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Bryophyte flora of the Apodi Plateau, Ceará, Brazil

Abstract

In the state of Ceará, bryophytes have been mainly sampled in humid and sub-humid enclaves, remnants of Atlantic rain forest, while studies in the Caatinga Domain are practically non-existent. The present work aimed to survey the floristic composition of bryophytes of the Apodi Plateau, a region predominantly covered by Caatinga. Collections were conducted as part of the field activities of the Program for Research in the Biodiversity of the Semiarid region of Brazil (PPBio). The briological material was collected in 2014, in the Apodi Plateau located within the territory of Ceará state. Fifty-nine bryophyte species were found: 27 liverworts - Marchantiophyta - distributed in seven families and 11 genera, and 32 mosses - Bryophyta - distributed in 14 families and 26 genera. Among the species found, 25 were new records for the state of Ceará and Riccia subplana is reported for the first time for the Northeast region of Brazil. Twenty-three are new records for the Caatinga Domain. Taxonomic comments are provided for the new records for the state of Ceará and for Caatinga, as well as an illustration of the species Weisiopsis bahiensis.

Key words:
Bryophyta; Marchantiophyta; taxonomy

Resumo

No estado do Ceará, as principais áreas amostradas com relação às briófitas são constituídas de encraves úmidos e subúmidos, remanescentes da Mata Atlântica. Estudos em áreas de Caatinga são praticamente inexistentes. O presente trabalho objetivou realizar um inventário florístico das briófitas da Chapada do Apodi, dentro dos limites do território do estado do Ceará, uma região com predominância do Domínio Caatinga. As coletas fizeram parte das atividades de campo do Programa de Pesquisa em Biodiversidade do Semiárido (PPBio). O material briológico foi coletado em 2014, na área da Chapada do Apodi localizada dentro do território do estado do Ceará. Foram encontradas 59 espécies de briófitas sendo 27 hepáticas - Marchantiophyta - distribuídas em sete famílias e 11 gêneros e 32 espécies de musgos - Bryophyta - distribuídas em 14 famílias e 26 gêneros. Das espécies encontradas, 25 se configuram como novos registros para o estado do Ceará e Riccia subplana está sendo citada pela primeira vez para a Região Nordeste. Estão sendo acrescentados ainda 23 novas ocorrências para a Caatinga. São fornecidos comentários taxonômicos para as novas ocorrências para o estado do Ceará e para a Caatinga. A espécie Weisiopsis bahiensis é ilustrada.

Palavras-chave:
Bryophyta; Marchantiophyta; taxonomia

Introduction

The northeastern region of Brazil occupies an area of ​​1,540,827 km2 (Nimer 1989Nimer E (1989) Climatologia do Brasil. 2ª ed. IBGE-SUPREN (Fundação IBGE-SUPREN, Recursos Naturais e Meio Ambiente) , Rio de Janeiro. 421p.), and has a semiarid climate in approximately 800,000 km2, corresponding to 10% of the Brazilian territory (Ab’Sáber 1974Ab’Sáber AN (1974) O domínio morfoclimático semi-árido das caatingas brasileiras. Série Geomorfologia. Vol. 43. Instituto de Geografia/USP, São Paulo. Pp. 1-39.). The relief has altitudes mostly below 500 m, among surfaces that reach 1,000 m in the Borborema, Araripe and Ibiapaba plateaus, up to 1,200 m in the Diamantina plateau. This altitudinal variation associated with the position in the relief, where the increasing altitude causes forced convective rains and lower temperatures, leads to the occurrence of different types of vegetation. The xerophilous Caatinga vegetation is dominant in the semiarid area, presenting physiognomic and floristic variations (Romariz 1974Romariz DA (1974) Aspectos da vegetação do Brasil. IBGE, Rio de Janeiro. 126p.; Ferri 1980Ferri MG (1980) Vegetação brasileira. Editora Itatiaia/ EDUSP, São Paulo. 157p.; Andrade-Lima 1981Andrade-Lima D (1981) The caatingas dominium. Revista Brasileira de Botânica 4: 149-163. ; Sampaio 1995Sampaio EVSB (1995) Overview of the Brazilian caatinga. In: Bullock SH, Mooney HA & Medina E (eds.) Seasonally dry tropical forests. University Press, Cambridge. Pp. 35-63.; Rizzini 1997Rizzini CT (1997) Tratado de fitogeografia do Brasil. 2a ed. Âmbito Cultural, Rio de Janeiro. 747p.).

