Museu de Biologia Prof. Mello Leitão (MBML). Av. José Ruschi, 4, Santa Teresa, Espírito Santo-ES, Brazil.

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1 Three new species of Pabstiella (Pleurothallidinae, Orchidaceae) from Brazilian Atlantic Forest Claudio Nicoletti de Fraga 1 and Ludovic Jean Charles Kollmann 2 Abstract. Three new species of Pabstiella from Brazilian Atlantic Forest in the State of Espírito Santo are described and illustrated. Comments on their ecology and distribution are also provided. The new species, Pabstiella lueriana, P. villosisepala, and P. muricatifolia, are most similar to P. pellifeloides and P. hypnicola. According to the criteria of the IUCN Red List of endangered plant species, P. muricatifolia is included in the Data Deficient category, while the other two species were included in the Vulnerable category. Resumo. São descritas e ilustradas três novas espécies de Pabstiella da Floresta Atlântica brasileira no Espírito Santo. As novas espécies Pabstiella lueriana, P. villosisepala e P. muricatifolia, são comparadas com P. pellifeloides e P. hypnicola. São ainda fornecidas informações sobre ecologia e distribuição geográfica das espécies. De acordo com os critérios da IUCN, P. muricatifolia foi incluída na categoria Dados Deficientes e as demais na categoria Vulnerável. Keywords: Pabstiella, Pleurothallidinae, Orchidaceae, Atlantic Forest, Brazil Palavras-chave: Pabstiella, Pleurothallidinae, Orchidaceae, Floresta Atlântica, Brasil The genus Pabstiella Brieger & Senghas was segregated from Pleurothallis R. Br. by Brieger and Senghas (1976) to accommodate a single species, Pabstiella mirabilis (Schltr.) Brieger & Senghas (originally described in Pleurothallis). Based on molecular evidence, Pridgeon and Chase (2001) proposed the genus Anthereon Pridgeon & M. W. Chase to accommodate six species also previously residing in Pleurothallis, including Pleurothallis mirabilis, the type of the genus Pabstiella, therefore rendering their genus illegitimate. Barros (2002) re-established Pabstiella and transferred all species treated in Anthereon to Pabstiella. Later, Luer (2006) removed two repent taxa from Central America and northern South America, Pabstiella aryter (Luer) F. Barros and Pabstiella determannii (Luer) F. Barros, and established the genus Ronaldella Luer. He also expanded the circumscription of Pabstiella by adding one more name, Pabstiella parvifolia (Lindl.) Luer. More recently, Luer (2007) expanded the circumscription of Pabstiella and proposed 67 combinations based on Brazilian species of Pleurothallis. As understood today, Pabstiella consists of about 80 species (several unpublished distributed mainly in Southern Brazil). A few species have a wider distribution, occurring in Bolivia, Amazonian Brazil, Colombia, Ecuador, Peru and Venezuela, with one species reaching Panama and Costa Rica (Luer, 2006). Pabstiella is most similar Anathallis Barb. Rodr., but it is mainly distinguished by its more or less fleshy sepals with keels, with the laterals connate, and by the usually thick lip with a pair of calli, without membraneous, basal lobules (Luer, 2009). After conducting fieldwork in the state of Espírito Santo, Brazil and identification of herbarium specimens housed at MBML (following Holmgren et al., 1990), the authors detected three new species of Pabstiella that are described and illustrated herein. We acknowledge Hélio de Queiroz Boudet Fernandes, Curator of MBML herbarium, Museu de Biologia Mello Leitão, Espírito Santo, for his assistance in the field and herbarium work, and Josiane Rossini for the line drawings. We greatly appreciate critical comments from Carlyle A. Luer, Antônio L.V. Toscano de Brito, and two anonymous reviewers, as well as Donna Tremonte s editorial support. 1 Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rua Pacheco Leão, 915, , Jardim Botânico, Rio de Janeiro, Brazil. cnfraga@jbrj.gov.br. Author for correspondence. 2 Museu de Biologia Prof. Mello Leitão (MBML). Av. José Ruschi, 4, Santa Teresa, Espírito Santo-ES, Brazil. ludovic@limainfo.com.br Harvard Papers in Botany, Vol. 15, No. 1, 2010, pp President and Fellows of Harvard College, 2010.

