Rodriguésia 62(2): 341-366. 2011 http://rodriguesia.jbrj.gov.br

Floristics and life-forms along a topographic gradient, central-western Ceará, Florística e formas de vida ao longo de um gradiente topográfico no centro-oeste do estado do Ceará, Brasil

Francisca Soares de Araújo1,2, Rafael Carvalho da Costa1, Jacira Rabelo Lima1, Sandra Freitas de Vasconcelos1, Luciana Coe Girão1, Melissa Souza Sobrinho1, Morgana Maria Arcanjo Bruno1, Sarah Sued Gomes de Souza1, Edson Paula Nunes1, Maria Angélica Figueiredo1, Luiz Wilson Lima-Verde1 & Maria Iracema Bezerra Loiola1

Abstract To test whether the flora is organized in discrete or continuous units along a topographic gradient, three physiognomies were assessed on different soil classes in a semi-arid region of northeastern Brazil: caatinga (xeric shrubland) at altitudes from 300 to 500 m, deciduous forest at altitudes from 500 to 700 m and carrasco (deciduous shrubland) at 700 m. In each physiognomy a species inventory was carried out, and were classified according to life- and growth-forms. Species richness was higher in the deciduous forest (250) than in the carrasco (136) and caatinga (137). The caatinga shared only a few species with the carrasco (6 species) and the deciduous forest (18 species). The highest species overlap was between the deciduous forest and the carrasco (62 species). One hundred and four species occurred only in the caatinga, 161 only in the deciduous forest and 59 only in the carrasco. Woody species predominated in physiognomies on sedimentary soils with latosol and arenosol: 124 species occurred in the deciduous forest and 68 in the carrasco. In the caatinga on crystalline basement relief with predominance of planosol, herbs showed the highest species richness (69). Comparing the biological spectrum of Brazilian life-forms, the caatinga stood out with higher proportion of therophytes and chamaephytes. Considering the flora of the three phytophysiognomies studied here, we can affirm that the caatinga is a discrete floristic unit. Key words: vegetation classification, biological spectrum, growth-form, phytoclimate, plant community. Resumo Para verificar se a composição florística constitui unidades discretas ou contínuas ao longo de um gradiente topográfico foram analisadas três fitofisionomias (caatinga sobre altitudes de 300 a 500 m, floresta decídua sobre altitudes de 500 a 700 m e carrasco sobre atitudes de 700 m) sobre classes de solos distintas no semi- árido setentrional do Nordeste do Brasil. Em cada fisionomia foi realizado o levantamento das espécies, as quais foram classificadas em formas de vida e de crescimento. A riqueza de espécies foi maior na floresta decídua (250) do que no carrasco (136) e na caatinga (137). A caatinga apresentou poucas espécies em comum com as fitofisionomias de carrasco ou de floresta decídua (6 e 18 espécies). A maior sobreposição de espécies ocorreu entre a floresta decídua e o carrasco, 62 espécies. Foram exclusivas da caatinga, floresta decídua e do carrasco, 104, 161 e 59 espécies, respectivamente. Quanto às formas de crescimento, nas fisionomias sobre relevo sedimentar com Latossolo e Arenosolo predominaram espécies lenhosas: 124 na floresta decídua e 68 no carrasco. Na caatinga sobre relevo do embasamento cristalino com predominância de Planossolo, a maior riqueza de espécies (69) foi de ervas. Na análise comparativa do espectro biológico com outras formações brasileiras, o de caatinga se destacou dos demais, constituindo uma unidade individualizada pela maior proporção de terófitos e caméfitos. Em relação à flora das três fisionomias, objeto deste estudo, pode-se afirmar que a da caatinga representa uma unidade discreta. Palavras-chave: classificação de vegetação; espectro biológico; forma de crescimento; fitoclima, comunidade vegetal.

1Universidade Federal do Ceará, Depto. Biologia, Centro de Ciências, bloco 906, Campus do Pici, 60455-760, Fortaleza, CE, Brazil. 2Correponding author: [email protected] 342 Araújo, F.S. et al. Introduction in different directions reflect environmental effects, At a global scale, the main environmental especially related to climate, on plant adaptations variables used to classify vegetation are climate observed in a community (Raunkiaer 1934). Hence, zones. A group of similar vegetation types that whereas the growth-form classification is used to occur in similar climate zones in different continents characterize community structure (because some is known as a vegetation-type or biome (Whittaker forms are dominant or more conspicuous), the life- 1975, 1978a, b; Box & Fujiwara 2005). form spectrum describes environmental adaptations Changes in topography or microclimate can of the species that compose that community affect the biology of the vegetation, leading to (Whittaker 1975; Raunkiaer 1934). Indirectly, this particularities that can be detected only at a local system provides information on local seasonality. scale (Spellerberg & Sawyer 1999). Gradual According to Whittaker (1975), life-forms are not a changes in climate related to topography or to structural attribute, but a floristic attribute: when distance from the ocean, at a small scale, result in the number of species is converted into percentage continuous vegetation units, which makes a of life-forms, this percentage would represent the classification based on floristic attributes difficult. spectrum of life-forms in this community or However, when a climate variable is associated geographic area. The fact that a given community with different soil types, the regional flora may be is characterized by particular life-forms indicates discontinuously distributed, forming discrete species convergence toward certain environmental communities, whose limits, along a topographic conditions; and this represents a functional gradient, can be determined by an analysis of attribute of the community. floristic composition and of the main growth- or In the present study, we assessed life-forms, life-forms of the plant species (Whittaker 1975; growth-forms and floristic composition of three Box & Fujiwara 2005). neighboring physiognomies that occur under To describe community types it is necessary different climates, soils and topographies. These to characterize plant forms, since physiognomy community attributes were determined for an area results from the dominant forms that compose a located in the semi-arid region of northeastern Brazil, community (Whittaker 1975). Classes or types of which comprises two geomorphological units: plant forms are called growth-forms; this sedimentary basin and crystalline basement. classification usually does not correspond to the Based on these data, we tested the following categories used by taxonomists to classify plants. predictions: i) the floras of the two geomorphological Height, woody or herbaceous habit, stem form, leaf units are different, and constitute two discrete units; form and intensity of leaf deciduousness are ii) the life-form spectrum varies according to altitude characteristics used to define the following types and soil type, probably as a consequence of of growth-forms (Whittaker 1975): trees, , differences in water availability, resulting mainly in lianas, epiphytes, herbs and thallophytes. the occurrence of phanerophytes in the sedimentary Instead of using a system of multiple basin and of therophytes in the crystalline basement. characteristics such as the growth-form system proposed by Whittaker (1975), the life-form system Material and Methods of Raunkiaer (1934) is based on a single Location and environmental characteristic: the relationship between the position characterization of the study area of the perennial tissue (meristem), which remains Serra das Almas Natural Reserve covers an inactive during the winter or dry season, and the area of 5,646 ha, and is located between the growth surface. The life-form of a species coordinates 5°15’–5°00’S and 40°15’–41°00’W represents a set of life history characteristics (Fig. 1). The study area has three physiognomies: selected by the environment. Raunkiaer (1934) i) caatinga (xeric shrubland) with an area of 17.10 classified plants into five life-forms: phanerophytes, km2 (29.19%), ii) seasonal deciduous forest with chamaephytes, hemicryptophytes, cryptophytes an area of 27.93 km2 (47.64%) and iii) carrasco and therophytes. (deciduous shrubland)(Rougerie 1988) with an The world spectrum, or normal spectrum, was area of 11.79 km2 (20.12%). calculated by Raunkiaer (1934) based on a The study area is located in two representative sample of all the vascular flora of geomorphological units: i) the crystalline basement the world. From that sample, the patterns recorded complex, with flat to slightly undulating relief and

Rodriguésia 62(2): 341-366. 2011 Floristics and life-forms along a topographic gradient 343 IMSEAR; Biodiversity inventories – Caatinga (PROBIO-MMA) and Edital Universal do CNPq / 476285/2003-8. In these studies, branches of angiosperms (five duplicates) in reproductive phase (flower buds, flowers and/or fruits) were collected on trails and inside the best-conserved fragments of each physiognomy. Vouchers were deposited in the Prisco Bezerra Herbarium (EAC), of Universidade Federal do Ceará. Botanical identification was carried out using analytical keys (Freire 1983; Barroso et al. 1978, 1984, 1986) and by comparison with the material present in the EAC Herbarium or, when necessary, by consulting specialists. The Figure 1 – Location of Serra das Almas Natural classification used was APG III (2009). Species Reserve, Crateús, state of Ceará, Brazil. names were updated considering the synonymy of Missouri Botanical Garden (Tropicos.org 2009); names and/or abbreviations of species low altitude (c. 400 m) and ii) the Meio Norte authors were written in accordance with sedimentary basin, on its eastern margin, which Brummitt & Powell (1992). forms an asymmetric cuesta, known as Ibiapaba Plateau (altitudes between 500 and 700 m). Growth- and life-forms The caatinga occurs in the crystalline Each species was classified into growth-forms basement complex, where the dominant classes of following Whittaker (1975). soils are: Solodic Planosol, Solodized Solonetz The classification of each species in life-forms (natric Planosols) and Lithic soils (Lithic Neosols) was done based on the protection level of growing at altitudes that vary from 300 to 500 m. tips and on the reduction of the aerial part during In the Meio Norte sedimentary basin, on the unfavorable season, following Raunkiaer (1934, Ibiapaba Plateau, the Latosol occurs on the eastern see also Cain 1950; Mueller-Dombois & Ellenberg hogback and quartz sand (quartzarenic neosols) 1974): therophytes (Th), cryptophytes (Cr), on the top and backside (Brasil 1972). The hemicryptophytes (H), chamaephytes (Ch) and deciduous forest occurs on the eastern hogback of phanerophytes (Ph). Woody lianas and cacti were the plateau, on Latosol, at altitudes between 500 considered as phanerophytes and non-woody and 700 m. The carrasco is present on the backside lianas were classified according to the level of of the plateau, on quartz sand, at altitudes of ca. reduction of their aerial part during the dry season 700 m. We emphasize that the Ibiapaba Plateau (according to Raunkiaer 1934). is a ‘cuesta’, with higher asymmetry in its southern part, our study area, where there is no Data analysis top, but an inverted V-shaped topography where Floristic data were organized as a list with the leeward on the backside exhibits a smooth families, species, vernacular names, life and growth- declivity. forms, physiognomy and collectors. We calculated Climate data were not available, because there species and family richness for the whole dataset and are no meteorological or pluviometric stations by physiognomy. To compare the richest families located on the cuesta, top and immediate backside between physiognomies, we used histograms with sites on the southern part of the Ibiapaba Plateau, the ten richest families in descending order. our study area. Floristic overlap between physiognomies was analyzed by calculating the frequency of species Floristic inventory and families in overlapping classes: occurrence in The flora of Serra das Almas Natural Reserve all physiognomies, in pairs of physiognomies was extensively sampled from 1999 to 2004, in (caatinga/carrasco, caatinga/deciduous forest, several projects: reserve management plan; long- carrasco/deciduous forest), and restricted to each term ecological research programs – Site Caatinga/ physiognomy (caatinga, carrasco or deciduous CNPq/PELD; Instituto do Milênio do Semiárido- forest). Results are presented in histograms.

