Five species of Lejeuneaceae, tribe Lejeuneae from state of Bahia, Brazil are described and illustrated. Data on geographic distributions, distribution in Brazil and habitat in Bahia are also given

Bastos, Cid José Passos

Bôas-Bastos, Silvana Brito Vilas

English

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45TROPICAL BRYOLOGY 19 (2000)Occurrence of some Lejeuneaceae in Bahia-Tropical Bryology 20: 45-54, 2000Occurrence of some Lejeuneaceae (Jungermanniophyta) in Bahia,BrazilCid José Passos Bastos & Silvana Brito Vilas Bôas-BastosUniversidade Federal da Bahia, Instituto de Biologia, Departamento de Botânica, Laboratóriode Taxonomia de Algas e Briófitas. Campus de Ondina, 40170-280 Salvador, Bahia, Brasil.E-mail: cjpbasto@ufba.br   -  svbbastos@hotmail.comAbstract. Five species of Lejeuneaceae, tribe Lejeuneae from state of Bahia, Brazil are describedand illustrated. Data on geographic distributions, distribution in Brazil and habitat in Bahia are alsogiven.INTRODUCTIONThe Lejeuneaceae in Brazil arerepresented by ca. 63 genera and 465 species(Yano 1996) and in Bahia by ca. 97 species intoca. 35 genera (data in part not published). In Bahiathe members of Lejeuneaceae grow in a greatdiversity of habitats, such as “campos rupestres”,“caatinga”, “cerrado”, ombrophilous forest,seasonal forest, mangroves and urban zones.Most species are generalists, growing on varioussubstrates.In Brazil the Lejeuneaceae are poorlystudied taxonomically in spite of a great numberof records. The most important study forneotropics, including Brazil, was by Gradstein(1994), but only for Ptychanthoideae and part ofLejeuneoideae (tribe Brachiolejeuneae). In Bahiawe have begun a study of the Lejeuneaceae andresults have been very interesting. The membersof Lejeuneaceae found in Bahia belong to foursubfamilies according to the concept acceptedhere: Bryopteridoideae, Ptychanthoideae,Lejeuneoideae and Cololejeuneoideae. In ouropinion, these four evolutionary trends are welldefined in Lejeuneaceae. The Lejeuneoideae arethe best represented in numbers of species. Inthis paper five species belonging to theLejeuneoideae, tribe Lejeuneae, are described andillustrated: Cyrtolejeunea holostipa (Spruce)Evans, Harpalejeunea uncinata Steph.,Leptolejeunea exocellata (Spruce) Evans,Leucolejeunea conchifolia (Evans) Evans andLeucolejeunea xanthocarpa (Lehm. & Lindenb.)Evans.46TROPICAL BRYOLOGY 19 (2000)Bastos & BastosMATERIALS AND METHODSSpecies deposited in Alexandre LealCosta Herbarium (ALCB) collected in “EstaçãoVeracruz”, a private ecological reserve belongingto “Veracel Celulose S.A.”, in the state of Bahia,Brazil, were studied. The “Estação Veracruz” isprimary Atlantic forest fragment situated insouthern Bahia and occupies ca. 6,000 habetween Porto Seguro and Santa Cruz de Cabráliacounties. The systematic arrangement followsSchuster (1980) (in part).RESULTSCyrtolejeunea holostipa (Spruce) Evans, Bull.Torrey Bot. Club 30: 553. 1903.Basionym: Lejeunea holostipa Spruce, Trans.Proc. Bot. Soc. Edinburgh 15: 171. 1884.Type-locality: Venezuela.(Figure 1).Gametophyte small, prostrate, branchesLejeunea-type. Stem in cross section with 7-8cortical cells and 3 medullary cells. leaves erect-spreading, contiguous to imbricate; lobe ovate,238.5-265mm long x 132.5-153.7mm wide,margin entire, apex rounded; cells oblong torounded, 14.5-23.7mm x 11.9-17.2mm, thick-walled, with large trigones and inconspicuousintermediate thickenings; oil-bodies not see;ocelli absent; lobule ovate, 106mm long x 74.2mmwide, strongly inflated, free margin plane, keelstrongly arched, gibbous, apical tooth long,hyaline papilla not seen. Underleaves small,85.8mm wide, unlobed, distant, orbicular. Ventralmerophytes two cells wide. Androecia on longlateral branches, bracts