2 172 Harvard Papers in Botany Vol. 15, No. 1 Pabstiella lueriana Fraga & L.Kollmann, sp. nov. TYPE: Brazil. Espírito Santo, Santa Teresa, Nova Lombardia, Augusto Ruschi Biological Reserve, 800 m, 16 October 2001, L. Kollmann & E. Bausen 4847 (fl.) (Holotype: MBML [15561]; Isotype: RB). Fig. 1. Figure 1. Pabstiella lueriana Fraga & L. Kollmann. A, habit; B, flower in side view; C, dorsal sepals; D, lateral sepals; E, petal; F, lip in side view; G, lip; H, column and ovary in side view; I, anther, from below; J, anther, from above. Drawn from the holotype specimen.

3 2010 Fraga and Kollmann, New species of Pabstiella from Brazil 173 Haec species Pabstiella pellifeloide (Barb. Rodr.) Luer similis, sed sepalo dorsali maiore, apice petalorum mucronato differt. Plant epiphitic, up to (6 )8 18 cm tall, caespitose, erect. Roots whitish, fasciculate, terete, glabrous. Ramicauls (1 )4 6 cm long, green, terete, enclosed by 2 3 thin, tubular sheaths; sheaths cm long, green, paleaceous when mature. Leaf cm including a petiole ca. 1 cm long, green, erect; blade thickly chartaceous, lanceolate to narrowly elliptic-lanceolate, the base attenuate to narrowly cuneate below into the petiole, the apex acute, 3-toothed. Inflorescence cm long, a loose, slightly flexuous, several-flowered raceme, emerging from near the apex of the ramicaul; floral bracts 3 mm long, green, tubular, acuminate. Flowers not resupinate, opening in succession, occasionally 2-flowers remain open simultaneously. Pedicel cm long; ovary mm long, greenish, terete, longitudinally sulcate. Dorsal sepal mm, yellowish with orange apex, obovate, 3-veined, convex and velvety towards the obtuse to rounded apex, glabrous externally, the base concave, purplishlilac veined, attenuate and pubescent within, the margins somewhat revolute above the middle. Lateral sepals connate, forming a synsepal, mm, with subacute apices, yellowish with purple grooves along the veins, concave below the middle, externally glabrous with two longitudinal keels towards the apex, internally glabrous at base with purplishlilac spots, the apical half orange, velvety with revolute margins from the middle to the tips. Petal mm, rhomboid, asymmetrical, rough in both surfaces, externally thick and papillate, green and attenuate at base, yellowish on the margins, with purplish-lilac spots and veins towards the mucronate, black apex. Lip mm, ca. 4.3 mm long expanded, white with purplish-lilac spots, triangular, thick, 3-lobed, the base subtruncate with a pair of minute lobules on both corners, the disc shallowly channeled centrally between a pair of thick, oblong, glandular-celullar calli, the lateral lobes, short, rounded, the apical papillate, acute. Column mm long, white lilac flecked, semi-terete, winged above the middle, denticulate at apex, prolonged at base into a short (ca. 1 mm long), thick, lilac foot; anther ca. 1 mm long, hooded, yellowishwhite with slightly lilac apex and margins; pollinia 2, yellow, obovoid. Capsule unknown. Etymology: the new species honors Carlyle A. Luer, for his outstanding contributions to the knowledge of Pleurothallidinae. Habitat and distribution: so far this species has only been found growing as epiphytes in remnants of Atlantic Forest in the Augusto Ruschi Biological Reserve and Santa Lúcia Biological Station at m, but probably also occurs in adjacent areas in the State of Espírito Santo (Fig. 2). Conservation status: due to the apparent rarity and restricted distribution of Pabstiella lueriana in the Atlantic Forest in the central region of Espírito Santo State and its apparent vulnerability to human activities, we include this species in the Vulnerable category of the IUCN Red List of endangered plant species according to the following criteria (IUCN, 2001): VU D2. Paratypes: Brasil, Espírito Santo, Mun. Santa Teresa: Nova Lombardia, Augusto Ruschi Biological Reserve, 03 December 2002, (fl.), R. R. Vervloet and E. Bausen 1405 (MBML); Nova Lombardia, Augusto Ruschi Biological Reserve, 8 December 2000 (fl.), L. Kollmann 3454 (MBML, MO); Nova Lombardia, Augusto Ruschi Biological Reserve, 24 October 2002 (fl.), R. R. Vervloet, E. Bausen and W. Pizziolo 1289 (MBML); Nova Lombardia, Augusto Ruschi Biological Reserve, 03 October 2002 (fl.), R. R. Vervloet, E. Bausen and W. Pizziolo 1155 (MBML, SP); Nova Lombardia, Augusto Ruschi Biological Reserve, 24 October 2001 (fl.), L. Kollmann, E. Bausen and W. Pizziolo 4911 (MBML); Valsugana Velha, Santa Lúcia Biological Station, 10 October 2000 (fl.), L. Kollmann and R. R. Vervloet 3114 (MBML); Valsugana Velha, Santa Lúcia Biological Station, 25 November 1998, L. Kollmann, E. Bausen and W. Pizziolo 1111 (MBML); Valsugana Velha, Santa Lúcia Biological Station, 01 April 2004, (fl.), L. Kollmann and M. Sobral 6585 (MBML). This new species is most similar to Pabstiella pellifeloides (Barb. Rodr.) Luer, from which it may be distinguished by the size of the dorsal sepal and shape of the petals. The dorsal sepal in Pabstiella lueriana is mm long and the petals have an externally thick, papillate, mucronate apex, whereas in P. pellifeloides the dorsal sepal is 3 4 mm long and the petals are acute.

