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Journal of Arid Environments

Journal of Arid Environments 68 (2007) 237247 www.elsevier.com/locate/jnlabr/yjare

Flora and life-form spectrum in an area of deciduous thorn woodland (caatinga) in northeastern, Brazil
R. Carvalho da Costaa, F. Soares de Araujob,, b L. Wilson Lima-Verde
s-graduaca em biologia vegetal, departamento de bota nica, Programa de po - o Universidade Estadual de CampinasUNICAMP. Caixa postal 6109, 13084-971, Campinas, Sa Paulo, Brasil o b rio , Departamento de Biologia, Laborato de Fitogeograa, Universidade Federal do Ceara , Campus do PiciBloco 906, 60455-760 Fortaleza, Ceara Brazil
a

Received 30 October 2002; received in revised form 4 May 2005; accepted 15 June 2006 Available online 7 August 2006

Abstract Caatinga, a deciduous thorny woodland vegetation, is encountered in the semi-arid region of northeastern Brazil. In view of the importance of the herbaceous component of caatinga plant communities, a characterization of the ora of the Nao Me Deixes Reserve in Ceara State, Brazil (41490 3400 S, 381590 0900 W, at 210 m a.s.l.) was undertaken. The reserve has 300 ha of caatinga vegetation, including dense tree steppe and open tree steppe. The mean annual rainfall is 732.8 mm, concentrated between February and May (78%). The ora was surveyed at monthly intervals between February 2000 and June 2001. We encountered 133 species belonging to 47 families. The herbaceous/woody ratio was 1.4. Based on eld observations, the life-form spectrum was characterized according Raunkiaers system, and compared with his normal spectrum. The lifeform spectrum observed was: therophytes (42.9%), phanerophytes (26.3%), camaephytes (15.8%), hemicryptophytes (12.8%), and cryptophytes (2.3%). Previous data on the caatinga herbaceous ora, as well as the present study, indicate that the oristic richness of this biome has been underestimated, and that the herbaceous/woody proportion varies according to its physiognomy and water status. r 2006 Elsevier Ltd. All rights reserved.
Keywords: Deciduous woodland; Caatinga; Semi-arid region; Life-form spectrum

Corresponding author. Fax: +55 85 3366 9806.

E-mail addresses: [email protected] (R. Carvalho da Costa), [email protected] (F. Soares de Araujo). 0140-1963/$ - see front matter r 2006 Elsevier Ltd. All rights reserved. doi:10.1016/j.jaridenv.2006.06.003

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238 R. Carvalho da Costa et al. / Journal of Arid Environments 68 (2007) 237247

