New species and combinations in Thelypteris subg. Goniopteris (Thelypteridaceae)

ALEXANDRE SALINO

Salino, A. (Departamento de BotaÃnica, Instituto de CieÃncias BioloÂgicas, Univ- ersidade Federal de Minas Gerais. Av. AntoÃnio Carlos, 6627, Pampulha, 31270- 901, Belo Horizonte, Minas Gerais, Brazil; email: [email protected]). New spe- cies and combinations in Thelypteris subg. Goniopteris (Thelypteridaceae). Brit- tonia 54: 331±339. 2002.ÐThree new species of Thelypteris are described and illustrated: T. littoralis and T. paranaensis, endemic to the rainforest of the southern Brazilian Atlantic coast, and T. multigemmifera, endemic to inland gallery forests of the SaÄo Paulo state. In addition, four new combinations are made: T. cutiataensis (Brade) Salino, T. iguapensis (C. Chr.) Salino, T. mono- sora (C. Presl) Salino, and T. schwackeana (H. Christ in C. Chr.) Salino. A new name is proposed for Dryopteris lugubris var. quadrangularis: T. montana Sal- ino. The species treated here belong to Thelypteris subg. Goniopteris due to the indument of furcate and stellate trichomes. Key words: Brazil, ferns, Goniopteris, pteridophytes, Thelypteridaceae, Thelyp- teris.

During a taxonomic survey of the genus teris (Smith, 1983, 1992), but are always Thelypteris in southeastern Brazil, I discov- present in the Brazilian species (Salino & ered three new species of and made several Melo, 2000). Thelypteris subg. Goniopteris combinations for Thelypteris, one of the comprises 80 to 100 neotropical species most poorly known fern genera in Brazil. (Smith, 1992). Of the 30 species I have As treated by Smith (1992), this genus identi®ed in Brazil, 22 occur in southeast- comprises about 875 species worldwide ern Brazil with at least 14 endemic to this with nearly 300 species in the New World. region. The species of southeastern Brazil Following Smith (1992), I have recognized occur in shaded habitats in lowland and ®ve subgenera and 86 species in Brazil: montane rainforests, from 0 to 1000 m el- subg. Amauropelta (Kunze) A. R. Sm. is evation. In this work, I provide descriptions represented by 33 species, subg. Cyclosorus of three new species and four new combi- (Link) C. V. Morton by eight species, subg. nations, and propose one new name. Geo- Goniopteris (C. Presl) Duek by 30 species, graphical distributions and habitats are dis- subg. Meniscium (Schreb.) C. F. Reed by cussed under each species. eight species, and subg. Steiropteris (C. Chr.) K. Iwats. by ®ve species. There are Thelypteris cutiataensis (Brade) Salino, two more species of uncertain subgeneric comb. nov. position in southeastern Brazil. Dryopteris cutiataensis Brade, Arch. Jar. Bot. Rio The species treated here belong to The- de Janeiro. 11: 27. 1951. Goniopteris cutiataensis lypteris subg. Goniopteris because of the (Brade) Brade, Bradea 1: 215. 1972. indument of furcate and stellate trichomes. TYPE: BRAZIL. Rio de Janeiro: Mangar- The presence of furcate or stellate tri- atiba, Ilha de CutiataÂ-AssuÂ, 12 May 1940, chomes is a synapomorphy of subg. Gon- A. C. Brade 16275 (HOLOTYPE: RB). iopteris. Such trichomes are apparently Distribution and habitat. This species is lacking in a few species of subg. Goniop- endemic to the Atlantic Rain Forest in the