The different physiognomies of the Caatinga hold a diversified biological heritage, with rare and endemic taxa (Giulietti et al. 2004Giulietti AM, Neta ALB, Castro AAJF, Gamarra-Rojas CFL, Sampaio EVSB, Virgínio JF, Queiroz LP , Figueiredo MA, Rodal MJN, Barbosa MR V & Harley RM (2004) Diagnóstico da vegetação nativa do bioma Caatinga. In: Silva JMC , Tabarelli M & Fonseca MT (orgs.) Biodiversidade da Caatinga: áreas e ações prioritárias para a conservação. Ministério do Meio Ambiente, Brasília. Pp. 1-44.) and an invaluable richness of plant and animal species (MMA 2004MMA - Ministério do Meio Ambiente (2004) Biodiversidade da caatinga: áreas e ações prioritárias para a conservação. Ministério do Meio Ambiente, Universidade Federal de Pernambuco, Brasília. 382p.). In contrast to other semiarid regions of the world, the biological diversity of the Caatinga is extremely significant (Luetzelburg 1974Luetzelburg PV (1974) Estudo botânico do Nordeste. (IFCS, Série I.A. Publicação, 57, v. 3.). Inspetoria federal de obras contra as secas, Rio de Janeiro. 283p. ; Andrade-Lima 1981Andrade-Lima D (1981) The caatingas dominium. Revista Brasileira de Botânica 4: 149-163. ; Araújo Filho & Carvalho 1997Araújo Filho JA & Carvalho FC (1997) Desenvolvimento sustentado da caatinga. Embrapa-CNPC Circular Técnica, 13. Embrapa-CNPC, Sobral. 19p.) and adds biological and economic value to Brazil. Despite such remarkable characteristics, the Caatinga is one of the least known domains in the country. This situation is a consequence of the belief that the Domain has a very low diversity, no endemic species, and vegetation that is strongly modified by anthropic actions (Giulietti et al. 2004Giulietti AM, Neta ALB, Castro AAJF, Gamarra-Rojas CFL, Sampaio EVSB, Virgínio JF, Queiroz LP , Figueiredo MA, Rodal MJN, Barbosa MR V & Harley RM (2004) Diagnóstico da vegetação nativa do bioma Caatinga. In: Silva JMC , Tabarelli M & Fonseca MT (orgs.) Biodiversidade da Caatinga: áreas e ações prioritárias para a conservação. Ministério do Meio Ambiente, Brasília. Pp. 1-44.). It is noteworthy that this vegetation is one of the most affected by human interference in Brazil; in 2008, approximately 45% of its original coverage had already been deforested (MMA 2010MMA - Ministério do Meio Ambiente (2010) Monitoramento dos biomas brasileiros: bioma Caatinga. MMA, Brasília. 46p.). According to Castelletti et al. (2004)Castelletti CHM, Silva JMC, Tabarelli M & Santos AMM (2004) Quanto ainda resta da Caatinga? Uma estimativa preliminar. In: Silva JMC, Tabarelli M & Fonseca MT (orgs.) Biodiversidade da Caatinga: áreas e ações prioritárias para a conservação. Ministério do Meio Ambiente, Brasília. 440p., the last intact native vegetation remnants are extremely fragmented.

The Northeast region of Brazil ranks third in number of bryophyte species, behind the South and Southeast regions (Costa & Peralta 2015Costa DP & Peralta DF (2015) Bryophytes diversity in Brazil. Rodriguésia 66: 1063-1071.). As for the bryophyte flora of the state of Ceará, this is still partially known. Most studies on the flora of Ceará have focused on flowering plants, for they represent the predominant vegetation. There are still few reports of bryophytes in Ceará, especially due to the lack of collections in native vegetation, although bryophytes can be found even in the most inhospitable places (Brito & Pôrto 2000Brito AERM & Pôrto KC (2000) Guia de estudos de briófitas: briófitas do Ceará. Edições UFC: 1-66.). The best explored areas to date are quite small and, considering the extension of the state, it is unlikely that the results obtained portray the real situation of the bryophyte flora of Ceará. The main areas sampled until present are humid and sub-humid enclaves, remnants of Atlantic rain forest (Oliveira & Alves 2007Oliveira HC & Alves MH (2007) Adições à brioflora do estado do Ceará, Brasil. Rodriguésia 58: 1-11.; Oliveira & Bastos 2009Oliveira HC & Bastos CJP (2009) Jungermanniales (Marchantiophyta) da Chapada da Ibiapaba, Ceará, Brasil. Acta Botanica Brasílica 23: 1202-1209., 2010aOliveira HC & Bastos CJP (2010a) Musgos pleurocárpicos da Chapada da Ibiapaba, Ceará, Brasil. Acta Botanica Brasílica 24: 193-204., bOliveira HC & Bastos CJP (2010b) Fissidentaceae (Bryophyta) da Chapada da Ibiapaba, Ceará, Brasil. Revista Brasileira de Botânica 33: 393-405.; Siqueira et al. 2011Siqueira SMC, Costa PS, Souza EB & Oliveira HC (2011) Briófitas de um remanescente de Mata Atlântica no Município de Ubajara, CE, Brasil. Hoehnea 38: 597-608.).

There are no works specifically addressing the bryophytes growing in the Caatinga in Ceará. The few studies carried out so far were floristic surveys in humid environments, in high altitudes such as the Ibiapaba Plateau (Oliveira & Bastos 2009Oliveira HC & Bastos CJP (2009) Jungermanniales (Marchantiophyta) da Chapada da Ibiapaba, Ceará, Brasil. Acta Botanica Brasílica 23: 1202-1209., 2010aOliveira HC & Bastos CJP (2010a) Musgos pleurocárpicos da Chapada da Ibiapaba, Ceará, Brasil. Acta Botanica Brasílica 24: 193-204.,bOliveira HC & Bastos CJP (2010b) Fissidentaceae (Bryophyta) da Chapada da Ibiapaba, Ceará, Brasil. Revista Brasileira de Botânica 33: 393-405.) and the Guaramiranga mountain (Yano & Pôrto 2006Yano O & Pôrto KC (2006) Diversidade das briófitas das matas serranas do Ceará, Brasil. Hoehnea 33: 7-40.). In this context, the Apodi Plateau, located in the eastern side of Ceará, represents a relevant area to the ​​study of bryophytes growing in the Caatinga, taking into account the lack of works in the region. The knowledge of the bryophyte flora of the Apodi Plateau is fundamental to understand the diversity of the ecosystem as a whole, aiming at environmental valuation and support to appropriate management measures.