in 3 pairs, bracteoles likeunderleaves, present throughout. Gynoecia onshort lateral branches, bracts with lobes obovate,margin entire, bracteole large, obovate, apexemarginate; perianth not found.Specimens examined: Brazil, Bahia: “EstaçãoVeracru” Ecological Reserve, between Eunápolisand Porto Seguro, ombrophilous forest, domainof Tropical Atlantic Rainforest, in “mussununga”vegetation, 10/6/1999, S.B. Vilas Bôas-Bastos &C. Bastos 681 (ALCB 41881).Geographic distribution: Tropical America.Distribution in Brazil: Espírito Santo (Schäfer-Verwimp 1991), Pernambuco (Pôrto 1990), Riode Janeiro (Spruce 1888), São Paulo (Visnadi1998).Substrate: tree trunk.Leucolejeunea conchifolia (Evans) Evans,Torreya 7: 229. 1907.Basionym: Archilejeunea conchifolia Evans,Mem. Torrey Bot. Club 8(2): 128. 1902.Type-locality: Orange Band.(Figure 2).Gametophyte robust, 1.4mm wide, green,prostrate, branches Lejeunea-type. Stem in crosssection with 14 cortical cells and 21 medullarycells. Leaves imbricate, spreading; lobe ovate,0.8-0.9mm long x 0.6mm wide, margin entire,apex obtuse to rounded, falcate; cells oblong, 25-35.6mm long x 21,1-22.4mm wide, thin-walled,trigones small to inconspicuous, intermediatethickenings absent; oil-bodies not seen; ocelliabsent; lobule very large, 392.2-450.5mm long x206.7-185.5mm wide, oblong-ovate, inflated, freemargin arched, keel weakly convex or straight,apical tooth long, falcate, hyaline papilla not seen.Underleaves unlobed, 0.5-0.6mm wide,contiguous to imbricate, rotundate. ventralmerophytes two cells wide. Androecia not found.Gynoecia on short lateral branches, bractsoblong-ovate, margin entire, bracteole unlobed,oblong; perianth obovate, 5-keeled.Specimens examined: Brazil, Bahia: “EstaçãoVeracruz” Ecological Reserve, betweenEunápolis and Porto Seguro, ombrophilousforest, domain of Tropical Atlantic Rainforest,in “mussununga” vegetation, 10/6/1999, S.B.Vilas Bôas-Bastos & C. Bastos 627, 629, 630,636, 637, 658, 676, 689, 717 (ALCB 41826,42828, 41829, 41835, 41836, 41858, 41876,41889, 41917).Geographic distribution: United States ofAmerica and Brazil.Distribution in Brazil: Espírito Santo (Schäfer-Verwimp 1991), São Paulo (Visnadi 1998).Substrate: tree trunk.Leucolejeunea xanthocarpa (Lehm. & Lindenb.)Evans, Torreya 7: 229. 1907.Basionym: Jungermannia xanthocarpa Lehm. &Lindenb. in Lehmann, Nov. Min. Cogn. Stirp.47TROPICAL BRYOLOGY 19 (2000)Occurrence of some Lejeuneaceae in Bahia-Figure 1. Cyrtolejeunea holostipa (Spruce) Evans. A) gametophyte with gynoecia, ventral view; B)lobule; C) apical tooth: D) leaf, dorsal view; E) laminal cells; F) gametophyte with androecia; G)transverse section of the stem.48TROPICAL BRYOLOGY 19 (2000)Bastos & BastosFigure 2. Leucolejeunea conchifolia (Evans) Evans. A) gametophyte, ventral view; B) leaf; C) leafapex; D) underleaves; E) apical teeth; F) transverse section of the stem; G) laminal cells.49TROPICAL BRYOLOGY 19 (2000)Occurrence of some Lejeuneaceae in Bahia-Pug. 5: 8. 1833.Type-locality: Brazil.(Figure 3).Gametophyte robust, 1.34mm wide,prostrate, branches Lejeunea-type. Stem in crosssections with 14 cortical cells and 21 medullarycells. Leaves imbricate, spreading; lobe oblong-ovate, 0.5mm long x 0.3mm wide, marginrevolute, apex rounded; cells oblong, 18.9-22mmlong x 16.8-17.6 wide, thick-walled, trigoneslarge, sometimes with intermediate thickenings;oil-bodies not seen; ocelli absent; lobule ovate-rectangular, 243.6-285.6mm long x 117.6-138.6mm wide, free margin involute for its entirelength, keel straight, apical tooth short, hyalinepapilla not seen. Underleaves unlobed, large,0.5mm wide, imbricate, reniform. Ventralmerophytes two cells wide. Androecia on longlateral branch, bracts in 7-8 pairs, bracteole onbase. Gynoecia on short