Acta bot. bras. 22(3): 878-890. 2008.

Liverworts of Alagoas State, Brazil

Lisi Dámaris Pereira Alvarenga1,2, Juliana Rosa do Pará Marques de Oliveira1, Mércia Patrícia Pereira Silva1, Sarah Oliveira da Costa1 and Kátia Cavalcanti Pôrto1

Received: August 17, 2007. Accepted: November 29, 2007

RESUMO – (Hepáticas do Estado de Alagoas, Brasil). No presente trabalho, são compiladas as espécies de hepáticas ocorrentes no Estado de Alagoas, resultantes de informações disponíveis em catálogos e trabalhos anteriores e do inventário inédito de uma importante unidade de conservação do Nordeste, a Estação Ecológica (EsEc) Murici (9º11’05” - 9º16’48”S; 35º45’20” - 35º55’12”O). São referidas para Alagoas 116 espécies de hepáticas, tendo 106 delas ocorrido na EsEc Murici. Setenta e oito espécies são genuinamente novas ocorrências para o Estado de Alagoas, sete das quais são novas também para a região Nordeste do Brasil. São fornecidos dados sobre a distribuição geográfica das espécies no Brasil e no mundo, e adicionalmente, comentários taxonômicos e ecológicos das espécies que são novas ocorrências para o Nordeste.

Palavras-chave: Hepáticas, Floresta Atlântica, Nordeste do Brasil, brioflora

ABSTRACT – (Liverworts of Alagoas State, Brazil). A list of liverworts from Alagoas State was compiled and is presented here. The list is based on catalogues and previous papers as well as the results of an unpublished survey carried out at the Murici Ecological Station (EsEc), an important protected area of the Brazilian Northeast (9º11’05” - 9º16’48”S; 35º45’20” - 35º55’12”W). One-hundred and sixteen liverworts have been recorded for the state of Alagoas, of which 106 occur at EsEc Murici. Seventy-eight are new occurrences for Alagoas, and seven of these are also new occurrences for northeastern Brazil. Data on geographic distribution in Brazil and worldwide is given here, in addition to ecological and taxonomic comments on the species that are new occurrences for the Northeast region.

Key words: liverworts, Atlantic Forest, northeastern Brazil, bryophyte flora

Introduction remnants of northeastern Brazil. EsEc Murici was created in 2001 with 6,116.43 ha and is considered a priority The bryophyte flora of Alagoas is one of the most area for biodiversity conservation in Brazil and worldwide poorly studied of the Brazilian Northeast. To date, no (Brasil – MMA 2000). The survey resulted in a list of systematic survey has been performed and all references 181 species, of which 106 are liverworts, 72 are mosses, come from sparse and sporadic collections whose results and three await special taxonomic treatment. This paper have been put together in Yano’s catalogues (1981; 1984; focuses only on liverworts, while mosses will receive 1989; 1995; 2006). attention in another publication. Recently, Pôrto et al. (2006) carried out an inventory This paper presents a compilation of liverwort in four Atlantic Rain Forest remnants, recording 76 species occurring in Alagoas State, bringing together species of bryophytes - liverworts (31 spp.) and mosses published information (catalogues) and results from the (45 spp.) - while Yano & Peralta (2007) increased this EsEc Murici survey. Moreover, it provides data about number by publishing 36 new occurrences (26 mosses the species’ geographic distribution in Brazil and in the and 10 liverworts) for Alagoas. Yet species richness in world, as well as taxonomic and ecological comments Alagoas is still low when compared to other states like on the new occurrences for the northeastern region. Pernambuco and Bahia, which house more than 300 spp. (Pôrto & Germano 2002; Bôas-Bastos, S.B.V., personal communication). Material and methods Additionally, Alagoas’ bryophyte flora is now EsEc Murici (9º11’05” - 9º16’48”S; 35º45’20” - increased by the survey carried out at the Murici 35º55’12”W) is part of Murici and Messias Ecological Station (EsEc), one of the main Atlantic Forest