Brittonia, 54(4), 2002, pp. 331±339. ISSUED: 16 April 2003 ᭧ 2002, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. 332BRITTONIA [VOL. 54 coastal region of the Brazilian States of SaÄo Thelypteris iguapensis resembles T. vivi- Paulo and Rio de Janeiro. It grows mainly para (Raddi) C. F. Reed, but differs in its along streams, from 0±20 m. gradually reduced pinnati®d lamina apex, more numerous pinnae, and glabrous spo- Additional specimens examined: BRAZIL: Rio de Janeiro: Campo Grande, Serra da Mendanha, 1 Jun rangia. Thelypteris vivipara has a conform 1958, Costa et al. 31 (R). SaÄo Paulo: Ubatuba, Parque lamina apex and glabrous sporangia or has Estadual da Ilha Anchieta, 7 Feb 1996, Salino 2563 unbranched trichomes on the sporangial (BHCB). stalks. According to Brade (1951), Thelypteris cutiataensis differs from T. riograndensis Thelypteris littoralis Salino, sp. nov. (Lindman) C. F. Reed by having more veins (Fig. 1) (5±10 pairs per segment) and in its bud po- TYPE: BRAZIL. SaÄo Paulo: PeruõÂbe, Es- sition. Brade also compared T. cutiataensis tacËaÄo EcoloÂgica da JureÂia, Arpoador, Trilha with Dryopteris vivipara var. platylepis do Imperador, 7 Jan 1999, J. Prado et al. (FeÂe)C.Chr.[ϭT. vivipara (Raddi) C.F. 977 (HOLOTYPE: SP; ISOTYPE: UC). Reed], but T. cutiataensis has more deeply incised pinnae (to half the distance to the Species nova subgeneri Goniopteridis (C. Presl) costae) and usually has two pairs of basal Duek, Thelypteride semirii Salino & Melo et T. sca- brae (C. Presl) Lellinger af®nis sed differt frondibus veins from adjacent segments joining below subdimorphicis, petiolo frondis fertilis quam petiolo the sinus. Thelypteris cutiataensis is prob- frondis sterilis longiore, rachidibus adaxialiter dense ably more closely related to T. abbiattii C. pilosis, pilis stellatis vel furcatis vel simplicibus or- F. Reed and T. riograndensis, but differs in natis, facies abaxialis costae pilis simplicibus ornatis, pili longiori 0.25±0.5 mm longi, pili breviores simpli- having less incised pinnae (lobed to serrate cibus 0.02±0.12 mm longi. Sori inframediani indusiati, margin), 5±10 vein pairs per segment (2±5 sporangiis glabris. in the other two species) and linear to lan- ceolate scales abaxially on rachis and cos- Stem ascending to erect; caudex narrow tae. Thelypteris abbiattii has buds or plant- (ca. 1 cm diam.); scales lanceolate, casta- lets in the axils of the proximal pinnae. neous, pubescent; trichomes simple, fur- cate, and stellate. Leaves several, clustered, Thelypteris iguapensis (C. Chr.) Salino, 37.7±56.5 cm long, slightly dimorphic, the comb. nov. fertile ones longer petioled. Petioles sulcate adaxially, 16.5±33.5 cm ϫ 0.14±0.24 cm, Dryopteris iguapensis C. Chr., Dansk Bot. Ark. with scales only at the base, these sparsely 9(3): 63. 1937 [as ``ignapensis'']. Goniopteris ig- uapensis (C. Chr.) Brade, Bradea 1: 215. 1972. to moderately pilose; trichomes furcate and stellate. Laminae 20.8±23.5 cm long, ob- TYPE: BRAZIL. SaÄo Paulo: Iguape, Mor- long or elliptic or ovate, herbaceous, not ro das Pedras, 1922, A. C. Brade 8232 (HO- verrucose, 1±pinnate-pinnati®d; apex grad- LOTYPE:C;ISOTYPES: BM, NY, R, UC, US). ually reduced, con¯uent, and pinnati®d. Distribution and habitat. This species is Buds and aerophores absent. Rachises known from SaÄo Paulo to the State of Santa densely pilose (especially on adaxial side); Catarina in the coastal region of Brazil. It trichomes simple, furcate, and stellate. Pin- grows in the Atlantic Rain Forest, usually nae 9 or 10 pairs per leaf, 8.2±9.8 cm ϫ near streams, from 0±70 m. 1.8±2 cm, perpendicular to ascending (ex- cept basal pinnae strongly de¯exed), lan- Selected specimens examined: BRAZIL. Santa Ca- tarina: Hammonia, Aug 1910, Luederwaldt s.n. ceolate to elliptic, incised ⅔ to ¾ their (BHCB, SPF). SaÄo Paulo: CubataÄo, 1909, Wacket 95 width, sessile to short-stipitate (to 0.8 mm (NY); GuarujaÂ, s.d., Usteri s.n. (SP); Ribeira, Dec long), the apex