Rodriguésia 62(2): 341-366. 2011 344 Araújo, F.S. et al.

To test for differences in the composition of life-forms among physiognomies, we calculated the life-form spectrum, which is the proportion of species of each life-form. We determined which life- form characterized each physiognomy by comparing our results with the normal spectrum proposed by Raunkiaer (1934). This spectrum represents the world flora and was used here as null hypothesis. At first, we tested for differences between the obtained and the normal spectrum using a χ2 test (Vieira 2004). When differences were significant, we calculated the relative contribution of each life-form’s deviation to the computed χ2 statistic. The life-form with higher contribution in each test was considered as characteristic of the physiognomy where it occurs. To test for similarities with other Brazilian vegetation types (in terms of life-forms), we compiled studies with spectra determined for Brazilian physiognomies (Tab. 1). We kept the names used by each author for the vegetation types of each study. To facilitate comparison, we used only the five main life-form classes of Raunkiaer (1934). Hence, epiphytes and woody lianas were included in the class phanerophytes, saprophytes in cryptophytes, and aerophytes in chamaephytes. We compared the life-form spectra found in Serra das Almas Natural Reserve with those from other studies with a detrended correspondence analysis – DCA (Jongman et al. 1995; Batalha & Martins Figure 2 – Species-richest families in the three physiognomies 2002); results were expressed in ordination of Serra das Almas Natural Reserve, Crateús, state of Ceará, diagrams with scores of each study and of each Brazil. Abbreviations for families: FAB–, EUP– life-form. Euphorbiaceae, CON- Convolvulaceae, MAL– Malvaceae, AST– Asteraceae, POA–Poaceae, APO–Apocynaceae, Results RUB – Rubiaceae, MYR–Myrtaceae, SAP–Sapindaceae, We recorded 419 species/morphospecies from ACA–Acanthaceae, PAS–Passifloraceae, COM– 72 families (Annex 1). Families (55) and species Commelinaceae, LAM–Lamiaceae, MAP–Malpighiaceae, richness (250) were higher in the deciduous forest. BRO–Bromeliaceae. Richness values of the carrasco (46 and 136) and caatinga (44 and 137) were similar to each other and lower than in the deciduous forest. Family overlap was about one third among all Fabaceae (86 species), Euphorbiaceae (38 physiognomies (Fig. 3). However, the carrasco and species) and Convolvulaceae (22 species) were the the deciduous forest shared the highest number of richest plant families in Serra das Almas Natural families, and had the highest (carrasco) and lowest Reserve. The richest families were different among (deciduous forest) number of exclusive families (Fig. 3). physiognomies (Fig. 2). The exception was the Species overlap was low, as only nine out of 419 family Fabaceae, which had the highest number of species occurred in all physiognomies (Fig. 3). The species in all three physiognomies (Fig. 2). However, carrasco and the deciduous forest had higher the representativeness of subfamilies varied, with floristic affinity with each other, since they shared higher richness of Papilionoidae in the deciduous more species (15%) and both had low overlap with forest (25 species) and of Caesalpinioidae in the the caatinga (1.3 % overlap with carrasco and 4.2% caatinga (12 species) and carrasco (15 species). with deciduous forest – Fig. 3).

Rodriguésia 62(2): 341-366. 2011 Floristics and life-forms along a topographic gradient 345 77 70 80 87 Ph 25,8 94,7 95,4 28,8 4 6 18 9,6 1,3 6,1 Ch 19,5 50,4 15,6 24,7 17,2 45,3 12,2 30,9 18,1 26,4 17,9 57,9 22,5 58,1 = grassland with hemicryptophytes, – 8 H 32 34 14 13 16 11 30 47 14,9 23,9 52,2 14,6 18,6 11,5 66,4 10,7 31,8 41,3 19,1 19,2 14,1 60,3 36,1 17,1 40,1 28,3 24,5 39,6 55,1 49,9 12,8 31,6 44,9 13,5 39,1 12,8 15,8 26,3 campo sujo 0 2 Cr 1,1 2,8 7,9 5,3 2,6 6,4 0,9 2,8 0,9 1,4 6,3 3,4 3,4 2,6 2,2 2,3 9 0 0 5 0 0 0 5 5 0 0 4 0 3 3 7 0 3 0 1,8 1,8 6,7 0,8 7,8 2,1 5,6 1,1 Th 7,5 4,6 5,4 3,7 0,7 1,8 40,5 16,8 44,2 47,9 17,2 14,6 42,9 cryptophytes, H –

r = grassland with scatered shrubs; therophytes, C – campo cerrado = open savanna; (2005) (2005) . (2007) . (2007) . (2007) . (2007) . (2007) . (2007) (1997) in Batalha & Martins (2002) et al et al. et al et al et al et al et al et al. . (2008) (2009) . (2007) (1956) (1956) . (1956) . (1956) cerrado aberto et al et al et al. et al et al et al. et al. et al arming (1892) in Batalha & Martins (2002) Meira Neto Costa Meira Neto Meira Neto Batalha & Martins (2004) Meira Neto Almeida JR Meira Neto Araújo Meira Neto Cain Cain Cain in Batalha & Martins (2002) Cain Batalha &Mantovani (2001) Batalha Mantovani (1983) in Batalha & Martins (2002) Reference This study Meira Neto W This study Batalha & Martins (2002) Ratter (1980) in Batalha & Martins (2002) This study Costa = dense savanna; cerrado fechado A = savanna; = sandy coastal plains. restinga das Emas, GO Almas, Crateús, CE Almas, Crateús, CE Almas, Crateús, CE cerrado sensu stricto ARNA Caravela, BA Betânia, PE Águas de Sta. Barbara, SP Águas de Sta. Barbara, SP Itirapina, SP Águas de Sta. Barbara, SP Itamaracá. PE Águas de Sta. Barbara, SP Quixadá, CE Águas de Sta. Barbara, SP Horto Botânico, Pelotas, RS Caiobá, PR Alto do Palmital, Foz Iguaçu, PR Mucambo, Belém, P Sta. Rita do Passa Quatro, SP Pirassununga, SP Mojiguaçu, SP Sa. das Site Sa. das Mucurí, BA Lagoa Santa, MG P Sa. das Brasília, DF Faz. Não me Deixes, Quixadá, CE = tall woodland savanna; res res res caa caa caa carr fl pl fl pl fl pl cl lp = deciduous shrubland; cp sj fl dec cer ss cer ss cerradão inselb cer ab cer ab cer ab cp cer cer fec cer fec cer fec fl temp cerradão phanerophytes. – Abbreviation carrasco = grassland; campo limpo = xeric shrubland; – Life-form spectra used for comparisons in a detrended correspondence analysis (DCA). Life-forms: Th ga chamaephytes, Ph n caatinga – able 1 egetation type restinga caatinga campo limpo campo sujo campo cerrado resti inselberg vegetation cerrado sensu strictu cerrado sensu strictu cerradão temperate forest pluvial forest pluvial forest pluvial forest cerrado fechado cerrado fechado cerrado aberto caatinga scatered shrubs; Note: Ch T V restinga carrasco cerrado aberto cerrado aberto deciduous forest cerrado fechado caatinga

Rodriguésia 62(2): 341-366. 2011 346 Araújo, F.S. et al.

Figure 3 – Proportion of families (white) and species Figure 4 – Life-form spectra of the three physiognomies (black) occurring in one, two or in all physiognomies (CA – caatinga, CR – carrasco, DF – deciduous forest) of (CA – caatinga, CR – carrasco, DF – deciduous forest) Serra das Almas Natural Reserve, Crateús, state of Ceará, of Serra das Almas Natural Reserve, Crateús, state of Brazil, compared to Raunkiaer’s normal spectrum (N). Ceará, Brazil. Values over each physiognomy bar indicate the number of species of each life-form. Species percentages of each life-form are expressed by the width of the bar. Life-forms: In the physiognomies on sedimentary relief, therophyte (Th), cryptophyte (Cr), hemicryptophyte (H), woody species (shrubs and trees) predominated, chamaephyte (Ch), phanerophyte (Ph). totaling 124 in the deciduous forest and 68 in the carrasco. In the caatinga, on the crystalline basement, the highest species richness (69) was represented by herbs. group, carrasco and deciduous forest exhibited The life-form spectra of the studied scores close to those of restinga and different from physiognomies differed significantly from the normal those of cerrado, apparently because of the lower spectrum (caatinga: χ2 = 159.33 p < 0.01 df = 4; proportion of cryptophytes (Fig. 5). The caatinga carrasco χ2 = 49.07 p < 0.01 df = 4; deciduous forest composed a well-defined group, which comprised χ2 = 120, p < 0.01 df = 4). In general, the carrasco spectra of other caatinga studies, including and the deciduous forest exhibited similar proportions vegetation on inselbergs. This group is associated of species of each life-form, whereas the caatinga with higher proportion of chamaephytes and exhibited a different spectrum (Fig. 4). Therophytes, therophytes (Fig. 5). hemicryptophytes and chamaephytes were the predominant life-forms in the caatinga (69 %), carrasco Discussion (53%) and deciduous forest (46%), respectively; thus, In general, in the semi-arid region of they characterize each physiognomy. northeastern Brazil, areas with higher annual average In the comparisons of life-form spectra among rainfall associated with higher altitudes exhibit physiognomies of Serra das Almas Natural Reserve higher species richness (Lima et al. 2009; Araújo et with other Brazilian vegetation types, the two first al. 2007; Ferraz et al. 1998; Gomes 1980). This pattern axes of the DCA corresponded to over 60% of the was also observed in the physiognomies of total inertia: 49.68% on the first axis and 13.30 % on deciduous forest and carrasco, both located at higher the second. In the ordination diagram three groups altitudes than the caatinga in Serra das Almas of life-form spectra stood out: i) spectra with scores Natural Reserve. Besides, deciduous vegetation on next to the ones of phanerophytes, ii) of sedimentary areas, even with rainfall indexes similar cryptophytes and iii) of chamaephytes and to the caatinga area of the crystalline basement, have therophytes (Fig. 5). The life-form spectra of the been pointed out in general as having higher species carrasco and the deciduous forest in Serra das richness (Silva et al. 2003), though there are some Almas Natural Reserve nearly overlapped in the exceptions (Rodal et al. 1998; Pereira et al. 2002). ordination space, in group 2, which also comprises These exceptions show that being sedimentary alone the restinga and cerrado spectra (Fig. 5). In this does not result in higher species richness; other

