Revista Brasil. Bot., V.33, n.4, p.563-573, out.-dec. 2010

The coastal restinga vegetation of Pará, Brazilian Amazon: a synthesis RACHEL MACEDO DA SILVA1, ULF MEHLIG2, JOÃO UBIRATAN MOREIRA DOS SANTOS3 and MOIRAH PAULA MACHADO DE MENEZES2,4 (received: December 09, 2009; accepted: September 16, 2010)

ABSTRACT – (The coastal restinga vegetation of Pará, Brazilian Amazon: a synthesis) The present article reviews studies (some unpublished) of the vegetation of coastal sandy soils (restinga) along the coast of Pará State, northern Brazil. A total of 411 higher plant species are reported; Fabaceae, Poaceae, Cyperaceae, Rubiaceae and Myrtaceae are the most species-rich families. Nearly half of the restinga species (48%) are terrestrial herbs; palms, trees and shrubs account for 39% of the species, the remainder being lianas and epiphytes. Species are frequently wide-spread and occur in coastal areas of Southeastern Brazil as well as at inland sites in the Amazon region. Only two species appear to be exclusively coastal; whereas other species appear to exhibit a preference for sandy soils. Plant assemblages are commonly classified by means of “formations” associated with certain habitats but current data do not allow the description of well-defined plant associations. The species composition at different sites along the Pará coast does not show any clear regional grouping pattern. Seasonal changes in the composition of restinga vegetation are most probably linked to variation in ground water level. Restinga forest is mostly low and open; among the dominant tree species are Humiria balsamifera Aubl., Pouteria ramiflora (Mart.) Radlk., Anacardium occidentale L., Byrsonima crassifolia (L.) Kunth, and Tapirira guianensis Aubl. Key words - coastal forest, dune vegetation, sandy coastal plain RESUMO – (Vegetação de restinga do Estado do Pará, Amazônia Brasileira: uma síntese). Este artigo apresenta uma revisão dos estudos (alguns não publicados) da vegetação de restinga da costa do Estado do Pará, na região norte do Brasil. Ao todo foram registradas 411 espécies de plantas vasculares, sendo as famílias Fabaceae, Poaceae, Cyperaceae, Rubiaceae e Myrtaceae as mais ricas em espécies. Dentre as espécies da restinga, 48% são ervas terrestres, 39% são palmeiras, árvores e arbustos, sendo o restante constituído por lianas e epífitas. As espécies são amplamente distribuídas ocorrendo inclusive em ambientes costeiros de outras regiões brasileiras, como a região sudeste, assim como em ambientes não costeiros da Amazônia. Apenas duas espécies parecem ser exclusivamente costeiras, já outras espécies parecem ter preferência por ambientes de solo arenoso em geral. Diferentes associações de plantas são descritas e agrupadas em diferentes tipos de “formações vegetais” associadas à certos habitats, mas os dados da literatura não permitem identificar com precisão tais associações em toda a costa. Análises estatísticas mostraram que a distribuição das espécies ao longo da costa não apresentam nenhum padrão de agrupamento. Mudanças na composição da vegetação de restinga nas estações seca e chuvosa são mais provavelmente ligadas à variação do nível do lençol freático. As florestas de restinga são, em sua maioria, abertas e de pequeno porte. Entre as espécies arbóreas dominantes estão: Humiria balsamifera Aubl., Pouteria ramiflora (Mart.) Radlk., Anacardium occidentale L., Byrsonima crassifolia (L.) Kunth e Tapirira guianensis Aubl. Palavras-chave - floresta costeira, planície arenosa costeira, vegetação de dunas