acuminate to long acumi- 1910, Brade 5115 (HB, SP); Serra do Mar, s.d., Wacket nate and the base truncate or oblique in the s.n. (SP). basal pair; costae lacking scales abaxially; Christensen (1937) described this species costae, costule, sinus, and laminar tissue as Dryopteris ignapensis. Brade (1972), pubescent near margin of segments; tri- when transferring this species to Goniop- chomes adaxially simple; trichomes abaxi- teris, corrected the epithet to iguapensis. ally moderate-to-dense (especially on cos- 2002] SALINO: THELYPTERIDACEAE 333

FIG.1. Thelypteris littoralis. A. Habit. B. Abaxial surface of segments. C. Adaxial surface of rachis. D. Adaxial surface of segments. (From Prado et al. 977, SP.) tae), with mixture of long (ca. 1 mm) and dusia reniform, entire, the surface pubes- short (ca. 0.1 mm) mostly simple trichomes, cent with simple trichomes; sporangia gla- a few stellate, some reddish; laminar tissue brous. on abaxial side with numerous, simple, 2± Distribution and habitat. Thelypteris 4 mm long trichomes. Segments 3.6±4 mm littoralis is rare and known only from the wide, arcuate, entire, rounded to acute; type collection. It grows along trails in the veins 9±12 pairs per segment, unbranched, coastal Atlantic Rain Forest, at sea level. the basal vein pairs from adjacent segments Thelypteris littoralis is most similar to T. running to the sinus. Sori inframedial; in- scabra (C. Presl) Lellinger (from Argenti- 334BRITTONIA [VOL. 54 na, Paraguay, and southeastern Brazil) and Thelypteris monosora resembles T. cu- T. semirii Salino & Melo (from Minas Ger- neata (C. Chr.) C. F. Reed in pinnae shape ais, Brazil) in its pinna shape, deeply in- and venation, but the latter has longer pet- cised pinnae, and inframedial sori, but it ioles, lacks scales on the rachises and cos- differs in having slightly dimorphic leaves tae, and has furcate trichomes on the indu- with the fertile leaves longer petioled and sia and sporangial stalks. In contrast, The- indument on both sides of the pinnae. The- lypteris monosora has small scales on the lypteris semirii has short-stipitate glandular rachises and costae abaxially, and simple trichomes on both sides of the laminar tis- trichomes on the indusia and sporangial sue; simple trichomes are frequently absent. stalks. Thelypteris scabra has abaxially glabrous or slightly pubescent pinnae with simple tri- chomes, whereas T. littoralis has pinnae Thelypteris montana Salino, nom. nov. that are abaxially moderately to densely pi- (Fig. 2D±F) lose with simple 2±4 mm long trichomes, Aspidium quadrangulare FeÂe, Crypt. Vasc. BreÂs. 1: long-acuminate pinna apices and inframe- 145. 1869. Dryopteris lugubris var. quadrangu- dial sori. Thelypteris littoralis is slightly laris (FeÂe) C. Chr., Kongel. Danske Vidensk. similar to T. lugubris (Kunze ex Mett.) R. Selsk. Skr., Naturvidensk. Math. Afd. Ser. 7, 10: 243. 1913. M. Tryon & A. F. Tryon, but the latter spe- cies has long-creeping stems, medial sori, TYPE: BRAZIL. Rio de Janeiro: Serra da smaller simple trichomes on the abaxial Estrella, 4 Apr 1867, A. F. M. Glaziou 962 side of the laminar tissue, and costal scales. (HOLOTYPE: P). The speci®c epithet refers to the ocur- Distribution and habitat. This species is rence of the species only along the coast of known only from southeastern Brazil in the the State of SaÄo Paulo. States of Espirito Santo, Rio de Janeiro, and Minas Gerais. It grows in montane forests, Thelypteris monosora (C. Presl) Salino, from 700±1500 m. comb. nov. Selected specimens examined: BRAZIL. Espirito Lastrea monosora C. Presl, Epim. 36. 