1 Universidade Federal de Pernambuco, Departamento de Botânica, Laboratório Biologia de Briófitas, CCB, Av. Prof. Moraes Rêgo s.n., Cidade Universitária, 50670-901 Recife, PE, Brasil 2 Corresponding Author: [email protected] Alvarenga, Oliveira, Silva, Costa & Pôrto: Liverworts of Alagoas State, Brazil 879 municipalities, 50 km from Maceió, capital of the state geographic distribution in Brazil and in the world (Tab. 1). of Alagoas (Moura 2006). The elevation of the reserve Most of the species had typically Neotropical (59%), ranges between 200-650 m, and the area has warm, Pantropical (16%) and African-American (7%) humid wheather, with a dry summer and a wet autumn- geographic distributions. Lejeuneaceae is by far the winter season. The mean annual precipitation and richest family in the state, with 83 species. This obviously temperature are 2,167.7 and 24 °C, respectively (National is because the flora presented here comes mainly from Institute of Meteorology, data for the period from 1961 Atlantic Rain Forest. It is known that this family used to to 1990). The vegetation cover is classified as be predominant in the flora of tropical regions, especially Submontane Open Humid Atlantic Rain Forest (Veloso in forest habitats (Gradstein et al. 2001). et al. 1991). Geographic distribution details, taxonomic and The botanical material studied consists of epiphyllic ecological comments and ilustrations of the main (leaf inhabiting), epiphytic (bark inhabiting) and epixyllic characters of the new occurrences for Northeast Brazil (decaying wood inhabiting) samples collected from the are presented below. understory and canopy. The material was examined in the laboratory; identification and taxonomic and ecological ANEURACEAE commentaries were based on the following literature: Aneura pinguis (L.) Dumort., Comment. Bot. 115. Fulford (1963; 1966; 1968; 1976), Stotler (1970), Kron 1822. (1988), Reiner-Drewald & Goda (2000), Gradstein & Fig. 1-2 Costa (2003), and Dauphin (2003). Occasionally the identification of specimens was confirmed by specialists. Material examined: BRAZIL. Alagoas: Murici, The following literature was consulted to compile the 2/XII/2004, Pôrto (UFP 45851); ib. 2/XII/2004, Pôrto information on geographic distribution: Yano (1984; (UFP 45850). 1989; 1995; 2004; 2005; 2006), Bastos & Yano (2004; Description: Schuster (1992). 2006), Câmara & Vital (2004), Câmara & Costa (2006), Comments: this species was placed within Riccardia Costa & Silva (2003), Costa et al. (2005), Ganacevich S.F. Gray genus until recent decades. However, it can & Mello (2006), Germano & Pôrto (2005), Gradstein & be distinguished from this genus because of at least two Costa (2003), Joyce et al. (2006), Molinaro & Costa clear characters on gametophytes: 1) thallus less ramified, (2001), Oliveira et al. (2006), Paixão & Mello (2006), darker and wider, reaching up to 8 mm while Riccardia Patrus & Starling (2006), Pôrto et al. (2006), Valente & spp. reaches no more than 3 mm wide and; 2) many Pôrto (2006a; 2006b), Vilas Bôas-Bastos, Bastos & minutely dispersed oil bodies inside the cells. Aneura Ballejos (2006), Yano & Bordin (2006), Yano & Câmara pinguis is easily recognized in the field even when sterile, (2004), Yano & Costa (2000), and Yano & Peralta (2005; due to its conspicuous and lustrous thallus, green to 2006; 2007). The taxonomic classification adopted was yellowish-green in color. Of the Aneura Dumort. species that presented by Crandall-Stotler & Stotler (2000). All occurring in Brazil, A. pinguis is the only one with samples studied are registered and deposited in the UFP- 10-18 (20) cell thickness in the middle (transversal Geraldo Mariz Herbarium, at the Federal University of section); the others are thinner, up to 9 cells thick. In Brazil, this species was only known for the North, Central Pernambuco. and Southeast regions. It is sub-cosmopolitan and occurs The Alagoas liverwort species listed that did not at altitudes that vary from sea level to 800 m. It was occur in EsEc Murici where found in Yano’s catalogues found growing on decaying wood in EsEc Murici. (1981; 1984; 1989; 1995; 2006), Gradstein & Costa (2003), and Pôrto et al. (2006). LEJEUNEACEAE