1. Introduction The Brazilian northeastern region covers 1,542,246 km2 (IBGE, 1998). Of this area, 750,000 km2 has a semi-arid climate (AbSaber, 1977) corresponding to Koeppens BSh climatic type (Andrade-Lima, 1981). This semi-arid region demonstrates varying degrees of edapho-climatic aridity, generally associated with its distance to the Atlantic coast, altitude, geomorphology, degree of dissection of the landscape, slope, wind exposure, as well as soil depth and its physical and chemical composition. Rainfall usually totals less than 750 mm/year in most of this domain, and it is concentrated in three consecutive months during the southern hemisphere summer or summer/autumn (November until June). Temperatures vary little, with an annual average of approximately 26 1C (Nimer, 1989). The seasonal xerophilous thorn woodland/shrubland, regionally denominated caatinga, prevails in the semi-arid lowlands on an extensive regional crystalline basement complex (Andrade-Lima, 1981; Sampaio, 1995). The term caatinga refers to xerophytic, woody, thorny, and deciduous physiognomies with a seasonal herbaceous layer (Veloso et al., 1991). It comprises a mosaic of vegetation types varying from dry thorn forest to open shrubby vegetation (Andrade-Lima, 1981). These variations have been attributed to largescale variations in the climate, orographic patterns, and small-scale variations in topography and soils (Andrade-Lima, 1981; Sampaio, 1995). Although many authors have stressed the importance of herbaceous species within caatinga physiognomies (Veloso et al., 1991; Sampaio, 1995; Rizzini, 1997), most oristic and phytosociological studies have focused only the woody component (Tavares et al., 1969a, b, 1970, 1974; Gomes, 1980; Figueiredo, 1987; Fonseca, 1991; Rodal, 1992; Araujo et al., 1995; Lima and Lima, 1998; Camacho, 2001; Lemos and Rodal, 2002; Pereira et al., 2002; Alcoforado-Filho et al., 2003; Nascimento et al., 2003). Few studies have been devoted to the structure and ora of the herbaceous layer in plant communities of caatinga (Figueiredo, 1983; Santos, 1987; Oliveira et al., 1988; Oliveira, 1995; Ferraz et al., 1998). Plant species and individuals can be grouped into different life-form classes based on structural and functional similarities (Mueller-Dombois and Ellenberg, 1974). Life-forms have close relationships with environmental factors (Mueller-Dombois and Ellenberg, 1974) and can be viewed as strategies for obtaining resources (Crosswhite and Crosswhite, 1984; Cody, 1986). Raunkiaer (1934) proposed a life-form classication system based on the manner in which plants protect their perennating buds during unfavourable seasons. According to this classication system, plant species can be grouped into ve main classes: phanerophytes, camaephytes, hemicryptophytes, cryptophytes, and therophytes. This sequence corresponds to an increasing protection of the perennating buds. Climatic types can be characterized by the prevailing life-forms in plant communities growing under a given climatic regime, using the proportions of species in each life-form class, or the biological spectrum (Raunkiaer, 1934; Cain, 1950; Mueller-Dombois and Ellenberg, 1974). Studies carried out in arid and semi-arid areas have shown that there is a high proportion of life-forms that lose their aerial shoots during the driest months (therophytes, hemicryptophytes, and cryptophytes) (van Rooyen et al., 1990). The importance of therophytes increases as rainfall decreases and becomes more irregular (Raunkiaer, 1934; Kovacs-Lang et al., 2000). Few studies have shown high herbaceous species richness in areas of caatinga (Silva, 1985; Santos, 1987). In these studies, species were classied subjectively according to their growth habit, without precisely stating their life-form. In the present study, we used

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R. Carvalho da Costa et al. / Journal of Arid Environments 68 (2007) 237247 239

Raunkiaers life-forms system to characterize the ora of the Nao Me Deixes Reserve and to examine the importance of the richness of the herbaceous stratum in this vegetation type in an area of semi-arid caatinga vegetation in northeastern Brazil. According Raunkiaers system, as well as data from literature, it would be expected that plant communities in areas with low annual rainfall levels, high mean temperatures, and severe periods of drought (such as the caatinga) would have high proportions of species in life-forms classes that afford high drought protection (mainly therophytes). 2. Materials and methods The present study was carried out in the Nao Me Deixes Reserve, in the state of Ceara, northeastern Brazil (41490 3400 S 381580 900 W, at 210 m a.s.l.). Climatic data (rainfall and temperature) were obtained from the Ceara Foundation of Meteorology and Water Resources (FUNCEME). The mean annual rainfall and temperature were 732.8 mm and 26.6 1C, respectively. Most rainfall (79.6%) was concentrated between February and May. Soils at the Nao Me Deixes Reserve are a mixture of planosols, solonetz, and regosols (BRASIL, 1972). The reserve has 300 ha of caatinga vegetation, classied as dense treesteppe/savanna and open tree-steppe/savanna according to the ofcial Brazilian system (RADAMBRASIL, 1983). These are the more widespread types of caatinga vegetation in semi-arid northeastern Brazil. From February 2000 to June 2001 the vascular ora of the Nao Me Deixes Reserve was surveyed monthly. All specimens collected were identied and subsequently incorporated into the EAC Herbarium collection. Data was organized listing the species, their families, and their life-forms. Observations were made on aerial shoot reduction during unfavourably dry conditions, and presence of subterranean reserve organs. Species were classied as phanerophytes, camaephytes, hemicryptophytes, cryptophytes, or therophytes according to Raunkiaer (1934). As our aim was to compare the caatinga life-form spectrum with Raunkiaers normal spectrum, Cactaceae species were considered phanerophytes (see Cain, 1950). As there is no specic life-form class in Raunkiaers original system for non self-supporting plants, these were classied according to the reduction of their aerial parts (Cain, 1950). We computed the proportion of species in each life-form class and compared these numbers to Raunkiaers normal spectrum using a w2 test (Mann, 1998). Finally, we compared the ratio of herb/woody species in the reserve with other caatinga study sites. For this comparison, therophytes, cryptophytes, and hemicryptophytes were considered herbs. 3. Results We recorded 133 species distributed among 103 genera and 47 families (Table 1). The families with the greatest number of species were Euphorbiaceae (16), Fabaceae (11), Asteraceae (7), and Convolvulaceae (7). Twenty two families (47%) were represented by only a single species. The herbaceous ora (hemicryptophytes, cryptophytes, and therophytes) comprised 77 species (57.9%), whereas the woody ora was represented by 56 species (42.1%) (Table 2), yielding a ratio of 1.4 between them. The biological spectrum of the Nao Me Deixes Reserve had a high proportion of therophytes (42.9%), followed by phanerophytes (26.3%), camaephytes (15.8%), hemicryptophytes (12.8%), and cryptophytes (2.3%). The w2 test demonstrated signicant differences between the Nao Me