Rodriguésia 62(2): 341-366. 2011 Floristics and life-forms along a topographic gradient 347 in woody growth-forms is related to the increase in aridity (lower rainfall and higher temperature). In previous studies, the replacement of non-woody life-forms by woody life-forms and the increase in richness along humidity gradients have been observed in arid areas (Pavón et al. 2000), tropical savannas (Williams et al. 1996), forests and temperate grasslands (Kovács-Lang et al. 2000). Considering woody and herbaceous flora together, the deciduous forest on the sedimentary basin exhibited higher richness than the caatinga located on the crystalline basement. Potentially, there must be higher humidity in the air and soil Figure 5 – Ordination diagram of the detrended resulting from the elevation; there must be also correspondence analysis (DCA), with scores of life-forms soils with permanent water availability in deep layers and compiled inventories, including the physiognomies of (latosols and quartz sands), which possibly Serra das Almas Natural Reserve. Abbreviations for contribute to the higher floristic richness observed. vegetation types follow Table 1. Life-forms: therophyte Comparing the carrasco and the deciduous (Th), cryptophyte (Cr), hemicryptophyte (H), forest located in the same sedimentary basin, the chamaephyte (Ch), phanerophyte (Ph). latter exhibited higher richness. In this case, humidity seems to be an important factor: the deciduous forest is located on the cuesta and the factors must also be considered, such as the position carrasco on the immediate backside. On the of the hogback, level of desiccation of the relief backside the air is probably drier and wind speed is and physiochemical composition of the soil. The higher, which causes more desiccation. Besides, deciduous forest of Serra das Almas Natural soil seems to play a role too, since carrasco soils are Reserve is located on the windward side, between sandier (Araújo & Martins 1999; Araújo et al. 1999). 500 m and 700 m, whereas the carrasco, though Despite the high species richness found in located at a higher altitude about 700 m, is located the region of Ibiapaba Plateau, it is important to on the leeward side and on sandier soils, which highlight the contribution of the non-woody results in a physiognomy of lower height, smaller component (herbs, subshrubs and herbaceous and slender plants and lower richness than in the lianas) to the total species richness of each deciduous forest. physiognomy. In the caatinga, on the crystalline Concerning the herbaceous component of the basement, non-woody plants were responsible for Brazilian semi-arid flora, studies carried out in the most of the floristic richness, that is expected in inter-plateau depression of the crystalline complex arid and semi-arid climates, due to the predominance indicate that the highest richness of the caatinga of therophytes in these environments. On the contrary, sensu stricto is in the herbaceous component in the carrasco and in the deciduous forest, woody (Sampaio 1995; Rodal et al. 2005; Costa et al. 2007; plants were responsible for the highest richness, since Mamede & Araújo 2008). Comparatively, studies in more humid climates phanerophytes predominate. carried out in sedimentary formations recorded low Higher water availability favors the richness of herbaceous flora (Rodal et al. 1999; establishment of life-forms that do not need large Figueirêdo et al. 2000). reductions of the aerial shoot system during the In Serra das Almas Natural Reserve, the unfavorable season (phanerophytes), which is a floristic richness of woody species increased at necessary strategy for the survival of most species high altitudes in areas of deciduous forest and in arid and semi-arid regions (see Raunkiaer 1934; carrasco, whereas the richness of herbaceous van Rooyen et al. 1990; Kovács-Lang et al. 2000). species decreased. The increase in richness of trees In the case of Serra das Almas Natural Reserve, and shrubs with altitude seems to be a general which is inserted in a semi-arid climatic domain, the pattern for vegetation of arid and semi-arid regions. increase in altitude may potentially favor high water In the Brazilian semi-arid region, the increase in availability on the windward side. Besides, soil must richness of herbaceous growth-forms and decrease be taken into account, since there are two different

Rodriguésia 62(2): 341-366. 2011 348 Araújo, F.S. et al. geological units: lowlands of the crystalline deciduous forest occurs on the windward side basement and the Meio Norte sedimentary basin. whereas the carrasco occurs on the leeward side), these Herbaceous or sub-woody plants (herbs, do not form discrete units. They form a continuum subshrubs and herbaceous lianas) are the life-forms represented by species overlap and by the same that exhibit the highest reduction of the aerial shoot biological spectrum, as emphasized by Austin (2005). system during the dry season (therophytes, When analyzing physiognomies on different cryptophytes, and hemicryptophytes; Raunkiaer geomorphological units, apart from the climate, the 1934). The biological spectrum of the caatinga soil component may determine discrete units; studied was characterized mainly by therophytes, communities that, according to Whittaker (1975), a life-form characteristic of arid and semi-arid can be delimited by floristic composition and life- regions (Raunkiaer 1934; van Rooyen et al. 1990; forms, such is the case of the difference found Kovács-Lang et al. 2000). Indeed, among the three between the caatinga and the complex deciduous physiognomies studied, the caatinga occurs on forest + carrasco. shallow soils in the lowlands of the crystalline In the comparative analysis with the basement, where temperature is potentially higher biological spectra from other Brazilian seasonal and rainfall is potentially lower than in mountain- vegetation types, the discrimination of the caatinga range areas, resulting in lower water availability. by higher proportion of therophytes and The physiognomies on the Ibiapaba plateau chamaephytes shows that this vegetation is (carrasco and deciduous forest) must occur under composed of species whose life-forms represent lower water restrictions, since higher altitude better the semi-arid climatic pattern, since the contributes to the potential occurrence of higher predominance of these life-forms is characteristic rainfall and lower temperature, which favor of vegetations of arid and semi-arid environments phanerophytes, a life-form characteristic of sites (Raunkiaer 1934; Cain 1950). The biological with lower water restriction. spectrum is similar to the spectrum of arid and semi- In addition to numeric differences in species arid climate zones of the world. richness, remarkable differences between the In summary, the two geomorphological units floristic complexes of each physiognomy were present in the study area have two distinct floristic observed in the present study. The two main complexes, characterized by the predominance of complexes (caatinga and carrasco + deciduous therophytes on the crystalline basement and of forest) are consistent with the soil types that occur phanerophytes on the sedimentary basin. These in the area, resulting from the type of source rock. results show that when implementing reserves in Although species overlap between deciduous Brazilian semi-arid areas, abiotic local factors, such forest and carrasco may be considered low (15%), as soils and relief, must be taken into account, differences are even larger when compared with because these factors seem to reflect regional caatinga, whose overlap is only 4%. Carrasco and floristic variation. The environmental heterogeneity deciduous forest are floristically more similar may result not only in high species diversity, but because both have a set of species that prefer sandy also in high functional diversity in the Brazilian soil with low pH, whereas caatinga differs from semi-arid domain, which, in the present study, may that floristic group by the presence of species be observed in differences in life-form spectra typical of soils originated from the crystalline among the three physiognomies analyzed. basement of the inter-plateau depression. The crystalline and sedimentary floras of northeastern Acknowledgements Brazil also differ at a broader scale, as it was The non-governmental organization Associação observed in analyses of data matrices created from Caatinga funded the management plan for the local inventories, carried out in several areas of the reserve, through which the floristic inventory of Brazilian semi-arid region (Araújo et al. 1998a, b; the area was carried out. Later, studies were carried Lemos & Rodal 2002; Alcoforado-Filho et al. 2003; out with funding from Ministério de Ciência e Araújo et al. 2005; Lima et al. 2009). Tecnologia, long-term ecological research programs As Andrade-Lima (1981) emphasized, in the (CNPq/PELD – Pesquisa Ecológica de Longa Duração Brazilian semi-arid region, when the predominant – site Caatinga), Instituto do Milênio do Semiárido variation is in climate, as observed in the two (IMSEAR-MCT/CNPq), of Edital Universal do physiognomies studied in the Ibiapaba Plateau (the CNPq (proc. n°. 476285/2003-8) and PROBIO/MMA

