Floristic composition of the Montane Forest in the Almadina-Barro Preto axis, Southern Bahia, BrazilAbstracts
The aim of this study is to survey the angiosperms of two montane forest remnants in the southern Bahia, Brazil: Corcovado (SCO) and Pedra Lascada (SPL). Both fragments are located in the municipality of Almadina and Barro Preto, respectively, and are 18 km distant from each other. We sampled 899 species of angiosperms distributed in 437 genera and 116 families. The SCO was the richest area with 678 species, distributed in 367 genera and 100 families. SPL showed 466 species in 269 genera and 88 families. The percentage of species identified was 85.8% and of this total, 37.7% are endemic to the Atlantic Forest, 11.2% are endemic to southern Bahia and northern Espírito Santo and 7% are disjunct between the Atlantic Forest and Amazon. The remaining percentages (44.3%) were of species widely distributed. The richest families in the two areas were Orchidaceae (10%), Rubiaceae (7%), Bromeliaceae (5.5%), Melastomataceae (4.2%) and Poaceae (4%). The richest genera were Psychotria (2%),Piper (1.8%), Ocotea (1.6%),Vriesea (1.5%) and Peperomia (1.4%). More than half of the recorded species showed non-arboreal habit, regarding life forms documented. That comes against the assertion that many authors in the tropical forests, where species richness in angiosperms is expected for non-woody species, especially in montane forests. Twelve species have been identified as new, but seven others already described from collections previously obtained in these two areas. Orchidaceae, Rubiaceae, Poaceae and Bromeliaceae showed significant richness in this study these families are commonly reported as the richest in other inventories in the Atlantic Forest in southern Bahia reinforcing their importance to the regional flora. The high levels of richness, endemism, and the growing numbers of new taxonomic discoveries from the SPL and SCO sites indicate the biological importance of these two forest remnants. The implementation of parks or other protected environmental reserves would be essential to the conservation of its species.
Atlantic Forest; Biodiversity conservation; Floristic survey and Remnant forests
O presente estudo objetivou inventariar as angiospermas de dois remanescentes florestais no sul da Bahia, Brasil. Os remanescentes se encontram nas Serras do Corcovado (SCO) e da Pedra Lascada (SPL), situados nos municípios de Almadina e Barro Preto, respectivamente, e distantes 18 km um do outro. Foram registradas 899 espécies de angiospermas distribuídas em 437 gêneros e 116 famílias. A SCO foi a área mais rica em espécies, com um total de 678 espécies distribuídas em 367 gêneros e 100 famílias, enquanto a SPL apresentou 466 espécies em 269 gêneros e 88 famílias. O percentual de espécies identificadas foi de 85,8%. Desse total, 37,7% são endêmicas da Floresta Atlântica e 11,2% são endêmicas do sul da Bahia e norte do Espírito Santo. A distribuição disjunta entre Florestas Atlântica e Amazônica foi constatada em 7% das espécies inventariadas. O percentual restante (44,3%) foi de espécies amplamente distribuídas no Brasil. As famílias mais ricas nas duas áreas foram Orchidaceae (com 10%), Rubiaceae (7%), Bromeliaceae (5,5%), Melastomataceae (4,2%) e Poaceae (4%). Já os gêneros mais ricos foramPsychotria, (com 2%), Piper (1,8%),Ocotea (1,6%), Vriesea (1,5%) ePeperomia (1,4%). Mais de metade das espécies registradas apresentaram hábito não-arbóreo com relação ès formas de vida documentadas. Isso vem de encontro com a afirmativa de diversos autores de que em florestas tropicais a grande riqueza nas angiospermas é esperada para as espécies não-lenhosas, especialmente na Floresta Montana. Até o momento, doze espécies foram apontadas como novas, além de outras sete já descritas a partir das coleções obtidas anteriormente nessas duas áreas. Orchidaceae, Rubiaceae, Bromeliaceae e Poaceae apresentaram significativa riqueza nesse estudo e são famílias comumente reportadas como as mais ricas em outros inventários na Floresta Atlântica no sul da Bahia comprovando sua importância na flora local. Os altos índices de riqueza, endemismo e o crescente número de novidades taxonômicas provenientes de ambas as áreas indicam a importância biológica desses dois remanescentes. A implementação de parques ou demais reservas ambientais protegidas seriam essenciais para a conservação de suas espécies.