lateral branch, bractsoblongs, margin revolute, entire, bracteoleoblong, strongly revolute; perianth obovate,rostrum long.Specimens examined: Brazil, Bahia: “EstaçãoVeracruz” Ecological Reserve, betweenEunápolis and Porto Seguro, ombrophilousforest, domain of Tropical Atlantic Rainforest,in “mussununga” vegetation, 10/6/1999, S.B.Vilas Bôas-Bastos & C. Bastos 628, 643, 648,686, 707, 701, 709, 719, 720 (ALCB 41827,41842, 41847, 41886, 41907, 41901, 41909,41919, 41920).Geographic distribution: Pantropical.Distribution in Brazil: Espírito Santo (Schäfer-Verwimp 1991), Pernambuco (Pôrto 1990), Riode Janeiro (Gottsche et al. 1844, Costa 1992,1999), Santa Catarina (Thiers 1982), São Paulo(Herzog 1927, Schiffner & Arnell 1964, Giancotti& Vital 1989, Visnadi et al. 1994, Vital & Visnadi1994, Visnadi 1998).Substrate: tree trunk.Harpalejeunea uncinata Steph. Hedwigia 35: 97.1896.Type-locality: Luqillo Montains, Puerto Rico.(Figure 4).Gametophyte small, 0.6mm wide, green,prostrate, branches Lejeunea-type. Stem in crosssection with 7 cortical cells and 4 medullary cells.Leaves contiguous to moderately imbricate; lobeovate, 307.4-371mm long x 180.2-206.7mmwide, falcate, margin entire, apex apiculate tocuspidate, formed by 4-6 cells; cells haxagonalto oblong, 15.8-27.7mm long x 18.4mm wide,thin-walled, trigones small, intermediatethickenings inconspicuous; oil-bodies not seen;ocelli 1-2, basal, large; lobule ovate, 132.5mmlong x 100.7mm wide, strongly inflated, freemargin involute, keel strongly arched, apicaltooth long, hyaline papilla on base of apical tooth.Underleaves small, 106mm wide, cordate, distant,sinus acute to lunate, lobes 4 cells wide. Ventralmerophytes two cells wide. Androecia andgynoecia not found.Specimens examined: Brazil, Bahia: “EstaçãoVeracruz” Ecological Reserve, betweenEunápolis and Porto Seguro, ombrophilousforest, domain of Tropical Atlantic Rainforest,S.B. Vilas Bôas-Bastos & C. Bastos 778 (ALCB41978).Geographic distribution: CaribbeanDistribution in Brazil: Rio de Janeiro (Oliveira eSilva & Yano 1998).Substrate: tree trunk.Leptolejeunea exocellata (Spruce) Evans, Bull.Torrey Bot. Club 29: 498. 1902.Basionym: Lejeunea exocellata Spruce, Trans.Proc. Bot. Soc. Edinburgh 15: 195. 1884.Type-locality: Peru(Figure 5).Gametophyte small, prostrate, branchesLejeunea-type. Leaves distant to contiguous,erect-spreading; lobe ovate, 344.5mm long x185.5mm wide, margin crenate, apex rounded toobtuse, formed by 1-2 cells; cells hexagonal tooblong, 19.8-25mm long x 15.8-17.1mm wide,thick-walled, trigones small, intermediatethickenings nodulose; oil-bodies small; ocellusbasal, large, partially hidden by lobule; lobuleovate to ovate-rectangular, inflated, 185.5mmlong x 132.5mm wide, free margin plane, keelmoderately arched, crenate, apical tooth short,hyaline papilla not seen. Underleaves small,81.8mm wide, distant, lobes filiform, formed ofa row of 2-3 cells. Ventral merophytes two cellswide. Androecia and gynoecia not seen.50TROPICAL BRYOLOGY 19 (2000)Bastos & BastosFigure 3. Leucolejeunea xanthocarpa (Lehm. & Lindenb.) Evans. A) gametophyte, ventral view; B)leaf; C) underleaves; D) lobule apices; E) laminal cells; F) gynoecia; G) gynoecial bracts; H) androecia;I) transverse section of the stem.51TROPICAL BRYOLOGY 19 (2000)Occurrence of some Lejeuneaceae in Bahia-Figure 4. Harpalejeunea uncinata Steph. A) gametophyte, ventral view; B) lobule; C) underleaf; D)leaves with ocelli, dorsal view; E) lobe apices; F) laminal cells; G) transverse section of the stem.52TROPICAL BRYOLOGY 19 (2000)Bastos & BastosFigure 5. Leptolejeunea exocellata (Spruce) Evans. A) gametophyte, ventral view; B) lobule; C)leaf with ocellus, dorsal view; D) lobe apices; E) laminal cells.53TROPICAL BRYOLOGY 19 (2000)Occurrence