Introduction Henriques (1984), who characterize restinga vegetation as the complex of plant communities occurring on the The term restinga is commonly used in Brazil to quaternary sandy coastal plains of Brazil. Restinga describe the landscape forming on coastal sandy soils as vegetation includes herbaceous plant communities as well as the associated vegetation. More specifically, it is well as assemblages dominated by shrubs or trees, with applied to sand bars forming between coastal lagoons and widely varying canopy coverage. Restinga vegetation the ocean (Suguio 1992). For our discussion of northern colonizes chêniers, active or fixed dunes and dune valleys Brazilian coastal vegetation, we follow Araújo & and is usually distinct from the neighbouring mangroves thriving on muddy sediments regularly inundated by the tides. Variety and composition of plant assemblages 1. Universidade Federal do Pará, Campus Universitário do Marajó- along the coastal plain depend on the local setting of Breves, Conj. Bandeirante s/n, 68800-000 Breves, PA, Brazil. 2. Instituto de Estudos Costeiros, Universidade Federal do Pará, marine and terrestrial environmental parameters (Alves Campus de Bragança, Al. Leandro Ribeiro s/n, 68600-000 Bragança, et al. 2007). Restinga vegetation plays an ecologically PA, Brazil. important role in stabilizing active dunes by reducing the 3. Museu Paraense Emilio Goeldi, Departamento de Botânica, Avenida Perimetral 1901, Terra Firme, 66000-000 Belém, PA, Brazil. movement of sand (Pfadenhauer 1978, Tsoar & Arens 4. Autor para correspondência: [email protected] 2003). 564 R. M. Silva et al.: The coastal restinga vegetation of Pará

Restingas occur along approximately 70% of the abstract collections, technical reports as well as graduate Brazilian coastline (Guedes et al. 2006), which exceeds monographs, master’s dissertations and doctoral studies. 8,000 km in length (Tessler & Goya 2005). However, Studies were included if the respective study sites were distribution of restinga areas is not continuous; they located along the coast of the Brazilian state of Pará (between reach their greatest extensions (up to 700 km) in Rio 0°42’ N, 50°06’ W and 1°00’ S, 46°00’ W). In this region, the climate is warm throughout the year (annual mean air Grande do Sul State in southern Brazil (Villwock 1987 temperature about 26 °C, with minimum temperatures above fide Araujo 2000). 18 °C). Precipitation is seasonal with a perhumid rainy Large areas of coastal vegetation suffer from human season between January and June. During the driest months impacts, leading to degradation of beaches and dunes (September to November), monthly precipitation is below along the entire coast of Brazil (Scarano et al. 2004). 50 mm. The annual precipitation average is about 2,500 mm Disorderly occupation of beaches along the Brazilian (INMET 1992). coast has caused loss of restinga vegetation cover Plant species lists were checked for synonyms using (Scherer et al. 2005). taxonomic databases (IPNI 2008, Tropicos 2008) and About 40% of the 1,500 km-long coastline of the checklists (Peterson et al. 2001, Zuloaga et al. 2003, Funk Amazon region belongs to the state of Pará (Suguio et al. 2007); taxonomic names from the analysed restinga articles were updated if necessary. The assignment of genera & Tessler 1984). Amazonian restingas are found to angiosperm families follows APG II (Stevens 2001, APG sporadically on dune ridges and beaches between Marajó 2003). Bay (Pará) and São Marcos Bay (Maranhão) (Seeliger Based on the species lists obtained from the literature, the 1992). In Pará State, the greatest increase in human distribution and habitat preferences of selected woody species population has been recorded for the north-eastern were investigated by analysing the respective information coastal region (Szlafsztein 2003), where restinga areas from the data base of the João Murça Pires Herbarium (MG) at are located. The state government of Pará is encouraging the Emilio Goeldi Museum of Pará, Belém. To detect possible tourism as an additional source of income for the local groupings among study sites, a dissimilarity matrix based population (Szlafstein & Stern 2007). In this context, on site-wise presence/absence lists of fully identified plant road infrastructure has been improved to facilitate tourist species was calculated. The applied dissimilarity measure was the Raup-Crick index (Raup & Crick 1979) which access to beach areas. Tourism is, however, restricted to is not sensitive to the number of species recorded at each a few sites (namely Marudá, Algodoal, Salinópolis and site, thus reducing the influence of site-specific differences Ajuruteua) within the so-called Salgado Paraense (Prost in sampling effort on the grouping procedure (e.g. Vellent & Rabelo 1996). Here and elsewhere, uncontrolled urban et al. 2007). Grouping was then analysed by non-metric expansion as well as exploitation of illegal extraction multidimensional scaling (NMDS). All calculations were sites for building materials (sand, loam) or unauthorized performed with GNU R 2.8.1 (R Development Core Team establishment of landfills has led to serious environmental 2008) with packages “vegan” (Oksanen et al. 2008) and problems within the protected coastal areas (Souza Filho “MASS” (Venables & Ripley 2002). 2001). In addition, wood cutting (Glaser et al. 2003, focussing on mangroves) and hunting (Pereira & Alvez Results and discussion 2006) impose further threats to coastal vegetation and the associated fauna. Study sites are shown in figure 1. The majority As destruction of this fragile ecosystem through of the reviewed studies are abstracts published in human impacts increases at an accelerating pace (Bastos conference proceedings (42% of the total number of et al. 2001), gathering the existing information about studies, n = 62). Articles published in scientific journals restingas and intensifying corresponding research efforts account for 26%, book chapters for 4% and doctoral, are crucial steps towards their conservation and recovery. master’s and graduate studies for 5, 10 and 10%, As a contribution towards this, we review the currently respectively; technical reports account for another 8% available literature on restinga vegetation of the coast of the reviewed studies. All consulted sources are in the of Pará and present an analysis of the current state of Portuguese language. knowledge. The first to mentionrestinga vegetation in Pará was Pires (1973), who described it as “the vegetation growing Material and methods on the white sand of the beaches and likewise on the dunes”. He further mentions Anacardium occidentale Data collection was based on articles published in L., Byrsonima crassifolia (L.) Kunth and Chrysobalanus scientific journals and books, conference proceedings with icaco L. as typical constituents of restinga vegetation. Revista Brasil. Bot., V.33, n.4, p.563-573, out.-dec. 2010 565