4 174 Harvard Papers in Botany Vol. 15, No. 1 Figure 2. Map showing the distribution of Pabstiella lueriana (black triangles), P. muricatifolia (black dots), and P. villosisepala (black squares) in Espírito Santo State, Brazil. Pabstiella muricatifolia Fraga & L.Kollmann, sp. nov. TYPE: BRAZIL. Espírito Santo: Águia Branca, São Pedro, CEIER roky, elev m, 19 01'22.2"S e 40 38'52.8"W, 26 April 2006, V. Demuner; L. F. S. Magnago; T. Cruz; E. Bausen 2246 (Holotype: MBML [26760]; Isotype: RB). Fig. 3. Species haec Pabstiella hypnicolae cui affinis, sed foliis obovatis ad oblongis coriaceis cum superficiebus muricatis, floribus resupinatis, sepalis apice papilloso, labelo trilobato, apice obtuso et papilloso differt. Plant epiphitic, up to cm tall, caespitose, erect. Roots whitish, faciculate, terete, glabrous, emerging near the base of the rhizome. Ramicauls cm long, green, cylindrical, enclosed by 2 3 thin tubular sheaths; sheaths cm long, green and paleaceous when mature, acute. Leaf cm including a petiole ca. 0.5 cm long; blade dark green, erect, thickly coriaceous, muricate on both surfaces, obovate to oblong, slightly sulcate on the abaxial surface, the base acute, the apex obtuse and apiculate, the margins lightly reflexed. Inflorescence 3 9 cm long, an erect to subflexuous, several-flowered raceme emerging from near the apex of the ramicaul; floral bracts ca. 2 mm long, purplish-lilac, tubular. Flowers resupinate, opening in succession, occasionally 2 flowers remain open simultaneously. Pedicels mm long, purplish-lilac; ovary mm long, purplish-lilac, terete, longitudinally sulcate. Dorsal sepal mm, purplish-lilac, white with purplish-lilac spots towards the base and with green margins towards the apex, elliptic-spathulate to obovate, acute, externally glabrous and convex towards the apex, internally glabrous, concave at the base, lightly convex and papillate towards the apex, the apical margins reflexed. Lateral sepals connate, forming a synsepal, mm, with subacute apices, purplishlilac, white with purplish-lilac spots towards the base and with green margins towards the apex, externally glabrous, carinate, convex towards the apex; internally concave, glabrous

5 2010 Fraga and Kollmann, New species of Pabstiella from Brazil 175 Figure 3. Pabstiella muricatifolia Fraga & L. Kollmann. A, habit; B, detail of the adaxial surface of the leaf; C, flower in side view; D, dorsal sepal; E, lateral sepals; F, petal; G, lip; H, lip in side view; I, column and ovary in side view; J, anther from below; K, anther, from above; L, pollinia. Drawn from the holotype specimen.