Floresta Atlântica; Conservação da biodiversidade; Documentação florística e Remanescentes florestais
Introduction
The Atlantic Forest covers the east coast of Brazil and is the second largest tropical rainforest on the American continent. It comprises 1% to 8% of the world's biodiversity (Silva and Casteleti 2005) and is the largest hotspot in the country, comprising 17,691 plant species, including algae, bryophytes, ferns, lycophytes, gymnosperms, and angiosperms, 40% of which are endemic (Forzza et al. 2012). The Atlantic Forest is considered one of the most important phytogeographic domains for biodiversity preservation worldwide because it is extremely diversified, covering regions with various levels of species abundance, composition, and endemism (Silva and Casteleti 2005).
The abundance and diversity are thought to have resulted in the isolation of two large South American forest blocks: the Amazonian Forest and the Andean Forests (Silva and Casteleti 2005). The Atlantic Forest and the vast Amazonian domain are separated by an open corridor formed by seasonal vegetation, including the Caatinga in the northeast semi-arid region of Brazil; the Cerrado in the midwest; and the Chaco, a region of dry vegetation located in the central lowlands of South America (Argentia, Bolivia, and Paraguay), which separates the Atlantic domain from the Andean forests (Rizzini 1997, Silva and Casteleti 2005). The transition from the Atlantic Forest to the Caatinga in the semi-arid regions is relatively abrupt and occurs in northeastern Brazil, where a narrow strip of coastal forests (less than 50 km) is delimited by an equally narrow strip of seasonal semi-deciduous forests (Oliveira-Filho and Fontes 2000). In southeastern Brazil, the transition from coastal forests to the Cerrado biome involves a much larger extension of semi-deciduous forests, which extends southward and forms complex mosaics with the Cerrado vegetation to the west. In the south, these semi-deciduous forests also extend along the Paraná river basin in eastern Paraguay and northeastern Argentina, where they make a transition to the Chaco biome (Oliveira-Filho and Fontes 2000).
The isolation of the South American forest blocks has resulted in consistent demographic changes in forest populations during the Pleistocene and Holocene eras, followed by climate changes during the late Quarternary era (Carnaval and Moritz 2008). These changes have had a greater impact on the southern portion of the Atlantic Forest and resulted in the evolution of a unique biota (Oliveira-Filho and Ratter 1995, Carnaval and Moritz 2008). The Atlantic Forest is considered one of the most unique biogeographical zones in South America. It shows great variations in topography, pluviometric regimes, and phytogeographic units because of its wide latitude (approximately 27°), longitude (from the coast to the interior), and altitude (from the sea level to altitudes of approximately 2700 m) (Silva and Casteleti 2005). These elements have led to the floristic and physiognomic heterogeneity present across the entire area (Pinto et al. 1996, Oliveira-Filho and Fontes 2000).
Special importance should be given to the forest altitude, which accounts for several environmental factors, including variations in the availability of solar energy, resources, and the forest's potential to serve as refuge for immigrant species; this potential may be higher in the lowlands and lower in the more isolated montane areas (Lomolino 2001). In addition, higher altitudes lead to a reduction in the number of species because of the occurrence of more severe environmental conditions with an increase in the altitude (e.g., edaphic factors, temperature, wind speed, and rainfall) (Lieberman et al. 1996, Pendry and Proctor 1996).
Taken together, these factors result in a wide variation in the species composition of the Atlantic Forest and make this area a heterogeneous unit with regard to studies on biodiversity conservation (Silva and Casteleti 2005). The difference between ombrophilous and semi-deciduous forests is consistent from a floristic point of view and is closely correlated with the rainfall (Oliveira-Filho and Fontes 2000). In this respect, the arboreal flora of semi-deciduous forests is largely a subset of the flora of ombrophilous forests and probably gives rise to species capable of withstanding more prolonged dry seasons (Oliveira-Filho and Fontes 2000, Oliveira-Filho et al. 2005). In addition, changes in the flora of semi-deciduous forests are associated with an increase in length of the dry season, which is caused by an increase in the distance from the ocean. Furthermore, altitude variations and the corresponding temperature variations are closely correlated with the internal floristic differentiation in both ombrophilous and semi-deciduous forests (Oliveira-Filho and Fontes 2000). Notably, the difference between deciduous and semi-deciduous forests is probably linked to a combination of chemical properties of the soil, rainfall, and variations in the altitude and latitude (Oliveira-Filho and Ratter 1995). Therefore, the definition of the Atlantic Forest should be as broad as that of the Amazonian formations (Oliveira-Filho and Fontes 2000).