Aphanolejeunea asperrima Steph., Sp. Hepat. 5: 859. Results and discussion 1916. One-hundred sixteen species of liverworts were Fig. 3-6 recorded for Alagoas. Seventy-eight species are new Material examined: BRAZIL. Alagoas: Murici, occurrences for Alagoas state, occurring exclusively at 2/XII/2004, Pôrto (UFP 45822); ib. 2/XII/2004, Pôrto EsEc Murici, and seven species are new occurrences (UFP 45823); ib. 2/XII/2004, Pôrto (UFP 45825); ib. for the Brazilian Northeast as well. The other 38 species 2/XII/2004, Pôrto (UFP 45829); ib. 2/XII/2004, Pôrto had been previously cited in catalogues and other papers. (UFP 45830); ib. 2/XII/2004, Pôrto (UFP 45833); ib. The taxa are listed in alphabetical order by family and 2/XII/2004, Pôrto (UFP 45845); ib. 2/XII/2004, Pôrto species and include locality, substrata and forest strata (UFP 45831); ib. 2/XII/2004, Pôrto (UFP 45842); ib. (understory or canopy) in the case of EsEc Murici, and 2/XII/2004, Pôrto (UFP 45844). 880 Alvarenga, Oliveira, Silva, Costa & Pôrto: Liverworts of Alagoas State, Brazil A, A, A, , P , P continue , MS, P . Sub-canopy A, PB, PE, A, PE, PR, A, PB, PE, , BA, CE , P , P , P , MS, MT , MS, MT A, PE, PR, RJ, RS, A, PE, RJ, RO, RR, , MT , MT , P , SP , P A, PE, RJ, RS, SP , SP , MS, MT A, PE, RJ, SP A, PE, RJ, RS, SP , PE, RJ, RS, SC, SP , ES, MS, P , P , PR, RS, RJ, SC, SP A, PE, MT AP AM, BA, ES, GO, MA, MT , MS, RJ, SP , BA, CE, ES, MG , MG AP AM, P AM, BA, CE, ES, MG AL, AM, AM, BA, CE, ES, GO, MG AM, BA, CE, ES, MG AM, BA, CE, ES, GO, MG AC, BA, CE, ES, MG AC, RJ, RO, RR, SP AL, DF AM, BA, GO, P PR, RJ, RS, SE, SP PR, RJ, RR, RS, SE, SP BA, ES, MG Distribution in Brazil AM, SC, SE, SP BA, CE, ES, GO, MG AL, RS, SC ES, GO, MG AM, MG RS, SC, SE, SP AL, BA, CE, ES, GO, MT AC, RJ, PE, RS, SP PE, PR, RJ, RR, RS, SC, SP AC, BA, ES, PE, RJ, SP AC, RJ, RS, SC, SE, SP ES, PE, RJ, RR, RS, SC, SE, SP AL, AL, AL, AC, PB, PE, PR, RJ, RS, SE, SP AC, America America America Africa and Brazil Pantropical Pantropical Pantropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical orld distribution W Subcomsmopolitan Africa and Africa and Africa and Asia, , , , , , , , , , canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy canopy canopy canopy canopy canopy canopy canopy canopy understory understory understory understory understory understory understory understory Forest strata sub-canopy understory understory understory understory understory understory understory V V V V , L , L , L , L V V V ------LB LB DW DW DW DW LB, L LB, L LB, L LB, DW LB, DW LB, DW Substrata LB, DW LB, DW LB, DW LB, DW , live bark - LB and decaying wood DW) forest strata occurrence concern only the EsEc Murici survey V Locality Junqueiro EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici Serra Grande Boca da Mata Marechal Deodoro Barra de São Miguel EsEc Murici, Messias, União dos Palmares, and Arnell A. Evans . .) Nees . et al . Gottsche (Meissn.) (Mont.) R.M. Schust. (Lindenb. & (Lehm. & ta .) Nees .) Nees (Spruce) S.W (Nees) Mont. Berk. (Reinw (Sw (Sw A Alagoas state. Information on substrate (Leaves - L (L.) Dumort. (Raddi) Ångstr Raddi Gottsche (Nees) Nees Nees teph. ensis S Gottsche (Lehm. & Lindenb.) dia amazonica rata euteliana YCACEAE Lindenb.) Schiffn. ullania apiculata Gottsche) Kruijt. yopteris diffusa Lehm. & Lindenb. yopteris filicina . ser . riojaneir . dusenii . brasiliensis . br . caulisequa . kunzei YOPTERIDACEAE *Aneura pinguis Fr Acrolejeunea torulosa Anoplolejeunea confer Acanthocoleus aberrans L. martiana F F F Riccar Cylindrocolea rhizantha Cronisia paradoxa Br F Lophocolea liebmanniana F F Br Frullania ericoides F ANEURACEAE BR axa able 1. Liverworts of T T MARCHANTIOPHYT JUBULACEAE LEJEUNEACEAE is defined here as strata above 4 meters high. New references for Northeast Brazil are preceded by an asterisk. CEPHALOZIELLACEAE CORSINIACEAE GEOCAL Acta bot. bras. 22(3): 878-890. 2008. 881 , PB, A, continue , P , MS, MT A, PE, PR, , P A, PE, RJ, A, PB, PE, PR, A, PE, PR, RR, , P , P , P , MS, MT A, PE, RJ, SC, SP A, PE, PR, RJ, RO, , P PR, SP A, PB, PE, RJ, RO, SP , , MS, MT , MT A, PE, RJ, RR, SP A , P , MT A, PE, RJ, RO, RR, RS, SP , MS, MT , RS, SP , MS, MT A, PE, RJ, RR , BA, CE, P , BA, CE, ES, GO, MG A, PE , MS, P A, PE, PR, RJ, SC, SP A, PR, RJ, SP AP AP , P , MS, MT , BA, MA, P , BA, CE, ES, P A, RJ AM, P AP , PE, RJ, RO, SP AP AM, AM, BA, ES, MG AM, , PR, RJ, RR, SP AM, BA, ES, GO, MG AM, BA, P , P A, PE A AL, BA, P AM, BA, ES, MG AL, AL, AL, AM, BA, CE, GO, MG AL, AL, AM, AM, AL, AM, BA, CE, ES, MT AM, BA, MG , SC A, PB, PE, PR, RJ, RR, SC, SE, SP A, PE, PR, RS, RO, RR, SC, SE SP SC, SP AC, AM, BA, ES, GO, MG AC, RJ, SP BA, P AC, BA, ES, MG AC, BA, ES, MG BA, MG P AM, AC, AM, MG AC, SC, SP BA, P PE, PR, RJ, RR, SP RJ, RO, RR, SC, SP AM AC, AM, BA, ES, GO, MG AC, AM, MG RS, SC, SP AL, AM, BA, ES, GO, MG AC, Distribution in Brazil MG AC, P AM, PE, SP AC, America America Brazil America and Pantropical Pantropical Pantropical Pantropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Costa Rica Pantropical Neotropical Neotropical Neotropical Neotropical orld distribution Brazil and Guianas Africa and Costa Rica, East W South Índias, Chocó, and Africa and , , , , , , , , , , , , , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy understory understory understory understory understory understory understory understory understory understory understory understory Forest strata understory understory understory understory understory understory understory understory understory understory understory understory V V V V V V V V V , L , L , L , L , L , L , L , L V V V , L LV LV LV LV LV LB LB LB LB LB DW LB, L LB, L LB, L DW LB, DW LB, DW LB, DW Substrata LB, DW LB, DW LB, DW LB, DW LB, DW LB, DW LB, DW LB, DW Locality EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici Serra Grande Serra Grande Serra Grande EsEc Murici, EsEc Murici, Serra Grande EsEc Murici and EsEc Murici and EsEc Murici and EsEc Murici and and Serra Grande and Serra Grande and Serra Grande and Serra Grande Marechal Deodoro, Marechal Deodoro, . & A. Evans Steph. (Gottsche) (Nees) Grolle (Mont.) (Hampe ex (Gottsche) Steph. ) Mizut. . Dauphin .) Grolle & E. Reiner et al. . diocarpa Horik. (Ångstr teph.) G (Nees & Mont.) Steph. Steph. (Nees) Schiffn. (Lehm. & Lindenb.) Kachr (Austin) R.M. Schust. (Spruce) Grolle & R.