scopulifer Silveira (1928: 98). Type: Brazil, Minas Gerais, In locis humidis humidosisque in Serra do Caparaó, Minas Geraes, altitudine 2700 m, April 1913, A. Silveira 611 (holotype R), synon. nov.

Paepalanthus aequalis (Vell.) J. F. Macbr. (Macbride 1931: 43.). Type: Original drawings, Tabula 85. Dupatya aequalis Vell. (Vellozo 1829: 36, tab 85). Paepalanthus loefgrenianus Ruhland (1903: 142). Type: Brazil, São Paulo, São Paulo, campo, S. J. da Boa Vista, June 1893, A. Löfgren & Edwall 2227 (holotype C; isotypes B, SP), synon. nov.

NOTES. Paepalanthus caparoensis was described by Ruhland (1903), based on two collections deposited in B (Schwacke 6712 and Moura 944), but without designation of the holotype. Schwacke’s collection bears the original description with Ruhland’s drawings and comments on the label. Therefore, this syntype is here designated as the lectotype. Paepalanthus caparoensis is characterised by the presence of glabrous linear leaves with a recurvate, acute apex. Silveira (1928) described P. scopulifer as presenting the same diagnostic characteristics, although with a smaller habit. After analysing several collections, we reached the conclusion that this variation is due to phenotypic plasticity. Therefore, the species described by Silveira (1928) is here considered as synonymous.

NOTES. Vellozo (1829) described Dupatya aequalis, which was collected near Rio de Janeiro. Macbride (1931) combined the species in Paepalanthus aequalis (Vell.) J. F. Macbr. In his original description, Vellozo

Paepalanthus flaccidus (Bong.) Kunth (1841: 511). Type: Brazil, São Paulo, “Brasilia”, L. Riedel 1034 (holotype LE; isotypes B, LE, P, U, UPS).

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NOMENCLATURAL AND TAXONOMIC CHANGES IN PAEPALANTHUS (ERIOCAULACEAE)

Eriocaulon flaccidum Bong. (Bongard 1831: 636, 646, tab. IV). Paepalanthus caldensis Silveira (1928: 186, tab 120), nom. illegit. non Malme (1901). Paepalanthus neocaldensis Moldenke (1946: 140). Type: Brazil, Minas Gerais, in humidis prope Poços de Caldas, April 1922, A. Silveira 727 (lectotype R, selected here), synon. nov. NOTES. The species Eriocaulon flaccidum Bong., previously described by Bongard (1831), was later combined in P. flaccidus by Kunth (1841). Originally, Bongard (1831) described a decumbent plant with a thin ramified stem and dimerous flowers, and collected in São Paulo. Silveira (1928) described Paepalanthus caldensis. However, this name was found to be a later homonym of a species described by Malme (1901), whereby Moldenke (1946) created a new one. The species described by Silveira (1928) is identical to P. flaccidus. In accordance with current circumscription, P. flaccidus is considered as a species with wide geographic distribution and high morphological variation, this including the whole range presented by P. neocaldensis. Consequently, we propose that P. neocaldensis is synonymous with P. flaccidus. Silveira (1908) referred to two batches of material to describe Paepalanthus caldensis, one collected in Poços de Caldas, Minas Gerais (Silveira 727) and the other in São Paulo (Toledo 12 in the herbarium “Com. Geogr. e Geol. de São Paulo”). Nevertheless, there is only one collection deposited in R, this bearing the original Silveira label. On the same label and below Silveira’s collector number, there is also Toledo’s collector number. Nevertheless, the specimen clearly came from a single collection. Since there is no indication of the original label of the Herbarium of the Comissão Geographica e Geológica de São Paulo, it is assumed that this material belongs to Silveira’s collection, therefore we designated it as the lectotype.

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notes, with duplicates in several herbaria. Consequently, we selected this as the lectotype. The species is characterised by the presence of hirsute linear leaves, long basal trichomes, and linear floral bracts. Silveira (1908) described Paepalanthus macrotrichus (A. Loefgren 1485), collected in São Paulo. As this presents the same diagnostic features and distribution as P. lundii, we consider them synonymous.

Paepalanthus oerstedianus Koern. (Koernicke 1863: 374). Type: Brazil, Minas Gerais, in humidis Serra de Cubatão, Oct. 1833, L. Riedel 1777 (lectotype B, selected here; isolectotypes G, K, L, M, P, UPS). NOTES. Koernicke (1863) described Paepalanthus oerstedianus based on two collections from Cubatão (Riedel 1777, Lund s.n.). The material collected by Lund was not found. The material collected by Riedel, and deposited in B, is accompanied by Koernicke’s original notes with duplicates in several herbaria. Thus, we designated this as the lectotype.

Paepalanthus striatus Ruhland (1903: 149). Type: Brazil, São Paulo, Campos da Bocaina, dans le bois près des sources du Rio Parahybo, 10 Feb. 1876, A. Glaziou 7994 (lectotype B, selected here; isolectotypes G, K, P). NOTES. Paepalanthus striatus was described by Ruhland (1903), based on two collections from the Bocaina Range and deposited in B (Glaziou 7994, Schwacke 1912). Schwacke’s collection consists of leaf-fragments and reproductive structures. The material collected by Glaziou is that of a complete specimen together with Ruhland’s illustrations. Therefore, this was chosen as the lectotype.

Paepalanthus lundii Koern. (Koernicke 1863: 385). Type: Brazil, São Paulo, “In humidis Araracoara et Vila Franca”, L. Riedel 2204 (lectotype B, selected here; isolectotypes G, K, L, LE, P, S, U, UPS). Paepalanthus macrotrichus Silveira (1908: 43). Type: Brazil, São Paulo, in campis prope Corrego Fundo, Nov. 1889, A. Loefgren 1485 (holotype R; isotypes P, SP), synon. nov.

Acknowledgements We are indebted to the curators of all the herbaria we visited, who obligingly permitted access to historical collections of Eriocaulaceae, and in particular to those of herbaria B, LE and R. We are also indebted to Leandro Assis, Maria Ana Farinaccio, and Thais Lavagnolli for critically reading the original manuscript. This work was financially supported by Capes, DAAD and IAPT (Marcelo Trovó — PhD studentship), and by CNPq (Paulo Takeo Sano — protocol n°. 309376/2006 – 8).

NOTES. Paepalanthus lundii was described by Koernicke (1863), based on two collections from São Paulo (Riedel 2204, Lund s.n.). The material collected by Lund was unavailable, whereas that by Riedel had been deposited in B together with Koernicke’s original

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