1849. Dryop- Santo: Cachoeiro do Itapemirim, 31 Aug 1948, Brade teris monosora (C. Presl) C. Chr., Biolog. Arbej- 19374(RB). Minas Gerais: Juiz de Fora, 29 Sep 1937, der Tilegn. Eug. Warming 84. 1911. Goniopteris Brade 15910 (RB). Rio de Janeiro: Itatiaia, 21 May monosora (C. Presl) Brade, Bradea 1: 216. 1972. 1935, Brade 14499 (NY, RB); PetroÂpolis, Spannagel TYPE: BRAZIL. Rio de Janeiro: s.d., J. 491 (GH,UC); Serra da Estrela, Nov 1916, Lutzelburg 6838 (UC, US); TeresoÂpolis, 28 Sep 1929, Brade 9490 HOLOTYPE ISO B. E. Pohl s.n. ( : PR, n.v.; - (UC). TYPE: BM). Distribution and habitat. This species is Christensen (1913) recognized four va- known from the coastal region of the Bra- rieties of Dryopteris lugubris [ϭT. lugub- zilian States of Rio de Janeiro and SaÄo Pau- ris] based on lamina and pinna size, quan- lo. It grows in the Atlantic rain forest, from tity of costal scales, presence of buds, num- 30±1200 m. ber of veins in the segments, and presence of stellate trichomes on the abaxial side of Selected specimens examined: BRAZIL. Rio de Ja- neiro: Angra dos Reis, 29 Jun 1935, Brade 14939 the laminar tissue. However, the majority of (GH); Parati, rd. to Praia do Sono, 15 Nov 1993, Sal- these characters are not constant in these ino 1910a (BHCB). SaÄo Paulo: Bananal, EstacËaÄo varieties; only the presence of stellate tri- EcoloÂgica de Bananal, 11 Sep 2001, Salino 7481 chomes on the abaxial side of laminar tissue (BHCB); Ilha Bela, Parque Estadual, rd. to Praia dos is constant in var. quadrangularis. Among Castelhanos, 02 Mar 1995, Salino 2098 (BHCB, UC); Serra do Itatins, Mar 1924, Brade 8509 (HB); SaÄo Pau- the three varieties (except var. quadrangu- lo, Ypiranga, 22 Jul 1910, Luederwaldt s.n. (SP); SaÄo laris) there are numerous intermediate Vicente, Vila SaÄo Jorge, Horto Municipal, 30 Sep forms, suggesting they represent forms of 1991, Mendez & Castro 05 (SJRP); Raiz da Serra, the same species. Dryopteris lugubris var. 1907, Wacket 200 (UC); Ubatuba, EstacËaÄo Experimen- tal do Instituto AgronoÃmico, 12 Nov 1993, Salino quadrangularis is here considered as a dis- 1899 (BHCB, UC); Picinguaba, Casa da Farinha, 30 tinct species from T. lugubris; it receives a Jan 1996, Salino 2460 (BHCB). new name because the epithet quadrangu- 2002] SALINO: THELYPTERIDACEAE 335

FIG.2. A±C. Thelypteris paranaensis. A. Habit. B. Abaxial surface of segments. C. Abaxial side of laminar tissue, showing furcate and stellate trichomes. (From Salino 1459, BHCB.) D±F. Thelypteris montana. D. Habit; note buds in axils of distal pinnae. E. Abaxial surface of segments. F. Detail of the abaxial side of laminar tissue, showing furcated and stellate trichomes. (From Brade 14499, RB.) 336BRITTONIA [VOL. 54

FIG.3. Thelypteris multigemmifera. A. Habit. B. Abaxial surface of segments. C. Adaxial surface of seg- ments. D. Adaxial side of pinnae base; note bud in pinna axil. E. Adaxial surface of rachis. (From Salino 1800, BHCB.) laris is not available at the species level in Thelypteris. Thelypteris multigemmifera Salino, sp. Thelypteris montana differs from T. lu- nov. (Fig. 3) gubris in having the laminar tissue densely TYPE: Brazil. SaÄo Paulo: AnalaÃndia, Ser- puberulous with stellate trichomes abaxially ra do Cuscuzeiro, Sitio Agua Fria, 28 Aug (Fig. 2F), and indusia bearing furcate and 1993, A. Salino 1800 (HOLOTYPE: BHCB; stellate trichomes. Thelypteris lugubris has ISOTYPES: UC, UEC). only sparse, simple trichomes on the lami- nar tissue abaxially and simple or rarely Species nova subgeneri Goniopteridis (C. Presl) furcate trichomes on the indusia. Duek, Thelypteride scabrae (C. Presl) Lellinger af®nis The speci®c epithet refers to the occur- sed facie pinnis abaxialiter trichomatibus stellatis vel furcatis destitutis, gemmis vel plantulis in axillis pin- rence of the species only in a mountaineous narum medianarum et distaliter costis adaxialiter, sinu region (Serra da Mantiqueira) of southeast- prominens, sori mediani, indusiis persistentibus reni- ern Brazil. formis pilosis et sporangiis glabris differt. 2002] SALINO: THELYPTERIDACEAE 337