-L. Zhu (Lehm.) R.M. Schust. (Mont.) Schiffn. (S (Mont.) R.M. Schust. (Nees & Mont.) Schiffn. (Lindenb.) Schiffn. (Reinw R.M. Schust. (Kunze) Grolle Dauphin (Nees. & Mont.) R.M. Schust. (Lehm. & Lindenb.) Bonner Lehm.) Fulford A. Evans R.M. Schust. *A. camillii A. truncatifolia Archilejeunea fuscescens Cheilolejeunea acutangula Caudalejeunea lehmaniana C. adnata C. clausa A. parviflora *Aphanolejeunea asperrima C. holostipa C. minuta C. laetefusca C. rigidula C. discoidea C. oncophylla C. confusa Ceratolejeunea coarina C. trifaria C guianensis C. falax C. cornuta C. dussiana C. cubensis Cololejeunea car *C. lanciloba C. obliqua axa able 1 (continuation) T T 882 Alvarenga, Oliveira, Silva, Costa & Pôrto: Liverworts of Alagoas State, Brazil A, , P continue , MS, MT A, RJ, SP , PE, PR, RJ, SC, SP , P A, PE, PR, RJ, RR, SP , RJ, SC, SP , MT A, SP , P A, PE, RJ, RR, SP , P A, RJ, SC, SP A, PE, RJ, RR, SP , MT , PB, PE, RR, SP AS, MS, MT , RJ, RR, SP , BA, CE, P A, PB, PE, RJ, SP A, PE, RJ, SP A, SP A, PE, RJ, SP AP AM, BA, CE, MT , MT , RJ, SP AM, BA, ES, MG A AM, BA, CE, ES, GO, MG AL, AM, AM, BA, ES, PB, PE, RJ, SC, SE, SP AM, GO, MT AM, MG AM, BA, P AL, BA, MG AC, BA, MS, SP PB, PE, PR, RJ, RR, RS, SP BA AC, MS, P AC, ES, PE, RJ, SP AC, BA, CE, ES, MS, MT AL, BA, CE, CE, PE, PR, SC, SP BA, SP AC, AL, BA, PE, RO AL, AC, AL, AL, BA, MG AC, CE, P AM, BA, MG AM, BA, P AC, SP AL, AC, BA, P AM, BA, ES, P AM, BA, ES, MS, PE, PR, RJ, RS, SC, SP AM, P RJ, SC, SP Distribution in Brazil gentina Ar America Andes and Brazil Brazil and Brazil and Brazil East Índias, Pantropical Pantropical Pantropical Pantropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical South Argentina, and Dominica, Peru, North orld distribution Guiana and Brazil Southeastern Brazil W Cuba, USA, , , , , , , , , , , canopy , canopy , canopy , canopy , canopy , canopy , canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy understory understory understory understory , sub-canopy understory , sub-canopy understory , sub-canopy understory , sub-canopy understory , sub-canopy Forest strata understory understory understory understory understory understory understory understory understory understory understory understory understory understory understory understory V V V V V , L , L , L V V V V V V V V , L , L ------LV LV LV LV LB LB DW LB, L LB, L LB, L LB, L LB, L LB, L LB, L LB, L DW DW LB, DW LB, DW LB, DW LB, DW Substrata LB, DW LB, DW LB, DW Locality EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici Serra Grande Serra Grande EsEc Murici, and Serra Grande and Serra Grande and Serra Grande Marechal Deodoro Marechal Deodoro Marechal Deodoro, EsEc Murici and Serra Grande A. Evans Tixier (Lehm. & teph.) Steph. (Nees & Mont.) Steph. (S A. Evans (Lehm. & Lindenb.) Nees & Mont. ayl.) (Spruce) Steph. Tixier (Nees & Mont.) Steph. (T Steph. Gottsche Tixier (Steph.) E. Reiner & Goda (Steph.) E. Reiner & Goda A. Evans Gottsche ex Steph. Steph. (Meissn.) Schiffn. .) Nees (Steph.) Bischl. (Besch. & Spruce) Steph. Bischl. (Spruce) Grolle (Lindenb. & Gottsche) Steph. Spruce (Lehm. & Lindenb.) Gottsche Spruce (Sw ucianella epanolejeunea bidens Lindenb.) van Slageren L. grossiretis L. flava L. filipes L. controversa L. elliotti L. cerina L. caulicalyx D. cobrensis D. pellucida D. cavifolia D. latipuensis Cololejeunea subcardiocarpa Colura tortifolia Diplasiolejeunea brunnea Lejeunea cancellata Cyclolejeunea convexistipa D. rudolphiana C. luteola C. submarginata Dr Harpalejeunea oxyphylla Frullanoides corticalis *H. tridens H. stricta D. fragilis D. mosenii *D. campanulata *D. cr axa able 1 (continuation) T T Acta bot. bras. 22(3): 878-890. 2008. 883 , continue A, PB, A, PE, A, PB, , P , P , P , MS, MT A, PE, RJ, SP A, RJ, RS, SP , MT , P A, PE, RJ, RS, SP , P , MS, MT A, PE, RR, RS, SC, SP A, PE, SP A, PB, PE, SP A, PE, RJ, SC, SP , P A, PE, PR, RJ, SC, SP , MS, P , PE, RJ, RS, SC, SP , BA, ES, MG A, PE A, PE, RJ, RS, SC, SP AP , P , PE, RJ, RS, SC, SP A, PE A, PE, RJ A, PE, PR, RO, RR, SP AM, BA, CE, ES, MS, MT AM, BA, ES, P AM, BA, CE, ES, GO, MG AM, BA, CE, ES, MS, P AM, BA, CE, ES, MG , PE, SP AM, , PE, PR, RJ, SC, SP , AP AM, BA, CE, ES, MG AL, AM, BA, ES, P AL, BA, CE, ES, MS, MT AL, AM, BA, CE, ES, P AL, AL, AM, RR AL, , BA, ES, MG A, PB, PE, RJ, RR, RS, SC, SE, SP AM, BA, P AC, PE, RJ, RO, RR, SC, SP AC, AP ES, MG BA, ES, MG BA, ES, MG PR, RR, RS, SP AC, AC, AM, BA, CE, ES, MG AM, CE, P AC, BA, ES, P BA, SP PE, RJ, RR, SP AL, AM, ES, P AC, MS, MT AL, BA, ES, MG AC, P BA, PE, SP AC, BA, PE, PR, RJ, SC, SP AC, AL, AC, Distribution in Brazil Africa gentina gentina gentina America America Ar Ar Ar America Brazil and Pantropical Pantropical Pantropical Pantropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical orld distribution northern northern Africa and Africa and Brasil, Paraguai, and Brazil and W Neotropics and North Neotropics and East , , , , , , , , , , , , , canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy , sub-canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy canopy understory understory understory understory understory understory understory understory understory Forest strata understory understory understory understory understory understory understory understory understory understory understory understory understory V V V V V V V V V , L , L , L , L , L , L , L V V V V V V , L , L -- LV LV LV LB LB LB LB, L LB, L LB, L LB, L LB, L LB, L DW DW LB, DW LB, DW LB, DW Substrata LB, DW LB, DW LB, DW LB, DW LB, DW LB, DW LB, DW Locality EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici Serra Grande Serra Grande Serra Grande Serra Grande Serra Grande Serra Grande EsEc Murici and EsEc Murici and EsEc Murici and EsEc Murici and EsEc Murici and Barra de São Miguel, EsEc Murici, Maceió, EsEc Murici and Penedo Penedo, and Serra Grande EsEc Murici and Junqueiro fn. fn. eber) fn. W A. Evans A. Evans eber) Schif (Nees) Gradst. A. Evans (Lehm. & eber) Schif illd. ex (Lindenb. & (W (Pôrto & Grolle) (Gottsche ex (Spruce) (W Bischl. (Gottsche) Grolle (W (Nees) Schiffn. (Nees) Grolle (Gottsche) (Lehm. & Lindeb.) (Steph.) E. Reiner & Goda (Gottsche) Evans