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240 R. Carvalho da Costa et al. / Journal of Arid Environments 68 (2007) 237247 Table 1 List of species, families, their life-forms, and collection numbers in the Nao me Deixes Reserve, Ceara State, Brazil. Life forms: Ph phanerophytes, Ch camaephytes, H hemicryptophytes, Cr cryptophytes, Th therophytes Species Acanthaceae Justicia strobilacea (Ness) Lindau Justicia schomburgkiana (Nees) V.A.W. Graham Dicliptera ciliaris Juss. Alismataceae Echinodorus subulatus Griseb. Amaranthaceae Alternanthera brasiliana (L.)Kuntze Alternanthera tenella Colla Anacardiaceae Myracrodruon urundeuva Allemao Apocynaceae Aspidosperma pyrifolium Mart. Allamanda blanchetii A.DC. Araceae Taccarum peregrinum Schott Asclepiadaceae Schubertia sp. Asteraceae Aspilia attenuata Baker Trichogonia sp. Melanthera latifolia (Gardner)Cabrera Porophillum ruderale Cass. Stilpnopappus sp. Aspilia sp. Blainvillea lanceolata Baker Bignoniaceae Arrabidaea subverticillata Bureau & K.Schum. Boraginaceae Cordia cf. globosa Humb., Bonpl.. & Kunth Cordia sp. Auxemma oncocalyx (Allemao) Taub. Auxemma glazioviana Taub. Heliotropium sp. Brassicaceae Brassica sp. Burseraceae Commiphora leptophloeos (Mart.) J.B. Gillett Cactaceae Cereus jamacaru DC. Capparaceae Cleome spinosa Jacq. Life-form Number