N

100 km 1 3 4 0°30’ S 5 6 2 7 8 9

1°00’ S 10 Marajó Bay Pará Belém 1°30’ S

48°00’ W 47°00’ W 46°00’ W

Figure 1. Restinga study sites along the coast of Pará State, Brazil (name of the corresponding administrative district in parentheses): 1. Bacurizal (Salvaterra). 2. Ilha Nova (São Caetano de Odivelas). 3. Ilha Romana (Curuça). 4. Crispim Beach (Marapanim). 5. Algodoal (Maracanã). 6. Atalaia (Salinópolis). 7. Ilha Canela (Bragança). 8. Vila Bonifácio (Bragança). 9. Urumajó (Augusto Corrêa). 10. Jabotitiua – Jatium (Viseu). Map based on the GMT coastline database (Wessel & Smith 1996).

Later, Braga (1979) suggested a phytogeographical 2008; Rubiaceae: Margalho 2008) or for all woody or subdivision of the Amazon region, where he considered herbaceous species found at a specific site (Amaral 1997, the restingas of Pará as poorly developed vegetation Costa-Neto 1999, Silva 2008). with low diversity. Studies specifically targetingrestinga Within the restinga landscape, authors differentiate vegetation were first conducted in the 1980s around between several plant formations; we use “formation” in Algodoal beach (site 5 in figure 1; Bastos 1988, Santos the sense of an assemblage of plants visibly discernible & Rosário 1988). from other assemblages by forming a distinctive structural From the 90s onward, the number of studies pattern. Formations are associated with certain habitat focussing on restinga vegetation has increased but the types and may be characterized by a distinctive set of plant more extensive studies concentrate on Algodoal, Crispim species. In denominating formations, the literature mainly beach (site 4, figure 1) and Vila Bonifácio (site 8, figure follows Araújo & Henriques (1984). The formations 1). Studies conducted at these sites considered mainly distinguished are: (1) halophile herbaceous salt-marsh floristic and phytosociological aspects of vascular plant vegetation subject to occasional tidal inundation (HSM); formations (Algodoal: Bastos 1996, Crispim: Costa- (2) psammophile herbaceous-rhizomatous vegetation of Neto et al. 1995, 2000, Amaral 1997, Costa-Neto 1999, active dunes, possibly involved in dune build-up and Bonifácio: Silva et al. 2007, Santos 2008, Silva 2008). fixation (HAD); (3) herbaceous dune valley vegetation The species assemblages at the remaining sites within the (HDV; campo de dunas); (4) patches of woody vegetation study area were characterized by “rapid assessment”-type within marshland (PWM; moitas, formação arbustiva surveys (Lisboa et al. 1993, Amaral et al. 2001, Santos aberta); (5) restinga forest (RF) of non-inundated areas et al. 2001, Bastos et al. 2002). Amaral et al. (2008) such as paleodunes. It is assumed that the influence of present a check-list of species reported at 10 sites along tidal inundation and salt spray from the sea decreases the Brazilian Amazon coast but do not consider the more from (1) to (5) but data on inundation frequency and soil recent studies at Vila Bonifácio (Silva et al. 2007, Santos pore water salinities are not available. Bastos (1996) and 2008, Silva 2008). A small number of studies provide Amaral (1997) further identify a (salt-)marsh formation local taxonomic keys for certain groups (Turneraceae: established behind fore-dunes next to the beach (brejo Vicente et al. 1999; Poaceae: Rocha et al. 2001a, b; herbáceo, herbaceous swamp: HS). HSM is described for Eriocaulaceae: Rocha & Bastos 2004; Myrtaceae: Crispim Beach only (Costa-Neto 1999) but it is possible Rosário et al. 