The present study aimed to carry out a floristic survey of the bryophytes of the Apodi Plateau, within the limits of the territory of the state of Ceará.

Material and Methods

Study area

The Apodi Plateau is located in the eastern side of the state of Ceará, also extending into the territory of Rio Grande do Norte (Fig. 1). In general, the natural vegetation is represented by hyperoxerophilic Caatinga, which is part of the dominant landscape of the ecosystem of the Apodi Plateau, associated to secondary forests with varied herbaceous strata. According to Filho et al. (2001)Filho PT, Silva JAA, Meunier IMJ & Ferreira RLC (2001) Fisionomias da cobertura vegetal da Floresta Nacional do Araripe, estado do Ceará. Brasil Florestal 20: 13-21. , different types of vegetation coverage can be observed in the region: Savanna (Cerrado); Forested savanna (Cerradão), currently named as xeromorphic tropical subcaducifolia forest; Carrasco; and a seasonal dense Ombrophilous Forest remnant, probably resulting from great environmental heterogeneity throughout many geological periods (Giulietti et al. 2004Giulietti AM, Neta ALB, Castro AAJF, Gamarra-Rojas CFL, Sampaio EVSB, Virgínio JF, Queiroz LP , Figueiredo MA, Rodal MJN, Barbosa MR V & Harley RM (2004) Diagnóstico da vegetação nativa do bioma Caatinga. In: Silva JMC , Tabarelli M & Fonseca MT (orgs.) Biodiversidade da Caatinga: áreas e ações prioritárias para a conservação. Ministério do Meio Ambiente, Brasília. Pp. 1-44.). Fernandes (1990)Fernandes AG (1990) Temas fisiográficos. Stylos Comunicações, Fortaleza. 116p. believes that the presence of a forest such as Cerradão in the state of Ceará was driven by climate change through geological time, which also caused progressions and regressions of this forest into the central region of Brazil. The predominant climate of the region is Hot Sub-humid Tropical, the soil is Yellow Dystrophic Latosol, and the mean annual precipitation and temperature is 1,061 mm and 25 ºC, respectively.

Figure 1
Location of the Apodi Plateau.

Collections of botanical material

Collections were part of the field activities of the Program for Research in the Biodiversity of the Semiarid region of Brazil (PPBio). Briological material was collected in 2014, in the Apodi Plateau located within the territory of Ceará, according to specialized techniques described in Frahm (2003)Frahm JP (2003) Manual of tropical bryology. Tropical Bryology 23: 1-196.. The material was collected manually or with the aid of a knife and stored in Kraft paper bags. In the field, altitudes and geographical coordinates were recorded at all points of collection with the aid of a GPS. The following data were also recorded: date and site of collection, type of ecosystem, habit, growth form, colonized substrate (bryocenological group), and other relevant ecological and/or taxonomic observations.

Samples were identified in the Plant Taxonomy Laboratory of the Universidade Estadual de Feira de Santana. For the macro and microscopic analysis, morpho-anatomical characteristics of gametophytes and sporophytes were observed with the aid of a stereomicroscope and an optical microscope. The classification system used was the one of Crandall-Stotler et al. (2009)Crandall-Stotler B, Stotler RE & Long DG (2009) Phylogeny and classification of the Marchantiophyta. Edinburgh Journal of Botany 66: 155-198. for liverworts, and Goffinet et al. (2009)Goffinet B, Buck WR & Shaw AJ (2009) Morphology and classification of the bryophyta. In: Goffinet B & Shaw AJ (eds.) Bryophyte biology, Cambridge University Press. 2: 55-138. for mosses. The geographical distribution of the species in Brazil was based on the BFG (2018)BFG - The Brazil Flora Group (2018) Brazilian Flora 2020: innovation and collaboration to meet Target 1 of the Global Strategy for Plant Conservation (GSPC). Rodriguésia 69: 1513-1527.. In order to identify species, the following works were used: Buck (1998)Buck WR (1998) Pleurocarpous mosses of the West Indies. Memoirs of The New York Botanical Garden 82: 1-400., Bordin & Yano (2013)Bordin J & Yano O (2013) Fissidentaceae (Bryophyta) do Brasil. Boletim do Instituto de Botânica 22: 1-168., Costa (2016)Costa DP (2016) A synopsis of the family Pottiaceae in Brazil. Phytotaxa 251: 1-69., Frahm (1991)Frahm JP (1991) Dicranaceae: Campylopodioideae, Paraleucobryoideae. Flora Neotropica, monograph 54: 1-237., Gradstein & Costa (2003)Gradstein SR & Costa DP (2003) The Hepaticae and Anthocerotae of Brazil. Memoirs of The New York Botanical Garden 87: 1-301., Lemos-Michel (2001)Lemos-Michel E (2001) Hepáticas epífitas sobre o pinheiro-brasileiro no Rio Grande do Sul. Editora da Universidade Federal do Rio Grande do Sul, Porto Alegre. 191p., Sharp et al. (1994)Sharp AJ, Crum H & Eckel PM (1994) The moss flora of Mexico. Memoirs of The New York Botanical Garden 69: 1- 1113., Yano & Peralta (2007)Yano O & Peralta DF (2007) Musgos (Bryophyta). In: Rizzo JA (coord.) Flora dos estados de Goiás e Tocantins: criptógamos. Vol. 6. Universidade Federal de Goiás, Goiânia. Pp. 1-333. and Zander (1993)Zander RH (1993) Genera of the Pottiaceae: mosses of harsh enviroments. Bulletin of the Buffalo Society of Natural Sciences 32: 1-378..