and the aerophores that are sometimes pre- Stem short- to long-creeping; scales lan- sent. However, T. multigemmifera has lon- ceolate, brownish, pubescent; trichomes ger stems, less prominent sinuses of the simple, furcate, and stellate. Leaves several, pinnae, medial sori, persistent and reniform clustered, 37±91 cm long, monomorphic. indusia, and glabrous sporangia, whereas T. Petioles sulcate adaxially, 17±53 cm ϫ scabra has prominent sinuses, inframedial 0.15±0.40 cm, glabrescent to short-pubes- sori, subpersistent and spathulate indusia, cent, with scales on proximal half. Laminae and simple or no trichomes on the sporan- 20 to 45 cm long, elliptic, chartaceous, not gial stalks. Thelypteris multigemmifera verrucose, 1±pinnate-pinnati®d; apex grad- slightly resembles T. lugubris (Kunze ex ually reduced, con¯uent, and pinnati®d. Mett.) R. M. Tryon & A. F. Tryon (from Buds and plantlets present in axils of me- southeastern Brazil) in having short- to dian and distal pinnae and on adaxial side long-creeping stems and a similar rachis in- of costae of proximal and median pinnae. dument, but T. multigemmifera differs in Aerophores absent or sometimes present. having proximal pinnae with an oblique or Rachises pulverulent to pilose, with red- cuneate base, 5±9 vein pairs per segment dish, simple, furcate and stellate trichomes, (vs. 9±25 in T. lugubris), and pinnae abax- especially on adaxial side. Pinnae 12±23 ially lacking furcate and stellate trichomes. pairs per leaf, perpendicular to ascending Thelypteris lugubris usually has proximal (except basal pinnae strongly de¯exed), pinnae with a truncate base, and furcate and sessile or short stipitate (to 1.3 mm), 6.5± stellate trichomes abaxially on the costae, 11 cm ϫ 1.4±2.2 (±3.5) cm, elliptic or lan- costules, and veins. ceolate, incised ca. ½ their width; apex acu- The speci®c epithet refers to the presence minate in distil pinnae; base truncate in dis- of many buds or plantlets on the laminae. tal pinnae and oblique or slightly cuneate in proximal pinnae; adaxially pilose with Thelypteris paranaensis Salino, sp. nov. simple or furcate trichomes on the costae, (Fig. 2A±C) sometimes with simple trichomes on the costules; abaxially pilose with redish simple TYPE: BRAZIL. ParanaÂ: ParanaguaÂ, Ilha trichomes on the costae, costules, veins, and do Mel, 11 Oct 1992, A. Salino 1459 (HO- segment margins. Segments 3±5 mm wide, LOTYPE: BHCB; ISOTYPES: MBM, UC, entire, the apex rounded; veins 5±9 pairs UEC). per segment, unbranched, the basal pairs from adjacent segments running to or below Species nova subgeneri Goniopteridis (C. Presl) the sinus. Sori medial; indusia perisistent, Duek, Thelypteride cuneatae (C. Chr.) C. F. Reed et T. monosorae (C. Presl) Salino af®nis sed pinnis abaxi- reniform, entire, pale to brownish, with aliter et indusiis dense pilosis, pilis pedicelatis furcatis simple trichomeson the margin; sporangia vel stellatis, venis 2±4 paribus basalibus ex segmentis glabrous. contiguis ad sinum conniventibus differt. Sori mediani Distribution and habitat. Thelypteris indusiati, sporangiis glabris vel pedicellis pilosis. multigemmifera is rare and known