2005; Fabaceae-Caesalpinioideae: Sousa that this formation was not clearly separated from other 566 R. M. Silva et al.: The coastal restinga vegetation of Pará types of marsh vegetation by other authors. Occurrence with patch size (between 0.7 and 339 m2) from 4 to 39, of PWM was documented for Crispim and Algodoal and that certain species do not occur when patch size (Amaral et al. 2008); the remaining formations have is too small. been observed along the entire coastline. Overall species richness is low when compared to The total number of species reported for restingas the southern Brazilian restinga flora (Rio de Janeiro State: from the coast of Pará amounts to 411, from 90 plant 1005 species, Araújo 2000, Bertioga/São Paulo State: families (Amaral et al. 2008); the most intensively 611 species, Martins et al. 2008). High species richness studied sites yielded the greatest number of species of southern Brazilian restingas has been explained by (273/77, 205/67 and 126/58 species/families at Algodoal, their proximity (both in space and species composition) Crispim and Bonifácio, respectively; Bastos 1996, to the Atlantic rainforest (Araújo 2000, Scarano 2002). Amaral 1997, Amaral et al. 2008, Santos 2008, Silva The relationship between restinga species from the 2008). The highest species richness was found in the Pará coast and the corresponding Amazonian rainforest Fabaceae family (53 species), followed by Poaceae (39), species has not been analysed so far. Large variation in Cyperaceae (35), Rubiaceae (20) and Myrtaceae (19). diversity can be expected, considering the differences Fabaceae (20) and Myrtaceae (≥ 17) featured the highest in size and habitat diversity between the extensive number of woody species (Bastos 1996, Amaral 1997, restingas of southern Brazil and those considered here. Silva 2008). Myrtaceae were important constituents Furthermore, both extension and habitat diversity may of woody restinga vegetation, as in other parts of the not be comparable among studies due to the individual country (Lemos et al. 2001, Araujo et al. 2004, Assis et researcher’s decisions regarding the delimitation of the al. 2004a; Assis et al. 2004b; Dornelles & Waechter 2004, respective study area. Scherer et al. 2005, Pimentel et al. 2007, Sacramento Table 1 shows a selection of those species reported et al. 2007, Martins et al. 2008). Dominating genera in from at least 50% of the reviewed sites with their Myrtaceae are Eugenia (≥ 7 species) and Myrcia (8). respective habitats (Costa-Neto et al. 1995, 2000, A special adaptation to restinga soils is attributed for Bastos 1996, Amaral 1997, Costa-Neto 1999, Amaral example to Myrcia cuprea (O. Berg) Kiaersk. (Ferreira et al. 2008). It is evident that there is plenty of overlap 2006). between species assemblages assigned to the different Almost one half (48%) of the plant species are habitat types. The occurrence of woody species within predominantly herbaceous, whereas 1, 17 and 21% are HS and HDV refers mostly to seedlings, which are palms, trees and shrubs, respectively; the remaining usually unable to establish themselves permanently species are lianas (11%) and epiphytes (2%). within the respective habitat (Bastos 1996), but may, Amaral (1997) was able to show that the number within HDV, succeed to form patches of woody of tree and shrub species forming patches of woody vegetation (PWM; Amaral 1997). Similarly, seedlings vegetation within marsh (PWM) at Crispim increases of mangrove tree species (Avicennia germinans (L.) L.,

