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Revista de Biología Tropical

On-line version ISSN 0034-7744Print version ISSN 0034-7744

Rev. biol. trop vol.63 n.3 San José Jul./Sep. 2015



Floristic similarity and dispersal syndromes in a rocky outcrop in semi-arid Northeastern Brazil

Similitud florística y síndromes de dispersión en un afloramiento rocoso de la región semiárida del noreste de Brasil

Elainne Cristina Silva Costa 1  

Sergio de Faria Lopes 2  

José Iranildo de Miranda Melo 2  

1Graduate Program in Ecology and Conservation, Universidade Estadual da Paraiba, Avenida das Baraunas, 351, Bairro Universitario, CEP 58429-500 Campina Grande, Paraiba, Brazil;

2Universidade Estadual da Paraiba, Graduate Program in Ecology and Conservation, Department of Biology, Center for Biological and Health Sciences, Avenida das Baraunas, 351, Bairro Universitario, CEP 58429-500 Campina Grande, Paraiba, Brazil;,


Floristic studies provide valuable information on species richness in a region, and are particularly important if these areas belong to less studied environments, such as rocky outcrops, that may increase our knowledge. An important aspect for species colonization includes the mechanisms of diaspores dispersal in each community; these are essential to understand its structure, dynamics, and the regeneration process, and constitute an important tool for conservation. We developed a floristic survey on a granite-gneiss outcrop with the objective to increase the knowledge on plant diversity, through a floristic similarity analysis and detection of dispersal syndromes of sampled species, in a semi-arid region of Brazil. The fieldwork included collection and observation of the botanical material in loco during a period of 12 months. A total of 161 species belonging to 127 genera and 50 families of angiosperms were recorded. Fabaceae, Asteraceae and Convolvulaceae were the most representative families in number of species. Allophylus quercifolius (Mart.) Radlk. (Capparaceae) and Lafoensia pacari A. St.-Hil. (Lythraceae) represented new records for the State of Paraiba. The autochoric syndrome was the most representative, with 51.5 % of the recorded species; the anemochory was the second most representative syndrome with 26.7 % of the species; and finally the zoochory, representing 22.3 % of the species. The floristic similarity dendrogram showed the formation of three well-defined groups, whose area with the highest value (J = 33.2) is located in a Caatinga region called Cariri Paraibano, while the lowest value observed (J = 5.2), occurred in a settled area in two geomorphological units, a crystalline complex and a plateau region. These results may be due to the varying topographic conditions and edaphic heterogeneity arising from the specific geological formation of the region. These results yet demonstrate that, in rocky outcrops, abiotic syndromes represent an effective dispersion of its diaspores, favoring plant specie's colonization dynamics.

Key words: inselbergs; floristic similarity; diasporas; xeric corridor; South America


Los estudios florísticos proveen información valiosa sobre la riqueza de especies de una región, y son particularmente importantes si estas áreas pertenecen a ambientes menos estudiados como los afloramientos rocosos. Un aspecto importante para la colonización de especies incluye los mecanismos de dispersión de diáspo-ras en cada comunidad; estos son esenciales para entender la estructura, dinámica y el proceso de regeneración, constituyendo una herramienta importante para la conservación. En este estudio se realizó un levantamiento florístico de un afloramiento de granito-gneis con el objetivo de ampliar el conocimiento sobre la diversidad vegetal a través del análisis de similitud florística con la de otros afloramientos rocosos del nordeste brasileño y la detección de los síndromes de dispersión de las especies muestreadas en un área de la región semiárida de Brasil. Los trabajos de campo incluyeron recolección y observación de los materiales vegetales 'i" loco' durante el periodo de 12 meses. Un total de 161 especies pertenecientes a 127 géneros y 50 familias de angiospermas fueron registradas. Fabaceae, Asteraceae y Convolvulaceae fueron las familias más representativas en número de especies. Allophylus quercifolius (Mart.) Radlk. (Sapindaceae) y Lafoensia pacari A.St.-Hil. (Lythraceae) representan nuevas citas para el estado de Paraíba. El síndrome autocórico fue lo más representativo, con 51 % de las especies registradas; la anemocoria el segundo, con 27.7 % de las especies y finalmente la zoocoria, representando el 22.3 %. El dendrograma de similitud florística ha demostrado la formación de tres grupos bien definidos, cuya área con mayor índice (J = 33.2) está ubicada en una región de Caatinga llamada Cariri Paraibano mientras el menor índice "in loco" (J = 5.2) ocurrió en un área ubicada en dos unidades geomorfológicas: un complejo cristalino y una región de Planalto. Estos resultados se deben a las variables condiciones topográficas y a la heterogeneidad edáfica proveniente de la formación geológica especifica de la región. Estos resultados demuestran todavía que en afloramientos rocosos, síndromes abióticos representan una dispersión eficaz de sus diásporas, favoreciendo la dinámica de colonización de las especies vegetales.

Palabras clave: inselbergs; similitud florística; diasporas; pasillo xérico; América del Sur

With a wide distribution in tropical areas, inselbergs feature a range of sizes and degrees of isolation associated with major biomes of the world, allowing experimental work, as well as being excellent elements to address different topics related to biodiversity and conservation (Porembski & Barthlott, 2000; Romer, 2005).

Due to its own characteristics, inselbergs form centers of diversity for certain functional groups of plants that are well adapted to extreme environmental conditions, and where the occurrence of seeds adapted to these conditions, becomes also greater in these formations than in the adjacent matrix (Hunter, 2003; Porembski, 2007). Its rocky structure is capable of forming xeric islands within a tropical rainforest matrix, resisting harsh soil edaphic and microclimate conditions (Sarthou, Larpinb, Fontyc, Pavoined, & Ponge, 2010).

Studies in outcrops of Brazilian semiarid region showed that, while the dominant climate is the semi-arid, the formation of semi-deciduous forests may happen, which characterizes the local occurrence of wetter environments, favoring the establishment of phanerophytes (Franga, Melo, & Santos, 1997). According to Araujo, Oliveira & Lima-Verde (2008), the vegetation that is established on rocky outcrops in the semi-arid region has a high species richness, demonstrating the relevance of studies related to the biology and ecology of these species, and the conservation actions for these areas (Scarano, 2007).

Fig. 1 Location map of the study area, Parque das Pedras, Pocinhos, Paraiba state, Brazil. 

Due to the absence of large substrate accumulations, water storage becomes incipient and much of it is rapidly lost by runoff. Due to the little substrate available and extreme microclimate conditions, the vegetation that is established in these environments is very different from its surroundings, favoring even the development of ephemeral vegetation (Sarthou & Villiers, 1998). Environmental conditions inherent to these formations are capable of providing niches to different taxa, depending on the surrounding vegetation and environmental pressure. In such harsh environments, the presence of effective dispersive mechanisms is of paramount importance for the development and establishment of seedlings (Willson & Traveset, 2000).

Based on the premise that short distances associated with abiotic syndromes may influence the floristic composition in rocky environments, this study aimed to verify the composition, and to perform a floristic similarity analysis and detect dispersion syndromes of the species of a granite-gneiss outcrop, situated in the semi-arid tropic in Northeastern Brazil. This study sought to answer the following questions: a) what is the degree of similarity/ dissimilarity of this area in relation to other areas with rocky environments already studied in Northeastern Brazil?; b) do the dispersal syndromes corroborate the patterns found in outcrops of other arid or semiarid regions?

Material and Methods

Study area: The outcrop selected for this study is located in the municipality of Pocinhos, Agreste mesorregion of Paraiba state, in the semi-arid Northeastern Brazil. It is located in a private estate with approximately 55 ha known as Parque das Pedras (07° 05' 14'' S - 36° 03' 56'' W), and, although it is open to tourism, it is visibly preserved (Fig. 1). The Agreste is one of mesorregion of the state of Paraiba. It is characterized as a climatic transition area where winds heated in the depression zone ascends and cools up, causing rainfall (Rodriguez, 2000), with annual average rates ranging from 600 to 1 000 mm (AESA, 2014). According to the new updated classification of Koppen-Geiger (Peel, Finlayson, & McMahon, 2007), the climate in this region is characterized as BSh, hot semi-arid, and the annual precipitation rates are close to littoral rates (Rodriguez, 2000). It presents well-developed soils with clayey characteristics (Oliveira & Oliveira, 2008). The vegetation of this unit consists of semidecidu-ous and deciduous forests, particular to areas of Agreste (Beltrao et al., 2005).

Floristic survey: For the record of the species that occur in the outcrop, 16 excursions were made in the period from December 2012 to November 2013, consisting of monthly collections in the dry season and bimonthly collections in the rainy season throughout the study area. Fertile branches (flowers and/or fruits) of individuals belonging to all strata of vegetation were obtained (Fig. 2 and Fig. 3). The specimens were processed according to the usual techniques (Judd, Campbel, Kellogg, Stevens, & Donoghue, 2009) and led to dry in the Herbarium Manuel de Arruda Camara (ACAM) of the State University of Paraiba (UEPB), Campina Grande, Paraiba state, Brazil. The taxonomic identification was based on the taxonomic literature or by comparison with specimens, identified by experts, belonging to the collections of herbaria in the state of Paraiba (EAN, JPB) and in the Virtual Herbarium of the Brazilian Flora (REFLORA). Acronyms of the herbaria were mentioned according to Holmgren, Holmgren and Barnett (1990). Specimens and scanned images also were sent to confirmation and/or identification by experts in different taxonomic groups in Brazil. Exsiccatae were incorporated into the collection of the Herbarium Manuel de Arruda Cámara (ACAM). The species were listed in alphabetical order by family, according to the 2). The spelling of the names of species and their respective authors were found in the database of the Missouri Botanical Garden (Tropicos, 2013) and in the online Species List of Brazilian Flora (Forzza et al., 2014). This work has been included exclusively the native species and for this reason, the only exotic species recorded in the study area was not included in the floristic list.

Fig. 2 Species recorded in the studied rocky outcrops: A) Jatropha mollissima. B) Tacinga inamoena. C) Cyrtopodium holstii. D) Lantana camara. E) Angelonia campestris. F) Lafoensia pacari

Dispersal syndromes: To determine dispersal syndromes, young and mature fruits of almost all species recorded in outcrop were collected, except for Lippia grata Schauer (Verbenaceae), Justicia aequilabris (Nees) Lindau (Acanthaceae), Mitracarpus salzman-nianus DC., Staelia virgata (Link ex Roem. & Schult.) K. Schum. (Rubiaceae), and Evolvulus ovatus Fernald (Convolvulaceae) and their syndromes' determinations were conducted through taxonomic literature consulted or with expert assistance. For the other species, the syndromes were classified according to the work by Van der Pijl (1972), being classified into three categories: I-Anemochoric: when diaspores presented some sort of appendix with wings, feathers or in the form of dust; Il-Zoochoric: when some attractive element, such as, for example, strong colors and/or a food source in its diaspores, is presented, as well as those with adhesive structures in the form of hooks, bristles, spines or mucilage; III-Autochoric: when the diaspores showed no characteristic that allowed their classification in the previous two categories.

Fig. 3 Species recorded in the studied rocky outcrops: A) Jatropha mollissima. B) Pilosocereus pachycladus. C) Cyrtopodium holstii. D) Marsdenia caatingae. E) Angelonia campestris. F) Sapium argutum.  

Fig. 4 Dendrogram of floristic similarity linking the study area with 11 studies on rocky outcrops and other areas of the surrounding matrix (Caatinga vegetation). A1 - Present study, A2 - Gomes et al. (2011), A3 - Porto et al. (2008), A4 -Araujo et al. (2008), A5 - Franga et al. (2005), A6 - Tolke et al. (2011), A7 - Gomes & Alves (2009), A8 - Machado-Filho (2011), A9 - Lima (2012), A10 - Araujo et al. (2005), A11 - Barbosa et al. (2005), A12 - Araujo et al. (Ined. 2013). 

According to Van der Pijl (1972), the auto-choric dispersion may occur in two main forms: active, when the plant expels the diaspores with a ballistic action associated with dehiscence of the fruit; and passive, when they are carried by the movement of the sediment or by small animals. In the latter, species with explosive dispersion or by gravity (barochoric) were grouped. Of the species recorded in the area, two did not have their dispersal syndromes identified due to the absence of fruiting material or because they had immature fruits. All fruits collected in the study area were classified according to Spjut (1994), and, for species that did not fruit during fieldwork, the classification was determined based on the specific literature and with expert advice.

Similarity analysis: To calculate the flo-ristic similarity, originally a list consisting of the compilation of 11 floristic references was compiled using Excel software version 7.0, generating a presence/absence binary matrix with all elements identified at the species level. The selected studies included studies in rocky environments inserted in areas of Caatinga in states from the Northern and Southern portion of Northeast of Brazil (Table 1).

To avoid repetition and synonyms, the current nomenclature of each species was used. The dendrogram was obtained with the Primer 6.0 software, using the Jaccard index, based on the Unweighted Pair Group Method using Arithmetic averages (UPGMA) method.


Floristic: In the study area, 161 species belonging to 127 genera and 50 families of angiosperms were recorded (Table 2, Fig. 2 and Fig. 3). Among the species, five were identified up to the genus level. The most representative family was Fabaceae (24 spp.), followed by Asteraceae and Convolvulaceae, with 12 and 11 species, respectively; together, they correspond to 29 % of all recorded species.

Table 1: Floristic studies in semi-arid Northeastern Brazil compiled from the elaboration of the matrix of presence/absence of species 

Table 2: Floristic list containing names of families and species, types of fruits and their dispersal syndromes of species recorded in the study area, Parque das Pedras, Pocinhos, Paraiba state, Brazil 

Allophylus quercifolius (Mart.) Radlk (Sap-indaceae) and Lafoensia pacari A. St.-Hil. (Lythraceae) constituted new records for the state of Paraiba. Regarding monocots, the Bro-meliaceae family should be highlighted; it was represented by five genera and seven species, being Tillandsia the most representative genus, with three species. Within Orchidaceae, represented by five species and three genera, Cyrto-podium is the most representative genus with two species. Poaceae, Cyperaceae and Araceae totaled 3.1 % of the total species recorded.

With regard to habits, the herbaceous stratum was the most representative, with 54 % of the total composition of the outcrop. Shrub (15 %) and subshrubs (12 %) components make up together 27% of all recorded species, and vines and epiphytes total 12.5 %. Three species of hemiparasites belonging to two families were recorded: Phoradendron affine Pohl ex DC., Phoradendron piperoides (Kunth) Trel. (Santalaceae) and Struthanthus margin-atus (Desr.) Blume (Loranthaceae).

Similarity analysis: The similarity dendrogram showed the formation of well-defined clusters and a block whose areas appear forming similar subgroups among them (Fig. 4). The first group consists of rocky outcrops located in the municipality of Puxinana (Araujo et al., 2013, unpublished data- A12, Tolke, Silva, Pereira & Melo, 2011 - A6) with an index of floristic similarity between areas of J = 30. The studied area (A1) demonstrated greater affinity with this group, being presented, in the dendrogram, as its subgroup. This floristic relationship can be explained by the fact that the whole work was done in areas geographically close together. The second group comprises the works of Machado-Filho (2011) - A8 and Lima (2012) - A9 (both with unpublished data), showing the highest similarity index (J = 33.2), being these works developed in a region called Cariri, located in Northeastern Brazil. The third group comprises the studies of Barbosa, Lima, Agra, Cunha & Pessoa (2005) - A11, held at Eastern Curimatau of Paraiba state and Gomes, Costa, Rodal & Alves (2011) - A2, in the semi-arid region of the state of Pernambuco.

Some areas settled in the Caatinga domain formed subgroups apart from the rest. These areas include rocky habitats (Franga et al., 2005; Araújo et al., 2008; Porto, Almeida, Pessoa, Trovao, & Félix, 2008; Gomes & Alves, 2009), and one study was conducted in the backlands ("sertao") of Ceará State by Araújo, Costa, Figueiredo & Nunes (2005), being the most dissimilar (J = 0.6) among selected studies when compared to this study.

Dispersal syndromes: With respect to the dispersion mode of the diaspores, the results showed a predominance of abiotic syndromes, where authocory was the most representative totaling 51.5 % (83 spp.) of total species. This mode of dispersion is the result of the plant's own mechanisms through explosive dehiscence of dried fruits or by gravity. The second most representative syndrome is anemochory, with 26.7 % (43 spp.) of the recorded species, followed by zoochory, comprising 22.3 % (36 spp.). In this category, plants depend on animals to disperse as far as possible from the mother-plant.


Bromeliaceae, Poaceae and Cyperaceae are very representative families in outcrops in South America (Sarthou & Villiers, 1998). However, Poaceae and Cyperaceae, in this study, were represented by two species each. Melinis minutiflora P. Beauv. (Poaceae) was one of the species found in the outcrop, but, because it is an exotic and invasive species, it was not included in the floristic list. Bro-meliaceae is represented by seven species, and the genera Encholirium and Tillandsia occur throughout the outcrop. This pattern of occurrence for the above mentioned genera was also observed by Franga, Melo and Gongalves (2006).

Corroborating the works of Araújo et al. (2005), Barbosa et al. (2005), Lima, Sampaio, Rodal and Araújo (2009) and Santos & Melo (2010), Fabaceae was the family best represented, with 24 species, corresponding to 15 % of species richness in the studied area. Considered the most taxonomically diverse of the Caatinga, this family is the third largest among angiosperms, whose economic importance makes it even more known (Judd et al., 2009).

Asteraceae was the second best represented family in number of species and its wide distribution, particularly common in open areas (Souza & Lorenzi, 2012), illustrates the predominance of its species in its different aspects, represented in the works of Lima (2012), Neves & Conceigao (2007) and Zappi et al. (2003).

Cactaceae, with five species, proved to be frequent in the study area, showing species with herbaceous to arborescent habits. Among the species found, Melocactus zehntneri (Britton & Rose) Luetzelburg is noteworthy because it is threatened by exploration and/or by anthropic pressure on their habitats (Fabricante, Andrade, & Marques, 2010). According to these authors, M. zehntneri plays a very important ecological role in the succession process, being able to colonize xeric environments and make them less harsh.

With regard to the habit, the tree component was the least representative, comprising 9 % of the total and, in that sense, Caiafa and Silva (2007) and Safford and Martinelli (2000) note that some outcrops showed shallow rock fractures, forming microenvironments characterized by the accumulation of 5 to 12 cm of soil, thus limiting the development of this stratum.

In the dendrogram, the highest value was found among the areas studied by Machado-Filho (2012) and Lima (2012), both settled in the Environmental Protection Area (EPA) of Cariri, in the semi-arid region of the state of Paraiba, Brazil. This close relation was expected, given that the studies were developed in the same geographical area between the municipalities of Cabaceiras and Boa Vista, Paraiba, whose climate and soil conditions are very similar and where rocky outcrops predominate.

The studies conducted in rocky environments settled in the city of Puxinana, in Agreste mesorregion of Paraíba (Tólke et al., 2011; Araújo et al., 2013, unpublished data), showed a floristic relation (J = 31) closer with each other rather than with the studied area (J = 19). However, it was expected, because they are geographically nearby areas, that the similarity value between them would be higher than the one recorded (Kunz, Ivanauskas, Martins, Silva, & Stefanello, 2009; Oliveira & Nelson, 2001). This dissimilarity can be explained partly by species recording, which occurred only in this study: Chloroleucon dumosum (Benth.) G.P. Lewis, Cuphea impatientifolia A. St.-Hil., Helicteres eichleri K. Schum., Jacquemontia corymbulosa Benth. andXimenia americana L.

The areas studied by Barbosa et al. (2005) and Gomes et al. (2011) formed a subgroup with a similarity value of J = 20, both located in the Agreste of the states of Paraíba and Pernambuco, respectively, showing weather conditions with erratic rainfall, varying from 664 mm to 1 054 mm, and similar frequency of rock formations. The most representative families in the two areas were Fabaceae and Euphorbiaceae, common in areas of caatinga (Cardoso & Queiroz, 2007; Sátiro & Roque, 2008), confirming the results obtained in the area under study.

The other areas had low similarity values (J < 20), in particular the study by Araújo et al. (2005), conducted in the Natural Reserve of Serra das Almas, CE, where three different formations were explored, contemplating areas from dense scrub vegetation to thorny deciduous trees and shrubs, with a dissimilarity between the flora of the surrounding matrix and the flora that is established on rocky outcrops. This vegetation is probably severely influenced by strict environmental aspects, demanding a greater adaptation of individuals, which contributed to this dissimilarity (Porembski &amp; Barthlott, 2000).

Species such as Jatropha mollissima (Pohl) Baill. and Cnidosculus urens (L.) Arthur -Euphorbiaceae-, recorded in this study, have a passive authocory mechanism (Leal, 2003). According to the same author, seeds released by gravity may be secondarily carried to other areas by ants. Sapium argutum (Müll. Arg.) Huber (Euphorbiaceae) presents voluminous seeds, with vivid colors, and, although not mentioned in the study, it is possible that its seeds also present passive authocory by dispersion done by ants or even by sediment carrying along the outcrop.

The anemochoric syndrome includes 26.7 % of the recorded species' total, corroborating the data exposed by the literature, where areas of greatest exposure with low water levels show a predominance of abiotic vectors (Butler, Green, Lamb, McDonald, &amp; Forster, 2007; Barbosa, Silva, &amp; Barbosa, 2002; Griz &amp; Machado, 2001; Machado, Barros, &amp; Sampaio, 1997). Van der Pijl (1972) points out those taxa with wind dispersal present various structures that facilitate its spread. In addition, regarding anemochoric species collected in the study area, winged and feathery seeds were the most frequently observed.

The microclimate aridity of these rocky islands in relation to the surrounding area reflects the predominance of abiotic syndromes, as they represent habitats with a greater exposure to winds and sunlight (Araújo et al., 2008). The zoochorous syndrome amounted to 22.3 %, with most species that falls into this category being collected in wetlands surrounding the studied rocky outcrop. These conditions reflect what was found in the study by Silva and Rodal (2009), after the detection of dispersal syndromes in three areas with different rainfall regimes in the state of Pernambuco, Northeastern Brazil. It can be inferred that there is a gradual change in the spread spectrum of wetlands, dominated by species with zoochorous syndromes.

Thus, conducting floristic surveys associated with ecological aspects, such as the recognition of syndromes, provide a better understanding of the dynamics of species colonization, mainly with regard to the Caatinga vegetation in Northeastern Brazil. The results obtained in this study, compared to other similar studies (Araújo et al., 2008; Barbosa et al., 2002; Lima, 2012; Silva &amp; Rodal, 2009), showed that abiotic syndromes (anemochory or authocory) are prevalent in outcrops and dry forests, supporting patterns found in outcrops in other regions of arid or semi-arid climates. Similar edaphic-climatic conditions are those that best explain the floristic similarity between the compared areas, especially those closer to each other. The combination of short distances and the predominance of abiotic syndromes suggest this affinity in the floristic composition, as observed in the works by Machado-Filho (2011) and Lima (2012).


To the Graduate Program (MSc) in Ecology and Conservation of the State University of Paraíba (PPGEC/UEPB) for the opportunity to carry out this work. To CAPES (Coordination for the Improvement of Higher Education Personnel) for granting a master's scholarship. To the specialists Jefferson Maciel and Carlos Alberto Garcia for determining Poaceae species; Teresa Buril for the identification of Convolvulaceae and to Amanda Coelho for the determination of Cactaceae species. J. I. M. Melo thanks CNPq (National Council for Scientific and Technological Development) for the Productivity Research Fellowship (PQ-2 Proc. no. 302751/2012-2).


Agencia Executiva de gestao das águas do estado da Paraíba (AESA). (2014). Geoportal Retrieved from http:// [ Links ]

The Angiosperm Phylogeny Group (APG). (2009). An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161105-12. [ Links ]

Araújo, F. S., Costa, R. C., Figueiredo, M. A., &amp; Nunes, E. P. (2005). Vegetacao e flora fanerogámica da área Reserva Serra das Almas, Ceará. In F. S. Araújo, M. J. N. Rodal, &amp; M. R. V. Barbosa (Eds.), Análise das variagoes da biodiversidade do bioma Caatinga (pp. 91-119). Recife, Brasil: Ministério do Meio Ambiente. [ Links ]

Araújo, F. S., Oliveira, R. F., &amp; Lima-Verde, L. W. (2008). Composicao, espectro biológico e síndromes de dis-persao da vegetacao de um inselbergue no domínio da caatinga, Ceará. Rodriguésia 59659-671. [ Links ]

Barbosa, D. C. A., Silva, P. G. G., &amp; Barbosa, M. C. A. (2002). Tipos de frutos e síndromes de dispersao de espécies lenhosas da caatinga de Pernambuco. In M. Tabarelli &amp; J. M. C. Silva (Eds.), Diagnóstico da Biodiversidade de Pernambuco (pp. 609-621). Recife, Brasil: Secretaria de Ciencia, Tecnologia e Meio Ambiente, Fundacäo Joaquim Nabuco e Editora Massangana. [ Links ]

Barbosa, M. R. V., Lima, R. B., Agra, M. F., Cunha, J. P., &amp; Pessoa, M. C. R. (2005). Vegetacao e flora fanerogámica do Curimataú, Paraíba. In F. S. Araújo, M. J. N. Rodal, &amp; M. R. V. Barbosa (Eds.), Análise das variagoes da biodiversidade do bioma Caatinga (pp. 122-138). Brasilia, Brasil: Ministério do Meio Ambiente. [ Links ]

Beltrao, B. A., Morais, F., Mascarenhas, J. C., Miranda, J. L. F., Junior, L. C. S., &amp; Mendes, V. A. (2005). Diagnóstico do municipio de Pocinhos. Projeto cadastro de fontes de abastecimento por água subterránea Estado da Paraiba Recife, Brasil: CPRM/ PRODEEM. [ Links ]

Butler, D. W., Green, R. J., Lamb, D., McDonald, W. J. F., &amp; Forster, P. I. (2007). Biogeography of seed-dispersal syndromes, life-forms and seed sizes among woody rain-forest plants in Australia's subtropics. Journal of Biogeography 341736-1750. [ Links ]

Caiafa, A. N., &amp; Silva, A. F. (2007). Structural analysis of the vegetation on a highland granitic rock outcrop in Southeast Brazil. Revista Brasileira de Botánica 30657-664. [ Links ]

Cardoso, D. B. O. S., &amp; Queiroz, L. P. (2007). Diversidade de Leguminosae nas caatingas de Tucano, Bahia: implicacöes para a fitogeografia do semi-árido do nordeste do Brasil. Rodriguésia 58379-391. [ Links ]

Fabricante, J. R., Andrade, L. A., &amp; Marques, F. J. (2010). Caracterizacao populacional de Melocactus zehntneri (Britton &amp; Rose) Luetzelburg (Cactaceae) ocorrente em um inselbergue da Caatinga paraibana. Biotemas 2361-67. [ Links ]

Forzza, R. C., Stehmann, J. R., Nadruz, M., Filardi, F. L. R., Costa, A., Carvalho-Junior, A. A., Peixoto, A. L., &amp; Walter, B. M. T. (2014). Lista de Espécies da Flora do Brasil. Rio de Janeiro, Brasil: Jardim Botánico do Rio de Janeiro. Retrieved from http://floradobrasil. ]

Franca, F., Melo, E., &amp; Santos, C. C. (1997). Flora de inselbergs da Regiao dos milagres, Bahia, Brasil: Caracterizacao da vegetacao e lista de espécies de dois inselbergs. Sitientibus 17163-184. [ Links ]

Franca, F., Melo, E., Santos, A. K. A., Melo, J. G. A. N., Marques, M., Silva-Filho, M. F. B., &amp; Machado, C. (2005). Estudos ecológicos e floristico em ilhas de vegetacao de um inselberg no semiárido da Bahia, Brasil. Hoehnea 3293-101. [ Links ]

Franca, F., Melo, E., &amp; Goncalves, J. M. (2006). Aspectos da diversidade da vegetacao no topo de um inselber-gue no semiárido da Bahia, Brasil. Sitientibus Série Ciencias Biológicas, 630-35. [ Links ]

Gomes, P., &amp; Alves, M. (2009). Floristic and vegetational aspects of an inselberg in the semi-arid region of Northeast Brazil. Edinburgh Journal of Botany 66, 329-346. [ Links ]

Gomes, P., Costa, K. C., Rodal, M. J. N., &amp; Alves, M. (2011). Checklist of Angiosperms from the Pedra Furada Municipal Park, Northeastern Brazil. Check List 7173-181. [ Links ]

Griz, L. M. S., &amp; Machado, I. C. S. (2001). Fruiting phenology and seed dispersal syndromes in caatinga, a tropical dry forest in the Northeast of Brazil. 303-321. [ Links ]

Holmgren, P. K., Holmgren, N. H., &amp; Barnett., L. C. (1990). Index Herbariorum 8The Herbaria of the worldNew York: The New York Botanical Garden. [ Links ]

Hunter, J. T. (2003). Persistence on inselbergs: the role of obligate seeders and resprouters. Journal of Biogeography 30497-510. [ Links ]

Judd, W. S., Campbel, C. S., Kellogg, E. A., Stevens, P. F., &amp; Donoghue, M. J. (2009). Sistemática Vegetal: Um enfoque filogenètico. Porto Alegre, Brasil: Artmed. [ Links ]

Kunz, S. H., Ivanauskas, N. M., Martins, S. V., Silva, E., &amp; Stefanello, D. (2009). Análise da similaridade florística entre florestas do Alto Rio Xingu, da Bacia Amazónica e do Planalto Central. Revista Brasileira de Botánica 32725-736. [ Links ]

Leal, I. R. (2003). Dispersao de sementes por formigas na Caatinga. In I. R. Leal, M. Tabarelli, &amp; J. M. C. Silva (Eds.). Ecologia e Conservagäo da Caatinga (pp. 593-624). Recife, Brasil: Universidade Federal de Pernambuco. [ Links ]

Lima, E. A. (2012). Estudo floristico da APA do Cariri, Paraiba, Brasil: Riqueza, similaridade e sindromes de dispersäo (Dissertacao de Mestrado). Universida-de Estadual da Paraíba, Paraíba, Brasil. [ Links ]

Lima, J. R., Sampaio, E. V. S. B, Rodal, M. J. N., &amp; Araú-jo, F. S. (2009). Composicao florística da floresta estacional decídua montana de Serra das Almas, CE, Brasil. Acta Botanica Brasilica 23756-763. [ Links ]

Machado, I. C. S., Barros, L. M., &amp; Sampaio, E. V. S. B. (1997). Phenology of Caatinga Species at Serra Tal-hada, PE, Northeastern Brazil. Biotropica 2957-68. [ Links ]

Machado-Filho, H. O. (2011). Estudo floristico de um ambiente rochosos da área de protegäo ambiental (APA) do Cariri, Paraiba: Riqueza, similaridade e fitogeografia (Dissertacao de Mestrado). Universida-de Estadual da Paraíba, Paraíba, Brasil. [ Links ]

Neves, S. P. S., &amp; Conceicao, A. A. (2007). Vegetacao em afloramentos rochosos na Serra do Sincorá, Chapada Diamantina, Bahia, Brasil. Sitientibus Ciencias Biológicas 736-45. [ Links ]

Oliveira, A. A., &amp; Nelson, B. W. (2001). Floristic relationships of terra firme Forest in the Brazilian Amazon. Forest Ecology and Management 146169-179. [ Links ]

Oliveira, I., &amp; Oliveira, C. (2008). Paraiba: meu espago, minha paisagem: Geografia Curitiba, Brasil: Base Editora. [ Links ]

Peel, M. C., Finlayson, B. L., &amp; McMahon, T. A. (2007). Updated world map of the Köppen-Geiger climate classification. Hydrology and Earth System Sciences Discussions 4439-473. [ Links ]

Porembski, S., &amp; Barthlott, W. (2000). Why study Inselbergs? In S. Porembski, &amp; W. Barthlott (Eds.), Inselbergs: biotic Diversity of Isolated Rock Outcrops in Tropical and Temperate Regions (pp. 1-6). New York: Springer. [ Links ]

Porembski, S. (2007). Tropical inselbergs: habitat types, adaptive strategies and diversity patterns. Revista Brasileira de Botánica 30579-586. [ Links ]

Porto, P. A. F., Almeida, A., Pessoa, W. J., Trovao, D., &amp; Félix, L. P. (2008). Composicao florística de um inselbergue no agreste paraibano, município de Esperanca, nordeste do Brasil. Revista Caatinga 21214-223. [ Links ]

Rodríguez, J. L. (2000). Atlas escolar da Paraiba. Espago geo-histórico e cultural Joao Pessoa, Brasil: Ed. Grafset. [ Links ]

Romer, W. (2005). The distribution of inselbergs and their relationship to geomorphological, structural and lithological controls in Southern Zimbabwe. Geomorphology 72156-176. [ Links ]

Safford, H. D., &amp; Martinelli, G. (2000). Southeast Brazil. In S. Porembski &amp; W. Barthlott (Eds.), Inselbergs: biotic Diversity of Isolated Rock Outcrops in Tropical and Temperate Regions (pp. 339-389).. New York: Springer [ Links ]

Sarthou, C., &amp; Villiers, J. F. (1998). Epilithic plant communities on inselbergs in French Guiana. Journal of Vegetation Science 9847-860. [ Links ]

Sarthou, C., Larpinb, D., Fontyc, É., Pavoined, S., &amp; Ponge, J. F. (2010). Stability of plant communities along a tropical inselberg ecotone in French Guiana (South America). Flora 205682-694. [ Links ]

Santos, A. C. J., &amp; Melo, J. I. M. 2010. Flora vascular de uma área de caatinga no estado da Paraíba-Nordeste do Brasil. Revista Caatinga 2332-40. [ Links ]

Sátiro, L. N., &amp; Roque, N. (2008). A familia Euphorbiaceae nas caatingas arenosas do médio rio Sao Francisco, BA, Brasil. Acta Botanica Brasilica 2299-118. [ Links ]

Scarano, F. R. (2007). Rocky outcrop vegetation in Brazil: a brief overview. Revista Brasileira de Botánica 30561-568. [ Links ]

Silva, M. C. N. A., &amp; Rodal, M. J. N. (2009). Padroes das síndromes de dispersao de plantas em áreas com diferentes graus de pluviosidade, PE, Brasil. Acta Botanica Brasilica 231040-1047. [ Links ]

Souza, V. C., &amp; Lorenzi, H. (2012). Botánica Sistemática: Guia ilustrado para identificagäo das familias de fanerógamas nativas e exóticas no Brasil, baseado em APG III. Nova Odessa, Brasil: Instituto Plantarum. [ Links ]

Spjut, R. W. (1994). A Systematic treatment of fruit types New York: The New York Botanical Garden. [ Links ]

Tölke, E. E. A. D., Silva, J. B., Pereira, A. R. L., &amp; Melo, J. I. M. (2011). Flora vascular de um inselbergue no estado da Paraíba, Nordeste do Brasil. Biotemas 2439-48. [ Links ]

Tropicos. (2013). Missouri Botanical Garden. Retrieved from [ Links ]

Van der Pijl, L. (1972). Principles of dispersal in higher plants . New York: Springer [ Links ]

Willson, M. F., &amp; Traveset, A. (2000). The ecology of seed dispersal. In M. Fenner, (Ed.), Seeds: the ecology of regeneration in plant communities (pp. 85-1102). Wallingford: Ed. CABI. [ Links ]

Zappi, D. C., Lucas, E., Stannard, B. L., Lughadha, E. N, Pirani, J. R., Queiroz, L. P., &amp; Carvalho, A. M. (2003). Lista das plantas vasculares de Catolés, Chapada Diamantina, Bahia, Brasil. Boletim de Botánica da Universidade de Sao Paulo 21345-398. [ Links ]

Received: September 16, 2014; Revised: May 12, 2015; Accepted: June 09, 2015

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