In this context, the Atlantic Forest of southern Bahia can be included among the wet forests of the Northeast, which extend from Pernambuco to northern Espírito Santo. It is represented by coastal forests that cover an area of approximately 100-200 km in width along the east coast of Brazil and by forests that become increasingly drier toward the interior. Therefore, open formations gradually change to ombrophilous forests, semi-deciduous forests, and seasonally dry deciduous forest as one moves from east to west (Gouvêa et al. 1976, Silva and Casteleti 2005). In these forests, the minimum annual rainfall of 1600 mm and a dry period of not more than 2 months per year reflect the separation between wet and seasonal forests, with wet forests being essentially evergreen and comprising less than 20% of deciduous trees (Thomas and Barbosa 2008). In addition, wet forests can be classified according to the altitude as follows: lowland (20-100 m), submontane (100-600 m), and montane (600-800 m) (Thomas and Barbosa 2008). However, Veloso (1992) defined other altitude ranges for these areas, with montane forests occurring at altitudes between 500 and 1500 m.
The coastal forests of southern Bahia may have provided refuge for the biodiversity present during the Pleistocene era. The continual identification of new species and the high level of endemism detected in other studies reinforce the evidence of the uniqueness of this relatively unexplored region (Mori et al. 1981, Thomas et al. 1998, Carnaval and Moritz 2008,Amorim et al. 2009). Of all the coastal areas of Brazil, coastal forests contain the highest number of endemic species of Myrtaceae (15 species), which are threatened with extinction (Carnaval and Moritz 2008, Murray-Smith et al. 2008).
To investigate the flora of the Montane Forest in southern Bahia, the present study aimed to produce an inventory of angiosperms in two vestigial areas of the Montane Forest located on the Almadina-Barro Preto axis in the cocoa-growing region of southern Bahia. Moreover, the present study aimed to investigate whether these vestigial areas differ in terms of abundance from other areas of vestigial forests located in nearby regions, for which floristic documentation exists. In southern Bahia, the abundance and endemism of vascular plants reportedly yields high levels of diversity (Thomas et al. 1998, Martini et al. 2007, Thomas et al. 2009, Amorim et al. 2009, Murray-Smith et al. 2008). Importantly, this floristic documentation provides more detailed information on the endemism, abundance, areas of occurrence of taxa, and identification of new species and may serve as the foundation for future studies on biome similarities, biogeography, and community structure (Giulietti et al. 2005, Funk 2006).
Material and Methods
Study areas
The study areas are located in the Montane Forest (Veloso, 1992) in the cocoa-growing region of the State of Bahia (Gouvêa, 1976), in the cities of Barro Preto [Serra da Pedra Lascada (SPL)] and Almadina [Serra do Corcovado (SCO)], which are approximately 18-km apart (Figure 1). The floristic richness of SLP has been reported previously (Amorim et al. 2009) and has been revised and extended in the present study.
Location of Serra da Pedra Lascada (SPL) and Serra do Corcovado (SCO) regions and their respective cities (Barro Preto and Almadina) in southern Bahia, Brazil.
SPL (14°46′S and 39°32′W) is a vestigial forest of approximately 300 ha that lies approximately 56 km from the coast, with altitudes ranging from 600 to 950 m above the sea level (Amorim et al. 2009). There is a rocky outcrop of the “inselberg” type on the eastern side of the slope. This outcrop is formed by granites and gnaisses from the Precambrian era, comprising ancient elements of the landscape (Porembski et al. 1998). SPL is primarily populated by rupicolous monocotyledons and exhibits particular edaphic and microclimatic conditions, revealing its importance from a geomorphological and topographical perspective (Figures 2B, C and D). This forest has some well-preserved areas, with trees 20 to 30 m in height, and a dense subforest. There is an abundance of epiphytes, particularly mosses, ferns, and lycophytes, which sometimes give the appearance of cloud forests as the altitude increases. The forest fragment surveyed has an irregular area and is sharply demarked by the adjacent cocoa plantations (Theobroma cacao L.) where the trees are cultivated under the cabruca system. This fragment also comprises regeneration areas and pasture lands (Amorim et al. 2009).