Table 1. Herbaceous and woody restinga plant species from the coast of Pará and their respective habitat types. Listed are only species present at least in 5 of 10 reviewed study sites. (HSM = herbaceous salt marsh; HAD = herbaceous vegetation of active dunes; HS = herbaceous swamp; HDV = herbaceous dune valley vegetation; PWM = patches of woody vegetation within marshland; RF = restinga forest.) Species are listed in sequence of supposed ability to thrive under marine influence, derived from occupied habitat types and the corresponding distance to the sea.

Sites Species HSM HAD HS HDV PWM RF %

Herbaceous Sesuvium portulacastrum (L.) L. Aizoaceae • • 90 Blutaparon portulacoides (A. St.-Hil.) Amaranthaceae • • • 50 Mears Sporobolus virginicus (L.) Kunth Poaceae • • • • 80 Fimbristylis cymosa (Lam.) R. Br. Cyperaceae • 70 Alternanthera tenella Colla Amaranthaceae • 70 Blutaparon vermiculare (L.) Mears Amaranthaceae • 60 Canavalia rosea (Sw.) DC. Fabaceae-Faboideae • 60 continue Revista Brasil. Bot., V.33, n.4, p.563-573, out.-dec. 2010 567 continuation Sites Species HSM HAD HS HDV PWM RF % Fimbristylis spadicea (L.) Vahl Cyperaceae • • 90 Ipomoea pes-caprae (L.) R. Br. Convolvulaceae • • 100 Ipomoea imperati (Vahl) Griseb. Convolvulaceae • • 90 Vigna luteola (Jacq.) Benth. Fabaceae-Faboideae • • 60 Cassytha filiformis L. Lauraceae • • • 80 Cyperus ligularis L. Cyperaceae • • • 80 Paspalum vaginatum Sw. Poaceae • • • 70 Ambrosia microcephala DC. Asteraceae • • • 70 Eclipta prostrata (L.) L. Asteraceae • • • 60 Sarcostemma clausum (Jacq.) Schult. Apocynaceae-Asclepiadoideae • • • 50 Centrosema brasilianum (L.) Benth. Fabaceae-Faboideae • • • • • 60 Eleocharis geniculata (L.) Roem. & Cyperaceae • • • 80 Schult. Rhynchospora riparia (Nees) Boeck. Cyperaceae • • 70 Cyperus polystachyos Rottb. Cyperaceae • • 50 Scoparia dulcis L. Plantaginaceae • • 50 Stylosanthes angustifolia Vogel Fabaceae-Faboideae • • • 50 Ludwigia hyssopifolia (G. Don) Exell Onagraceae • • • 70 Desmodium barbatum (L.) Benth. & Oerst. Fabaceae-Faboideae • • • 50 Comolia villosa (Aubl.) Triana Melastomataceae • • • 50 Chamaecrista ramosa (Vogel) H.S. Irwin Fabaceae-”Caesalpinoideae” • • 50 & Barneby Crotalaria retusa L. Fabaceae-Faboideae • • 60 Turnera melochioides A. St.-Hill. & Camb. Turneraceae • 70 Cissus biformifolia Standl. Vitaceae • 70 Cissus verticillata (L.) Nicolson & C.E. Vitaceae • 70 Jarvis Woody Chrysobalanus icaco L. Chrysobalanaceae • • • • • 90 Entada polystachya (L.) DC. Fabaceae-Mimosoideae • • 60 Rhabdadenia biflora (Jacq.) Müll. Arg. Apocynaceae • • 50 Dalbergia ecastaphyllum (L.) Taub. Fabaceae-Faboideae • 90 Rhizophora mangle L. Rhizophoraceae • • 50 Laguncularia racemosa (L.) C.F. Gaertn. Combretaceae • • 50 Conocarpus erectus L. Combretaceae • • 50 Byrsonima crassifolia L. Malpighiaceae • • • 50 Annona glabra L. Annonaceae • • 70 Guettarda angelica Mart. ex Müll. Arg. Rubiaceae • • 60 Tapirira guianensis Aubl. Anacardiaceae • • • 70 Anacardium occidentale L. Anacardiaceae • • • 90 Cassipourea guianensis Aubl. Rhizophoraceae • • • 50 Myrcia cuprea (O. Berg) Kiaersk. Myrtaceae • • • 70 Eugenia biflora (L.) DC. Myrtaceae • • • 50 Himatanthus articulatus (Vahl) Woodson Apocynaceae • • • 70 Vismia guianensis (Aubl.) Choisy Clusiaceae • • • 70 Clusia grandiflora Splitg. Clusiaceae • • • 60 Licania octandra (Hoffmanns. ex Roem. Chrysobalanaceae • • • 50 & Schult.) Kuntze Copaifera martii Hayne Fabaceae-”Caesalpinoideae” • • • 60 Protium heptaphyllum (Aubl.) Marchand Burseraceae • • • 70 Pouteria ramiflora (Mart.) Radlk. Sapotaceae • • 50 Humiria balsamifera Aubl. Humiriaceae • • 50 Astrocaryum vulgare Mart. Arecaceae • • 60 568 R. M. Silva et al.: The coastal restinga vegetation of Pará