6 176 Harvard Papers in Botany Vol. 15, No. 1 at base, papillate above, acute. Petal mm, purplish-lilac, base white, margins green towards the apex, deltoid, asymmetrical, margins lightly deflexed, externally lightly carinate, apex acute and apiculate. Lip cm, greenish with purplish-lilac spots and grooves, oblong, 3-lobed, base truncate with a pair of white, minute, rounded lobules at both corners, the disc shallowly channeled between a pair of purplish-lilac, longitudinal, papillate, thick calli, the lateral lobes white with purplishlilac spots, the apical lobe lightly concave, papillate, externally glabrous and carinate, the apex purplish-lilac, obtuse. Column mm long, translucent white with purplish-lilac spots, semiterete, winged above the middle, denticulate at the apex, the anther, rostellum and stigma, ventral, the foot thick, ca.1 mm long. Anther mm, yellowish with lightly purplish-lilac apical margins, hooded, gibbous; pollinia 2, yellow, obovoid. Capsule mm, green, cylindrical, asymmetrical, recurved. Etymology: the specific epithet derives from the Latin muricatus, muricate, and folium, a leaf, in reference to the muricate surfaces of the leaves, the most distinguishing character of this species. Habitat and distribution: so far this species is only known from two collections in the northwest of the Espírito Santo State to inselbergs covered with dry seasonal forests. However A. L. V. Toscano de Brito (pers. comm.) informs the occurrence of this species in the Atlantic Forest of Domingos Martins, in direction to the south of the State (Fig. 2). Conservation status: due to the insufficient information on the abundance and distribution, we include this species in the Data Deficient category of the IUCN Red List of endangered plant species (IUCN, 2001): DD. Paratypes: Brazil, Espírito Santo: Pancas, Lajinha, 3 April 2007, (fl., cult.) L. Kollmann (MBML) This new species is similar to Pabstiella hypnicola and Pabstiella villosisepala from which it is distinguished by the shape, texture, and size of the leaves, and floral morphology. In Pabstiella muricatifolia the leaves are obovate to oblong, thick, coriaceous, muricate on both surfaces, and less than 2.7 cm long, the flowers are resupinate, the sepals are papillate on the apical portion, and the lip is distinctly 3-lobed, obtuse and papillate at apex, whereas in Pabstiella hypnicola and in Pabstiella villosisepala the leaves are lanceolate to ellipticlanceolate, chartaceous, more than 4 cm long, the flowers are non-resupinate, the sepals are velvety on the apical portion, and the lip is entire or indistinctly 3-lobed with a rounded, smooth to verruculose apex. Pabstiella villosisepala L.Kollmann & Fraga, sp. nov. TYPE: BRAZIL. Espírito Santo: Santa Teresa, Nova Lombardia, Augusto Ruschi Biological Reserve, 25 March 2004 (fl.), L. Kollmann 6571 (Holotype: MBML [22235]; Isotype: RB). Fig. 4. Haec species Pabstiella hypnicolae (Lindl.) Luer affinis, sed foliis angustioribus et apice sepalorum villoso differt. Plant epiphitic, up to cm tall, caespitose, erect. Roots whitish, fasciculate, terete, glabrous. Ramicauls cm long, green, terete, enclosed by 1 2 thin, tubular sheaths; sheaths cm long, green and paleaceous when mature. Leaf cm, including a petiole cm long, green, erect; blade thickly chartaceous, lanceolate to narrowly elliptic-lanceolate, the base long acuminate to subpetiolate, the apex acute, 3-toothed. Inflorescence 5 12 cm long, a loose, slightly flexuous, several-flowered raceme, emerging from near the apex of the ramicaul; floral bracts, ca. 2 mm long, green, tubular, acuminate. Flowers not resupinate, opening in succession, occasionally 2 flowers remain open simultaneously. Pedicels cm long; ovary mm long, greenish, subterete, longitudinally sulcate. Dorsal sepal mm, yellowish towards the apex and translucent with purplish-lilac spots towards the base, elliptic-spathulate to obovate, base attenuate, apex obtuse, concave basally and convex towards the apex, externally glabrous, internally glabrous at base and villous towards the apex, margins somewhat reflexed. Lateral sepals connate, forming a synsepal, mm, with subacute apices, translucent at the widest portion, white in the center from the base to the apex, translucent yellow near the margins with purplish-lilac spots at base, glabrous externally, internally glabrous with the revolute, lateral margins villous from about the middle towards the tips, the base slightly cuneate, the tips obtuse Petal mm,

7 2010 Fraga and Kollmann, New species of Pabstiella from Brazil 177 Figure 4. Pabstiella villosisepala L. Kollmann & Fraga. A, habit; B, flower in side view; C, dorsal sepal; D, lateral sepals; E, petal; F, lip in side view; G, lip; H, column and ovary in side view; I, anther in side view; J, anther, from above. Drawn from the holotype specimen.