Rodriguésia 62(2): 341-366. 2011 Floristics and life-forms along a topographic gradient 349 (Biodiversity inventories – Caatinga). Marcelo Austin, M.P. 2005. Vegetation and environment: Oliveira Teles de Menezes helped us make Figure discontinuities and continuities. In: Maarel, E.V.D. 1. Reviewers contributed for improving the final (ed.). Vegetation ecology. Blackwell Publishing, version of the manuscript. Oxford. Pp. 52-84. Barroso, G.M.; Guimarães, E.F.; Ichaso, C.L.F.; Costa, C.G.; Peixoto, A.L. & Lima, H.C. 1978. Sistemática References de angiospermas do Brasil. Vol. 1. EdUSP, São Alcoforado-Filho, F.G.; Sampaio, E.V.S.B. & Rodal, Paulo. 255p. M.J.N. 2003. Florística e fitossociologia de um Barroso, G.M.; Guimarães, E.F.; Ichaso, C.L.F.; Costa, C.G.; remanescente de vegetação caducifólia espinhosa Peixoto, A.L. & Lima, H.C. 1984. Sistemática de arbórea em Caruaru, . Acta Botanica angiospermas do Brasil. Vol. 2. UFV, Viçosa. 377p. Brasilica 17: 287-303. Barroso, G.M.; Guimarães, E.F.; Ichaso, C.L.F.; Costa, C.G.; Almeida Jr., E.B; Pimentel, R.M.M. & Zickel, C.S. 2007. Peixoto, A.L. & Lima, H.C. 1986. Sistemática de Flora e formas de vida em uma área de restinga no angiospermas do Brasil. Vol. 3. UFV, Viçosa. 326p. litoral norte de Pernambuco, Brasil. Revista de Batalha, M.A. & Martins, F.R. 2002. Life-form spectra Geografia 24: 19-34. of Brazilian cerrado sites. Flora 197: 452-460. Andrade-Lima, D. 1981. The caatingas dominium. Batalha, M.A. & Martins, F.R. 2004. Floristic, frequency, Revista Brasileira de Botânica 4: 149-153. and life-form spectra of a cerrado site. Brazilian APG III. 2009. An update of the Angiosperm Phylogeny Journal of Biology 64: 203-209. Group classification for the orders and families of Box, E.O. & Fujiwara, K. 2005. Vegetation types and flowering plants: APG III. Botanical Journal of the their broad-scale distribution. In: Maarel, E.V.D. Linnean Society 161: 105–121. (eds.). Vegetation ecology. Blackwell Publishing, Oxford. Pp. 106-128. Araújo, F.S. & Martins, F.R. 1999. Fisionomia e organização da vegetação do carrasco no planalto da Ibiapaba, estado Brasil. 1972. Mapa exploratório-reconhecimento de do Ceará. Acta Botanica Brasilica 13: 1-14. solos: estado do Ceará, escala 1:600.000. SUDENE. Brumitt, R.K. & Powell, C.E. 1992. Authors of plant names. Araújo, F.S.; Martins, F.R. & Shepherd, G.J. 1999. Richmond, Kew Royal Botanic Gardens. 732p. Variações estruturais e florísticas do carrasco no planalto da Ibiapaba, estado do Ceará. Revista Cain, S.A. 1950. Life forms and phytoclimate. Botanical Brasileira de Biologia 59: 663-678. Review 16: 1-32. Cain, S.A.; Castro, G.M.O.; Pires, J.M. & Silva, N.T. Araújo, F.S.; Gomes, V.S.; Silveira, A.P.; Figueiredo, 1956. Application of some phytosociological M.A.; Oliveirra, R.F.; Bruno, M.M.A.; Lima-Verde, techniques to Brazilian rain forest. American Journal L.W.; Silva, E.F.; Otutumi, A.T. & Ribeiro, K.A. of Botany 43: 911-941. 2007. Efeito da variação topoclimática na fisionomia Costa, R.C.; Araújo, F.S. & Lima-Verde, L.W. 2007. e estrutura da vegetação da serra de Baturité, Ceará. Flora and life-form spectrum in an area of deciduous In: Oliveira, T.S. & Araújo, F.S. (orgs.). Diversidade thorn woodland (caatinga) in northeastern, Brazil. e conservação da biota da serra de Baturité, Ceará. Journal of Arid Environments 68: 237-247. Seri&A Gráfica, Fortaleza. Pp. 73-136. Costa, K.C.; Lima, A.L.A.; Fernandes, C.F.M.; Silva, Araújo, F.S.; Rodal, M.J.N.; Barbosa, M.R.V. & Martins, M.C.N.A.; Silva, A.C.B.L.& Rodal, M.J.N. 2009. F.R. 2005. Repartição da flora lenhosa no domínio Flora vascular e formas de vida em um hectare de da caatinga. In: Araújo, F.S.; Rodal, M.J.N. & caatinga no nordeste brasileiro. Revista Brasileira Barbosa M.R.V. (orgs.). Análise das variações da de Ciências Agrárias 4: 48-54. biodiversidade do bioma caatinga: suporte a Ferraz, E.M.N.; Rodal, M.J.N.; Sampaio, E.V.S.B. & estratégias regionais de conservação. Ministério do Pereira, R.C.A. 1998. Variação florística ao longo Meio Ambiente, Brasília. Pp. 15-33. de um gradiente altitudinal no alto vale do Pajeú, Araújo, F.S.; Sampaio, E.V.S.B.; Figueiredo, M.A.; Rodal, Pernambuco. Revista Brasileira de Botânica 21: 7-13. M.J.N. & Fernandes, A.G. 1998a. Composição Figueirêdo, L.S.; Rodal, M.J.N. & Melo, A.L. 2000. florística da vegetação do carrasco, Novo Oriente, Florística e fitossociologia de uma área de vegetação CE. Revista Brasileira de Botânica 21: 105-116. arbustiva caducifólia espinhosa no município de Araújo, F.S.; Sampaio, E.V.S.B.; Rodal, M.J.N. & Buíque – Pernambuco. Naturalia 25: 205-224. Figueiredo, M.A. 1998b. Organização comunitária do Freire, C.V. 1983. Chaves analíticas para a determinação componente lenhoso de três áreas de carrasco em Novo das famílias das plantas pteridófitas, gimnospermas e Oriente - CE. Revista Brasileira de Biologia 58: 85-95. angiospermas brasileiras ou exóticas cultivadas no Araújo, F.S.; Oliveira, R.F. & Lima-Verde, L.W. 2008. Brasil. v. CCC. Coleção Mossoroense, Mossoró. 366p. Composição, espectro biológico e síndromes de Gomes, M.A.F. 1980. A vegetação dos Cariris Velhos, dispersão da vegetação de uma inselbergue no domínio no estado da Paraíba. Vegetalia – escritos e da caatinga, Ceará. Rodriguésia 59: 659-671. documentos 14 (UNESP).

Rodriguésia 62(2): 341-366. 2011 350 Araújo, F.S. et al.

Jongman, R.H.G.; Ter Braak, C.J.F. & van Tongeren, O.F.R. Rodal, M.J.N.; Lins e Silva, A.C.B.; Pessoa, L.M. & 1995. Data analysis in community and landscape Cavalcanti, A.D.C. 2005. Vegetação e flora ecology. Cambrige University Press, Cambridge. 299p. fanerogâmica da área de Betânia, Pernambuco. In: Kovács-Lang, E.; Kroel-Dulay, G.; Kertész, M.; Fekete, Araújo, F.S.; Rodal, M.J.N. & Barbosa, M.R.V. G.; Bartha, S.; Mika, J.; Dobi-Wantuch, I.; Redei, (orgs.). Análise das variações da biodiversidade do T.; Rajkai, K. & Hahn, I. 2000. Changes in bioma caatinga: suporte a estratégias regionais composition of sand grasslands along a gradient in de conservação. Ministério do Meio Ambiente, Hungary and implications for climate change. Brasília. Pp. 91-119. Phytocoenologia 30: 385-407. Rodal, M.J.N.; Nascimento, L.M. & Melo, A.L. 1999. Lemos, J.R. & Rodal, M.J.N. 2002. Fitossociologia do Composição florística de um trecho de vegetação componente lenhoso de um trecho da vegetação de arbustiva caducifólia, no município de Ibimirim, PE, caatinga no parque nacional da Serra da Capivara, Brasil. Acta Botanica Brasilica 13: 1: 15-28. Piauí, Brasil. Acta Botanica Brasilica 16: 23-22. Rougerie, G. 1988. Géographie de la biosphére. Armand Lima, J.R.; Sampaio, E.V.S.B.; Rodal, M.J.N. & Araújo, Colin Editeur, Paris. 288p. F.S. 2009. Composição florística da floresta estacional Sampaio, E.V.S.B. 1995. Overview of the Brazilian decídua montana da Serra das Almas, Ceará, Brasil. caatinga. In: Bullock, S.H.; Mooney, H.A. & Acta Botanica Brasilica 23: 756-763. Medina, E. Seasonally dry tropical forest. Cambridge Mamede, M.A. & Araújo, F.S. 2008. Effects of slash University Press, Cambridge. Pp. 35-63. and burn practices on a soil seed bank of caatinga Silva, R.A.; Santos, A.M.M. & Tabarelli, M. 2003. vegetation in Northeastern Brazil. Journal of Arid Riqueza e diversidade de plantas lenhosas em cinco Environments 72: 458-470. unidades de paisagem da Caatinga. In: Leal, I.R.; Meira Neto, J.A.A.; Souza A.L.; Lana, J.M. & Valente, Tabarelli, M. & Silva, J.M.C. (eds.). Ecologia e G.E. 2005. Composição florística espectro biológico conservação da caatinga. Ed. Universitária da e fitofisionomia da vegetação de muçununga nos UFRPE, Recife. Pp. 337-365. municípios de Caravelas e Mucurí, . Revista Spellerberg, I.F. & Sawyer, J.W.D. 1999. An introduction Árvore 29: 139-150. to applied biogeography. Cambridge University Meira Neto, J.A.A.; Martins, F.R. & Valente, G.E. 2007. Press, Cambridge. 243p. Composição florística espectro biológico na Estação van Rooyen, M.W.; Theron, G.K. & Grobbelaar, N. Ecológica de Santa Bárbara, estado de São Paulo, 1990. Life forms and spectra of flora of Namaqualand, Brasil. Revista Árvore 315: 907-922. South Africa. Journal of Arid Environments 19: Mueller-Dombois, D. & Ellenberg, H. 1974. Aims and 133-145. methods of vegetation ecology. John Willey and Vieira, S. 2004. Bioestatística, tópicos avançados – testes Sons, New York. 547p. não-paramétricos, tabelas de contingência e análise Pavón, N.P.; Hernandez-Trejo, H. & Rico-Gray, V. 2000. de regressão. 2ed. Elsevier, Rio de Janeiro. 216p. Distribution of life forms along an altitudinal gradient Whittaker, R.H. 1975. Communities and ecosytems. 2ed. in the semi-arid valley of Zapotitlán, Mexico. Macmillan Publishing, New York. 385p. Journal of Vegetation Science 11: 39-42. Whittaker, R.H. 1978a. Approaches to classifying Pereira, I.M.; Andrade, L.A.; Barbosa, M.R.V. & vegetation. In: Whittaker, R.H. (ed.). Classification Sampaio, E.V.S.B. 2002. Composição florística e of plant communities. Dr. W. Junk Publishers, The análise fitossociológica do componente arbustivo- Hague. Pp 3-31. arbóreo de um remanescente florestal no agreste Whittaker, R.H. 1978b. Dominace-types. In: Whittaker, R.H. Paraibano. Acta Botanica Brasilica 16: 241-369 (ed.). Classification of plant communities. Dr. W. Junk Raunkiaer, C. 1934. The life forms of plants and statistical Publishers, The Hague. Pp. 65-79. plant geography. Clarendon Press, Oxford. 632p. Williams, R.J.; Duff, G.A.; Bowman, D.M.J.S & Cook, Rodal, M.J.N.; Andrade, K.V.A.; Sales, M.F. & Gomes, G.D. 1996. Variation in the composition and A.P.S. 1998. Fitossociologia do componente lenhoso structure of tropical savannas as a function of rainfall de um refúgio vegetacional no município de and soil texture along a large-scale climatic gradient Buíque, Pernambuco. Revista Brasileira de Biologia and soil texture in the Northern Territory, Australia. 58: 517-526. Journal of Biogeography 23: 747-756.