Vouchers of the identified material were deposited in the collection of the Herbarium of the Universidade Estadual de Feira de Santana (HUEFS).

Results

Fifty-nine bryophyte species were recorded, consisting of 27 liverworts - Marchantiophyta - distributed in seven families and 11 genera, and 32 mosses - Bryophyta - distributed in 14 families and 26 genera (Tab. 1). Among the species found, 25 are new records for Ceará, and Riccia subplana Steph. is reported for the first time to the Northeast region of Brazil. Twenty-one are new occurrences for the Caatinga Domain.

Table 1
Bryophyte species found in the Apodi Plateau, Ceará, Brazil. Substrates (Sub.): CO = corticolous; EX = epixylous; EF = epiphyllous; RU = rupicolous; TE = terrestrial. Domains: AM = Amazon; AT = Atlantic rain forest; SA = Savannah; CA = Caatinga; PA = Pampa; PL = Pantanal. The number of genera and species is given inside parentheses, next to each family name. ¨ = new record for the Caatinga Domain; * = new record for the state of Ceará; ** = new record for the Northeast of Brazil.

The predominant community, regarding the colonized substrate, was the corticolous (32 spp.), followed by rupicolous (12 spp.), terrestrial (10 spp.) and epixylous (6 spp.).

A list of all the new records for the state of Ceará, for the Caatinga, and for the Northeast region, with taxonomic comments for each species, is provided in the list below, in alphabetical order of families. An illustration of Weisiopsis bahiensis (Müll. Hal.) Broth. is provided because this is a species with restricted distribution in Brazil, cited only for the states of Bahia (site where the type was collected) and Piauí, and because there are few illustrations of the species available in the literature.

Marchantiophyta

Cephaloziellaceae Douin

Cylindrocolea rhizantha (Mont.) R.M.Schust., New Hedwigia 22 (1-2): 175. 1971 [1972].

The illustrations of this species is in Gradstein & Costa (2003)Gradstein SR & Costa DP (2003) The Hepaticae and Anthocerotae of Brazil. Memoirs of The New York Botanical Garden 87: 1-301..

It is characterized by bilobed leaves with acute to obtuse apex and subquadratic cells. It resembles Cylidrocolea planifolia, but presents relatively larger gametophytes (0.5-1 mm long). It was found on live bark. It is a new record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (AC), Northeast (BA, PE), Midwest (GO), Southeast (ES, RJ, SP).

Lejeuneaceae Cavers

Archilejeunea fuscescens (Hampe ex Lehm.) Fulford, The Bryologist 45: 174. 1942.

The illustrations of this species is in Gradstein & Costa (2003)Gradstein SR & Costa DP (2003) The Hepaticae and Anthocerotae of Brazil. Memoirs of The New York Botanical Garden 87: 1-301..

It is characterized by creeping habit, green to brown coloration, entire and imbricate underleaves, oblong-orbicular leaves with rectangular lobules, which have an apical tooth consisting of 1-2 cells. It was collected on rotting log. It is one of the most common epiphytic liverworts in the Amazon forest and is reported here for the first time for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (AC, AM, PA, RR), Northeast (AL, BA, PE), Southeast (ES, MG, RJ).

Ceratolejeunea cornuta (Lindenb.) Steph., Pflanzenw. Ost-Afrikas C: 65, 1895.

The illustrations of this species is in Dauphin (2003)Dauphin G (2003) Ceratolejeunea. Flora Neotropica Monograph 90: 1-86..

It is distinguished by inflated, spherical lobules present in some parts or in the whole plant, and perianths with four horns. It was found on live bark. This is the first record for the Caatinga Domain. Brazilian geographical distribution: North (AC, AM, AP, PA, RO, RR), Northeast (BA, CE, PE), Southeast (MG, RJ, SP), South (PR, SC).

Ceratolejeunea laetefusca (Austin) R.M.Schust., J. Elisha Mitchell Sci. Soc. 72 (2): 306. 1956.

The illustrations of this species is in Dauphin (2003)Dauphin G (2003) Ceratolejeunea. Flora Neotropica Monograph 90: 1-86..

It is characterized by the presence of caducous leaves and frequent reduced lobules. It was found on live bark. This is the first record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (AC, AM, PA, RR), Northeast (BA, PE), Midwest (GO), Southeast (ES, MG, RJ, SP).

Cheilolejeunea acutangula (Nees) Grolle, J. Hattori Bot. Lab., 45: 173. 1979.

The illustrations of this species is in Gradstein & Costa (2003)Gradstein SR & Costa DP (2003) The Hepaticae and Anthocerotae of Brazil. Memoirs of The New York Botanical Garden 87: 1-301..

It is distinguished by clearly mammillose leaf cells and the presence of 2-5 oil bodies per cell. It was collected on rotting log. This is the first record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (AM, PA, RR), Northeast (AL, BA, PE), Southeast (ES, MG, RJ, SP), South (RS, SC).

Cheilolejeunea adnata (Kunze ex Lehm.) Grolle, J. Bryol. 9 (4): 529. 1977Grolle R (1977) Miscellanea hepaticologica. Journal of Bryology 9: 529-538..

The illustrations of this species is in Grolle (1977)Grolle R (1977) Miscellanea hepaticologica. Journal of Bryology 9: 529-538..