of some Lejeuneaceae in Bahia-Specimens examined: Brazil, Bahia: “EstaçãoVeracruz” Ecological Reserve, betweenEunápolis and Porto Seguro, ombrophilousforest, domain of Tropical Atlantic Rainforest,28/10/1999, S.B. Vilas Bôas-Bastos & C. Bastos2035, 2036 (ALCB 44602, 44603).Geographic distribution: Neotropical.Distribution in Brazil: Acre, Amazonas, Paraná,Rio de Janeiro, Santa Catarina (Bischler 1969),São Paulo (Schiffner & Arnell 1964).Substrate: tree trunk.The material examined was insufficientfor the preparation of a transverse section of thestem, and the underleaves were inadequate fordrawing. According to Reiner-Drehwald (1995),L. exocellata has 7 cortical cells and 3 medullarycells in the stem. The underleaves are distant, withfiliform lobes (3-4 cells long and two cells wide).DISCUSSIONThe species studied were collected insouthern Bahia in ombrophilous forest, domainof Atlantic Forest, a vegetation type occurringon the Brazilian coast with characteristics of thetropical rain forest. The Atlantic Forest ofsouthern Bahia has certain floristic similaritieswith Amazonian forests. It is true for bothbryophytes and vascular plants, and suggestsancient links between Amazonian and Atlanticforests, as was advocated by Andrade-Lima(1966), who found 388 Amazonian speciesranging southern Bahia to northern Espírito Santo(fide Rizzini 1979).Most of the species studied in this paperare restricted to southeastern and northeasternBrazil, and only Leptolejeunea exocellatarepresents an Amazonian element. Leucolejeuneaconchifolia and Harpalejeunea uncinata have amore restricted distribution in Brazil, and theformer has not been reported for other SouthAmerican countries, being reported only from thecoastal plain of southeastern USA by Schuster(1980). According to Delgadillo (1995) thisdisjunctive pattern also occurs in severalneotropical mosses, whose distribution includesNorth America and Brazil, and is not easilyexplained by a range dissection hypothesis. Thelack of adequate collecting may be the source ofthese patterns. Leucolejeunea xanthocarpa is aPantropical species, occurring in Asia(Indomalaya to China and Japan) and central andsouthern Africa, and widespread in the NewWorld (fide Schuster 1980), with largerdistribution in South America. Cyrtolejeuneaholostipa, Leucolejeunea conchifolia andleucolejeunea xanthocarpa in the collecting sitewere found in “mussununga” vegetation, whichis a similar vegetation to “restinga” (coastal shrubwith halophyte and xeromorphic vegetation) intoombrophilous forest. In other areas out side ofBahia these species were found mainly in Atlanticforest areas.The bryophyte flora of the southern Bahiais somewhat similar that of the southeasternBrazil, but with some Amazonian elements. It islikely due to floristic reasons, since the Atlanticforest ranges from northeastern (Pernambucostate) to southeastern (Santa Catarina state)Brazil, although extensive areas have alreadybeen destroyed, and according to Costa (1999)of the original areas (ca. 1.6 million km2) only 1-6% persists as isolated fragments, and theirbryophyte flora has been less studied than thevascular plant flora. Nevertheless, climatic causesand distribution patterns must be undervalued.There are several approaches to explaindifferent disjunctive patterns among bryophytes,including step-by-step migrations, geologicalevents, climatic changes and long range dispersal,as discussed by various authors (see Gradstein1983 and Delgadillo 1995). However, theapparent disjunct distribution may be representlack of complete floristic catalogues for theneotropical area, mainly for Lejeuneoideae. TheBrazilian bryophyte flora is in need of moreextensive fieldwork as well as monographictreatments of many genera or families (as