eutelli A. Steph. Spruce (Nees & Mont.) Schif Nees & Motn. Nees & Mont. Lindenb. & Gottsche A. Evans A. Evans Spruce A. Evans ginuliflora oloba olejeunea br teph.) Lindenb.) S Schiffn. R.M. Schust. & Giancotti Schiffn. Gottsche) italianthus bischlerianus . macr tictolejeunea squamata axilejeunea obtusangula L. laetevirens Lopholejeunea subfusca L. immersa Microlejeunea epiphylla Neur Odontolejeunea lunulata Lejeunea grossitexta Rectolejeunea berteroana Prionolejeunea denticulata Leucolejeunea xanthocarpa P L. tapajosensis L. phyllobola R. emar Pycnolejeunea contigua Lepidolejeunea involuta Leptolejeunea elliptica L. puiggariana R. flageliformis L. magnolia S V Schiffneriolejeunea polycarpa T Symbiezidium barbiflorum axa able 1 (continuation) T T 884 Alvarenga, Oliveira, Silva, Costa & Pôrto: Liverworts of Alagoas State, Brazil , A, PR, , P , MS, MT , PE, PR, RJ, , MT , ES, MG , MS, MT A, PE, PR, RJ, SP A, PE, RJ, RO, RR, SP A, PR, RR, SP A, PE, PR, RJ, RS, SP A, PE, PR, RJ, RS, SC, SP , P , P , RJ , CE, DF , P , ES, MG AP , PB, PE, RJ, RS, SP , BA, ES, P , BA, ES, MG , PE, PR, RJ, RS, SP , RJ, RS, SC, SP A, PE, RJ, RS, SP AP AP AM, , PE, RJ, SC, SP , PE, PR, RJ, RS, SC, SP A, PE, MG AM, AM, BA, ES, P AL, AM, BA, MG AL, P AM, AL, CE, DF AM, BA, CE, ES, GO, MA, MS, PB, PE, A, PB, PE, RJ, RR, RS, SC, SP PI, RN, SE, TO ES, MG AC, AM, BA, MA, MG CE, ES, MG AL, GO, MG AC, BA, CE, ES, MG ES, PE, MG P AM, P RS, SC, SP AC, AC, MG AC, AL, AC, RJ, RS, SC, SP AL, PR, RJ, RS, SC AC, Distribution in Brazil America gentina America Ar Brazil America Pantropical Pantropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical Neotropical orld distribution Subcomsmopolitan Africa and Brazil and W Costa Rica and Brazil Neotropics and subtropical , , , Neotropics and North , sub-canopy , sub-canopy , sub-canopy canopy canopy canopy canopy canopy understory understory understory understory understory understory understory understory understory understory understory Forest strata understory understory understory V V V , L , L , L -- LV LV LV LB LB LB DW DW DW LB, DW LB, DW LB, DW LB, DW LB, DW Substrata LB, DW LB, DW Locality Arapiraca EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici EsEc Murici Serra Grande Serra Grande EsEc Murici and EsEc Murici and EsEc Murici, Marechal Deodoro and Serra Grande EsEc Murici and Serra Grande (Mont.) Howe (Nees & Mont.) (Steph.) Fulford (Lehm. & .) Nees Spruce (Lehm. & Lindenb.) Lindenb. Lindenb. & Gottsche Lindenb. Lindb. ayl. (Sw Jovet-Ast. Lindb. Nees (Nees) Lindenb. T Gottsche ex Steph. Steph. tiana ecubans X.-L. He & Grolle Lindenb.) Lindenb. . distinctifolia . gymnocalycina . mar . montagnei Bazzania heterostipa Arachniopsis diacantha Xylolejeunea crenata Metzgeria albinea M. conjugata M. dichotoma Plagiochila disticha M. myriopoda P P P P Radula flaccida Riccia vitalii R. kegelii R. ligula R. r axa able 1 (continuation) T T LEPIDOZIACEAE METZGERIACEAE PLAGIOCHILACEAE RADULACEAE RICCIACEAE Acta bot. bras. 22(3): 878-890. 2008. 885