It is characterized by leaf cells presenting trigones, lobules never reduced and with long apical and curved teeth, caducous leaves, and marginal rizhoids on leaves. It was found on live bark. This is the first record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (AM, PA), Northeast (BA, PE), Midwest (MT), South (PR, SC).

Cheilolejeunea discoidea (Lehm. & Lindenb.) R.M.Schust. & Kachroo, J. Linn. Soc., Bot. 56 (368): 509. 1961.

The illustrations of this species is in Bastos (2017)Bastos CJP (2017) The genus Cheilolejeunea (Spruce) Steph. (Lejeuneaceae, Marchantiophyta) in the Americas. Pesquisas. Botânica 70: 5-78..

It is distinguished by the small size of the gametophytes (up to 0.65 mm wide), being one of the smaller species of the genus. The reduced and weakly delimited trigones are also diagnostic characteristics. It was collected on live bark. This is the first record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (PA), Northeast (BA, SE), Midwest (DF, GO, MS), Southeast (ES, MG, SP).

Cheilolejeunea xanthocarpa (Lehm. & Lindenb.) Malombe, Acta Bot. Hung. 51 (3-4): 326, 2009.

The illustrations of this species is in Gradstein & Costa (2003)Gradstein SR & Costa DP (2003) The Hepaticae and Anthocerotae of Brazil. Memoirs of The New York Botanical Garden 87: 1-301..

The striking features of this species are the leaf lobes with strongly involute margins. It was found growing on rotting log. This is the first occurrence for the Caatinga. Brazilian geographical distribution: Northeast (BA, CE), Southeast (ES, MG, SP).

Lejeunea aphanes Spruce, J. Bot. 19: 36. 1881.

The illustrations of this species is in Bastos & Yano (2004)Bastos CJP & Yano O (2004) New records of Lejeuneaceae (Marchantiophyta) for the Brazil. Acta Botanica Malacitana 29: 13-21., as L. filipes Spruce.

The diagnostic feature of the species is the underleaf that grows attached to the stem and which is difficult to visualize due to its very thin cell walls which are usually collapsed. It was collected on live bark. This is the first record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: Northeast (BA).

Lejeunea oligoclada Spruce, Bull. Soc. Bot. France 36 (Suppl. Congress Bot. 1889): cxcix. 1889 [1890].

The illustrations of this species is in Bastos & Yano (2009)Yano O, Bordin J & Peralta DF (2009) Briófitas dos estados do Ceará, Maranhão, Paraíba, Piauí e Rio Grande do Norte (Brasil). Hoehnea 36: 387-415..

It may be confused with Lejeunea phyllobola; however, the smaller size of the gametophyte and the cells finely papillose in Lejeunea oligoclada distinguishes the two species. It was found on live bark. It is endemic to Brazil and this is the first occurrence for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: Northeast (AL, BA, PE), Southeast (ES, MG, RJ, SP), South (PR, SC).

Metzgeriaceae H. Klinggr.

Metzgeria albinea Spruce, Bull. Soc. Bot. France 36: 201. 1889.

The illustrations of this species is in Gradstein & Costa (2003).Gradstein SR & Costa DP (2003) The Hepaticae and Anthocerotae of Brazil. Memoirs of The New York Botanical Garden 87: 1-301.

It is characterized by the presence of two rows of cells on both surfaces (dorsal and ventral) of the midrib in transverse section, with about 10-15 medullary cells and two hairs per marginal cell. It was collected on live bark. This is the first record for the Caatinga Domain. Brazilian geographical distribution: Northeast (BA, CE, PE), Midwest (GO), Southeast (ES, MG, RJ, SP), South (PR, RS, SC).

Plagiochillaceae Müll. Frib. & Herzog

Plagiochila cristata (Sw.) Lindenb., Sp. Hepat. (fasc.1): 33. 1839.

The illustrations of this species is in Heinrichs (2002Heinrichs J (2002) A taxonomic revision of Plagiochila sect. Hylacoetes, sect. Adiantoidea and sect. Fuscoluteae in the Neotropics with a preliminary subdivision of neotropical Plagiochilaceae into nine lineages. Bryophytorum Bibliotheca 58: 1-184.).

It is distinguished by leaves that are longer than wider, with 15 to 50 teeth, two long teeth at the apex, and dentate ventral margin. This is the first occurrence for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (AM, PA), Northeast (BA, RN), Southeast (ES, MG, RJ, SP).

Plagiochila patula (Sw.) Lindenb., Sp. Hepat. (fasc.1): 21. 1839.

The illustrations of this species is in Gradstein & Costa (2003)Gradstein SR & Costa DP (2003) The Hepaticae and Anthocerotae of Brazil. Memoirs of The New York Botanical Garden 87: 1-301..

The dichotomous branching pattern, weakly dentate leaves with a high shoulder and with long-decurrent and entire ventral base are diagnostic characteristics of the species. Heinrichs et al. (1998)Heinrichs J, Gradstein SR & Grolle R (1998) A revision of the neotropical species of Plagiochila (Dumort.) Dumort. (Hepaticae) described by Olof Swartz. The Journal of the Hattori Botanical Laboratory 85: 1-32. also mention the presence of a high dorsal wing in the perianth. This is the first record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (AC, PA), Northeast (BA, PE), Southeast (MG, RJ, SP), South (PR).

Ricciaceae Rchb.