only Stem short-creeping, the scales lanceo- from the type collection. It grows in par- late, brownish, with numerous stipitate, fur- tially shaded areas, in wet clayey soil, near cate trichomes. Leaves clustered, 80±160 a waterfall in the rain forest, at 1000 m. cm long, monomorphic. Petiole sulcate Thelypteris multigemmifera is character- adaxially, 35±70 cm ϫ 4±7 mm, with stip- ized by having simple trichomes on the ab- itate stellate trichomes and lanceolate, pi- axial side of the costae, costules, veins, and lose scales. Lamina 40±90 cm long, char- segment margins; aerophores (often) abax- taceous, ovate, 1±pinnate-pinnati®d, the ially in pinna axils; and many buds and apex abruptly reduced and similar (con- plantlets adaxially in pinna and costa axils. form) to lateral pinnae. Buds and sometimes This species is probably related to T. scabra plantlets present in the axils of distal pin- (from Argentina, Paraguay, and southeast- nae. Aerophores absent. Rachises with ern Brazil) because of its creeping stem, the sparse to moderately dense, stipitate, fur- position of furcate and stellate trichomes, cate trichomes. Pinnae 8±12 pairs per leaf, 338BRITTONIA [VOL. 54 sessile to often stalked to 1.7 mm, 12±30 ϫ southern Mexico, Guatemala to Panama, 1.8±4.0 cm, elliptic, incised ca. 1/3±2/3; Venezuela, Colombia to Peru, and Brazil) is apex acuminate; basal pinnae sometimes similar to T. paranaensis in having the de¯exed; suprabasal pinnae cuneate at the dense pubescence on the abaxial side of the base with several pairs of reduced seg- pinnae and in having furcate trichomes on ments; median pinnae truncate at the base. the sporangia stalks, but T. paranaensis dif- Segments 5±9 mm wide, entire, obtuse; fers in having proximal veins running to the veins 6±14 pairs per segment, sometimes sinuses, conform apical pinnae, 8±12 pairs furcated, often unbranched, the proximal 2± of lateral pinnae, the presence of proliferous 4 pairs from adjacent segments connivent buds in the axils of the distal pinnae, the at the sinuses. Indument abaxially: on cos- lack of anchor shaped trichomes, and the tae of 3 types (trichomes moderately dense presence of stipitate stellate trichomes of to dense, stipitate, furcate and stellate; tri- laminar tissue. Thelypteris biolleyi,onthe chomes sparse, simple; scales hairlike); on other hand, has proximal veins that unite veins and laminar tissue trichomes moder- below the sinuses with an excurrent vein ately dense to dense, stipitate, furcate and running to the sinus; lacks a conform apical stellate. Indument adaxially: on costae, seg- pinnae; lacks proliferous buds in the axils ment margins, and sometimes costules tri- of distal pinnae; has 15±32 pinnae pairs; chomes sparse, stipitate, furcate or simple. has a lamina apex that is gradually reduced, Sori medial; indusia entire to repand, pinnati®d, and con¯uent; has anchor shaped brownish, densely pilose, the trichomes trichomes on the abaxial side of the costae stipitate, furcate and stellate; sporangia gla- and laminar tissue (such trichomes are often brous or with 1-furcate (stipitate) or un- absent in Brazilian specimens); and has ses- branched trichomes on the sporangial sile stellate trichomes of the laminar tissue. stalks. The speci®c epithet refers to the occur- Distribution and habitat. Thelypteris rence of the species only in the Brazilian paranaensis is endemic to the northern state of ParanaÂ. coast of the State of Parana (including Ilha do Mel) in Brazil. It grows in forests (At- Thelypteris schwackeana (H. Christ in C. lantic forest), from 30 to 900 m. Chr.) Salino, comb. nov.