Description: Gradstein & Costa (2003). genera for the Neotropics (pers. comm.). The only Comments: this species is characterized by well reliable distinction between them is that C. lanciloba developed lobules in most leaves, reaching 2/5 to 2/3 of has narrowly ligulate to acute lobules usually with 1-2 lobe length and with the first lobule tooth falcate, lateral teeth while C. latilobula always has broad, obtuse, consisting of two slightly elongated cells; the surface of ligulate lobules. Cololejeunea lanciloba is Pantropical lobes, lobules, and perianths is covered with conical and grows on leaves in areas with altitudes up to 50 m. papillae. The marginal cells of lobes and lobules are In Brazil it was restricted to Amazonas state, where it conically protuberant. Aphanolejeunea asperrima was recorded by the first name, Cololejeunea tonkinensis resembles A. microscopica (Tayl.) A. Evans, but the Steph. It was found growing on leaves in the understory latter has greater lobules that reach 2/3 to 4/5 of lobe at EsEc Murici. length. Aphanolejeunea asperrima has only been recorded for Patagonia and Brazil, where it has been Drepanolejeunea campanulata (Spruce) Steph., Sp. found growing on leaves at altitudes that vary from Hepat. 5: 328. 1913. 1450-2000 m. In Brazil it was previously known to the Fig. 17-21 South and Southeastern regions. It was found growing Material examined: BRAZIL. Alagoas: Murici, on leaves in the understory at EsEc Murici. 2/XII/2004, Pôrto (UFP 45826). Description: Bischler (1964). Aphanolejeunea camillii (Lehm.) R.M. Schust., Hepat. Comments: this species is characterized by the Anthocer. N. Amer. 4: 1297. 1980. presence of a conspicuous tooth 2-3 cells long on the Fig. 7-10 ventral lobe margin, positioned just above the junction Material examined: BRAZIL. Alagoas: Murici, with the keel. This species resembles Drepanolejeunea 2/XII/2004, Pôrto (UFP 45844); ib. 2/XII/2004, Pôrto lichenicola (Spruce) Steph., but the pre-apical lobule teeth (UFP 45827). of the latter are longer, curved, and usually cross the Description: Gradstein & Costa (2003). apical teeth, whereas in D. campanulata the teeth are Comments: this species is characterized by elliptic, short and inconspicuous. Drepanolejeunea campanulata rhombic, or lanceolate leaves that are wider in the middle, was previously known to Ecuador, Peru, and Southeast nearly symmetrical, and with an acute apex that ends in Brazil. It was found growing on leaves and bark in the one cell. The lobules are reduced in most leaves, but are understory, sub-canopy, and canopy at EsEc Murici. occasionally developed and can reach 1/4 of lobe length. Aphanolejeunea camillii resembles A. paucifolia Drepanolejeunea crucianella (Tayl.) A. Evans, Bull. (Spruce) E. Reiner, but it is clearly distinguished by the Torrey Bot. Club 30: 33. 1903. leaves - those of the latter are asymmetrical, wider at the Fig. 22-26 base, and with an apex that ends in a row of 3-4 cells; in Material examined: BRAZIL. Alagoas: Murici, addition, the lobules are always reduced. It is common 2/XII/2004, Pôrto (UFP 45822); ib. 2/XII/2004, Pôrto in the Neotropics, occurring from Mexico to Paraguay (UFP 45824); ib. 2/XII/2004, Pôrto (UFP 45833); ib. and northern Argentina. In Brazil it is known to occur in 2/XII/2004, Pôrto (UFP 45841); ib. 2/XII/2004, Pôrto the North and Southeast regions. It was found growing (UFP 45845). on leaves in the understory in the EsEc Murici. Description: Bischler (1964). Cololejeunea lanciloba Steph., Hedwigia 34: 250. 1895. Comments: this species can be easily recognized Fig. 11-16 due to the presence of conspicuous teeth that are 2-6 cells long and 1-3 cells wide at the base in both the Material examined: BRAZIL. Alagoas: Murici, dorsal and ventral leaf margins. This species resembles 3/XII/2004, Pôrto (UFP 45828). Drepanolejeunea palmifolia (Nees) Steph., but the latter Description: Tixier (1991), as Cololejeunea evansii has teeth only on dorsal margins. Drepanolejeunea P. Tixier. crucianella generally occurs in lowland forests, growing Comments: this species is characterized by leaves on leaves. It has a Neotropical distribution and in Brazil with a hyaline margin of rectangular cells and with plane it was previously restricted to Amazonas and Pará, in lobules parallel to the stem frequently bilobate and with the North region. It was found growing on leaves in the a rounded apex. Cololejeunea lanciloba has highly understory at EsEc Murci. variable gametophytic characters, even concerning lobule shape; the latter fact causes problems in distinguishing Harpalejeunea tridens (Besch. & Spruce) Steph., Sp. it from Cololejeunea latilobula (Herzog) Tixier, Hepat. 5: 263. 1913. according Dr. Tamás Pócs who is presently revising the Fig. 27-30 886 Alvarenga, Oliveira, Silva, Costa & Pôrto: Liverworts of Alagoas State, Brazil