Serra do Corcovado (SCO) and Serra da Pedra Lascada (SPL) in the Montane Forest, southern Bahia, Brazil
A: General view of SCO from the urban center of Almadina. B: General view of SPL from the access road. C: Secondary vegetation on one of the slopes of SPL. D: Interior of a mature forest in SPL showing the base of the inselberg. E: Vegetation at the top of the slope showing one of the exposed sides of the inselberg in SCO. Photos A and E were taken by André Paviotti. Photos B, C, and D were taken by André Amorim.
SCO (14°42′S and 39°36′W) is located at altitudes ranging from 400 to 1040 m above the sea level. It comprises an area of approximately 2500 ha and is located at a distance of 65 km from the coast; its slopes contain springs that form part of the river basin of Almada, the main river in the region. These springs ultimately provide water supply to the city of Almadina (Figure 2A). A part of the slope of SCO comprises a steep rocky outcrop of the “inselberg” type, which gives it a unique geomorphology. This outcrop is populated by a large number of vascular plants, particularly rupicolous flora such as Bromeliaceae and Orchidaceae (Figure 2E) (Porembski et al. 1998). The forest contains dense subforests, trees up to 35 m in height, and an abundance of epiphytes, giving it the appearance of a cloud forest at altitudes of 800 m above the sea level. There is an abundance of rupicolous species, particularly mosses, and the presence of various species of Cyatheaceae. Cocoa plantations, with trees cultivated under the cabruca system, and pasture lands are also common around SCO.
The climate of the region is warm and wet, with a dry season of the Af (Köppen) type (Peel et al. 2007). The average annual rainfall ranges between 1500 and 1750 mm, and the average daily rainfall varies between 50 and 100 mm. The average annual temperature varies between 23°C and 24°C, with a thermal range of 10°C to 14°C. The annual total potential evapotranspiration varies from 1200 to 1300, mm and the relative humidity is less than 80%.
Floristic Surveys
Eight field trips were conducted between July 2011 and June 2012, each lasting for 2 to 3 days. The surveys prioritized SCO because it is an area with large gaps in floristic documentation. These field trips, when added to the 12 previous trips (five in SCO and seven in SPL) conducted by various collaborators between 2004 and 2010, enabled the collections to be distributed throughout the year and increased the documentation of fertile specimens. Preliminary data from SPL used in this study had been previously published by Amorim et al. (2009).
The documentation of angiosperms was conducted through collection along the trails and access roads by careful visual examination with the aid of binoculars, with the aim of collecting the largest number of fertile species possible. Some trees were surveyed using climbing techniques to collect arboreal specimens and document the epiphytic flora. In addition, fallen trees and canopy branches were examined in detail. Sterile specimens were collected whenever their identification in the field was possible.
The material collected was prepared according to Fidalgo and Bononi (1989) and deposited in the CEPEC Herbarium. Duplicates were sent to the HUEFS and RB archives. Species were identified by literature search, comparison with the material deposited in the CEPEC, and consultation with specialists. The material identified was standardized in morphotypes and classified as proposed by APG III (2009). The specific epithets and citations of the authors of the species were standardized on the basis of the Lista de Espécies da Flora do Brasil (2012) [List of Species of the Brazilian Flora (2012)] and on the website The Plant List (2012). The occurrence of species and endemism were verified in the Lista de Espécies da Flora do Brasil (2012), Amorim et al. (2009), Thomas et al. (2003), and Stehmann et al. (2009). Endangered species were searched in the lists of Biodiversitas (2009) and MMA (2008).
Classification of life forms into arboreal, arbustive, epiphytic, hemiepiphytic, parasitic, and hemiparasitic followed the standard used by Amorim et al. (2009) and was obtained by field observation and, in some cases, from exsiccate labels of each taxon. Rupicolous speces are indicated in Table 1 with an asterisk (*). The percentage of contribution of these species was calculated and compared with that obtained in previous survey (Amorim et al. 2009) conducted in the Atlantic Forest in southern Bahia, which used a similar methodology and for which sample material is accessible in scientific archives such as CEPEC and RB.