Ch Ph Ch H Th Th Ph Ph H Cr Ch Th Th Th Th Th Th Ch Ph Ph Ph Ph Ph Ch H Ph Ph Ch

259

340 171 40 341 337 336 290

247 221 83 185 262 159

255 12 301 201 204

22 17

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R. Carvalho da Costa et al. / Journal of Arid Environments 68 (2007) 237247 Table 1 (continued ) Species Caesalpiniaceae Chamaecrista calycioides Greene Chamaecrista cf. duckeana (P. Bezerra & Afr. Fern.) H.S. Irwin Barneby Caesalpinia bracteosa Tul. Caesalpinia ferrea var. glabrescens Benth. Senna trachypus (Benth.) H.S. Irwin & Barneby Bauhinia cheilantha (Bong.) D. Dietr. Cochlospermaceae Cochlospermum vitilfolium (Willd.) Spreng. Combretaceae Combretum leprosum Mart. Comelinaceae Commelina cf. virginica L. Aneilema brasiliense C.B. Clarke Callisia cf. filiformis (M. Martens & Galeotti) D.R. Hunt Convovulaceae Aniseia heterantha Choisy Evolvulus ovatus Fernald. Evolvulus sp. Ipomoea bahiensis Willd. ex Roem. & Schult. Ipomoea rosea Choisy Jacquemontia cf. velutina Choisy Jacquemontia sp. Cucurbitaceae Cayaponia cf. racemosa Cogn. Cyperaceae Cyperus uncinulatus Schrad. ex Nees Euphorbiaceae Caperonia palustris (L.) A.St.Hil. Croton adenocalx Baill. Croton glandulosus L. Croton moritibensis Baill. Croton blanchetianus Baill. Croton sp. Euphorbia heterophylla L. Dalechampia pernambucensis Baill. Jatropha mollissima Baill. Phyllanthus caroliniensis Walter Phyllanthus orbiculatus Rich. Phyllanthus sp. Sebastiania corniculata (Vahl.) Mull.Arg. Sebastiania macrocarpa Mull. Arg. Sebastiania sp. Tragia cf. volubilis L. Fabaceae Arachis dardani Krapov. & W.C. Greg. Macroptilium martii (Benth.) Marechal & Baudet Life-form Number 241

Ch Ch Ph Ph Ph Ph Ph Ph H Th Th Th Th Th Ch Ch H H Th Th Th Ph Th Ph Ph Th Th Th Ph Th Th Th Th Ph Ph H Th Th

198 327 323 343 344 269 338 333 126 235 117 230

312 249 335 141 329 307 143 281 280 63 41 304 316 95 50 94 111 46 275 20 1 241

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242 R. Carvalho da Costa et al. / Journal of Arid Environments 68 (2007) 237247 Table 1 (continued ) Species Macroptilium sp. Chaetocalyx scandens (L.) Urb. Crotalaria holosericea Nees & Mart. Desmodium sp. Dioclea grandiora Mart. ex Benth Aeschynomene sp. Canavalia brasiliensis Mart. ex Benth. Stylosanthes humilis Kunth Galactia striata Urb. Iridaceae Ebertia sp. Lamiaceae Hyptis suaveolens (L.) Poit. Marsypianthes chamaedrys (Vahl) Kuntze Liliaceae Hippeastrum sp. Lythraceae Cuphea circaeoides Sm. ex Sims Cuphea sp. Cuphea campestris Mart. ex Koehne Pleurophora anomala Koehne Loganiaceae Spigelia anthelmia L. Malpighiaceae Heteropterys trichanthera A. Juss. Stigmaphyllon auriculatum A. Juss. Malvaceae Sida ciliaris L. Herissantia crispa (L.) Briz. Herissantia tiubae (K.Schum.) Brizicky Pavonia cancellata (L.) Cav. Wissadula contracta (Link) R.E. Fr. Wissadula amplissima (L.) R.E. Fr. Mimosaceae Mimosa tenuiora (Willd.) Poir. Anadenanthera colubrina var. cebil (Griseb.)Altschul Mimosa caesalpiniifolia Benth. Piptadenia stipulacea (Benth.) Ducke Piptadenia viridiora (Kunth) Benth. Molluginaceae Mollugo verticillata L. Nyctaginaceae Boerhavia diffusa L. Guapira sp. Oxalidaceae Oxalis sp. Life-form Th H Ph Th Ph H Ch Th H Cr Th Th Cr Th H Th Th Th Ph Ph Th Ch Ch Ch H Th Ph Ph Ph Ph Ph Th Ch Ph Th Number 245 258 244 26 318 326 147 251 103 324 150 284 299 178 248 291 131 306 287 115

238 184 273 272 330 345 271 298

283 264

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R. Carvalho da Costa et al. / Journal of Arid Environments 68 (2007) 237247 Table 1 (continued ) Species Passioraceae Passiora foetida L. Poaceae Urochloa mollis (Sw) Morrone & Zuloaga Digitaria sp. Panicum trichoides Sw. Paspalum scutatum Nees ex. Trin Paspalum sp. Setaria sp. Polygalaceae Polygala aff. lancifolia A. St.-Hil. Portulacaceae Portulaca cf. halimoides L. Portulaca sp. Talinum sp. Rhamnaceae Crumenaria decumbens Mart. Zizyphus joazeiro Mart. Rubiaceae Diodia teres Walter Diodia rigida Cham. & Schltdl. Mitracarpus hirtus (Sw.) DC. Spermacoce vegeta (Standl. & Steyerm.) C.D. Adams Staelia virgata K. Schum. Sapindaceae Cardiospermum corindum L. Scrophulariaceae Angelonia biora Benth. Angelonia pubescens Benth. Scoparia dulcis L. Tetraulacium sp. Sterculiaceae Waltheria indica L. Waltheria macropoda Turcz. Tiliaceae Corchorus sp. Turneraceae Turnera pumilea L. Turnera sp. Violaceae Hybanthus ipecacuanha Baill. Verbenaceae Stachytarpheta coccinea Schau Stachytarpheta sessilis Moldenke Lantana camara L. Life-form Number 243

Ch Th Th Th Th Th Th Th H H H Th Ph Th Th Th Th Th Ch H Th Ch Th Ch Ch Th Ch Ph H Th Th Ph 160 292 193 191 149 153 174 151 56 175 89 311 310 319 163 15 176 232 139 152 328

274 13 267 309 308

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244 R. Carvalho da Costa et al. / Journal of Arid Environments 68 (2007) 237247 Table 2 Results of w2 tests of the Nao me Deixes Reserve and Raunkiaers normal spectra Ph Nao Me Deixes, Ceara, Brazil (No. of species) Nao Me Deixes, Ceara, Brazil (% of species) Raunkiaers normal spectrum (% of species) w2 35 26.3 46 2.87 Ch 21 15.8 9 1.72 H 17 12.8 26 1.39 Cr 3 2.3 6 0.25 Th 57 42.9 13 4.67 Total 133 100.0 100 10.89

Ph Phanerophytes, Ch Camaephytes, H Hemicryptophytes, Cr Cryptophytes, Th Therophytes.

Deixes ora and Raunkiaers normal spectra (N 100, po0:05, d.f. 4) (Table 2). Therophytes had the highest individual value obtained from w2 test, followed by phanerophytes (Table 2). 4. Discussion Studies of the herbaceous ora and the structure of caatinga areas are scarce, and this study demonstrates the importance of herbaceous plants in caatinga species richness and physiognomy. Most published studies indicate that Leguminosae (Caesalpinioidae, Mimosoidae, and Papilionoidae), Euphorbiaceae, and Cactaceae are among the most species rich families in the caatinga, when only woody species are considered (Sampaio, 1995). When herbaceous species are included, other families become oristically important. Asteraceae and Convolvulaceae were among the richest families in the present study, and were represented mainly by herbaceous species (hemicryptophytes, cryptophytes, and therophytes). If species richness is compared among different studies (Table 3), it can be seen that the ratios between herbaceous and woody species are quite variable. Woody species may either be the richest (Alcoforado-Filho, 1993) or the poorest (Santos, 1987) components in caatinga plant communities. In the Nao Me Deixes Reserve herbaceous and woody plant species showed nearly the same proportion. The relative importance of each component can vary as a result of large-scale variations in climate, orography, and soils, as pointed out by Andrade-Lima (1981) and Sampaio (1995). In fact, Table 3 demonstrates that the sites with the lowest mean annual rainfall (Santos, 1987) had the highest herb/woody species ratios (Alcoforado-Filho, 1993). The relationship between climate and physiognomy becomes very evident when using the lifeform approach. Raunkiaers system was useful in characterizing the ora of a caatinga site in the present study. The high proportion of therophytes at the Nao Me Deixes reserve is in agreement with the predictions for areas with Koppens BSh climate (Cain, 1950), which corresponds to a therophytic phytoclimate (Raunkiaer, 1934). The predominance of therophytes reects an effective strategy for avoiding water losses due to humidity extremes and water deciencies (Van Rooyen et al., 1990). A high proportion of therophytes is a common feature of the biological spectra on hot steppes with a BSh climate (Table 4), although differences in other life-forms are difcult to interpret in terms of large-scale climatic patterns. The data presented here demonstrates the importance of the herbaceous component in caatinga plant communities. Herbaceous plants may represent up to 2/3 of the plant species richness at a given location. The predominance of the BSh climate results in high

Table 3 Total, woody and herbaceous species richness, herbaceous/woody species ratio, mean annual rainfall, and elevation among different studies undertaken in areas of caatinga vegetation Elevation (m) 246590 400 210 600 317-542 530 300900 459.7659.7 579.2585.4 732.8 689 631.8650.9 585 694 10 8 7 1 1 4 3 1 B A All All A A A All 32 43 52 56 56 56 58 88 Annual rainfall No. of (mm) locations Inclusion Woody species Herbaceous Ratio herb/ Total species woody 136 77 16 2.6 1.4 0.2 32 43 188 133 56 56 58 114

Reference

Location

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Gomes (1980) Figueiredo (1987) Santos (1987) This study Lemos and Rodal (2002) Rodal (1992) Araujo et al. (1995) Alcoforado-Filho (1993)

Cariris Velhos/PB Salineira/RN region Parnamirim/PE Quixada/CE Serra da Capivara/PI Floresta & Custodia/PE Floresta & Custodia/PE Caruaru/PE

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Inclusion: A plants with stem diameter at soil level X3 cm; B plants with stem diameter at soil level X5 cm.

245

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246 R. Carvalho da Costa et al. / Journal of Arid Environments 68 (2007) 237247 Table 4 Life-form spectra of sites with a BSh climate Locality Nao Me Deixes, Ceara, Brazil Whitehill, South Africa Timbuctu, Africa Tripoli, North Africa Cyrenaica, North Africa Madeira Islands, lowlands Source Present study Admson (1939) in Cain (1950) Hagerup (1930) in Cain (1950) Raunkiaer (1934) Raunkiaer (1934) Raunkiaer (1934) Ph 26.3 10 24 9 8 15 Ch 15.8 42 36 13 14 7 H 12.8 2 9 19 19 24 Cr 2.3 18 6 11 8 3 Th 42.9 23 25 51 50 51

proportions of herbaceous life-forms that avoid unfavourable conditions by losing their aerial portions (hemicryptophytes, cryptophytes, and therophytes). Although it is difcult to make generalizations from the limited data available in the literature, the variations in herb/woody species ratios seen among different studies seem to represent differences in small-scale factors such as the soil and micro-climate at each site. More comparative studies will lend new insights into physiognomic variations in the caatinga. Acknowledgements The authors would like to thank the Fundac ao Cearense de Amparo a Pesquisa - (FUNCAP) for partial funding of this study, as well as the Conselho Nacional de Desenvolvimento Cient co de Tecnologico (CNPq) for grants provided by the PIBIC and M.A. Figueiredo and M.J.N. Rodal for comments on an early version of this manuscript. References
AbSaber, A.N., 1977. Espacos ocupados pela expansao dos climas secos na America do Sul, por ocasiao dos per odos glaciais quaternarios. Instituto de Geograa/USP, Sao Paulo (Serie Paleoclimas, 3). Alcoforado-Filho, F.G., 1993. Composicao or stica e tossociologia de uma area de caatinga arborea no - munic pio de CaruaruPE. M.Sc. Thesis, Universidade Federal Rural de Pernambuco, Recife, Pernambuco, Brazil. Alcoforado-Filho, F.G.A., Sampaio, E.V.S.B., Rodal, M.J.N., 2003. Flor stica e tossociologia de um remanescente de vegetacao caducifolia espinhosa arborea em Caruaru, Pernambuco. Acta Botanica Brasilica - 17 (2), 287303. Andrade-Lima, D., 1981. The caatingas dominium. Revista Brasileira de Botanica 4, 149153. Araujo, E.L., Sampaio, E.V.S.B., Rodal, M.J.N., 1995. Composicao or stica e tossociologia de tres areas de - caatinga de Pernambuco. Revista Brasileira de Biologia 55 (4), 595607. BRASIL, 1972. Mapa exploratorio-reconhecimento de solos: estado do Ceara, escala 1:600.000, SUDENE, Recife, Brazil. Cain, S.A., 1950. Life forms and phytoclimate. Botanical Review 16, 132. Camacho, R.G.V., 2001. Estudo tosiograco da caatinga do serido- Estacao Ecologica do Serido, RN. Ph.D. - Thesis, Universidade de Sao Paulo, Sao Paulo, Brazil. Cody, M.L., 1986. Structural niches in plant communities. In: Diamond, J., Case, T. (Eds.), Community Ecology. Harper and Row, San Francisco, pp. 381405. Crosswhite, F., Crosswhite, C., 1984. A classication of life forms of the Sonoran desert with emphasis on seed plants and their survival strategies. Desert Plants 5, 131161. Ferraz, E.M.N., Rodal, M.J.N., Sampaio, E.V.S.B., Pereira, R.C.A., 1998. Composicao or stica em trechos de - vegetacao de caatinga e brejo de altitude na regiao do Vale do Pajeu, Pernambuco. Revista Brasileira de - Botanica 21 (1), 715. Figueiredo, M.A., 1983. A regiao dos Inhamuns-CE no dom nio das caatingas. Colecao Mossoroense B-411. -

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R. Carvalho da Costa et al. / Journal of Arid Environments 68 (2007) 237247 247 Figueiredo, M.A., 1987. A microregiao salineira norte-riograndense no dom nio das caatingas. Colecao - Mossoroense 353. Fonseca, M.R., 1991. Analise da vegetacao arbustivo-arborea da caatinga hiperxerola do noroeste do estado de - Sergipe. Ph.D. Thesis, Universidade Estadual de Campinas, Sao Paulo, Brazil. Gomes, M.A.F., 1980. A vegetacao dos Carir s Velhos no estado da Para ba. Vegetalia 14. - IBGE, 1998. Fundacao Instituto Brasileiro de Geograa e Estat stica. Anuario Estat stico do Brasil. IBGE, Rio - de Janeiro, Brazil. Kovacs-Lang, E., Kroel-Dulay, G., Kertesz, M., Fekete, G., Bartha, S., Mika, J., Dobi-Wantuch, I., Redei, T., Rajkai, K., Hahn, I., 2000. Changes in composition of sand grasslands along a gradient in Hungary and implications for climate change. Phytocoenologia 30 (34), 385407. Lemos, J.R., Rodal, M.J.N., 2002. Fitossociologia do componente lenhoso de um trecho da vegetac ao de caatinga - no parque nacional da Serra da Capivara, Piau , Brasil. Acta Botanica Brasilica 16 (1), 2223. Lima, P.C., Lima, J.L.S., 1998. Composicao or stica e tossociologia de uma area de caatinga em Contendas do - Sincora, Bahia, microregiao homogenea da Chapada Diamantina. Acta Botanica Brasilica 12 (3), 431440. Mann, P.S., 1998. Introductory Statistics. Wiley, New York. Mueller-Dombois, D., Ellenberg, H., 1974. Aims and Methods of Vegetation Ecology. Wiley, New York. Nascimento, C.E.S., Rodal, M.J.N., Cavalvanti, A.C., 2003. Phytossociology of the remaining xerophytic woodland associated to an environmental gradient at the banks of the Sao Francisco river-Petrolina, Pernambuco, Brazil. Revista Brasileira de Botanica 26 (3), 271287. Nimer, E., 1989. Climatologia do Brasil, second ed. IBGE-SUPREN (Fundacao IBGE-SUPREN), Recursos - Naturais e Meio Ambiente, Rio de Janeiro. Oliveira, M.E.A., 1995. Vegetacao e ora de uma area de transicao caatinga-carrasco Padre Marcos. Piau . M.Sc. - - Thesis, Universidade Federal de Pernambuco, Recife, Brazil. Oliveira, J.G.B., Quesado, H.L.C., Nunes, E.P., Figueiredo, M.A., Bezerra, C.L.F., 1988. Vegetac ao da estacao - - ecologica de Aiuaba, Ceara. Colecao Mossoroense 537B. - Pereira, I.M., Andrade, L.A., Barbosa, M.R.V., Sampaio, E.V.S.B., 2002. Composicao or stica e analise - tossociologica do componente arbustivo-arboreo de um remanescente orestal no Agreste Paraibano. Acta Botanica Brasilica 16 (3), 241369. RADAMBRASIL. 1983. Folhas SC 24/25. Aracaju/RecifeGeologia, geomorfologia, pedologia, vegetacao, uso - potencial da terra, RADAMBRASIL, Rio de Janeiro, Brazil. Raunkiaer, C., 1934. The Life forms of Plants and Statistical Plant Geography. Clarendon Press, Oxford. Rizzini, C.T., 1997. Tratado de Fitogeograa do Brasil. Ambito Cultural, Rio de Janeiro, Brazil. Rodal, M.J.N., 1992. Fitossociologia da vegetacao arvbustivo-arborea em quatro areas de Caatinga em - Pernambuco. Ph.D. Thesis, Universidade Estadual de Campinas, Campinas, Brazil. Sampaio, E.V.S.B., 1995. Overview of the Brazilian caatinga. In: Bullock, S.H., Mooney, H.A., Medina, E. (Eds.), Seasonally Tropical Dry Forests. Cambridge University Press, Cambridge. Santos, M.F.A., 1987. Caracter sticas dos solos e da vegetacao em sete areas de Parnamirim, Pernambuco. M.Sc. - Thesis, Universidade Federal Rural de Pernambuco, Recife, Brazil. Silva, A.Q., 1985. Flora e vegetacao das depressoes inundaveis da regiao de Ouricuri-PE. M.Sc. Thesis, - Universidade Federal Rural de Pernambuco, Recife, Brazil. Tavares, S., Paiva, F.A.F., Tavares, E.J.S., Lima, J.L.S., 1969a. Inventario orestal do Ceara-Estudo preliminar das matas remanescentes do munic pio de Quixada. Boletim de Recursos NaturaisSUDENE 7 (14), 93111. Tavares, S., Paiva, F.A.F., Tavares, E.J.S., Carvalho, G.H., e Lima, J.L.S., 1969b. Inventario orestal Pernambuco-Estudo preliminar das matas remanescentes do munic pio de Sao Jose de Belmonte. Boletim de Recursos NaturaisSUDENE 7 (14), 113139. Tavares, S., Paiva, F.A.F., Tavares, E.J.S., Carvalho, G.H., e Lima, J.L.S., 1970. Inventario orestal Pernambuco-Estudo preliminar das matas remanescentes dos munic pios de Ouricur , Bodoco, Santa Maria da Boa Vista e Petrolina. Boletim de Recursos NaturaisSUDENE 8, 14. Tavares, S., Paiva, F.A.F., Tavares, E.J.S., Lima, J.L.S., 1974. Inventario orestal do Ceara-estudo preliminar das matas remanescentes do munic pio de Taua. Boletim de Recursos NaturaisSUDENE 12 (2), 519. van Rooyen, M.W., Theron, G.K., Grobbelaar, N., 1990. Life forms and spectra of ora of Namaqualand, South Africa. Journal of Arid Environments 19, 133145. Veloso, H.P., Rangel-Filho, A.L.R., Lima, J.C.A., 1991. Classicac ao da vegetacao brasileira, adaptada a um - - sistema universal. IBGEDepartamento de Recursos Naturais e Estudos Ambientais, Rio de Janeiro.