Riccia albopunctataJovet-Ast, Cryptog. Bryol. Lichénol. 12 (3): 237, 1991Jovet-Ast S (1991) Riccia (Hépatiques, Marchantiales) d’Amérique Latine. Taxons du Sous-Genre Riccia. Cryptogamie, Bryologie Lichénologie 12: 189-370..

The illustrations of this species is in Jovet-Ast (1991)Jovet-Ast S (1991) Riccia (Hépatiques, Marchantiales) d’Amérique Latine. Taxons du Sous-Genre Riccia. Cryptogamie, Bryologie Lichénologie 12: 189-370..

It is characterized by the numerous white spots, which are idioblastic cells visible through the epidermis, densely areolated distal spore surface and with tuberculated angles. It was collected on soil. This is the first citation for the state of Ceará. Brazilian geographical distribution: Northeast (BA), Midwest (MS, MT), Southeast (SP), South (PR, RS, SC).

Riccia subplana Steph., Symb. Antill. (Urban) 3 (2): 275, 1902.

The illustrations of this species is in Ayub et al. (2014)Ayub DM, Costa DP & Santos RP (2014) Additions to the Ricciaceae flora of Rio Grande do Sul, including two remarkable records for the Brazilian liverwort flora. Phytotaxa 161: 294-300..

It is characterized by stems 6-7 mm long, round apex, entire margin, absence of papillae or cilia, and presence of pink scales. It was collected on soil. The species has a disjunct distribution and had been recorded only for the Amazon forest and Rio Grande do Sul in Brazil (Ayub et al. 2014Ayub DM, Costa DP & Santos RP (2014) Additions to the Ricciaceae flora of Rio Grande do Sul, including two remarkable records for the Brazilian liverwort flora. Phytotaxa 161: 294-300.). It was found on soil. This is the first citation for the Northeast region of Brazil and for the Caatinga Domain. Brazilian geographical distribution: North (AM), South (RS).

Bryophyta

Bryaceae Schwägr.

Brachymenium exile (Dozy & Molk.) Bosch & Sande Lac., Bryol. Jav. 1: 139, 1860.

The illustrations of this species is in Sharp et al. (1994)Sharp AJ, Crum H & Eckel PM (1994) The moss flora of Mexico. Memoirs of The New York Botanical Garden 69: 1- 1113..

It is distinguished by the ligulate-lanceolate leaves, equally arranged in the stem, with acute to acuminate apex, rhomboid-hexagonal cells, entire and bordered margins, and long-excurrent costa ending in a wing tip. It was found on soil. It is endemic to Brazil and this is the first record for the state of Ceará. Brazilian geographical distribution: Northeast (BA, PE), Midwest (GO, MS), Southeast (RJ, SP).

Rosulabryum densifolium (Brid.) Ochyra, Biodivers. Poland 3: 162. 2003Ochyra R, Żarnowiec J & Bednarek-Ochyra H (2003) Census catalogue of Polish mosses. Biodiversity of Poland 3: 1-372..

The illustrations of this species is in Ochyra et al. (2003)Ochyra R, Żarnowiec J & Bednarek-Ochyra H (2003) Census catalogue of Polish mosses. Biodiversity of Poland 3: 1-372..

It is characterized by the robust size of the gametophytes, up to 10 cm long, narrow leaves, 6.5 mm long, with serrate margins. It was found on soil. This is the first record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: Northeast (BA, PE), Midwest (DF), Southeast (ES, MG, RJ, SP), South (PR, RS, SC).

Calymperaceae Kindb.

Syrrhopodon parasiticus (Sw. ex Brid.) Besch., Ann. Sci. Nat., Bot., Ser. 8, 1 (5-6): 298, 1895.

The illustrations of this species is in Sharp et al. (1994)Sharp AJ, Crum H & Eckel PM (1994) The moss flora of Mexico. Memoirs of The New York Botanical Garden 69: 1- 1113..

It is characterized by dimorphic leaves with unipapillose cells, filamentous propagules in the median region of the leaf, and cancellinae ending in acute angles. It was collected on live bark. This is the first record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (AC, AM, PA, RO, RR), Northeast (BA, PE), Midwest (DF, GO, MS, MT), Southeast (ES, MG, RJ, SP), South (PR, SC).

Fissidentaceae Schimp.

Fissidens perfalcatus Broth., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 Afd. 3 (7): 13. 1900.

The illustrations of this species is in Sharp et al. (1994)Sharp AJ, Crum H & Eckel PM (1994) The moss flora of Mexico. Memoirs of The New York Botanical Garden 69: 1- 1113..

It is characterized by oblong-ovate leaves, with vaginant laminae completely covering the stem, usually extending beyond the stem, limbidium occupying all the extension or up to 3/4 of the vaginant lamina in all leaves, and unipapillose cells. It was collected on live bark. This is the first record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (TO), Northeast (BA, PB, PE), Midwest (GO, MT), Southeast (ES).

Meteoriaceae Kindb.

Meteorium nigrescens (Sw. ex Hedw.) Dozy & Molk., Musc. Frond. Ined. Archip. Ind. 5: 160, 1848.

The illustrations of this species is in Buck (1998)Buck WR (1998) Pleurocarpous mosses of the West Indies. Memoirs of The New York Botanical Garden 82: 1-400..

It is distinguished by gametophytes with cylindrical branches and blackish apex, weakly concave and triangular-ovate leaves, with gradually acuminate and never piliferous apex, cordate base, crenulated margin, costa extending up to 1/2 of the leaf, and sub-quadratic and smooth alar cells. It was found on live bark. This is the first record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (PA), Northeast (BA, PE), Midwest (DF, GO, MS), Southeast (ES, MG, RJ, SP), South (PR, RS, SC).