List of species sampled on Serra do Corcovado and Serra da Pedra Lascada in the Montane Forest, southern Bahia, Brazil. Arb. = Arboreal, Shr. = Shrub, Epip. = Epiphyte, Hemiep.= Hemiepiphytes , Hemi-par. = Hemi-parasitic, Herb. = Herbaceous, Holopar. = Holoparasitic, Sub-shr. = Sub-shrub, Clim. = Climber; Collectors: AA = André Amorim, AF = André Fontana, Adriana Lobao AL =, DC = Domingos Cardoso, DM = Daniele Monteiro JJ = Jomar Jardim, JP = José Paixão, LD = Daneu Lucas, MC = Macielle Coelho, ML = Mardel Lopes, PF = Pedro Fiaschi, PO = Patricia Oliveira, RB = Rafael Borges, RP = Ricardo Perdiz, WT = Thomas Wayt. Domains: AM = Amazonian, CA = Caatinga CE = Cerrado, AF = Atlantic Forest, PA = Pampa, PAN = Pantanal. Category of threat: CR = Critically Endangered, EN = Endangered, LC = Least concern, NT = Near Threatened, VU = Vulnerable. * = rupicolous
Results
Floristic Survey
In the SCO and SPL areas, a total of 899 species of angiosperms, distributed in 437 genera and 116 families, was documented (Table 1). SCO was the area with a greater abundance, with 678 species distributed in 367 genera and 100 families. SPL comprised 466 species in 269 genera and 88 families. The percentage of species identified was 85.8% (772 species), 14% (124 species) and 0.5% (5 species) of which were identified only at the genus and family level, respectively.
Of the total species documented, 37.7% (291 species) are endemic to the Atlantic Forest and 11.2% (101 species) are endemic to southern Bahia and northern Espírito Santo (Figure 3). The remaining 44.3% species are widely distributed in Brazil. Most endemic species were arboreal (36%), followed by epiphytic (23.3%), arbustive (11%), climbing (10.7%), and herbaceous (7.6%) species. In total, 81 species (7%) had a disjunct distribution between the Atlantic and Amazonian forests.
Endemic species
A and B: Portea filifera (Bromeliaceae), an endemic species of southern Bahia. C: Chamaecostus cuspidatus(Costaceae), an endemic species of the Atlantic Forest. D:Bertolonia bullata (Melastomataceae), an endemic species of southern Bahia. E: Dorstenia hirta(Moraceae), an endemic species of the Atlantic Forest. F:Notopleura tapajozensis (Rubiaceae), a disjunct species between the Amazonian and Atlantic forests. Photos A, B, D,E and F were taken by André Amorim. Photo C was taken by Macielle Coelho.
In SCO and SPL, the five most abundant families were Orchidaceae (10%; 91 species), Rubiaceae (7%; 63 species), Bromeliaceae (5.5%; 50 species), Melastomataceae (4.2%; 38 species), and Poaceae (4%; 37 species). These families accounted for 30.7% of the documented species (Figure 4). In terms of the number of species, the five most abundant families in SCO were Orchidaceae (69 species), Rubiaceae (46 species), Bromeliaceae (33 species), Fabaceae (30 species), and Melastomataceae (28 species). In SPL, the families with the largest number of species were Orchidaceae (42 species), Rubiaceae (41 species), Bromeliaceae (30 species), Melastomataceae (27 species), and Poaceae (23 species).
The most abundant angiosperm families in the vestigial forests of Serra do Corcovado (SCO) and Serra da Pedra Lascada (SPL) located in southern Bahia, Brazil
In these two areas, the most abundant genera were Psychotria (18 species), Piper (17 species), Ocotea (15 species), Vriesea (14 species), and Peperomia(13 species) (Figure 5). In SCO, the number of genera represented by a single species totaled 34.2% (233 genera), and in SPL, it totaled 40% (185 genera). When both the areas were analyzed together, the percentage was only 29.4% (265 genera). In terms of the number of species, the most abundant genera in SCO were Psychotria (16 species),Piper (13 species), Peperomia andSolanum (11 species each), and Anthurium,Ocotea, and Vriesea (10 species each), whereas the most abundant genera in SPL were Psychotria (12 species), Piper (11 species), Ocotea andPeperomia (9 species each), and Aechmea,Leandra, Miconia, andVriesea (8 species each).