8 178 Harvard Papers in Botany Vol. 15, No. 1 spathulate-obovate to rhomboid, asymmetrical, rough on both surfaces, attenuate at base, yellow with purplish-lilac spots at the base, dark purple towards the acute, papillate and thick externally apex. Lip cm, mm wide expanded, white in the center with purplish-lilac spotted margins, elliptic to oblong, 3-lobed, thick, the base subtruncate with a pair of white, minute, rounded lobules at both corners, the disc shallowly channeled centrally between a pair of thick, oblong, glandular-cellular calli the lateral lobes short, rounded, the apical lobe elliptic, papillose, acute. Column 3 4 mm long, white with purplish-lilac spots and, semiterete, winged above the middle, denticulate at the apex, prolonged at base into a short (ca. 1 mm long), thick, purplish-lilac foot, the anther, rostellum and stigma ventral. Anther 0.8 mm long, yellowish with lightly purplish-lilac apex and margins, hooded, the apex papillose; pollinia 2, yellow, obovoid. Capsule unknown. Etymology: the specific epithet derives from the Latin villosus, villous, and sepalum, a sepal; in reference to the adaxially villous apical portion of the sepals, the most distinguishing character of this species. Habitat and distribution: so far this species has only been found in two localities in the município of Santa Teresa: the Augusto Ruschi Biological Reserve, in Nova Lombardia, and Mata Fria, in São Lourenço. It is an epiphyte in patches of Atlantic Forest at ca. 700 m, but probably also occurs in adjacent areas in the State of Espírito Santo (Fig. 2). Conservation status: due to the apparent rarity and restricted distribution of Pabstiella villosisepala in the Atlantic Forest in central region of Espírito Santo State and its apparent vulnerability to human activities, we include this species in the Vulnerable category of the IUCN Red List of endangered plant species according to the following criteria (IUCN, 2001): VU D2. Paratypes: Brazil: Espírito Santo, Mun. Santa Teresa: Nova Lombardia, Augusto Ruschi Biological Reserve, 11 April 2002 (fl.), R. R. Vervloet and E. Bausen 109 (MBML); Nova Lombardia, Augusto Ruschi Biological Reserve, July 1976 (fl.), W. Emerich s.n. (MBML); Nova Lombardia, Augusto Ruschi Biological Reserve, 27 May 2003 (fl.), R. R. Vervloet and W. Pizziolo 2498 (MBML, RB). São Lourenço, Mata Fria, 18 August 2004 (fl.), L. Kollmann and W. Pizziolo 6954 (MBML). This new species is similar to Pabstiella hypnicola, from which it may be distinguished by the leaf width and the indumentum of sepals. The leaves in Pabstiella hypnicola are usually more than 1.5 cm wide and the sepals are velvety on the apical portion, whereas in P. villosisepala the leaves are less than 1 cm wide and the sepals are shaggy on the apical portion. Barros, F Notas nomenclaturas em Pleurothallidinae (Orchidaceae), principalmente brasileiras. Bradea 8: Brieger, F. G., and K. Senghas Pabstiella, eine neue Orchideengattung aus Brasilien. Die Orchidee 27: Holmgren, P. K., N. H. Holmgren, and L. C. Barnett, (eds.) Index Herbariorum Part I: The herbaria of the world. 8 ed. New York Botanical Garden, New York. 704p. IUCN IUCN Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission. IUCN, Gland, Switzerland, and Cambridge, United Kingdom. Luer, C.A Icones Pleurothallidinarum XXVIII. A reconsideration of Masdevallia, Systematics Literature Cited of Specklinia and vegetatively similar taxa. (Orchidaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 105: Icones Pleurothallidinarum XXIX. A Third Century of Stelis of Ecuador, Systematics of Apoda-Prorepentia, Systematics of Miscellaneous Small Genera, Addenda New Genera, Species and Combinations (Orchidaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 112: (2009). Icones Pleurothallidinarum XXX. Addenda: Miscellaneous new combinations. Monogr. Syst. Bot. Missouri Bot. Gard 115: Pridgeon, A. M., and M. W. Chase A Phylogenetic reclassification of the Pleurothallidinae (Orchidaceae). Lindleyana 16: