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Biological spectrum and dispersal syndromes in an area of the semi-arid region of north-eastern Brazil/Espectro biologico e sindromes de dispersao em uma area do semiarido do nordeste brasileiro.

Introduction

The strong seasonality in semi-arid regions requires that species adopt strategies for their survival during the driest periods. For instance, certain plant species adapt temporary life forms which produce vegetative buds that are protected until environmental conditions allow the development of the aerial parts (MANTOVANI; MARTINS, 1988). This close relationship between environmental conditions, prevalent life-forms and a given area has generated a system of vegetation classification based on the degree of protection of vegetative buds and their position on the plant's principal axis with regard to the substrate.

Proposed by Raunkiaer (1934), the system classifies plants according to forms used to protect their perennating buds during unfavorable seasons, and groups them into five main classes: therophytes, cryptophytes, hemicryptophytes, chamaephytes and phanerophytes, according to their different resistance strategies to environmental conditions.

Studies carried out in the caatinga region have demonstrated the predominance of therophytes over other life forms, at a proportion close to 40%, as underscored by Costa et al. (2006) and Porto et al. (2008). A high proportion of phanerophytes and chamaephytes, close to 30 and 15%, respectively, were also registered.

Dispersal syndromes comprise a series of morphological characters of fruit or seeds that, associated with the biology of the disperser, determines a specific mode of dispersal, which in turn is an adaptation that promotes the establishment, development and endurance of the vegetative species in a given environment (VAN DER PIJL, 1982; RENNER, 1987; VASCONCELOS et al., 2010). Fruiting and dispersal are important stages of the reproductive cycle of angiosperms, with the dispersal process being the most sensitive and critical stage in a plant life history (BAWA et al., 1989).

From an ecological viewpoint, the dispersal of diaspores is a process of great importance for plant species because it enables them to expand their area of occurrence, decreases intraspecies competition and allows increase of their genetic variability into the population (HOWE; MIRITI, 2004). Efficiency of the dispersal mode also translates into increased recruitment of seedlings, by reducing competition and predation of seeds (HOWE, 1993).

According to Van der Pijl (1982), the richness and spatial distribution of plant populations are determined by dispersal modes and the frequency the diaspores reach favorable locations for the establishment of seedlings. In this context, Terborgh et al. (2002) found that the geographic distribution patterns of species and community structures depended on interactions between the community's biotic and abiotic components. The natural dispersal of diaspores has also an important role in the natural regeneration of plant ecosystems, enabling the re-colonization of degraded areas (DEMINICIS et al., 2009). According to Gentry (1983), information on dispersal ecology represents an important step in the understanding of the structure and function of plant communities in Neotropical forests, as well as in the restoration of degraded areas.

The frequency of different dispersal strategies is influenced by several factors, among which water availability in the environment is prominent, as observed by Silva and Rodal (2009).

Data on the dispersal syndromes in semi-arid regions of northeast Brazil have been expanded over the last decade. Since the first studies it has been observed that anemochoric species predominate in dry forests, while in humid forests zoochory is more important (HOWE; SMALLWOOD, 1982; GENTRY, 1983). This pattern was also observed in studies conducted by Barbosa et al. (2002), Griz and Machado (2001) and Griz et al. (2002).

Current study describes the floristic biological spectrum and dispersal syndromes of the flora in a conservation area (the Cariri Environment Conservation Area) in the semi-arid region of the northeastern state of Paraiba, Brazil, to verify whether the syndromes of the studied area are similar to those registered in other areas of the Brazilian northeast Caatinga region.

Material and methods

Study area

Current study was developed on the Fazenda Salambaia, a rural area in the municipality of Boa Vista, in the central region of the Borborema plateau in the northeastern state of Paraiba, Brazil. According to Koeppen and Geiger (1928), the region's climate is Bshw or semi-arid hot climate, with a 8-11-month dry season, an average temperature of 26[degrees]C and average annual rainfall not exceeding 600 mm (MOREIRA, 1988). The soils of the region are of the brown soil without calcium type, with dystrophic regosol areas (EMBRAPA, 2006).

The farm lies within the Cariri conservation area, specifically established to protect the scenic beauty of the region. The area comprises many rock formations where archaeological sites containing cave paintings and other records of prehistoric civilizations have been discovered. The vegetation of the studied area is characterized by sparse trees, shrubs and sappy thorny plants, or rather, predominantly dry shrub forest with xerophytic characteristics. Although it was not clear-cut in the last 20 years (according to information from a local resident), the local vegetation has been and is still impacted by goats, sheep and cattle grazing and trampling.

Field and laboratory procedures

Samples, collected every two weeks, between July 2010 and October 2011, consisted of fertile branches from individual trees, shrubs and herbaceous plants during walks through the area under analysis. After taxonomic identification, all species were classified by life form, according to a modified version (MUELLER-DOMBOIS; ELLENBERG, 1974) of Raunkiaer's (1934) classification system. So that the floristic biological spectrum of the vegetation could be determined, species were also classified according to fruit type, based on the morphological characteristics described by Van der Pijl (1982) and by Barroso et al. (1999), as well as the dispersal strategy adopted by each species, using specialized literature. The floristic list was prepared according to APG III (2009) and the species names and respective authors were verified in Forzza et al. (2014).

Following Van der Pijl (1982), the syndromes were classified into three groups: (a) zoochorous, when diaspores are dispersed by animals; for example, those with sweet flesh, and seeds with arils; species dispersed by insects, vertebrates and man are included in this group; (b) anemochorous, when the diaspores are adapted to wind dispersal, featuring structures such as feathers and wings; (c) autochorous, when the plants have their own dispersion mechanisms: seeds are either launched on the surrounding areas by any particular mechanism or they are simply released by the plant directly on the ground, or barochoric (gravity) which comprise species with explosive dispersal or dispersal by gravity.

Based on this dispersal syndrome classification, the community profile of a given area may be described according to the relative proportion of each dispersal strategy. The profile corresponds to the dispersion spectrum. However, it must be underscored that the object in current study is only the primary dispersal of diaspores. In other words, secondary events after departure from the parent plant were not observed. Following Barroso et al. (1999), a morphological classification of the fruit of the species that fruited during the period of data collection period was performed to characterize the syndromes.

Results

One hundred and sixty-six species, belonging to 123 genera and 41 families of angiosperms, were identified. Further, eight taxa were identified only at genus level and four taxa only at family level (Table 1). The richest family was Fabaceae, with 27 species recorded in this study.

In terms of the floristic biological spectrum among the 166 species, therophytes (46) were predominant over other life forms, representing 27.7% of the total, whereas phanerophytes (39) and chamaephytes (37) respectively represented 23.5 and 22.3% (Figure 1). Fifteen species of hemicryptophytes have also been registered. They represent 9% of the cryptophytes species, or 0.6% of all the species recorded in the study area. The life form could not be determined in the remaining 28 species (16.9% of all recorded species).

When the dispersal modes of diaspores are taken into account (Figure 2), anemochorous is the predominant syndrome in the area, with 47 species, or 28.3% of total; followed by autochorous (43), or 25.9% of total, in which dispersal is the result of the plants' own mechanisms, such as explosive dehiscence of dry fruits or gravity. Only 39 species are zoochorous, with 23.5% of total. This factor depends on the activity of animals so that dispersal may occur within an adequate distance from the parent plant. Dispersal syndromes were not identified in 37, or 22.3% of the 166 species. This was due to the absence of fruit on the vegetative material collected which identified the type of fruit and consequently, the dispersal form adopted.

Twenty-two out of the 39 zoochorous species had fleshy fruit, such as the Cactaceae, whereas 17 species produced dry fruit, such as Boerhavia diffusa (Nyctaginaceae). Among the autochorous species, most had dry fruit, such as the Fabaceae species that produced legume or loment fruit types. All anemochorous species had dry fruits, featuring dispersal units (whether fruit or seeds) with tissue expansions such as wings or other structures that facilitated transport by wind, such as Serjania glabrata (Sapindaceae), Schinopsis brasiliensis (Anacardiaceae) and Cochlospermum vitifolium (Bixaceae).

Discussion

Results on dispersal syndromes show a predominance of abiotic vectors in dry areas and biotic vectors in more humid areas (FRANCKIE et al., 1974; HOWE; SMALLWOOD, 1982; GENTRY, 1983). According to the last author, there is a defined pattern in the frequency of dispersal syndromes where zoochory is the most frequent form in the species of Neotropical rainforests. This fact may be related to several factors, such as conditions of greater humidity and water availability in the environment, or the presence of different species of frugivorous animals, such as birds, rodents, bats and an array of invertebrate species, especially insects, that eat the fruit and disperse the seeds in the adjacent areas. In arid and semi-arid regions, the abiotic syndromes (anemochory and autochory) grow in importance, as several studies carried out on the caatinga have demonstrated (MACHADO et al., 1997; GRIZ; MACHADO, 2001; BARBOSA et al., 2002, 2003). In current study, the abiotic syndromes, autochory and anemochory, together represent 69.7% of the identified syndromes and corroborated the results of the previously mentioned investigations.

The pattern of dispersal syndromes is related to a greater frequency of dry fruit (dispersed on the ground by autochory or anemochory) in drier areas with marked seasonality, already detected in the Brazilian savannah (Cerrado) and reported by Batalha and Mantovani (2000), Figueiredo (2008) and Oliveira (1998). However, another study for the Cerrado by Costa et al. (2004) underscored a predominance of zoochorous species over other dispersal modes.

There is evidence that a gradual change in the dispersal spectrum exists, from humid areas, where the predominant species have zoochorous syndromes, to drier ones, where syndromes of autochory or anemochory are more common. The above has been reported by Silva and Rodal (2009) who documented the dispersal syndromes in three areas with different rainfall levels in northeast Brazil. Vicente et al. (2005) also registered differences in the frequencies of dispersal syndromes between humid and semi-arid locations.

Barbosa et al. (2002) studied the phenology of woody species in the caatinga and reported the existence of a close relationship between the rainy season and the predominance of zoochorous species, whereas a higher percentage of autochorous and anemochorous species were reported in the dry season. According to Gentry (1983), differences in the amount and temporal distribution of rainfall are the most outstanding difference between wet and dry tropical forests, reflected in dispersal ecology.

In the case of the exclusive occurrence of small-and medium-sized phanerophytes in the studied area, similar data have been reported by Van Rooyen et al. (1990) for a semi-arid region in South Africa. According to these authors, the biological spectrum of species in an area indicates the survival strategy adopted by the local flora, while a high percentage of therophytes species represents an effective method for controlling water loss since the plant dies at the beginning of the dry season and avoids the water stress. Further, the seeds produced during its short life remain in the environment protected from desiccation by the seed coat and will grow when environmental conditions become favorable again, featuring an efficient escape strategy.

With regard to the life forms in the area under analysis, the therophytes form predominated and confirmed records in the literature showing the importance of this life form in the biological spectrum of other caatinga areas (ARAUJO et al., 2005; RODAL et al., 2005; COSTA et al., 2006). It should be emphasized that the marked predominance of small- and medium-sized phanerophytes has been recorded in several studies on caatinga areas (ARAUJO et al., 1995; ALCOFORADO-FILHO et al., 2003; FERRAZ et al., 2003).

Conclusion

Results of current investigation corroborate the few studies that address the biological spectrum and dispersal syndromes in caatinga areas and pointed the need for species to adopt strategies for their survival during the driest periods. It may also be inferred that both the biological spectrum and the dispersal syndromes are of fundamental importance for understanding the structure and function of this phytocenose and in particular the caatinga vegetation of northeast Brazil, perhaps one of the areas on the planet which is most threatened by anthropic action and yet one of the least known in terms of autoecology.

Doi: 10.4025/actascibiolsci.v37i1.23141

Acknowledgements

The authors would like to thank the State University of Paraiba (UEPB) for funding the project Vascular Flora of APA of Cariri, state of Paraiba, Brazilian northeast (Process no. 2011/057) coordinated by J.I.M. Melo. Thanks are also due to the National Council for Scientific and Technological Development (CNPq) for the granting of a Fellowship for Research Productivity (Process n. 302751/2012-2) to J.I.M. Melo, and to the referees for their valuable suggestions for the improvement of the text.

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Received on February 27, 2014.

Accepted on September 8, 2014.

License information: This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Elimar Alves de Lima * and Jose Iranildo Miranda de Melo

Departamento de Biologia, Centro de Ciencias Biologicas e da Saude, Universidade Estadual da Paraiba, Rua Baraunas, 351, 58429-500, Campina Grande, Paraiba, Brazil. * Author for correspondence. E-mail: tournefort@gmail.com
Table 1. Floristic list containing fruit types, respective dispersal
syndromes and life forms. Key: Fruit types. Cap = capsule; Ach =
achene; Fol = follicle; Utr = utricle; Dru = drupe; Sam = samara;
Cyp = cypsela; Ber = berry; Cer = ceratium; Sch= schizocarp;
Leg= legume; Lom= loment; Car= caryopsis; Coc = coccarium. Dispersal
syndromes types. Aut = autochory; Ane= anemochory; Zoo = zoochory.
Life forms. Cha = chamaephyte; The = therophyte; Hem = hemicryptophyte;
Pha = phanerophyte; Cry = cryptophyte. Collector: E.A. Lima.

                                                          Dispersal
Family/ Species                            Fruit type     syndrome

Acanthaceae
Dicliptera ciliaris Juss.                      Cap           Aut
Ruellia asperula (Mart. & Nees) Lindau         Cap           Aut
Ruellia bahiensis (Nees) Morong                Cap           Aut
Ruelliageminiflora Kunth                       Cap           Aut
Ruellia paniculata L.                          Cap           Aut
Alismataceae
Echinodorusgrandiflorus                        Ach           Ane
  (Cham. & Schltdl.) Micheli
Hydrocleys modesta Pedersen                    Fol            -
Amaranthaceae
Alternanthera brasiliana (L.) Kuntze           Utr           Ane
Alternanthera pungens Kunth                    Utr           Ane
Alternanthera tenella Colla                    Utr           Ane
Amaranthus spinosus L.                        Utri           Ane
Froelichia humboldtiana                       Utri           Ane
  (Roem. & Schult.) Seub.
Gomphrena vaga Mart.                           Cap           Ane
Anacardiaceae
Myracrodruon urundeuva Allemao                 Dru           Ane
Schinopsis brasiliensis Engl.                  Sam           Ane
Spondias tuberosa Arruda                       Dru           Zoo
Apocynaceae
Allamanda blanchetii A.DC.                     Fol           Ane
Aspidosperma pyrifolium Mart.                  Fol           Ane
Mandevilla tenuifolia                           -            Ane
  (J.C. Mikan) Woodson
Araceae
Pistia stratiotes L.                           Utr           Zoo
Asteraceae
Acanthospermum hispidum DC.                    Cyp           Zoo
Acmella uliginosa (Sw.) Cass.                  Cyp           Ane
Ageratum conyzoides L.                         Cyp           Ane
Bidens pilosa L.                               Cyp           Zoo
Centratherum punctatum Cass.                   Cyp           Ane
Chrysanthellum indicum DC.                      -             -
Conocliniopsisprasifolia (DC.)                 Cyp           Ane
  R.M. King & H. Rob.
Eclipta prostrata (L.) L.                      Cyp           Ane
Emilia sonchifolia (L.) DC.                    Cyp           Ane
Tilesia baccata (L.) Pruski                    Dru           Zoo
Bignoniaceae
Handroanthus impetiginosus                     Cap           Zoo
  (Mart. ex DC.) Mattos
Tabebuia aurea (Silva Manso) Benth.            Cap           Ane
  & Hook. f. ex S. Moore
Bixaceae
Cochlospermum vitifolium                       Cer           Ane
  (Willd.) Spreng.
Boraginaceae
Euploca procumbens (Mill.)                     Sch           Zoo
  Diane & Hilger
Heliotropium angiospermum Murray               Sch           Zoo
Varronia dardani (Taroda) J.S. Mill.           Dru           Zoo
Bromeliaceae
Bromelia baniosa Mart. ex Schult. f.           Cap           Ane
Encholirium spectabile Mart.                   Cap           Ane
  ex Schult. f.
Neoglaziovia variegata (Arruda) Mez.           Cap           Ane
Tillandsia recurvata (L.) L.                   Cap           Ane
Tillandsia streptocarpa Baker                  Cap           Ane
Burseraceae
Commiphora leptopHoeos (Mart.) Gillet.         Dru           Zoo
Cactaceae
Cereus jamacaru DC.                            Ber           Zoo
Melocactus ernestii Vaupel                     Ber           Zoo
Melocactus zehntneri                           Ber           Zoo
  (Britton & Rose) Luetzelb.
Pilosocereus gounellei (F.A.C.Weber)           Ber           Zoo
  Byles & G.D.Rowley
Pilosocereuspachydadus F. Ritter               Ber           Zoo
Tacinga inamoena (K. Schum.)                   Ber           Zoo
  N.P. Taylor & Stuppy
Tacinga palmadora (Britton & Rose)             Ber           Zoo
  N.P. Taylor & Stuppy
Capparaceae
Cynophala flexuosa (L.) J. Presl               Ber           Aut
Physostemon guianensis (Aubl.) Malme           Cer           Aut
Physostemon lanceolata                         Cer           Aut
  (Mart. & Zucc.) D.R. Hunt.
Tarenaya spinosa (Jacq.) Raf.                  Cer           Aut
Commelinaceae
Callisia filiformis (M. Martens                 -            Zoo
  & Galleotti) D. R. Hunt.
Callisia repens L.                             Cap           Aut
Commelina erecta L.                            Cap           Aut
Commelina obliqua Vahl                         Cap           Aut
Convolvulaceae
Evolvulusfdipes Mart.                          Cap            -
Evolvulus glomeratus Nees & C. Mart.           Cap            -
Ipomoea subincana (Choisy) Meisn.              Cap            -
Jacquemontia multiflora Haller f.              Cap            -
Operculina macrocarpa (L.) Urb.                Cap            -
Cyperaceae
Cyperus ligularis L.                           Ach           Aut
Cyperus odoratus L.                            Ach            -
Cyperus surinamensis Rottb.                    Ach            -
Cyperus uncinulatus Schrad. ex Nees            Ach            -
Eleocharis elegans (Kunth)                     Ach            -
  Roem. & Schult.
Eleocharis geniculata (L.)                     Ach           Zoo
  Roem. & Schult.
Eleocharis interstincta (Vahl)                 Ach            -
  Roem. & Schult.
Eragrostis ciliaris (L.) R.Br.                 Ach            -
Fimbristylis cymosa R. Br.                     Ach            -
Pycreus macrostachyos (Lam.) J. Raynal         Ach            -
Cucurbitaceae
Momordica charantia L.                         Cap           Zoo
Euphorbiacae
Cnidoscolus quercifolius Pohl                  Coc           Aut
Cnidoscolus urens (L.) Arthur                  Cap           Aut
Croton blanchetianus Baill.                    Coc           Aut
Croton heliotropiifolius Kunth                 Cap           Aut
Jatropha mollissima (Pohl) Baill.              Sch           Aut
Jatropha ribfolia (Pohl) Baill.                Sch           Aut
Manihot catingae Ule                           Sch           Aut
Fabaceae
Anadenanthera colubrina (Vell.) Brenan         Fol           Aut
Bauhinia cheilantha (Bong.) Steud.             Leg           Aut
Bauhinia subclavata Benth.                     Leg            -
Poincianella pyramidalis                       Leg           Aut
  (Tul.) L.P. Queiroz
Centrosema brasilianum (L.) Benth.             Leg           Aut
  var. angustifolium Amshoff
Centrosema brasilianum (L.) Benth.             Leg           Aut
  var. brasilianum Benth.
Centrosema virginianun (L.) Benth.             Leg           Aut
Chamaecrista rotundifolia                      Leg           Aut
  (Pers.) Greene
Desmodiumglabrum (Mill.) DC.                   Lom           Zoo
Dioclea grandiflora Mart. ex Benth.            Leg           Zoo
Dioclea violacea Mart. ex Benth.               Leg           Zoo
Erythrina velutina Willd.                      Fol           Zoo
Indigofera blanchettiana Benth.                Leg            -
Indigofera suffruticosa Mill.                  Fol           Aut
Libidibia ferrea (Mart. ex Tul.)               Leg           Aut
  L.P. Queiroz
Mimosa tenuiflora (Willd.) Poir.               Leg           Aut
Piptadenia stipulacea (Benth.) Ducke           Leg           Aut
Poincianella pyramidalis (Tul.)                Leg           Aut
  L.P. Queiroz
Senna macranthera (DC. ex Collad.)             Leg           Aut
  Irwin & Barneby
Senna martiana (Benth.) Irwin & Barneby        Leg           Ane
Senna obtusfolia (L.) Irwin & Barneby           -             -
Senna splendida (Vogel) Irwin & Barneby        Leg           Aut
Stylosanthes viscosa (L.) Sw.                  Lom           Zoo
Vigna adenantha (G.Mey) Marechal,              Leg            -
  Mascherpa & Stainer
Zorniagemella (Willd.) Vogel                   Lom           Zoo
Zornia leptophylla (Benth.) Pittier            Lom           Zoo
Zornia myriadena Benth.                        Lom           Zoo
Gentianaceae
Schultesia pohliana Progel                     Cap           Aut
Hydrocharitaceae
Apalanthegranatensis (Bonpl.) Planch.          Cap            -
Lamiaceae
Hyptis fruticosa Salzm. ex Benth.              Sch           Ane
Leonotis nepetfolia (L.) R.Br.                  -             -
Raphiodon echinus Schauer                       -             -
Loranthaceae
Struthanthus syringifolius (Mart.) Mart.       Ber           Zoo
Lythraceae
Cuphea campestres (Mart.) Koehne               Ber           Ane
Malvaceae
Ceiba glaziovii (Kuntze) K. Schum.             Cap           Ane
Herissantia tiubae (K. Schum.) Brizicky        Sch           Zoo
Melochia pyramidata L.                         Cap           Zoo
Melochia tomentosa L.                          Cap           Ane
Sidagalheirensis Ulbr.                         Sch           Aut
Sida Unifolia Cav.                             Sch           Aut
Sidastrum paniculatum (L.) Fryxell             Sch           Aut
Waltheria rotundifolia Schrank.                Cap            -
Waltheria tomentosa H.St.-John                  -             -
Molluginaceae
Mollugo verticillata L.                        Cap            -
Nyctaginaceae
Boehravia diffusa L.                           Ach           Zoo
Nymphaeaceae
Nymphaea ampla (Salisb.) DC.                   Cap            -
Onagraceae
Ludwigia leptocarpa (Nutt.) Hara               Cap           Aut
Ludwigia octovalvis (Jacq.) P.H. Raven         Cap           Aut
Oxalidaceae
Oxalis divaricata Mart. ex Zucc.               Cap            -
Passifloraceae
Passiflora foetida L.                          Ber           Zoo
Piriqueta duarteana (Cambess.) Urb.            Cap           Aut
Turnera subulata Sm.                           Cap           Aut
Plantaginaceae
Angelonia biflora Benth.                       Cap            -
Angelonia campestres Nees & Mart.              Cap            -
Angelonia cornigera Hook.                      Cap            -
Stemodia maritima L.                           Cap            -
Plumbaginaceae
Plumbago scandens L.                           Cap           Aut
Poaceae
Anthephora hermaphrodita (L.) Kuntze           Car           Ane
Chloris inflata Link                           Car           Ane
Chloris orthonothon Doll.                      Car           Ane
Dactylactenium aegyptium (L.) Willd.           Car           Ane
Digitaria eriantha Steud.                      Car           Ane
Eragrostis ciliaris L.                         Car           Ane
Eragrostis tenella (L.) P.Beauv.               Car           Ane
  ex Roem. & Schult.
Leptochloa fascicularis (Lam.) A. Gray         Car           Ane
Paspalum scutatum Nees ex Trin.                Car           Ane
Polygalaceae
Polygala violacea Aubl.                        Cap            -
Pontederiaceae
Eichhornia paniculata (Spreng.) Solms.         Cap            -
Heteranthera limosa (Sw.) Willd.               Cap            -
Heteranthera oblongifolia                      Cap            -
  C. Mart. ex Roem.
Portulacaceae
Portulaca elatior Mart. ex Rohrb.              Cap           Ane
Portulaca halimoides L.                        Cap           Ane
Rhamnaceae
Ziziphus joazeiro Mart.                        Dru           Zoo
Rubiaceae
Borreria verticillata (L.) G. Mey              Cap           Zoo
Mitracarpus frigidus (Willd.                   Cap           Ane
  ex Roem. & Schult.) K. Schum.
Richardiagrandiflora (Cham.                    Cap           Aut
  & Schlecht.) Steud.
Sapindaceae
Cardiospermum corindum L.                      Cap           Ane
Serjania glabrata Kunth                        Sam           Ane
Sapotaceae
Syderoxylon obtusfolium (Roem.                 Dru           Zoo
  & Schult.) T.D. Penn.
Solanaceae
Nicotiana glauca Graham                        Cap           Ane
Physalis angulata L.                           Ber           Ane
Solanum agrarium Sendtn.                       Ber           Zoo
Verbenaceae
Lantana camara L.                              Dru           Zoo
Lippiagracilis Schauer                         Dru           Zoo
Stachytarpheta elatior                         Dru           Zoo
  Schrad. ex Schult.

Family/ Species                             Life form      Voucher

Acanthaceae
Dicliptera ciliaris Juss.                      Cha            -
Ruellia asperula (Mart. & Nees) Lindau         Cha            -
Ruellia bahiensis (Nees) Morong                Cha            -
Ruelliageminiflora Kunth                       Cha           72
Ruellia paniculata L.                          Cha           110
Alismataceae
Echinodorusgrandiflorus                         -            75
  (Cham. & Schltdl.) Micheli
Hydrocleys modesta Pedersen                     -            112
Amaranthaceae
Alternanthera brasiliana (L.) Kuntze           The           223
Alternanthera pungens Kunth                     -            39
Alternanthera tenella Colla                    Hem           142
Amaranthus spinosus L.                          -            148
Froelichia humboldtiana                         -            08
  (Roem. & Schult.) Seub.
Gomphrena vaga Mart.                           Hem            -
Anacardiaceae
Myracrodruon urundeuva Allemao                 Pha            -
Schinopsis brasiliensis Engl.                  Pha            -
Spondias tuberosa Arruda                       Pha            -
Apocynaceae
Allamanda blanchetii A.DC.                     Hem           177
Aspidosperma pyrifolium Mart.                  Pha            -
Mandevilla tenuifolia                          Cry           129
  (J.C. Mikan) Woodson
Araceae
Pistia stratiotes L.                            -            140
Asteraceae
Acanthospermum hispidum DC.                    The           147
Acmella uliginosa (Sw.) Cass.                  The           71
Ageratum conyzoides L.                         The           187
Bidens pilosa L.                               The           67
Centratherum punctatum Cass.                   The           155
Chrysanthellum indicum DC.                      -            185
Conocliniopsisprasifolia (DC.)                 The           69
  R.M. King & H. Rob.
Eclipta prostrata (L.) L.                       -            144
Emilia sonchifolia (L.) DC.                    The            -
Tilesia baccata (L.) Pruski                     -            123
Bignoniaceae
Handroanthus impetiginosus                     Pha           131
  (Mart. ex DC.) Mattos
Tabebuia aurea (Silva Manso) Benth.            Pha            -
  & Hook. f. ex S. Moore
Bixaceae
Cochlospermum vitifolium                       Pha           121
  (Willd.) Spreng.
Boraginaceae
Euploca procumbens (Mill.)                     The            -
  Diane & Hilger
Heliotropium angiospermum Murray               The           132
Varronia dardani (Taroda) J.S. Mill.           Pha           153
Bromeliaceae
Bromelia baniosa Mart. ex Schult. f.           Cha            -
Encholirium spectabile Mart.                   Cha            -
  ex Schult. f.
Neoglaziovia variegata (Arruda) Mez.           Cha            -
Tillandsia recurvata (L.) L.                   Pha           26
Tillandsia streptocarpa Baker                  Pha            -
Burseraceae
Commiphora leptopHoeos (Mart.) Gillet.         Pha            -
Cactaceae
Cereus jamacaru DC.                            Pha            -
Melocactus ernestii Vaupel                     Hem            -
Melocactus zehntneri                           Hem            -
  (Britton & Rose) Luetzelb.
Pilosocereus gounellei (F.A.C.Weber)           Pha            -
  Byles & G.D.Rowley
Pilosocereuspachydadus F. Ritter               Pha            -
Tacinga inamoena (K. Schum.)                   Cha            -
  N.P. Taylor & Stuppy
Tacinga palmadora (Britton & Rose)             Cha            -
  N.P. Taylor & Stuppy
Capparaceae
Cynophala flexuosa (L.) J. Presl               Pha            -
Physostemon guianensis (Aubl.) Malme           The            -
Physostemon lanceolata                         The            -
  (Mart. & Zucc.) D.R. Hunt.
Tarenaya spinosa (Jacq.) Raf.                  Cha            -
Commelinaceae
Callisia filiformis (M. Martens                The           33
  & Galleotti) D. R. Hunt.
Callisia repens L.                             The           214
Commelina erecta L.                            The            -
Commelina obliqua Vahl                         The            -
Convolvulaceae
Evolvulusfdipes Mart.                          The            -
Evolvulus glomeratus Nees & C. Mart.           The           25
Ipomoea subincana (Choisy) Meisn.              Cha           118
Jacquemontia multiflora Haller f.               -            57
Operculina macrocarpa (L.) Urb.                 -            216
Cyperaceae
Cyperus ligularis L.                           Hem           106
Cyperus odoratus L.                             -            113
Cyperus surinamensis Rottb.                    Hem           194
Cyperus uncinulatus Schrad. ex Nees            The           90
Eleocharis elegans (Kunth)                      -            136
  Roem. & Schult.
Eleocharis geniculata (L.)                     The            -
  Roem. & Schult.
Eleocharis interstincta (Vahl)                  -            137
  Roem. & Schult.
Eragrostis ciliaris (L.) R.Br.                 The            -
Fimbristylis cymosa R. Br.                     Hem           139
Pycreus macrostachyos (Lam.) J. Raynal         The           212
Cucurbitaceae
Momordica charantia L.                          -             -
Euphorbiacae
Cnidoscolus quercifolius Pohl                  Pha            -
Cnidoscolus urens (L.) Arthur                  Cha            -
Croton blanchetianus Baill.                    Pha           182
Croton heliotropiifolius Kunth                 Pha           221
Jatropha mollissima (Pohl) Baill.              Pha            -
Jatropha ribfolia (Pohl) Baill.                Cha           152
Manihot catingae Ule                           Cha            -
Fabaceae
Anadenanthera colubrina (Vell.) Brenan         Pha            -
Bauhinia cheilantha (Bong.) Steud.             Pha           225
Bauhinia subclavata Benth.                     Pha           175
Poincianella pyramidalis                       Pha            -
  (Tul.) L.P. Queiroz
Centrosema brasilianum (L.) Benth.             Hem           101
  var. angustifolium Amshoff
Centrosema brasilianum (L.) Benth.             Hem           231
  var. brasilianum Benth.
Centrosema virginianun (L.) Benth.              -            178
Chamaecrista rotundifolia                      Hem           189
  (Pers.) Greene
Desmodiumglabrum (Mill.) DC.                   Hem           79
Dioclea grandiflora Mart. ex Benth.            Pha           135
Dioclea violacea Mart. ex Benth.                -            116
Erythrina velutina Willd.                      Pha            -
Indigofera blanchettiana Benth.                Cha            -
Indigofera suffruticosa Mill.                  Cha           227
Libidibia ferrea (Mart. ex Tul.)               Pha           97
  L.P. Queiroz
Mimosa tenuiflora (Willd.) Poir.               Pha            -
Piptadenia stipulacea (Benth.) Ducke           Pha            -
Poincianella pyramidalis (Tul.)                Pha           127
  L.P. Queiroz
Senna macranthera (DC. ex Collad.)             Pha            -
  Irwin & Barneby
Senna martiana (Benth.) Irwin & Barneby        Pha           180
Senna obtusfolia (L.) Irwin & Barneby          Cha           103
Senna splendida (Vogel) Irwin & Barneby        Pha           226
Stylosanthes viscosa (L.) Sw.                   -            65
Vigna adenantha (G.Mey) Marechal,              Hem           230
  Mascherpa & Stainer
Zorniagemella (Willd.) Vogel                   The           192
Zornia leptophylla (Benth.) Pittier            The           204
Zornia myriadena Benth.                        The           29
Gentianaceae
Schultesia pohliana Progel                     The           99
Hydrocharitaceae
Apalanthegranatensis (Bonpl.) Planch.           -             -
Lamiaceae
Hyptis fruticosa Salzm. ex Benth.              The           145
Leonotis nepetfolia (L.) R.Br.                  -            146
Raphiodon echinus Schauer                       -            188
Loranthaceae
Struthanthus syringifolius (Mart.) Mart.       Pha            -
Lythraceae
Cuphea campestres (Mart.) Koehne               The           215
Malvaceae
Ceiba glaziovii (Kuntze) K. Schum.             Pha            -
Herissantia tiubae (K. Schum.) Brizicky        Cha           76
Melochia pyramidata L.                         Cha           143
Melochia tomentosa L.                          Cha           31
Sidagalheirensis Ulbr.                         Cha           32
Sida Unifolia Cav.                             The           228
Sidastrum paniculatum (L.) Fryxell             Cha            -
Waltheria rotundifolia Schrank.                Cha           229
Waltheria tomentosa H.St.-John                  -            141
Molluginaceae
Mollugo verticillata L.                        The           170
Nyctaginaceae
Boehravia diffusa L.                           Cha           126
Nymphaeaceae
Nymphaea ampla (Salisb.) DC.                    -            62
Onagraceae
Ludwigia leptocarpa (Nutt.) Hara               The           198
Ludwigia octovalvis (Jacq.) P.H. Raven         The            -
Oxalidaceae
Oxalis divaricata Mart. ex Zucc.               Cha           130
Passifloraceae
Passiflora foetida L.                          Cha           232
Piriqueta duarteana (Cambess.) Urb.            Cha           20
Turnera subulata Sm.                           Cha           233
Plantaginaceae
Angelonia biflora Benth.                       Hem           22
Angelonia campestres Nees & Mart.               -            179
Angelonia cornigera Hook.                      The            -
Stemodia maritima L.                           The           43
Plumbaginaceae
Plumbago scandens L.                           Cha           56
Poaceae
Anthephora hermaphrodita (L.) Kuntze           The           206
Chloris inflata Link                           The           210
Chloris orthonothon Doll.                      The           199
Dactylactenium aegyptium (L.) Willd.           The           200
Digitaria eriantha Steud.                      The           202
Eragrostis ciliaris L.                         The           195
Eragrostis tenella (L.) P.Beauv.               The           209
  ex Roem. & Schult.
Leptochloa fascicularis (Lam.) A. Gray         The           211
Paspalum scutatum Nees ex Trin.                The           203
Polygalaceae
Polygala violacea Aubl.                        The           224
Pontederiaceae
Eichhornia paniculata (Spreng.) Solms.          -             -
Heteranthera limosa (Sw.) Willd.               Cha            -
Heteranthera oblongifolia                       -            134
  C. Mart. ex Roem.
Portulacaceae
Portulaca elatior Mart. ex Rohrb.              The           124
Portulaca halimoides L.                        Hem           125
Rhamnaceae
Ziziphus joazeiro Mart.                        Pha            -
Rubiaceae
Borreria verticillata (L.) G. Mey              Cha            -
Mitracarpus frigidus (Willd.                   Cha           98
  ex Roem. & Schult.) K. Schum.
Richardiagrandiflora (Cham.                    Cha           234
  & Schlecht.) Steud.
Sapindaceae
Cardiospermum corindum L.                      Cha           161
Serjania glabrata Kunth                        Pha           222
Sapotaceae
Syderoxylon obtusfolium (Roem.                 Pha           235
  & Schult.) T.D. Penn.
Solanaceae
Nicotiana glauca Graham                        Pha           36
Physalis angulata L.                           Cha           54
Solanum agrarium Sendtn.                        -             -
Verbenaceae
Lantana camara L.                              Pha           128
Lippiagracilis Schauer                         Cha           55
Stachytarpheta elatior                         The           63
  Schrad. ex Schult.

Figure 1. Percentage of life forms identified between species in the
study area, Fazenda Salambaia, APA of Cariri, Paraiba State,
Northeast, Brazil.

Cryptophytes         0.6%
Hemiciyptophytes       9%
Indeterminate       16.9%
Chamaephytes        22.3%
Phanerophytes       23.5%
Therophytes        127.7%

Source: The authors.

Note: Table made from bar graph.

Figure 2. Percentage of dispersal syndromes recorded in the study
area, Fazenda Salambaia, APA of Cariri, Paraiba State, northeast,
Brazil.

Anemochorous     28.3%
Autochorous      25.9%
Zoochorous       23.5%
No identified    22.3%

Note: Table made from bar graph.
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Author:de Lima, Elimar Alves; de Melo, Jose Iranildo Miranda
Publication:Acta Scientiarum. Biological Sciences (UEM)
Date:Jan 1, 2015
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