Orthotrichaceae Arn.

Groutiella tomentosa (Hornsch.) Wijk & Margad., Taxon 9 (2): 51, 1960.

The illustrations of this species is in Sharp et al. (1994)Sharp AJ, Crum H & Eckel PM (1994) The moss flora of Mexico. Memoirs of The New York Botanical Garden 69: 1- 1113..

It is characterized by leaves with fragile and brittle apex, and marginal elongated basal cells contrasting with the rounded hexagonal cells of the lamina. It was found on live bark. This is the first record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (AM, PA, RO), Northeast (BA, PE, SE), Midwest (MT), Southeast (RJ, SP).

Schlotheimia jamesonii (Arn.) Brid., Bryol. Univ. 1: 742. 1826.

The illustrations of this species is in Allen (2002)Allen BH (2002) Moss flora of Central America, Part 2. Encalyptaceae-Orthotrichaceae. Monographs in Systematic Botany from the Missouri Botanical Garden: 1-699..

It is distinguished by the glossy, green to brown coloration, densely tomentose gametophytes, oblong and smooth leaves with abruptly acuminate apex. It was found on rock. This is a new record for the Caatinga Domain and for the state of Ceará. Brazilian geographical distribution: North (AC), Northeast (BA, MA, PE), Midwest (DF, GO, MS), Southeast (ES, MG, RJ, SP), South (PR, RS, SC).

Pottiaceae Schimp.

Barbula indica (Hook.) Spreng., Nomencl. Bot. 2: 72. 1824.

The illustrations of this species is in Sharp et al. (1994)Sharp AJ, Crum H & Eckel PM (1994) The moss flora of Mexico. Memoirs of The New York Botanical Garden 69: 1- 1113.

It is characterized by elliptic ovate leaves with flat or weakly curved margins in the medial portion, presence of masses of green, ovoid propagules, with three or more cells, produced in the axils of the leaves. It was found on rock. This is the first record for the state of Ceará. Brazilian geographical distribution: North (AC, AM, PA), Northeast (BA, PE, RN, SE), Midwest (DF, GO, MS, MT), Southeast (ES, MG, RJ, SP), South (PR, RS).

Weisiopsis bahiensis (Müll. Hal.) Broth., Nat. Pflanzenfam. 10: 271. 1924. Fig. 2

Figure 2
a-e. Weisiopsis bahiensis - a. gametophyte; b. leaf; c. cross section of the leaf; d. leaf apex; e. basal portion of the leaf.

It is characterized by small gametophytes, tubular leaves with flat to curved margins, flattened guide cells, absent or single peristome. It was found on rock. It is endemic to Brazil and this is the first report for the state of Ceará. Brazilian geographical distribution: Northeast (BA, PI).

Sematophyllaceae Broth.

Sematophyllum adnatum (Michx.) E. Britton, Bryologist 5: 65, 1902.

The illustrations of this species is in Buck (1998)Buck WR (1998) Pleurocarpous mosses of the West Indies. Memoirs of The New York Botanical Garden 82: 1-400..

Gametophytes with erect-patent, ecostate, flat stem leaves, with gradually acuminate apex, weakly inflated and yellow-orange alar cells, quadrate supra alar cells, few in number (2-3), measuring 1/2 of the length of the alar cells. It was collected on live logs. This is the first occurrence for the state of Ceará. Brazilian geographical distribution: North (AM, PA, TO), Northeast (BA, MA, RN), Midwest (DF, GO, MS, MT), Southeast (ES, MG, RJ, SP).

Discussion

The 59 bryophyte species found in the Apodi Plateau correspond to 30% of the total number species registered in the state of Ceará. This number can be considered high when compared to other surveys carried out in Caatinga areas, such as those where were conducted the studies by Porto et al. (1994)Pôrto KC, Silveira M FG & Sá PSA (1994) Briófitas de Caatinga. Vol. 1. Estação experimental do IPA. Caruaru, PE. Acta Botanica Brasílica 8: 77-85. in Caruaru county, where 17 species were registered, and Porto & Bezerra (1996)Pôrto KC & Bezerra MFA (1996) Briófitas de Caatinga. Vol. 2. Agrestina, Pernambuco, Brasil. Acta Botanica Brasílica 10: 93-102., who reported 13 bryophyte species for the municipality of Agrestina, also in Pernambuco.