The most abundant angiosperm genera in the vestigial forests of Serra do Corcovado (SCO) and Serra da Pedra Lascada (SPL) located in southern Bahia, Brazil
Among the life forms documented in SCO, 35.2% of the species were arboreal, 21% were epiphytic/hemiepiphytic, 16.5% were arbustive/subarbustive, 14,6% were herbaceous, 11.5% were climbing, and 1.3% were hemiparasitic (Figure 6). In SPL, 33.7% of the species were arboreal, 23.8% were epiphytic/hemiepiphytic, 18,5% were arbustive/subarbustive, 13.4% were herbaceous, 10% were climbing, and 0.6% were hemiparasitic. In total, 17 species were rupicolous, and most of them belonged to the family Piperaceae. More than 50% of the species recorded were nonarboreal (64.8% in SCO and 66.3% in SPL). These values were similar to those found in previous studies performed in the same region (Figure 6, Table 2).
Life forms found in distinct areas of the Atlantic Forest in southern Bahia: Serra do Corcovado (SCO), Serra da Pedra Lascada (SPL), Serra Bonita (SBO), Serra das Lontras (SLO), Una Biological Reserve (UNA), and Serra do Teimoso Natural Reserve [Patrimônio Natural Serra do Teimoso (TMS)]
Comparison of taxonomic diversity and life forms from four montane areas of southern Bahia and other areas of Atlantic Forest. SCO = Corcovado Mountain, SPL = Pedra Lascada Mountain, SBO = Serra Bonita Mountain, SLO = Lontras Mountain, UNA = Una Biological Reserve, TMS = Teimoso Mountain. Epi./Hemiepiph. = Epiphytes and hemiepiphytes, Climb. = Climbers, Herb. = Herbaceous, Árb./Shr. = Trees and shrubs. (Adapted from Amorim et al. 2009 with updates numbers).
In SCO and SPL, the most abundant families in terms of liana species were Malpighiaceae (13 species), Asteraceae (12 species), Sapindaceae (9 species), Celastraceae (8 species), and Bignoniaceae, Fabaceae, and Cucurbitaceae (7 species each), accounting for 59.4% of the liana species recorded. The Orchidaceae family consisted of the highest number of epiphytic species (74 species), followed by Bromeliaceae (47 species), Araceae (25 species), and Piperaceae (12 species), accounting for 82.7% of the epiphytic species recorded. The Myrtaceae family had the highest number of arboreal species (25 species), followed by Fabaceae (23 species), Rubiaceae (22 species), Lauraceae (21 species), Melastomataceae (18 species), Annonaceae (10 species), and Euphorbiaceae and Sapindaceae (9 species each), accounting for 34.6% of the arboreal species sampled.
In addition to the seven angiosperm species recently described following the first field trips to SPL and SCO, 12 other species have been identified by nine specialists till date. Some of these species are in the description phase, including one species each of Philodendron (Araceae),Vriesea (Bromeliaceae), Dichorisandra(Commelinaceae), Bertolonia (Melastomataceae),Ichnanthus (Poaceae), Myrsine(Primulaceae), Faramea and Psychotria(Rubiaceae), Cupania (Sapindaceae), andSymplocos (Symplocaceae) and two species ofPeperomia (Piperaceae).
Discussion
From a floristic perspective, the abundance and percentage of endemic Atlantic Forest species found in the two areas surveyed (SCO and SPL) are corroborated in previous studies indicating that this environment is one of the richest ecosystems in Brazil with high levels of endemism (Mori et al. 1981, Gentry 1992, Martini et al. 2007, Murray-Smith et al. 2008, Amorim et al. 2009, Forzza et al. 2012). Southern Bahia is considered one of the three regions of endemism in the Atlantic Forest and one of the six regions with high levels of endemic plants threatened with extinction (Murray-Smith et al. 2008). However, research and conservation measures focused on this region are still inadequate (Carnaval and Moritz 2008, Amorim et al. 2009). The percentage of endemism among the species in southern Bahia and northern Espírito Santo found in this study (11.2%) was similar to that obtained byAmorim et al. (2009) in the Montane Forest (7.4%) but lower than that obtained in areas of lowland ombrophilous forests (18.9%-28.1% of the total flora) (Thomas et al. 1998, Amorim et al. 2008). However, the endemism in the Montane Forest appears to be more local, with species recorded only in these elevation zones, as exemplified by the recent addition of species to genera such as Bertolonia,Dichorisandra, Macrocarpaea,Quesnelia, and others.