Artigo recebido em 09/05/2010. Aceito para publicação em 18/12/2010.

Rodriguésia 62(2): 341-366. 2011 Floristics and life-forms along a topographic gradient 351 1 asconcelos, 8 asconcelos, 9 Araújo, 151 V V Araújo, 1400 Araújo, 1330 Araújo, 1436 Araújo, 1505 Araújo, 1539 Araújo, 1547 Araújo, 1377 Araújo, 1576 Araújo, 1458 Araújo, 1490 Araújo, 1442 Araújo, 1593 . . .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. (CR) and deciduous forest Probio, 400 F M.S. Sobrinho, 125 F F F M.S. Sobrinho, 124 F R.C. Costa 269 F F F F F F S.F F S.F F Collector carrasco x x x x x x DF (CA), x x x x x x x caatinga x x x x x x x x x x x x x x x x CA CR Phytophysiognomy Ph Cr Th Ch Th Ch Th Ph Ph Ch Th Cr Ch Ph Cr Ch H Ch Ch FV tre her her sub her sub her tre shr shr her her sub shr her sub sub sub sub FC aroeira cebola-brava mamona-brava cabeça-branca melosa-de-bode, melosa quebra-panela, pimentinha Common name (Cr). Gilg

Allemão

(L.) Kuntze Lindau Mart. cryptophyte (Nees) Morong (Nees) Lindau L. Juss. (Benth.) all. ex Clarke Moench undeuva (Tussac) Herb. W brasiliana sp. Lindau sp.

sp. (Moq.) Kuntze uon ur bahiensis sp. – List of families and species found, with respective growth-forms life-forms, in three phytophysiognomies, sp. cf. odr villosa . Gomphrena demissa Anacardiaceae Myracr Amaryllidaceae Hippeastrum var Froelichia lanata Lophothecium Ruellia villosa Alternanthera Achariaceae Lindackeria ovata Ruellia paniculata Justicia Ruellia Alstroemeriaceae Alstroemeria Justicia strobilacea Amaranthaceae Alternanthera brasiliana Justicia fragilis Bomarea edulis Dicliptera ciliaris Elytraria Acanthaceae Anisacanthus trilobus Families/species (DF), of the Natural Reserve Serra das Almas, Ceará State, deposited in EAC Herbarium Universidade Federal do Ceará. x = presence species phytophysionogmy. Growth-form (FC) = tree (tre), (shr), sub-shrub (sub), liana (lia), herb (her). Life-form (FV) phanerophyte (Ph), chamaephyte (Ch), hemicryptophyte (H), therophyte (Th), Annex 1

Rodriguésia 62(2): 341-366. 2011 352 Araújo, F.S. et al. erde, 1091 Araújo, 1290 Araújo, 1352 Araújo, 1543 Araújo, 1335 Araújo, 1323 Araújo, 1479 Araújo, 1590 Araújo, 1503 Araújo, 1497 Araújo, 1407 Araújo, 1379 . Lima-V .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. Collector Probio, 214 J.R. Lima, 18 J.R. Lima, 19 Probio, 403 F F J.R. Lima, 16 M.S. Sobrinho, 15 J.R. Lima, 89 F F M.S. Sobrinho, 245 F R.C. Costa, 97 F F F F J.R. Lima, 13 R.C. Costa, 358 F R.C. Costa, 441 R.C. Costa, 436 F L.W M.S. Sobrinho, 52 DF x x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy x x x x x x x x x x x x Ph Ph Ph FV Ph Ph Ph Ph Ph Ph Ph Ch H Cr Ph Th Ph Th Th Ch Cr Th Th Cr Th Cr Th tre tre tre FC lia tre lia tre tre tre shr lia lia her lia her shr her her lia her her her her her her her camucá canela-de-velho, caneleiro piquiá Common name pereiro-preto, pereiro pereiro-branco cravo-de-cachorro piquiá condurú bananinha pente-de-macaco, grão-de-porco milho-de-cobra agrião bamburral K.Schum.

A. DC. oodson . Blake W . . . Blake ex Pittier A. DC. Mart. ex . g. . . A. DC R. E. Fr Ar Mart. Cass. A. DC. um

A. DC. .) Cass.

(Schott) Engl. Hook. Baker (Gardner) S. F A. DC Brongn. ex Schott (J. C. Mikan) (Kunth) S.F Müll R. E. Fr (Gardner) Baker (L. f.) DC. (L. f.) DC. R. E. Fr (Hoffmanns. ex Roem. & Schult.) E. Fourn. (Sw subincanum Fontella & Morillo cf. egrinum chelii tenuifolia

per attenuata

cf. um estonia bahiensis assadia bur abernaemontana catharinensis accar Aspidosperma multiflor Families/species Annonaceae Duguetia riedeliana Secondatia floribunda Aspidosperma discolor Pr Aspidosperma cuspa Ephedranthus pisocarpus Aspidosperma pyrifolium Aspidosperma Matelea harleyi T Blainvillea lanceolata Rollinia leptopetala Apocynaceae Allamanda blanchetii T Aspilia bonplandiana Mandevilla Aspilia Mandevilla scabra Blainvillea latifolia Araceae Scaphispatha gracilis Asteraceae Acmella uliginosa Blainvillea ligulata T Spathicarpa hastifolia Blainvillea rhomboidea

Rodriguésia 62(2): 341-366. 2011 Floristics and life-forms along a topographic gradient 353 181 erde, 1 asconcelos, 4 asconcelos, 12 asconcelos, s/n V V V Araújo, 1329 Araújo, 1478 . Lima-V Araújo, 1497 Araújo, 1450 Araújo, 1560 Araújo, 1287 Araújo, 1422 Araújo, 1467 . . . .S. .S. .S. .S. .S. .S. .S. .S. F F R.C. Costa, 404 L.W S.F J.R. Lima, 23 Observada M.S. Sobrinho, 231 M.S. Sobrinho, 31 J.R. Lima, 20 R.C. Costa, 95 J.R. Lima, 21 R.C. Costa, 320 F F J.R. Lima, 85 F F M.S. Sobrinho, 236 M.S. Sobrinho, 84 M.S. Sobrinho, 109 S.F Probio, s/n S.F F F R.C. Costa, 440 R.C. Costa, 456 Collector x x x x x x x x x x x x x x x DF x x x x x x x x x x x x x x x x CA CR Phytophysiognomy FV Ph Ph Ph Ph Ph Ph Ph Ph Ph Ph Ph Ph Ph Ph Ch Th Th Ph Ph Th Th Th Th Th Th Th Th Th lia shr tre shr tre tre tre lia lia lia tre lia lia shr sub her her lia sub her her her her her her her her her FC grão-de-galo pau-branco maria-preta pacotê pau-d’arco pau-d’arco-roxo pente-de-macaco jacarandá Common name tandl. erl. V A.H.Gentry A.H. Gentry tandl. (L. f.) Baker (L.) Gardner Mart. ex DC. (Willd.) Spreng. Cass. aroda um (Mart. ex DC.) S (Thunb.) Sandwith (S. Moore) A.Gentry T Mart. ex DC. (Jacq.) Sandwith (Gardner) B. L. Rob. Gardner . (Gardner) Cabrera (Cham.) S Bureau ex K. Schum. A. DC. Less. uciger Allemão

( Humb. & Bonpl.) menthifolia Cav enaria Gardner sp. (Lag.) Less (L.) Kuntze sp. cf. ar f. tia af ufescens dia r dia leucomalloides dia oncocalyx rabidaea chica rabidaea caudigera edelia hookeriana edelia villosa ernonia obscura ernonia richogonia ournefor abebuia ochracea abebuia impetiginosa Families/species T Cor Cor Boraginaceae Cor Bixaceae Cochlospermum vitifolium T T Arrabidaea dispar Jacaranda jasminoides Pithecoctenium cr Arrabidaea corallina Ar V W Bignoniaceae Anemopaegma ataidei V W Ar T Stilpnopappus Pithecoseris pacourinoides Melanthera latifolia Melampodium camphoratum Lagascea mollis Jaegeria hirta Dissothrix imbricata Delilia biflora Centratherum punctatum

Rodriguésia 62(2): 341-366. 2011 354 Araújo, F.S. et al. 111 erde, 1 erde, 981 erde, 1222 erde, 1216 erde, 983 Araújo, 1393 Araújo, 1404 Araújo, 1473 Araújo, 1516 . Lima-V Araújo, 1401 . Lima-V . Lima-V . Lima-V . Lima-V .S. .S. .S. .S. .S. R.C. Costa, 395 F R.C. Costa, 367 F F F Probio, 326 L.W Probio, 204 Probio, 327 M.S. Sobrinho, 292 J.R. Lima 100 Probio, 563 Observada Observada Observada J.R. Lima, 48 L.W L.W L.W L.W F Collector x x x x DF x x x x x x x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy Th Th Th Th Ph Ph Ph Ph Ph Th Ph Ph Ph Ph Ph Ph Ph Ch Ch Ch Ch Th FV her her her her shr shr shr tre tre her tre tre tre shr tre shr tre her her her her her FC catinga-branca mofumbo catinga-branca cipaúba mirindiba oiticica trapiá feijão-bravo mandacaru rabo-de-raposa imburana-de-espinho macambirinha macambira-de-flexa macambira croatá Common name Eichler

Mart. (Mart.) J.B. Gillett L. B. Sm. (Aubl.) Standl. Pohl ex Eichler ahl) Eichler Mart. L. (Hook. f.) Fritsch (L.) J. Presl (V L.B.Sm. Ule Mart. ex Schult. f. (E. Morren ) L.B.Sm. (Britton & Rose) Luetzelb. DC. (M. Martens & Galeotti) D. R. Hunt sp. L. sp. sp. Dichorisandra Dichorisandra hexandra Commelina nudiflora Commelinaceae Callisia filiformis Combretum mellifluum Combretum leprosum Combretum lanceolatum Combretaceae Buchenavia capitata Cleomaceae Cleome microcarpa Combretum glaucocarpum Chrysobalanaceae Licania sclerophylla Celastraceae Maytenus Crateva tapia Capparaceae Cynophalla flexuosa Cereus jamacaru Cactaceae Cereus albicaulis Burseraceae Commiphora leptophloeos Bromelia laciniosa Bromelia plumieri Bromeliaceae Bromelia auriculata Encholirium erectiflorum Families/species Brassicaceae Brassica