Among the liverworts, the most representative families were Lejeuneaceae (12 spp.) and Plagiochilaceae (4 spp.). As for mosses, the families with the highest number of species were Bryaceae and Pottiaceae (4 spp.). In the present study, it was observed that the majority of the most frequent species have adaptations to thrive in restrictive environmental conditions, such as xeric environments, being considered generalists or typically adapted to solar incidence, such as Frullania caulisequa (Nees) Nees, Riccia vitallii Jovet-Ast, Bryum argenteum Broth., Octoblepharum albidum Hedw., Campylopus savannarum (Müll. Hal.) Mitt. and Hyophilla involuta (Hook.) A. Jaeger (Ochi 1980Ochi H (1980) A revision of the Neotropical Bryoideae, Musci. First Part. The Journal of the Faculty of Education Tottori University, Natural Science 29: 49-154.; Frahm 1991Frahm JP (1991) Dicranaceae: Campylopodioideae, Paraleucobryoideae. Flora Neotropica, monograph 54: 1-237.; Gradstein et al. 2001Gradstein SR, Churchill SP & Salazar Allen N (2001) Guide to the bryophytes of tropical America. Memoirs of The New York Botanical Garden 86: 1-585.). As regards the family Lejeuneaceae, this is one of the largest families of liverworts and, although better represented in humid forests, they occur also quite frequently in open and xeric environments (Gradstein 1994Gradstein SR (1994) Lejeuneaceae: Ptychantheae, Brachiolejeuneae. Flora Neotropica 61: 1-216.; Gradstein et al. 2001Gradstein SR, Churchill SP & Salazar Allen N (2001) Guide to the bryophytes of tropical America. Memoirs of The New York Botanical Garden 86: 1-585.). The Lejeuneaceae species recorded in this study present a considerable wide tolerance or are pioneers. In Brazil, they have also been found in disturbed forests, savannas, sandbanks, Caatinga and even plantations and pastures (Gradstein & Costa 2003Gradstein SR & Costa DP (2003) The Hepaticae and Anthocerotae of Brazil. Memoirs of The New York Botanical Garden 87: 1-301.; Forzza et al. 2010Forzza RC, Leitman PM, Costa AF, Carvalho Jr. AA, Peixoto AL, Walter BMT, Bicudo C, Zappi D, Costa DP , Lleras E, Martinelli G, Lima HC, Prado J, Stehmann JR, Baumgratz JFA, Pirani JR, Sylvestre L, Maia LC, Lohmann LG, Queiroz LP, Silveira M, Coelho MN, Mamede MC, Bastos MNC, Morim MP, Barbosa MR, Menezes M, Hopkins M, Secco R, Cavalcanti TB & Souza VC (2010) Catálogo de plantas e fungos do Brasil. 2 vols. Andrea Jakobsson Estúdio, Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. 1699p.).

The results of this study reinforce the importance of the Apodi Plateau for the preservation of bryophyte species in Caatinga areas and for the recognition of hot spots in order to guide conservation efforts to areas with high diversity of species and endemism (Kier et al. 2009Kier G, Kreft H, Lee TM, Jetz W, Ibisch P, Nowicki C, Mutke J & Barthlott W (2009) A global assessment of endemism and species richness across island and mainland regions. Proceedings of the National Academy of Sciences 106: 9322-9327.). According to Costa & Peralta (2015)Costa DP & Peralta DF (2015) Bryophytes diversity in Brazil. Rodriguésia 66: 1063-1071., Brazil has 242 endemic bryophyte species, of which only one is cited for the state of Ceará. The present study report three new records of endemic Brazilian species for the state of Ceará: Weisiopsis bahiensis (Müll. Hal.) Broth., Lejeunea oligoclada Spruce and Brachymenium exile (Dozy & Molk.) Bosch & Sande Lac., the first two are also new records for the Caatinga Domain.

Weisiopsis bahiensis seems to have some inconsistencies in the information available in literature. According to the original work (Müller 1855Müller C (1855) De muscis novis, incomplete descripts, neglectis criticisve. Botanische Zetuing 13: 764., as Pottia bahiensis Müll. Hal.) The species type is from the state of Bahia, however, at the BFG (2018)BFG - The Brazil Flora Group (2018) Brazilian Flora 2020: innovation and collaboration to meet Target 1 of the Global Strategy for Plant Conservation (GSPC). Rodriguésia 69: 1513-1527. platform, it is cited only for the state of Piauí, which is probably wrong. Yano et al. (2009)Yano O, Bordin J & Peralta DF (2009) Briófitas dos estados do Ceará, Maranhão, Paraíba, Piauí e Rio Grande do Norte (Brasil). Hoehnea 36: 387-415. cited W. bahiensis from Oeiras, Goiás state, although Oeiras is actually located in Piauí state. Crosby et al. (1999)Crosby MR, Magill RE, Allen B & He S (1999) A checklist of the mosses. Missouri Botanical Garden, St. Louis. 325p. ranked it as a species for which no new information could be found in the post-1962 literature, however, Gradstein et al. (2001)Gradstein SR, Churchill SP & Salazar Allen N (2001) Guide to the bryophytes of tropical America. Memoirs of The New York Botanical Garden 86: 1-585. considered this species as valid. Costa (2016)Costa DP (2016) A synopsis of the family Pottiaceae in Brazil. Phytotaxa 251: 1-69. is the most recent citation of this species in Brazil, however, as well as Yano et al. (2009)Yano O, Bordin J & Peralta DF (2009) Briófitas dos estados do Ceará, Maranhão, Paraíba, Piauí e Rio Grande do Norte (Brasil). Hoehnea 36: 387-415., with herbarium samples collected in 1978. Therefore, this work represents a rediscovery of W. bahiensis in nature and the third record in Brazil and in the world.

The high number of new species reported for the state of Ceará and for the Caatinga Domain underscores the importance of continuous floristic surveys, especially in areas to which there is a gap of knowledge such as the Apodi Plateau, part of the Caatinga Domain in the Semiarid region of Brazil.

  • Area Editor: Dra. Lana Sylvestre

Acknowledgements

The authors thank the Program for Research in the Biodiversity of the Semiarid region of Brazil (PPBio/CNPq) for the support in the field work and Thamara Rodrigues dos Reis for the collection of the samples.

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Publication Dates

  • Publication in this collection
    20 Dec 2019
  • Date of issue
    2019

History

  • Received
    24 Feb 2018
  • Accepted
    20 July 2018
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