Additonal specimens examined: BRAZIL. ParanaÂ: Dryopteris schwackeana H. Christ in C. Chr., Kon- Antonina, Areia Branca, 7 Mar 1974, Hatschbach gel. Danske Vidensk. Selsk. Skr., Naturvidensk. 34295 (MBM); Campina Grande do Sul, Serra do Es- Math. Afd., ser. 7, 10: 243, f. 37. 1913. Goniop- pia, 1 Sep 1962, Hatschbach 9230 (MBM, LP); Serra teris schwackeana (H. Christ in C. Chr.) Brade, do Capivari Grande, 2 Aug 1984, Kummrow 2496 Bradea 1: 217. 1972. (MBM, UC); Estrada Engenheiro Lange a Prainhas, Serra do Mar, 13 Feb 1951, Tessmann 470 (MBM, TYPE: BRAZIL. Minas Gerais: Ouro RB); Morretes, 2 Sep 1969, Hatschbach 22121 (MBM, Preto, locis humidis, 22 Aug 1902, C. A. UC); Morro Grande, 22 Nov 1971, Hatschbach 28122 (UC); ParanaguaÂ, Ilha do Mel, 18 Sep 1953, Tessmann W. Schwacke 14892 (LECTOTYPE: here des- 1220 (MBM), 14 Feb 1986, Silva 198 (BHCB), 23 Feb ignated, BM). 1986, Britez 403 (BHCB); Volta Grande, Serra do Mar, Distribution and habitat. This species is 29 Jul 1914, DuseÂn 741a (BM, F, GH, NY, US). known from the Brazilian States of SaÄo In aspect, Thelypteris paranaensis resem- Paulo and Minas Gerais. It grows in mesic bles T. monosora and T. cuneata, but dif- forests from 400±900 m. fers in having many stellate and furcate tri- Selected specimens examined: BRAZIL. Minas chomes on the costae, costules, veins, lam- Gerais: Caldas, 25 Feb 1876, MoseÂn 4616 (S); Furnas, inar tissue, and indusia. Thelypteris mono- 8 Jul 1995, Salino 2208 (BHCB); SabaraÂ, 15 Mar sora and T. cuneata have sparse, furcate 1983, Vaz et al. s.n. (BHCB). SaÄo Paulo: Atibaia, Ser- trichomes on the costae, costule, veins, and ra do Itapetinga, 26 Jun 1914, Brade & Tamandare indusia, but lack them on the laminar tissue, s.n. (SP); Brotas, Fazenda Santa Eliza, 10 Aug 1991, Salino 1004 (UC, UEC); Campinas, BaraÄo Geraldo, 16 and T. monosora has pilose scales on the Mar 1992, Salino 1333a (BHCB); GaÂlia, EstacËaÄo Ecol- rachis and costa abaxially. Thelypteris biol- oÂgica de Caitetus, 13 Jul 1994, Salino 1958a (BHCB); leyi (H. Christ) Proctor (from Jamaica, Nova Itapirema, Monte Belo, Aug 1988, Salino 534 2002] SALINO: THELYPTERIDACEAE 339

(BHCB, UEC); SaÄo Manuel, Jul 1912, Luederwaldt Morato Duarte for preparing the illustra- s.n. (SP). tions. I also thank the curators of GH, UC, Thelypteris schwackeana resembles T. NY, and US for access to specimens and jamesonii (Hook.) R. M. Tryon, but the lat- hospitality during visits. ter species has stems erect to decumbent; the pinnae are moderately to densely pilose Literature Cited with only spreading, setiform trichomes Brade, A. C. 1951. Filices Novae Brasiliensis VII. adaxially; and furcate trichomes are lacking Arch. Jar. Bot. Rio de Janeiro 11: 21±36. ÐÐÐ. 1972. O geÃnero Dryopteris (Pteridophyta) no on the costae and costules. Thelypteris Brasil e sua divisaÄo taxonoÃmica. Bradea 1: 191± schwackeana has creeping stems, and the 261. pinnae are sparsely pilose adaxially toward Christensen, C. 1913. A monograph of the genus the edge of the segments. This species is Dryopteris I. The tropical American pinnati®d-bip- innati®d species. Kongel. Danske Vidensk. Selsk. also related to T. lugubris but differs mainly Skr., Naturvidensk. Math. Afd., ser. 7. 10: 53±282. in lacking scales on the costae abaxially and ÐÐÐ. 1937. Taxonomic fern studies. V. Dansk Bot. in having fewer veins per segment (6±7 Ark. 9(3): 63. pairs). Salino, A. & L. C. N. Melo. 2000. A new species of the fern genus Thelypteris (Thelypteridaceae) from southeastern Brazil. Novon 10: 74±77. Acknowledgments Smith, A. R. 1983. Polypodiaceae-Thelypteridoideae. In: G. Harling & B. Sparre, editors. Flora of Ec- I thank Alan R. Smith, Robbin C. Moran, uador 18: 18±148. ÐÐÐ. 1992. Thelypteridaceae. In: R. M. Tryon & and David B. Lellinger for their helpful R. G. Stolze, editors. Pteridophyta of Peru. Part III. comments on the manuscript, and Miryan Fieldiana Bot. n.s. 29: 1±80.