Rodriguésia 62(2): 341-366. 2011 Floristics and life-forms along a topographic gradient 355 ., 7 erde, 1078 erde, 1092 asconcelos, 5 V Araújo, 1509 Araújo, 1523 Araújo, 1515 Araújo, 1395 Araújo, 1351 Araújo, 1424 Araújo, 1486 Araújo, 1522 Araújo, 1420 Araújo, 1521 Araújo, 1370 Araújo, 1372 Araújo, 1363 . Lima-V . Lima-V . asconcelos, S. F .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. J.R. Lima, 83 M.S. Sobrinho, 283 F F M.S. Sobrinho, 268 F F F F J.R. Lima, 25 Collector F R.C. Costa, 444 R.C. Costa, 448 F R.C. Costa, 92 V Costa, R. C., 453 F F F S.F M.S. Sobrinho, 183 F J.R. Lima, 106 R.C. Costa, 361 L.W L.W Probio, 199 F x x x x DF x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy x x x x x x x x x x x Ch Ch H Ch Th Th Ch Ch Th Ph FV Ch Ch Ch Th Th Ch Ch Th Ch Th Ph Ch Ch H Th H H Th Th sub sub her sub her her sub lia her lia FC lia lia lia lia her sub lia lia lia her lia lia lia her her her her her her jitirana-da-folha-pequena Common name pimenteira jitirana jitirana batata-de-purga barba de bode . Don . Don .) G Mart. (Choisy) Hallier f. (Jacq.) G .) Roem. & Schult. Dammer (Desr Ker Gawl. Moric. Rottb. Meisn. Schrank. Roem. & Schult. (Mill.) Cogn. Schrad. ex Nees L. Meisn. (Willd.) Endl. Meins. (L.) Urb. Willd. ex Roem. & Schult. (Desr Fernald Urb. Mart. Choisy sp. Lam. latifolius (L.) Roth sp. cf. ericaefolius

rosea subincana sp. aggregatus

Evolvulus macroblepharis Evolvulus pterocaulon Evolvulus ovatus Evolvulus Ipomoea asarifolia Evolvulus filipes Ipomoea bahiensis Ipomoea brasiliana Evolvulus Evolvulus Families/species Convolvulaceae Evolvulus elaeagnifolius Ipomoea hederifolia Ipomoea Merremia aegyptia Ipomoea nil Jacquemontia nodiflora Jacquemontia pentantha Cucurbitaceae Cayaponia racemosa Ipomoea polymorpha Ipomoea Ipomoea sericophylla Operculina alata Jacquemontia gracillima Cyperaceae Cyperus Cyperus laxus Cyperus surinamensis Kyllinga Rhynchospora Cyperus uncinulatus

Rodriguésia 62(2): 341-366. 2011 356 Araújo, F.S. et al. oton 14 1 199 Cr erde, 1 erde, 1 erde, 952 erde, 952 Araújo, 1346 Araújo, 1310 Araújo, 1454 Araújo, 1280 Araújo, 1356 Araújo, 1331 Araújo, 1294 Araújo, 1309 Araújo, 1342 . Lima-V Araújo, 1472 Araújo, 1339 . Lima-V Araújo, 1365 Araújo, 1322 Araújo, 1306 Araújo, 1498 Araújo, 1482 . Lima-V . Lima-V .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. F J.R. Lima, 79 Probio, 208 F F F F F F F F L.W L.W F F L.W L.W F F F R.C. Costa, 89 F R.C. Costa, 55 R.C. Costa, 366 F M.S. Sobrinho, 267 Collector x x x x x x x x x x DF x x x x x x x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy FV Ph Ph Ph Th Ph Ph Ph Ph Ph Ph Ph Ph H Ph Ph Ph Ch Th Ph Ph Ph Th Cr Cr Ch Cr shr shr shr her shr shr shr shr shr shr shr shr her shr shr shr sub her shr shr shr her lia lia lia lia FC velame canelinha marmeleiro marmeleiro-preto, marmeleiro branco catinga branca cansanção pirunga Common name . g Ar g. . Ar g. Mart. Ar Peyr Baill. g. O. E. Schulz Plowaman (L.) Small Müll O. E. Schulz Kunth Ar Plowman (Mill.) Pohl Baill. Müll. L. (Klotzsch) Müll. Baill. Baill. Baill. ell. Huber Müll. Baill. V sp. bezerrae vacciniifolium

sp. ophylloides sp.2 sp.1 multicaulis gyr

ea ovata echioides cordiifolius

dia sidoides oxylum nummularia oton oton ar ythr Families/species Croton heliotropiifolius Croton glandulosus Croton grewioides Cr Croton Croton blanchetianus Croton betaceus Cr Croton adenocalyx Cnidoscolus vitifolius Er Erythroxylum stipulosum Chamaesyce hyssopifolia Bernar Actinostemon Erythroxylum Erythroxylum laetevirens Euphorbiaceae Acalypha Erythroxylum Erythroxylum barbatum Erythroxylaceae Erythroxylum amplifolium Eriocaulaceae Syngonanthus Dioscorea Dioscorea Dioscoreaceae Dioscor Dilleniaceae Davilla cearensis

Rodriguésia 62(2): 341-366. 2011 Floristics and life-forms along a topographic gradient 357 1 erde, 1203 erde, 1044 erde, 077 asconcelos, s/n Araújo, 141 V Araújo, 1397 Araújo, 1569 Araújo, 1563 Araújo, 1321 Araújo, 1531 Araújo, 1470 Araújo, 1305 Araújo, 1318 Araújo, 1461 Araújo, 1428 Araújo, 1376 Araújo, 1325 Araújo, 1325 . Lima-V . Lima-V . Lima-V . .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. F F Probio, 408 J.R. Lima, 44 F F F M.S. Sobrinho, 54 Probio, 14 F F Probio, 273 L.W F F M.S. Sobrinho, 8 R.C. Costa, 350 J.R. Lima, 27 J.R. Lima, 29 S.F F F Probio, 40 F L.W R.C. Costa, 350 F L.W Probio 393 F Collector x x x x x x x x x x x x x DF x x x x x x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy FV Ph Ph Ph Ph Ph Ph Ch Ph Ph Th Th Ph Ph Ph Ph Ph Ph Ph Ph Th Ch Ch Ph Ph Ph Th Th Ph Ph Ph tre tre shr tre tre shr lia shr tre her her tre shr shr shr tre shr tre shr her lia sub shr shr shr her her shr shr shr FC mororó mororó mororó burra-leiteira maniçoba maniçoba pinhão canelinha marmeleiro-cravinho Common name teud. g. Ar g. Ar g.) Huber g. g. Baill. ogel ex S V ahl) Griseb. Ar Ar Ar g. g. (V Müll.

Ar (L.) Klotzsch & Garcke Ar .) (Baill.) Müll. Müll. ell. Müll. Ule (Müll. . Don. ell. (Bong.) Steud. (Pohl) Baill. Baill. (Bong.) Baill. V Moric. Baill. G Benth. V L. Lam. Pohl Müll. Pax & K. Hoffm. (Didr Müll. tiana pernambucensis

sp3. sp2. sp1. sp. dubia acuruana cf. lesser mollissima

udolphianus cf. ostachys corniculata oton lundianus oton r oton odontadenius ragia Families/species Bauhinia pentandra Bauhinia Bauhinia cheilantha Bauhinia ungulata Bauhinia pulchella T Fabaceae Caesalpinioideae Bauhinia Stillingia trapezoidea Sapium lanceolatum Poinsettia heterophylla Micr Maprounea Manihot palmata Manihot glaziovii Jatropha Manihot anomala Gymnanthes Gymnanthes Euphorbia insulana Gymnanthes Euphorbia comosa Dalechampia Croton zehntneri Croton jacobinensis Croton moritibensis Croton urticifolius Cr Croton nepetifolius Cr Cr

Rodriguésia 62(2): 341-366. 2011 358 Araújo, F.S. et al. erde, 1083 asconcelos, s/n V Araújo, 1566 Araújo, 1382 Araújo, 1538 Araújo, 1555 Araújo, 1387 Araújo, 1383 Araújo, 1390 Araújo, 1526 Araújo, 1410 Araújo, 1484 Araújo, 1368 Araújo, 1492 Araújo, 1388 Araújo, 1573 . Lima-V Araújo, s/n . .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. L.W R.C. Costa, 319 R. C. Costa, 562 R.C. Costa, 165 F R.C. Costa, 308 Probio, 365 F F R.C. Costa, 291 J. R. Lima, 46 F R.C. Costa, 401 J.R. Lima, 50 F F F M.S. Sobrinho, 57 F M.S. Sobrinho, 112 F F F F S.F R.C. Costa, 442 F Probio, 176 F F Collector x x x x x x x DF x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy FV Ph Ph Ph Ph Ph Ph Ch Ph Ph Ph Ph Ph Ph Ph Ph Ph Ph Ph Ch Ch Hh Hh Ch Ch Ch Ch H H Ch Ch tre tre tre shr shr shr sub shr shr shr shr tre tre tre tre tre shr tre sub sub her her sub sub sub sub her her sub sub FC arapiraca angico besouro besouro besouro besouro besouro besouro besouro catingueira jatobá-de-veia jucá, pau-ferro jatobá-de-porco, jatobá-batinga pau d’óleo canafístula-brava Common name . Grimes W Altschul .Fern.) H.S. Irwin & Barneby .Queiroz Afr (Griseb.) .Queiroz Greene

.Queiroz aub.) H.S. Irwin & Barneby (T . cebil ul.) L.P Moench

(Pers.) Greene T (Benth.) H.S.Irwin & Barneby (Benth.) L.P var ul.) L.P .Bezerra & (Ducke) Barneby & J. (Collad.) ogel) H. S. Irwin & Barneby (P ogel) H.S.Irwin & Barneby (T (Mart. ex Hayne) Benth. (L.) Greene (L.) (V (Nees & C. Mart.) H.S. Irwin Barneby (H. S. Irwin & Barneby) (Benth.) Britton & Rose ex Killip H. S. Irwin & Barneby (V Benth. ( DC.ex Collad.) H. S. Irwin & Barneby . Fern. Ducke ogel) H.S.Irwin & Barneby (L.) H. S. Irwin & Barneby Hayne dneriana . Don) H. S. Irwin & Barneby Afr (Mart. ex Benth.) H. S. Irwin & Barneby (V (Benth.) H. S. Irwin & Barneby (Mart. ex epens (G ea (Rich.) Willd. r ensis ugosa splendida

oleucon acacioides Families/species Inga ingoides Chlor Anadenanthera colubrina Senna trachypus Senna Senna r Senna macranthera Senna obtusifolia Senna lechriosperma Senna gardneri Senna cear Poincianella gar Poincianella bracteosa Peltogyne confertiflora Libidibia fer Hymenaea velutina Hymenaea eriogyne Chamaecrista zygophylloides Copaifera martii Chamaecrista tenuisepala Chamaecrista supplex Chamaecrista rotundifolia Chamaecrista ramosa Chamaecrista r Chamaecrista nictitans Chamaecrista duckeana Chamaecrista diphylla Chamaecrista calycioides Chamaecrista belemii Chamaecrista barbata

