Liverworts of Alagoas State , Brazil

(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 The list is based on catalogues and previous papers as well as the results of an unpublished survey carried out at the Murici Ecological Station (EsEc), an important protected area of the Brazilian Northeast (9o11’05” 9o16’48”S; 35o45’20” 35o55’12”W).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.


Introduction
The bryophyte flora of Alagoas is one of the most poorly studied of the Brazilian Northeast.To date, no systematic survey has been performed and all references come from sparse and sporadic collections whose results have been put together in Yano's catalogues (1981;1984;1989;1995;2006).
Additionally, Alagoas' bryophyte flora is now increased by the survey carried out at the Murici Ecological Station (EsEc), one of the main Atlantic Forest remnants of northeastern Brazil.EsEc Murici was created in 2001 with 6,116.43 ha and is considered a priority area for biodiversity conservation in Brazil and worldwide (Brasil -MMA 2000).The survey resulted in a list of 181 species, of which 106 are liverworts, 72 are mosses, and three await special taxonomic treatment.This paper focuses only on liverworts, while mosses will receive attention in another publication.
This paper presents a compilation of liverwort species occurring in Alagoas State, bringing together published information (catalogues) and results from the EsEc Murici survey.Moreover, it provides data about the species' geographic distribution in Brazil and in the world, as well as taxonomic and ecological comments on the new occurrences for the northeastern region.

Material and methods
EsEc  is part of Murici and Messias municipalities, 50 km from Maceió, capital of the state of Alagoas (Moura 2006).The elevation of the reserve ranges between 200-650 m, and the area has warm, humid wheather, with a dry summer and a wet autumnwinter season.The mean annual precipitation and temperature are 2,167.7 and 24 °C, respectively (National Institute of Meteorology, data for the period from 1961 to 1990).The vegetation cover is classified as Submontane Open Humid Atlantic Rain Forest (Veloso et al. 1991).

Results and discussion
One-hundred sixteen species of liverworts were recorded for Alagoas.Seventy-eight species are new occurrences for Alagoas state, occurring exclusively at EsEc Murici, and seven species are new occurrences for the Brazilian Northeast as well.The other 38 species had been previously cited in catalogues and other papers.The taxa are listed in alphabetical order by family and species and include locality, substrata and forest strata (understory or canopy) in the case of EsEc Murici, and geographic distribution in Brazil and in the world (Tab.1).
Most of the species had typically Neotropical (59%), Pantropical (16%) and African-American (7%) geographic distributions.Lejeuneaceae is by far the richest family in the state, with 83 species.This obviously is because the flora presented here comes mainly from Atlantic Rain Forest.It is known that this family used to be predominant in the flora of tropical regions, especially in forest habitats (Gradstein et al. 2001) Description: Schuster (1992).
Comments: this species was placed within Riccardia S.F.Gray genus until recent decades.However, it can be distinguished from this genus because of at least two clear characters on gametophytes: 1) thallus less ramified, darker and wider, reaching up to 8 mm while Riccardia spp.reaches no more than 3 mm wide and; 2) many minutely dispersed oil bodies inside the cells.Aneura pinguis is easily recognized in the field even when sterile, due to its conspicuous and lustrous thallus, green to yellowish-green in color.Of the Aneura Dumort.species occurring in Brazil, A. pinguis is the only one with 10-18 (20) cell thickness in the middle (transversal section); the others are thinner, up to 9 cells thick.In Brazil, this species was only known for the North, Central and Southeast regions.It is sub-cosmopolitan and occurs at altitudes that vary from sea level to 800 m.It was found growing on decaying wood in EsEc Murici.Steph., Sp. Hepat. 5: 859. 1916.Fig. 3 Description: Gradstein & Costa (2003).