Figures 1-10. 1-2. Aneura pinguis (L.) Dumort. 1. Habit, dorsal view. 2. Cross section of main axis. Figures 3-6. Aphanolejeunea asperrima Steph. 3. Habit, ventral view. 4, 6. Leaves. 5. Median cells of leaf. Figures 7-10. Aphanolejeunea camillii (Lehm.) R.M. Schust. 7. Habit, ventral view. 8, 9. Leaves. 10. Median cells of leaf. (Bars = Fig.1: 3 cm; Fig. 2: 96 µm; Fig. 3,7: 200 µm; Fig. 4,6,8,9: 48 µm; Fig 5,10: 20 Ám) Acta bot. bras. 22(3): 878-890. 2008. 887

Figures 11-21. 11-16.Cololejeunea lanciloba Steph. 11. Habit, ventral view. 12. Leaf, ventral view. 13. Marginal cells of leaf. 14, 15. Lobules. 16. Median cells of leaf. Figures 17-21. Drepanolejeunea campanulata (Spruce) Steph. 17. Habit, ventral view. 18. Leaf, dorsal view. 19. Leaf, ventral view. 20. Median cells of leaf. 21. Underleaf. (Bars = Fig. 11: 600 µm; Fig. 12: 200 µm; Fig. 17: 100 µm; Fig. 13,14,15,18,19: 48 µm; Fig. 21: 30 µm; Fig. 16,20: 20 µm). 888 Alvarenga, Oliveira, Silva, Costa & Pôrto: Liverworts of Alagoas State, Brazil

Figures 22-30. 22-26. Drepanolejeunea crucianella (Tayl.) A. Evans. 22. Habit, ventral view. 23. Leaf, ventral view. 24. Leaf, dorsal view. 25. Median cells of leaf. 26. Underleaf. Figures 27-30. Harpalejeunea tridens (Besch. & Spruce) Steph. 27. Habit, ventral view. 28. Leaf, dorsal view. 29. Detailed gametophyte, ventral view. 30. Median cells of leaf. (Bars: Fig. 22: 200 µm; Fig. 23,24,28,29: 48 µm; Fig.25,26,30: 20 µm; Fig. 27: 168 µm). Acta bot. bras. 22(3): 878-890. 2008. 889

Material examined: BRAZIL. Alagoas: Murici, Dauphin, G. 2003. Ceratolejeunea. Flora Neotropica, monograph 2/XII/2004, Pôrto (UFP 45849); ib. 25/III/2006, 90: 1-86. Evans, A. 1902. Hepaticae of Puerto Rico. III. Harpalejeunea, Alvarenga (UFP 45846); ib. 25/III/2006, Alvarenga (UFP Cyrtolejeunea, Euosmolejeunea and Trachylejeunea. Bulletin 45847); ib. 25/III/2006, Alvarenga (UFP 45848). of the Torrey Botanical Club 30: 544-563. Description: Evans (1902), as Harpalejeunea Fulford, M.H. 1963. Manual of the leafy Hepaticae of Latin America – Part I. Memoirs of The New York Botanical Garden 11: heterodonta A. Evans. 1-172. Comments: this species has leaf margins typically Fulford, M.H. 1966. Manual of the leafy Hepaticae of Latin America crenate, with an acute to piliferous apex that ends in a – Part II. Memoirs of The New York Botanical Garden 11: 173-276. row of 3-4 cells; there is also one or more additional Fulford, M.H. 1968. Manual of the leafy Hepaticae of Latin America teeth along the dorsal margin. This species was found – Part III. Memoirs of The New York Botanical Garden 11: growing on bark and leaves in lowland rain forests. In 277-392. Fulford, M.H. 1976. Manual of the leafy Hepaticae of Latin America Brazil it was previously referred only to São Paulo, in the – Part IV. Memoirs of The New York Botanical Garden 11: Southeast region. It occurred on bark and decaying wood 393-535. in the understory, sub-canopy and canopy at EsEc Murici. Ganacevich, N.A. & Mello, Z.R. 2006. Briófitas da Biquinha de Anchieta, São Vicente, São Paulo, Brasil. Boletim do Instituto de Botânica 18: 75-84. Acknowledgements Germano, S.R. & Pôrto, K.C. 2005. A Bryophyte Checklist of the Ecological Reserve of Gurjaú, Pernambuco, Brazil. Tropical Bryology 26: 1-12. The authors would like to thank the following Gradstein, S.R. & Costa, D.P. 2003. The Hepaticae and Anthocerotae specialists for help in specimen identification: Dr. Anna of Brazil. Memoirs of the New York Botanical Garden 87: Luiza Ilkiu-Borges (Museu Paraense Emílio Goeldi); Dr. 1-318. Gradstein, S.R.; Churchill, S.P. & Salazar Allen, N. 2001. Guide to the Denise Pinheiro da Costa (Jardim Botânico do Rio de Bryophytes of Tropical America. Memoirs of the New York Janeiro); Dr. Alfons Schäfer-Verwimp, Dr. Elena Reiner- Botanical Garden 86: 1-577. Drehwald, and Dr. Stephan Robert Gradstein (Göttingen Joyce, M.V.; Mello, Z R. & Yano, O. 2006. Briófitas da Ilha de Palmas, Guarujá, São Paulo, Brasil. Boletim do Instituto de University, Germany); Dr. Támas Pócs (Ezsterhazy Botânica 18: 101-109. Teachers’ College, Hungary); and Dr. Olga Yano (Instituto Kron, K.A. 1988. Taxonomic study of Venezuelan members of de Botânica de São Paulo). We would also like to thank Frullania Raddi subg. Meteoriopsis Spruce. The Journal of the Hattori Botanical Laboratory 64: 347-358. the Fundação O Boticário de Proteção à Natureza and Molinaro, L.C. & Costa, D.P. 2001. Briófitas do arboreto do Jardim CNPq for financial support. Botânico do Rio de Janeiro. Revista do Jardim Botânico do Rio de Janeiro 52: 107-124. Moura, F.B.P. 2006. A Mata Atlântica em Alagoas. Maceió, Editora References da UFAL. Oliveira, J.R.M.; Alvarenga, L.D.P. & Pôrto, K.C. 2006. Briófitas da Bastos, C.J.P. & Yano, O. 2004. New records of Lejeuneaceae Estação Ecológica de Águas Emendadas, Distrito Federal, material (Marchantiophyta) for the Brazil. Acta Botanica Malacitana coletado por Daniel Moreira Vital. Boletim do Instituto de 29: 13-21. Botânica 18: 181-195. Bastos, C.J.P. & Yano, O. 2006. Briófitas de restinga das regiões Paixão, R.C. & Mello, Z.P. 2006. Brioflora ao longo da Trilha do Metropolitana de Salvador e Litoral Norte do Estado da Bahia, Costão, Peruíbe, São Paulo, Brasil. Boletim do Instituto de Brasil. Boletim do Instituto de Botânica 18: 198-205. Botânica 18: 85-90. Bischler, H. 1964. Le genre Drepanolejeunea Steph. en Amérique Patrus, P. & Starling, M.F.V. 2006. Briófitas da Serra do Curral, Belo Centrale et Méridionale. Revue Briologique et Lichénoloque Horizonte, Minas Gerais, Brasil. Boletim do Instituto de 33: 15-133. Botânica 18: 123-129. Brasil - MMA. 2002. Avaliação e identificação de áreas e ações Pôrto, K.C. & Germano, S.R. 2002. Biodiversidade e importância das prioritárias para a conservação, utilização sustentável e briófitas na conservação dos ecossistemas naturais de Pernambuco. repartição dos benefícios da biodiversidade nos biomas Pp. 125-152. In: M. Tabarelli & J.M.C. Silva (eds.). Diagnóstico brasileiros. Brasília, Ministério do Meio Ambiente/SBF. da Biodiversidade de Pernambuco. SECTMA - Secretaria de Câmara, P.E.A.S. & Vital, D.M. 2004. Briófitas do Municipio de Ciência, Tecnologia e Meio Ambiente, Recife. Poconé, Pantanal de Mato Grosso, MT, Brasil. 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The Hepaticae and Anthocerotae of North state, Brazil. The Journal of the Hattori Botanical America. Chicago, Field Museum of Natural History, v. 5. Laboratory 98: 259-298. Stotler, R.E. 1970. The Genus Frullania in Latin America. Nova Crandall-Stotler, B. & Stotler, R.E. 2000. Morphology and Hedwigia 18: 397-555. classification of the Marchantiophyta. Pp. 21-70. In: A.J. Shaw Tixier, P. 1991. Bryophyta exótica – 9. Quelques Lejeuneaceae & B. Goffinet (orgs.) Bryophyte Biology. Cambridge, Cambridge (Hepatiques) nouvelles pour l’Amérique du Sud. Candollea 46: University Press. 267-302. 890 Alvarenga, Oliveira, Silva, Costa & Pôrto: Liverworts of Alagoas State, Brazil