Rodriguésia 62(2): 341-366. 2011 Floristics and life-forms along a topographic gradient 359 197 erde, 1 asconcelos, 17 V Araújo, 1589 . Lima-V Araújo, 1371 Araújo, 1518 Araújo, 1502 Araújo, 1298 Araújo, 1426 Araújo, 1567 Araújo, 1369 Araújo, 1544 Araújo, 1441 Araújo, 1476 Araújo,1328 . .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. Probio, 277 Probio, 172 F M.S. Sobrinho, 266 Probio, 157 L.W F M.S. Sobrinho, 271 F R.C. Costa, 451 Probio, 304 R.C. Costa, 369 M.S. Sobrinho, 285 M.S. Sobrinho, 202 F S.F Probio, 335 F M.S. Sobrinho, 195 F F R.C. Costa, 286 F F F M.S. Sobrinho, 27 R.C. Costa, 399 F M.S. Sobrinho, 240 F Collector x x x x x x x x x x x DF x x x x x x x x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy FV Ch Th Th Ph Ph Ch Ch Ch Th H Ph Th Ph Ph Ch Th Ph Ph Ph Ph Ph Ph Ph Ch Ph Ph Ch Ch Ph Ph sub her her tre lia sub sub sub her lia tre her tre tre sub her tre tre shr tre tre tre tre sub shr shr lia lia tre tre FC feijão-de-rolinha sucupira mondubim imburana-de-cheiro cumarú, jurema-de-bode catanduva jurema-branca faveira jurema-preta malícia sabiá Common name . (DC.) Barneby .Bezerra . Macbr A.C. Sm. . .C. Gregory Benth. Splitg. ex Pulle

Benth. W

Benth. leptocarpa (Benth.) Luckow & Jobson . . Fern. & P . & . (L.) Mart. ex Benth. Benth. Kunth (Aubl.) J.F Poir illd.) Poir Afr var (Benth.) Seigler & Ebinger (Benth.) Ducke (DC.) Britton & Rose

(Bondt) Benth. (L.) Britton & Rose (Mart.) (Allemão) Ducke Ker Gawl. Benth. tum (W Krapov L. Mart. ex Benth. ensis Mart. Mart. ex Colla gracilis

sp. 3 sp. 2 sp. 1 distor

danii Families/species Desmodium Crotalaria vitellina Dalbergia cearensis Desmodium Desmodium Cratylia mollis Desmodium Cranocarpus Centrosema pascuorum Centrosema brasilianum Bowdichia virgilioides Arachis dar Andira surinamensis Amburana cear Aeschynomene marginata Papilionoideae Aeschynomene histrix Senegalia tenuifolia Senegalia polyphylla Senegalia langsdorffii Pityrocarpa moniliformis Piptadenia stipulacea Parkia platycephala verrucosa Mimosa ursina Mimosa quadrivalvis Mimosa invisa Mimosa sensitiva Mimosa caesalpiniifolia Mimosa acutistipula

Rodriguésia 62(2): 341-366. 2011 360 Araújo, F.S. et al. erde, 1055 asconcelos,16 Araújo, 1535 Araújo, 1586 Araújo, 1564 Araújo, 1419 Araújo, 1501 Araújo, 1406 Araújo, 1289 Araújo, 1481 Araújo, 1421 Araújo, 1375 Araújo, 1570 V . Lima-V . .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. Rabelo, J. L., 37 Collector F R.C. Costa, 328 Probio, 303 M.S. Sobrinho, 228 F J.R. Lima, 49 M.S. Sobrinho, 286 L.W F R.C. Costa, 417 F Probio, 300, 213 M.S. Sobrinho, 13 M.S. Sobrinho, 181 Probio, 418 M.S. Sobrinho, 51 R.C. Costa, 340 F M.S. Sobrinho, 118 F S.F M.S. Sobrinho, 219 F F M.S. Sobrinho, 293 F F F x DF x x x x x x x x x x x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy x x x x x x Ph Ph FV Ph Ph Ch Ph Ph Ph Ph Ph Ph Ch Ph Ph Ch Ch Ch Th Ph Th Ph Th Th Ph Ch Cr Cr Th Th lia shr FC lia tre sub lia tre tre tre tre tre lia tre shr sub sub sub her tre her tre her her tre sub her her her her mucunã Common name mucunã mulungu pau-mocó violete violete candeia banha-de-galinha mama-cachorro jacarandá, alfazema-brava, alfazema ogel ahl) Kuntze V .) DC.

(V Benth. ogel ys (Sw (Allemão) Ducke V (DC.) Benth. Mill. ogel ogel Benth. (Aubl.) Moldenke (Benth.) Ducke Kunth V Mart. ex. Benth. Kunth V Benth. (Schrader) Benth. Rolfe (Benth.) Harley Raddi (L.) Poit. Tul. Mart. ex Benth. Moldenke Willd. Epling chamaedr

ocarpa . sp sp. salzmannii

ea macr itex schaueriana atair tylosanthes capitata Dioclea megacarpa Families/species Dioclea grandiflora Erythrina velutina Harpalyce brasiliana Galactia jussiaeana Indigofera suffruticosa Lonchocarpus araripensis Luetzelburgia auriculata Machaerium stipitatum Ormosia fastigiata Machaerium acutifolium Periandra coccinea Plathymenia reticulata Platypodium elegans Rhynchosia phaseoloides Sesbania marginata S Hyptis platanifolia V V Swartzia flaemingii Hypenia Stylosanthes humilis Lamiaceae Amasonia campestris Marsypianthes Nemastylis Hyptis simulans Hyptis suaveolens Iridaceae Herbertia

Rodriguésia 62(2): 341-366. 2011 Floristics and life-forms along a topographic gradient 361 Araújo, 1408 Araújo, 1324 Araújo, 1338 Araújo, 1343 Araújo, 1444 Araújo, 1578 Araújo, 1373 Araújo, 1550 Araújo, 1536 Araújo, 1514 Araújo, 1320 Araújo, 1437 Araújo, 1553 Araújo, 1561 Araújo, 1559 .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. Probio, 01 Collector R.C. Costa, 433 F F F F M.S. Sobrinho, 289 Probio, 331 F M.S. Sobrinho, 251 M.S. Sobrinho, 94 R.C. Costa, 80 F F M.S. Sobrinho, 43 F F F F M.S. Sobrinho, 252 F J.R. Lima, 90 M.S. Sobrinho, 273 Probio, 270 F F F DF x x x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy x x x x x x x x x x x x x Ph FV Th Th Ph Th H Th Ph Th Ph Ph Ph Ph Ph Ph Ch Ph Ch Ph Ph Ch Ch Th Ch Ch Ph Ch lia FC her her lia her her her tre her lia tre lia lia lia shr lia shr sub tre shr sub sub her sub sub tre sub Common name prega-prega mutamba murici saca-rolha tingui saca-rolha embiratanha Robyns . A .) Cambess . & . t-Hil. Juss A. Juss. (A.S . (A. Juss.) B. Gates A. Juss. Anderson. R. E. Fr Goldberg (A. Juss.) B. Gates. A. Juss. .

(Griseb) B. Gates .R. L.B. Sm. t.-Hil. (Griseb.) Cuatrec. Mart. Mart. ex Sm. ex Sims W Koehne Lam. ginatum ahl (L.) Cav L. (A. Juss.) Griseb. L. Huber V A. S

mar dneriana

L. longidentata ys trichanthera cf. sp.3 sp.2 sp.1 janusioides

opter ythraceae Families/species Loasaceae Mentzelia fragilis Cuphea circaeoides Malpighiaceae Banisteriopsis angustifolia Banisteriopsis lutea L Cuphea campestris Cuphea silvestris Loganiaceae Spigelia anthelmia Guazuma ulmifolia Banisteriopsis stellaris Banisteriopsis oxyclada Malvaceae Corchorus hirtus Helicteres heptandra Peixotoa jussieuana Heter Janusia Byrsonima gar Mascagnia rigida Luehea uniflora Helicteres muscosa Sida ciliaris Pseudoabutilon spicatum Pavonia Melochia Pavonia Pseudobombax Pavonia cancellata Pavonia

Rodriguésia 62(2): 341-366. 2011 362 Araújo, F.S. et al. 102 108 erde, 1 erde, 1 erde, 988 Araújo, 1594 . Lima-V . Lima-V Araújo, 1391 Araújo, 1392 Araújo, 1291 Araújo, 1459 Araújo, 1524 Araújo, 1582 Araújo, 1434 . Lima-V .S. .S. .S. .S. .S. .S. .S. .S. J.R. Lima, 34 R.C. Costa, 318 F L.W L.W M.S. Sobrinho, 264 L.W J. R. Lima, 73 F R.C. Costa, 241 F R.C. Costa, 14 F J. R. Lima, 61 Probio, 306 Probio, 15 J. R. Lima, 31 F Probio, 159 F M.S. Sobrinho, 261 M.S. Sobrinho, 254 F R.C. Costa, 454 M.S. Sobrinho, 56 F Collector x x x x x x x x x x DF x x x x x x x x x x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy FV Ph Ph Ph Ph Ph Ph H Ph Ph Ph Ph Ph Ph Ph Ph Ch Ph Cr Th Ch Ph Ch Ch Th Ch Ch shr shr shr shr shr shr her shr shr tre tre tre tre shr tre lia shr her her sub shr sub sub her sub sub FC joão-mole pega-pinto goiabinha goiabinha canela-de-veado jacaré guabiraba inharé paco-paco malva Common name . illd. (Aubl.) Griseb. t.–Hil. urcz. W DC. urcz. Trécul T .) . T A. S

(Link) R.E.Fr O. Berg. (Schmidt) Lundell Cambess. Mill. (Kunth) DC. DC. . (Aubl.) DC. (Sw (Lam.) DC. (O. Berg) Kiaersk. Ulbr A. Juss. O. Berg.