pointedout by Schäfer-Verwimp 1996) in order toexplain the real geographic distribution patterns.AcknowledgmentsThe authors are grateful to Veracel CeluloseS.A. for promoting access to the area and for supportto the collectors.54TROPICAL BRYOLOGY 19 (2000)Bastos & BastosLITERATURE CITEDAndrade-Lima, D. 1966. Contribuição ao estudo doparalelismo da flora amazonico-nordestina.Instituto de Pesquisas Agronômicas, BoletimTécnico 19: 3-30.Bischler, H. 1969. Le genre Leptolejeunea (Spruce)Steph. en Amérique. Nova Hedwigia 17:265-350.Costa, D.P. 1992. Hepáticas do Pico da Caledônia,Nova Friburgo, Rio de Janeiro, Brasil. ActaBotanica Brasilica 6(1): 3-39.Costa, D.P. 1999. Epiphytic bryophyte diversity inprimary and secondary lowland rainforestsin southeastern Brazil. The Bryologist102(2): 320-326.Delgadillo, C.M. 1995. Neotropical moss floras:species common to North and SouthAmerica. Tropical Bryology 10: 1-6.Giancotti, C. & Vital, D.M. 1989. Flora briofítica daReserva Biológica do Alto da Serra deParanapiacaba, São Paulo. I. Lejeuneaceae(Hepaticopsida). Acta Botanica Brasilica 3:169-177.Gottsche, C.M., Lindenberg, J.B.G. & Nees vonEsenbeck, C.G. 1844. SynopsisHepaticarum. J.A. Meissmer Ec., Hambur-ger.Gradstein, S.R. 1994. Lejeuneaceae: Ptychantheae,Brachiolejeuneae. Flora Neotropica,Monograph 62: 1-216.Gradstein, S.R., Pócs, T. & Vána, J. 1983. DisjunctHepaticae in Tropical America and Africa.Acta Botanica Hungarica 29(1-4): 127-171.Herzog, T. 1927. Zwei Bryophyten sammlungen ausSüdamerika. Hedwigia 67: 249-268.Oliveira e Silva, M.I.M.N. & Yano, O. 1998.Ocorrências novas de briófitas para o Brasil.Revista Brasileira de Botânica 21(2): 125-134.Pôrto, K.C. 1990. Bryoflore d’une forêt de plain etd’une fôret d’altitude moyene dans l’etat dePernambuco (Brésil): analyse floristic.Cryptogamie, Bryologie-Lichénologie11(2): 109-161.Reiner-Drehwald, M.E. 1995. Las Lejeuneaceae(Hepaticae) de Misiones, Argentina III.Drepanolejeunea  y Leptolejeunea. Tropi-cal Bryology 10: 21-27.Rizzini, C.T. 1979. Tratado de Fitogeografia do Brasil.Vol. 2. HUCITEC, Ed. da Universidade deSão Paulo, São Paulo.Schäfer-Verwimp, A. 1991. Contribution toknowledge of the bryophyte flora of EspíritoSanto, Brazil. Journal of the HattoriBotanical Laboratory 69: 147-170.Schäfer-Verwimp, A. 1992. New or intersting recordsof Brazilian bryophytes, V. Candollea 51(2):283-302.Schiffner, V. & Arnell, S. 1964. Ergebnisse der Bo-tanischen Expedition der kaiserlichen Aka-demie der Wissenschaften nach Südbrasilien1901. Österreichischen Akademie Wissen-schaften Mathematisch-Naturwissenschaft-liche Klass, Denkschriften 111: 1-156.Schuster, R.M. 1980. The Hepaticae andAnthocerotae of North America east of theHundredth Meridian, vol. IV. ColumbiaUniversity Press, New York.Spruce, R. 1888. Hepaticae in prov. Rio de Janeiro aGlaziou lectae, a R. Spruce determinatae.Revue Bryologique 15: 33-34.Thiers, B.M. 1982. Type studies in the LejeuneaceaeI. Archilejeunea pabstii, Hygrolejeuneapacifica and Hygrolejeunea spongia.Brittonia 34(2): 294-298.Visandi, S.R. 1998. Briófitas em ecossistemascosteiros do Núcleo Picinguaba do ParqueEstadual da Serra do Mar, Ubatuba - SP.Tese de Doutorado, Universidade EstadualPaulista, São Paulo.Visnadi, S.R., Matheus, D.R. & Vital, D.M. 1994.Occurrence of bryophytes in areas pollutedwith orhganopollutants and on nearbyvegetation, preliminary notes. Journal of theHattori Botanical Laboratory 77: 315-323.Vital, D.M. & Visnadi, S.R. 1994. Briófitas de umtrecho de restinga da Estação Ecológica daJuréia, Peruíbe, estado de São Paulo, Brasil.Anais do III Simpósio de Ecossistemas daCosta Brasileira, ACIESP, Serra Negra, SãoPaulo 3: 153-157.Yano, O. 1996. A checklist of the Brazilianbryophytes. Boletim do Instituto de Botânica10: 47-232.