In the two study areas, Psychotria, Piper,Ocotea, Vriesea, andPeperomia were the most abundant genera in terms of the number of species; these were also the most representative genera in previous study conducted in similar areas, with the exception of Vriesea (Amorim et al. 2009), which is epiphytic. In addition, the genera represented by a single species accounted for 29.4% of the total, which was close to the percentage (23.6%) obtained by Amorim et al. (2009).
In terms of the number of species, the families Orchidaceae, Rubiaceae, Bromeliaceae, and Poaceae were abundant not only in SCO and SPL but also in other areas of the Atlantic Forest (Pabst and Dungs 1975, Soderstrom et al. 1988, Giulietti et al. 2005, Martinelli et al. 2008, Amorim et al. 2009). Furthermore, Melastomataceae is another family with a high number of species in the areas surveyed and constitutes an important Atlantic Forest group, considering the species abundance (Oliveira-Filho and Fontes 2000, Rocha and Amorim 2012).
The abundance of Asteraceae, Melastomataceae, and Solanaceae tends to increase with altitude in the Atlantic Forest (Oliveira-Filho and Fontes 2000). According to Amorim et al. (2009), these families are among the most abundant families in the mountainous areas of southern Bahia, including SCO and SPL reported in the present study. In contrast, the abundance of Chrysobalanaceae, Rutaceae, and Sapotaceae tends to decrease with an increase in the altitude (Oliveira-Filho and Fontes 2000). In the present study, the latter three families contained five, seven, and six species, respectively, and were among the least abundant families. On the other hand, in the study of Amorim et al. (2005), a high abundance of Rutaceae was detected in a montane region.
One of the main differences in floristic documentation between the present study and previous studies is related to the families Burseraceae and Combretaceae. These families have been reported to be generally highly abundant in the lowlands of southern Bahia (Amorim et al. 2005). In contrast, we did not identify the family Burseraceae and found a low representativeness of the family Combretaceae in the present study.
With regard to the life forms documented, more than 50% of the species recorded in the two areas were nonarboreal (64.8% in SCO and 66.3 in SPL). This is in accordance with previous findings that in tropical forests, a high abundance of angiosperms is expected for nonwoody species (Gentry and Dodson 1987, Gentry 1988, Webster 1995), particularly in the Montane Forest. In addition, these values are very similar to those found in PARNA Serra das Lontras and in RPPN Serra Bonita and, to a lesser extent, in RPPN Serra do Teimoso and REBIO de Una (Amorim et al. 2009). However, the latter two regions represent very different physiognomies than SCO and SPL: RPPN Serra do Teimoso has a strong seasonal influence, whereas REBIO de Una is situated in a submontane area near the coast.
With regard to the Atlantic Forest regions located in southeast Brazil, our results were very similar to those of Lima and Guedes-Bruni (1997) in Macaé de Cima and, to a lesser extent, to the inventory of the Juréia Mountains (Mamede et al. 2001), indicating that this pattern of life forms is similar to that observed in the Atlantic Forest. Subtle differences in the life forms can be observed, for example, the greater abundance of climbing species in SCO than in SPL and the greater abundance of epiphytic species in PARNA Serra das Lontras in contast to the greater abundance of herbaceous species in REBIO de Una and RPPN Serra do Teimoso. However, the homogeneity in the patterns of life forms is evident in these distinct locations, as described by Amorim et al. (2009) in a floristic study conducted in three montane areas in southern Bahia.
According to the Lista de Espécies da Flora do Brasil (2012) [List of Species of the Brazilian Flora (2012)], three species were not found in the Atlantic Forest: Cattleya elongata (Orchidaceae), M. leuconeura (Primulaceae), and Passiflora nitida (Passifloraceae). Of all the species sampled, 47 are new occurrences in northeast Brazil, four are new occurrences in southern Bahia, and eight are new occurrences in Bahia. Of the 47 new occurrences,Vanilla cf. bicolor (Orchidaceae) andPiper subglabrifolium (Piperaceae) were known only in the State of Amazonas in northern Brazil. In addition, it should be emphasized that 32 species found in SCO and SPL have been classified into various categories of threat of extinction, including Bactris pickelli (Arecaceae),Rhipsalis baccifera subsp. hileiabaiana(Cactaceae), Abarema cochliacarpos and Inga grazielae (Fabaceae), Heteropterys bullata(Malpighiaceae), and C. warneri (Orchidaceae), all of which are in the vulnerable category (MMA 2008, Biodiversitas 2009).