Aphanolejeunea asperrima
Comments: this species is characterized by well developed lobules in most leaves, reaching 2/5 to 2/3 of lobe length and with the first lobule tooth falcate, consisting of two slightly elongated cells; the surface of lobes, lobules, and perianths is covered with conical papillae.The marginal cells of lobes and lobules are conically protuberant.Aphanolejeunea asperrima resembles A. microscopica (Tayl.) A. Evans, but the latter has greater lobules that reach 2/3 to 4/5 of lobe length.Aphanolejeunea asperrima has only been recorded for Patagonia and Brazil, where it has been found growing on leaves at altitudes that vary from 1450-2000 m.In Brazil it was previously known to the South and Southeastern regions.It was found growing on leaves in the understory at EsEc Murici.
Comments: this species is characterized by elliptic, rhombic, or lanceolate leaves that are wider in the middle, nearly symmetrical, and with an acute apex that ends in one cell.The lobules are reduced in most leaves, but are occasionally developed and can reach 1/4 of lobe length.Aphanolejeunea camillii resembles A. paucifolia (Spruce) E. Reiner, but it is clearly distinguished by the leaves -those of the latter are asymmetrical, wider at the base, and with an apex that ends in a row of 3-4 cells; in addition, the lobules are always reduced.It is common in the Neotropics, occurring from Mexico to Paraguay and northern Argentina.In Brazil it is known to occur in the North and Southeast regions.It was found growing on leaves in the understory in the EsEc Murici.Steph., Hedwigia 34: 250. 1895. Fig. 11-16 Material examined: BRAZIL.Alagoas: Murici, 3/XII/2004, Pôrto (UFP 45828).
Comments: this species is characterized by leaves with a hyaline margin of rectangular cells and with plane lobules parallel to the stem frequently bilobate and with a rounded apex.Cololejeunea lanciloba has highly variable gametophytic characters, even concerning lobule shape; the latter fact causes problems in distinguishing it from Cololejeunea latilobula (Herzog) Tixier, according Dr. Tamás Pócs who is presently revising the genera for the Neotropics (pers.comm.Description: Bischler (1964).
Comments: this species is characterized by the presence of a conspicuous tooth 2-3 cells long on the ventral lobe margin, positioned just above the junction with the keel.This species resembles Drepanolejeunea lichenicola (Spruce) Steph., but the pre-apical lobule teeth of the latter are longer, curved, and usually cross the apical teeth, whereas in D. campanulata the teeth are short and inconspicuous.Drepanolejeunea campanulata was previously known to Ecuador, Peru, and Southeast Brazil.It was found growing on leaves and bark in the understory, sub-canopy, and canopy at EsEc Murici.
Comments: this species can be easily recognized due to the presence of conspicuous teeth that are 2-6 cells long and 1-3 cells wide at the base in both the dorsal and ventral leaf margins.This species resembles Drepanolejeunea palmifolia (Nees) Steph., but the latter has teeth only on dorsal margins.Drepanolejeunea crucianella generally occurs in lowland forests, growing on leaves.It has a Neotropical distribution and in Brazil it was previously restricted to Amazonas and Pará, in the North region.It was found growing on leaves in the understory at EsEc Murci.

Table 1 .
Liverworts of Alagoas state.Information on substrate (Leaves -LV, live bark -LB and decaying wood -DW) and forest strata occurrence concern only the EsEc Murici survey.Sub-canopy is defined here as strata above 4 meters high.New references for Northeast Brazil are preceded by an asterisk.
AM, AP, BA, CE, ES, MG, PA, PE, PR, RJ, RS, SC, SE, SP JUBULACEAE Frullania apiculata (Reinw.et al.) Nees , BA, CE, ES, GO, MG, MS, MT, PA, PB, PE, PR, RJ, RS, SE, SP F. kunzei (Lehm.& Lindenb.) ).The only reliable distinction between them is that C. lanciloba has narrowly ligulate to acute lobules usually with 1-2 lateral teeth while C. latilobula always has broad, obtuse, ligulate lobules.Cololejeunea lanciloba is Pantropical and grows on leaves in areas with altitudes up to 50 m.In Brazil it was restricted to Amazonas state, where it was recorded by the first name, Cololejeunea tonkinensis Steph.It was found growing on leaves in the understory at EsEc Murici.