Valente, E.B. & Pôrto, K.C. 2006a. Briófitas do afloramento rochoso Yano, O. 2004. Novas ocorrências de briófitas para vários estados do na Serra da Jiboia, municipio de Santa Teresinha, Bahia, Brasil. Brasil. Acta Amazonica 34: 559-576. Boletim do Instituto de Botânica 18: 207-211. Yano, O. 2005. Adição às Briófitas da Reserva Natural da Vale do Rio Valente, E.B. & Pôrto, K.C. 2006b. Novas ocorrências de hepáticas Doce, Linhares, Espírito Santo, Brasil. Boletim do Museu de (Marchantiophyta) para o Estado da Bahia, Brasil. Acta Botanica Biologia Mello Leitão, Nova Série, 18: 5-48. Brasilica 20: 195-201. Yano, O. 2006. Novas adições as briófitas brasileiras. Boletim do Veloso, H.P.; Rangel Filho, A.L.R. & Lima, J.C.A. 1991. Instituto de Botanica 18: 229-233. Classificação da vegetação brasileira adaptada a um Yano, O. & Costa, D.P. 2000. Flora dos Estados de Goiás e Tocantins: sistema universal. Rio de Janeiro, Instituto Brasileiro de Criptógamos: Briófitas. v.5. Goiânia, Editora da UFG. Geografia e Estatística. Yano, O. & Câmara, P.E. 2004. Briófitas de Manaus, Amazonas, Vilas Bôas-Bastos, S.B.; Bastos, C.J.P. & Ballejos, J. 2006. Novas Brasil. Acta Amazonica 34: 445-457. ocorrências de hepáticas (Marchantiophyta) para o Estado da Yano, O. & Bordin, J. 2006. Novas ocorrências de briófitas para o Rio Bahia, Brasil. Boletim do Instituto de Botanica 18: 142-147. Grande do Sul, Brasil. Boletim do Instituto de Botânica 18: Yano, O. 1981. A checklist of Brazilian mosses. The Journal of the 111-122. Hattori Botanical Laboratory 50: 279-456. Yano, O. & Peralta, D.F. 2005. Hepáticas (Marchantiophyta) de Yano, O. 1984. Checklist of Brazilian liverworts and hornworts. The Mato Grosso, Brasil. Hoehnea 32: 185-205. Journal of the Hattori Botanical Laboratory 56: 481-548. Yano, O. & Peralta, D.F. 2006. Briófitas coletadas por Daniel Moreira Yano, O. 1989. An additional checklist of Brazilian bryophytes. The Vital no Estado da Bahia, Brasil. Boletim do Instituto de Journal of the Hattori Botanical Laboratory 66: 371-434. Botânica 18: 33-73. Yano, O. 1995. A new additional annotated checklist of brazilian Yano, O. & Peralta, D.F. 2007. Novas ocorrências de briófitas para os bryophytes. The Journal of the Hattori Botanical Laboratory Estados de Alagoas e Sergipe, Brasil. Arquivos do Museu 78: 137-182. Nacional 64: 287-297.

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