Lindl. L. DC. opoda Cav uginea sp. r uvalha dysenterica ensis gaudichaudii obtecta

graciliflora aff. aff.

sp. cf. issadula contracta altheria macr altheria indica altheria fer altheria brachypetela richilia elegans Families/species Myrcia Myrcia guianensis Myrcia multiflora Guapira Myrcia Myrcia acutiloba Eugenia Nyctaginaceae Boerhavia coccinea Eugenia punicifolia Eugenia piauhiensis Eugenia ligustrina Eugenia flavescens Eugenia Myrtaceae Campomanesia aromatica Moraceae Brosimum Menispermaceae Cissampelos Meliaceae T Marantaceae Calathea villosa W W W W W Sida galheir Sida jussieuana Sida glomerata

Rodriguésia 62(2): 341-366. 2011 Floristics and life-forms along a topographic gradient 363 193 erde, 1 erde, 1210 Araújo, 1507 . Lima-V Araújo, 1493 Araújo, 1491 Araújo, 1340 Araújo, 1336 Araújo, 1389 Araújo, 1283 Araújo, 1349 Araújo, 1480 Araújo, 1460 Araújo, 1540 . Lima-V .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. R.C. Costa, 357 R.C. Costa, 368 F Probio, 202 L.W F F F F F R.C. Costa, 66 F L.W F F F M.S. Sobrinho, 88 M.S. Sobrinho, 294 F Probio 312 J.R. Lima, 55 Probio 285 J.R. Lima, 54 J.R. Lima, 53 Collector x x x x x DF x x x x x x x x x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy FV Th Th Th Th Ch Th Th Ch Ch Ch Ch Ph Ch Ch Ph Ch Ch Ph Th Ph Ph Ph Ph Ph her her her her sub her her sub sub sub sub shr lia sub lia sub sub tre her shr tre shr tre tre FC vassourinha chanana chanana cheira-raposa, maracujá-de- raposa, maracujá-de-estalo maracujá pau-marfim ameixa Common name ettst. W A. Juss. & Cambess.) Urb. . alter (Benth.) Radlk ex. W t.-Hil. & Rich. Miers ex Benth. & Hook. f. Urb. Engl. Mast. N. E. Br Hook. Tiegh L. (A. S L. L. Mart. ex Zucc. L. Sessé & Moc. ex DC. Sm. L. oliniensis L. ulea disticha sp. niruri car

parvifolia sp. erecta (L.) H. Hara sp. aff. cf. frutescens

urnera subulata urnera pumilea urnera blanchetiana urnera coer Families/species Phyllanthus orbiculatus Phyllanthus Phyllanthus Phyllanthaceae Phyllanthus Scoparia dulcis Dizygostemon floribundum Plantaginaceae Angelonia cornigera T T T T Piriqueta sidifolia Passiflora foetida Passifloraceae Passiflora cincinnata Piriqueta guianensis Oxalis Oxalidaceae Oxalis divaricata Opiliaceae Agonandra brasiliensis Ximenia americana Onagraceae Ludwigia Schoepfia Ochnaceae Ouratea Ouratea Olacaceae Heisteria

Rodriguésia 62(2): 341-366. 2011 364 Araújo, F.S. et al. erde, 1205 erde, 1200 asconcelos, 1402 asconcelos, 1 asconcelos, s/n asconcelos, s/n V V V Araújo, 1403 V Araújo, 1440 Araújo, 1364 Araújo, 1541 Araújo, 1412 Araújo, 1384 Araújo, 1572 Araújo, 1307 Araújo, 1385 Araújo, 1499 . Lima-V . Lima-V . . . . .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. Collector M.S. Sobrinho, 107 M.S. Sobrinho, 66 M.S. Sobrinho, 93 S.F M.S. Sobrinho, 96 R.C. Costa, 433 S.F R. C. Costa, 354 S. F M.S. Sobrinho, 92 R. C. Costa, 368 L.W F F S.F F F L.W M.S. Sobrinho, 38 F F F F R.C. Costa, 396 F F DF x x x x x x x x x x x x x x x x CA CR Phytophysiognomy x x x x x x x x x x x x x x x x x FV Ch Th Th Th Th Th Th Th Th Th Th Th Th Th Th Th Ch Th Th Th Th Th Ph Th Th Th FC sub her her her her her her her her her her her her her her her her her her her her her lia her her her Common name capim capim beldroega ebster W . . illd. . illd. . W . W Desv .) R.D. .) Willd. Jacq. . Steud. (L.) R. Br (Jacq.) Gaertn. Nees Mikan ex Trin. Michx (Sw

(Sw L. (Jacq.) L. . Beauv Sw Ekman sp. e P Hitchc. Linden ex Herrm. Aubl.

Ness L. J.C. Kunth L. (Rich.) Desv ciliaris maximum cf. tenax cf. cf. ostis eptostachys asperifolia ochloa fasciculata alinum paniculatum alinum triangular tr Families/species Plumbaginaceae Plumbago scandens Poaceae Cenchrus ciliaris Chaetium festucoides Lasiacis anomala Setaria pauciflora Eragr Pseudechinolaena Panicum sellowii Setaria geniculata Panicum Panicum trichoides Paspalum plicatulum Paspalum faveolatum Portulacaceae Portulaca pilosa T T Setaria rariflora Rhamnaceae Pontederiaceae Heteranthera limosa Polygala violacea Polygala gracilis Polygalaceae Bredemeyera floribunda Ur Polygala paniculata Setaria S Steirachne diandra

Rodriguésia 62(2): 341-366. 2011 Floristics and life-forms along a topographic gradient 365 161 erde, 1 Araújo, 1354 Araújo, 1577 Araújo, 1452 Araújo, 1399 Araújo, 1587 Araújo, 1591 Araújo, 1299 Araújo, 1380 Araújo, 1301 Araújo, 1350 Araújo, 1592 Araújo, 1358 Araújo, 1360 . Lima-V Araújo, s/n .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. M.S. Sobrinho, 248 Collector F J. R. Lima, 102 F Probio, 231 F M.S. Sobrinho, 242 M.S. Sobrinho, 75 Probio, 201 F J.R. Lima, 104 F J.R. Lima, 91 F F F M.S. Sobrinho, 257 F F F L.W F F F DF x x x x x x x x x x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy x x x x Ph FV Ph Ph Ph Ph Th Ch H Ph Th Ph H Ph Ch Ch Ch Ph Ph Ph Ph Ph Ch Ph Ph shr FC tre tre shr lia her sub her tre her tre her shr sub hemip sub shr tre tre tre tre lia shr shr Common name juazeiro espinho-judeu jenipapo-bravo genipapo-bravo chá-de-conhã jaborandi tingui-de-bola espinho-de-judeu pau-prá-tudo, laça-vaqueiro, limãozinho mama-cachorro g. (Delprete & E.B. Souza) L. (Cham. & Schltdl.) Kuntze Ar Turcz. t.-Hil. L. Cham. & Schltdl. (Cham. & Schltdl.) Steud. Radlk. Huber Reissek Aubl. Reissek Aubl. A.S Spruce ex K. Schum. Holmes Müll.

(Clos) Eichler Mart. (Cham. & Schltdl.) K. Schum. sp. tiana sericeus (Willd. ex Roem. & Schult.) Cham. Schltdl. verticillata sp.

trifoliata cf. f. barbeyana sp. af aylor & E.B. Souza cf. T olubrina cordifolia ocoyena formosa C Families/species Rubiaceae Alibertia myrciifolia Ziziphus joazeiro Gouania colurnifolia Diodia Chomelia mar Diodia radula Spermacoce Rutaceae Galipea Faramea Spermacoce T C.M. Spermacoce scabiosoides Richardia grandiflora Margaritopsis carrascoana Guettarda viburnoides Matayba guianensis Santalaceae Phoradendron Pilocarpus spicatus Magonia pubescens Salicaceae Xylosma ciliatifolia Zanthoxylum stelligerum Sapindaceae Allophylus Cardiospermum corindum

Rodriguésia 62(2): 341-366. 2011 366 Araújo, F.S. et al. 139 180 erde, 1 erde, 1 erde, 1253 Araújo, 1427 Araújo, 1304 Araújo, 1597 Araújo, 1510 Araújo, 1386 Araújo, 1575 Araújo, 1580 Araújo, 1579 Araújo, 1312 Araújo, 1557 Araújo, 1588 Araújo, 1398 Araújo, 1292 . Lima-V . Lima-V . Lima-V .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. .S. Collector F F J.R. Lima, 35 M.S. Sobrinho, 290 F F L.W R.C. Costa, 374 M.S. Sobrinho, 234 F L.W Probio, 348 R.C. Costa, 523 L.W F F F R.C. Costa, 370 F F F F F DF x x x x x x x x x x x x x x x x x x x CA CR Phytophysiognomy x x x x x Ch Ph Ph FV Ph Ph Ph Ph Ph Ph Ch Ph H Th Ph Ph Ph Ph Ph Ph Ch Ph Ch Th lia lia lia FC lia tre tre lia lia shr lia lia her her shr tre tre shr shr shr sub tre sub her Common name sabonete pitomba pepaconha jurubeba jurubeba camará, chumbinho ahl V (Rich.) Allemão (L.) Baill. t.-Hil.) Radlk. t.-Hil. Huber Cambess. L. Somner & Ferrucci . Silva (L.) Chew (Burk ex Baker) Planch. Lam. enarium Kunth T (A. S (L.) Blume L. A. S sp. Mart. Radlk. Schauer Cambess. elegans sp. cf. cearensis

fucata Lindl. ysophyllum ar ema micrantha erbenaceae rigoniaceae rigonia nivea alisia esculenta tachytarpheta cayennensis Paullinia Paullinia Families/species Cissus tinctoria Sapindus saponaria T Serjania glabrata Serjania lethalis Urvillea laevis Vitaceae Cissus gongylodes Urticaceae Laportea aestuans T T Solanum crinitum Lippia gracilis Solanaceae Solanum baturitense Manilkara Ulmaceae Tr Sapotaceae Chr Violaceae Hybanthus ipecacuaha Chrysophyllum V Lantana camara Lippia magentea S Lantana

Rodriguésia 62(2): 341-366. 2011