The presence of disjunct taxa between the Amazonian and Atlantic forests (6.8%, represented by 78 species) reinforces the idea of possible floristic connections between southern Bahia and the Amazonian Forest during the Quaternary era (Prance 1979, Oliveira-Filho and Ratter 1995, Carnaval and Moritz 2008). Although the percentage of species typically found in the Amazonian Forest is lower in the Montane Forest than in lowland areas, the disjunct distribution between taxa of the Atlantic and Amazonian forests has been previously reported (Andrade-Lima 1953, Prance 1979, Mori et al. 1981, Gentry 1982,Rizzini 1997, Amorim et al. 2008, 2009). The presence of common species among Bahia, Espírito Santo, and Rio de Janeiro (40 species; 3.5%) indicates another pattern in taxa distribution, which is in agreement with the study of Oliveira-Filho et al. (2005), which suggests the occurrence of a continuous gradient in the ombrophilous forests, from Rio de Janeiro to the south of Bahia.
The percentage of species not yet described (1.3%) was similar to that reported in studies by Amorim et al. (2005) (1.8%) andAmorim et al. (2009) (3.5%). These figures when added to the recently published species collected in SCO and SPL, such as A. viridipetala A.F.Costa and Amorim (Bromeliaceae), M. atlantica J.R.Grant and V.Trunz and M. orbiculataJ.R.Grant and V.Trunz (Gentianaceae), B. bullata Baumgratz, Amorim, and A.B.Jardim and M. dorsaliporosa R.Goldenb and Reginato (Melastomataceae), Myrcia lascada Sobral (Myrtaceae), andCarrapichea lucida J.G.Jardim and Zappi (Rubiaceae), together with the records of other species previously unknown in southern Bahia (7.9%) indicate the lack of floristic studies in this region (Amorim et al. 2009). In addition to the rare and unknown angiosperm species, the presence of species of ferns and lycophytes also deserves attention, particularly in SCO, where the Diplazium fimbriatum(Athyriaceae) (Mynssen and Matos 2012) type originates and where Adiantum diphyllum (Pteridaceae), a rare and endemic species of Bahia, was detected during the initial field trips to the area (Sundue and Prado 2005).
Therefore, floristic surveys that contribute to the description of new species and help delimit the distribution of occurrence of taxa constitute rich sources of information for the studies on biodiversity and conservation because these surveys will enable the identification of hotspots that should be prioritized in future conservation projects, at both the state and national levels. Till date, there has been no effective action for the establishment of legal protection units in SCO and SPL, which is a cause for concern, considering the abundance of vascular plants and the high number of new and endemic species documented, in addition to the large number of threatened species. Morever, the results of the present study revealed the ecological importance of the vestigial Montane Forest in southern Bahia and the need for further surveys in these areas. We believe that the data presented here will be useful in developing future conservation strategies in these areas, while serving as the fountation for future ecological, phylogenetic, and taxonomic studies, thereby complementing studies on local biodiversity, which are necessary for the preservation of these ecosystems.
Acknowledgment
The authors are thankful to the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior [Coordination for the Improvement of Higher Education Personnel (CAPES)] for the Master's degree scholarship granted to the first author; the Conselho Nacional de Desenvolvimento Científico e Tecnológico [National Council for Scientific and Technological Development (CNPq)] for the productivity scholarship (process 306992/2012-4) granted to the second author and for the funding obtained in the Edital (call for research proposals) (process No. 481592/2009-1) and Edital Reflora (process No. 563548/2010-0), which assisted in the field activities; the Fundação de Amparo a Pesquisa do Estado da Bahia (FAPESB) (Bahia Research Foundation) for the funding obtained in the Edital de Pesquisas (process APP0041/2009) in the initial phase of this research and the postgraduate program in Botany of the Universidade Estadual de Feira de Santana. We also thank the various taxonomists who assisted in the identification of the material; the technicians L. H. Daneu, L. C. Gomes, and J. L. Paixão; the collaborators who participated in previous field trips to the study areas; and R. Ramos for producing the map.
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Publication Dates
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Publication in this collection
08 Apr 2014 -
Date of issue
Jan-Mar 2014
History
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Received
03 May 2013 -
Reviewed
11 Aug 2013 -
Accepted
19 Jan 2014