A checklist of the Brazilian bryophytes.

Briófitas de um trecho de restinga da Estação Ecológica da Juréia, Peruíbe, estado de São Paulo,

Briófitas em ecossistemas costeiros do Núcleo Picinguaba do Parque Estadual da Serra do Mar, Ubatuba - SP. Tese de Doutorado, Universidade Estadual Paulista, São Paulo.

Bryoflore d’une forêt de plain et d’une fôret d’altitude moyene dans l’etat de Pernambuco (Brésil): analyse floristic.

Contribuição ao estudo do paralelismo da flora amazonico-nordestina. Instituto de Pesquisas Agronômicas,

Contribution to knowledge of the bryophyte flora of Espírito Santo,

Epiphytic bryophyte diversity in primary and secondary lowland rainforests in southeastern Brazil.

Ergebnisse der Botanischen Expedition der kaiserlichen Akademie der Wissenschaften nach Südbrasilien

Flora briofítica da Reserva Biológica do Alto da Serra de Paranapiacaba, São Paulo.

Hepaticae in prov. Rio de Janeiro a Glaziou lectae, a R. Spruce determinatae.

Hepáticas do Pico da Caledônia, Nova Friburgo,

Las Lejeuneaceae (Hepaticae) de Misiones, Argentina III. Drepanolejeunea y Leptolejeunea.

Le genre Leptolejeunea (Spruce) Steph. en Amérique.

Neotropical moss floras: species common to North and South America.

New or intersting records of Brazilian bryophytes,

Occurrence of bryophytes in areas polluted with orhganopollutants and on nearby vegetation, preliminary notes.

Ocorrências novas de briófitas para o Brasil.

The Hepaticae and Anthocerotae of

Tratado de Fitogeografia do Brasil.

Type studies in the Lejeuneaceae I. Archilejeunea pabstii, Hygrolejeunea pacifica and Hygrolejeunea spongia.

Zwei Bryophyten sammlungen aus Südamerika.

Occurrence of some Lejeuneaceae (Jungermanniophyta) in Bahia, Brazil

Occurrence of some Lejeuneaceae (Jungermanniophyta) in Bahia, Brazil

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