Acanthaceae-Avicennioideae; Laguncularia racemosa species restricted to restinga vegetation of northern Brazil (L.) C. F. Gaertn., Combretaceae; Rhizophora mangle (Freire & Monteiro 1993, Costa-Neto 1999). Widespread L., Rhizophoraceae) are reported in HS and PWM but along the coast of Brazil (and other tropical coasts) do not develop to full size (Bastos 1996, Santos et al. are the HAD/HSM species Sesuvium portulacastrum 2001, Bastos et al. 2002). (L.) L. (Aizoaceae), Ipomoea pes-caprae (L.) R. Br. A number of species are widespread and occur (Convolvulaceae), Sporobolus virginicus (L.) Kunth also in southern Brazilian restingas, for example the (Poaceae) and Blutaparon portulacoides (A. St.-Hil.) trees Anacardium occidentale (Anacardiaceae; Silva & Mears (Amaranthaceae), which are considered to be an Oliveira 1989, Freire 1990, Freire & Monteiro 1993, important stabilizer of active dunes (Pfadenhauer 1978, Matias & Nunes 2001, Pereira & Alvez 2006, Queiroz Cordazzo 1985, Costa-Neto 1999). 2007), Guapira opposita (Vell.) Reitz (Nyctaginaceae; It was not possible within the scope of this review, Silva & Oliveira 1989, Danilevicz et al. 1990, Rossoni to verify the identification of species by consulting & Batista 1994, Menezes & Araujo 1999, Lemos et al. all original collections deposited in the local herbaria 2001, Moreto & Mondin 2002, Sá 2002, Sztutman & MG and IAN (both in Belém, Brazil). A small Rodrigues 2002, Dornelles & Waechter 2004, Batista et number of corrections of previous misidentifications al. 2005, Neto et al. 2005, Scherer et al. 2005, Sonehara discussed by Rocha et al. (2001a, b) and Margalho 2005, Klein et al. 2007, Pimentel et al. 2007, Martins et (2008) are silently applied in table 1. Due to the less al. 2008) and Protium heptaphyllum (Aubl.) Marchand taxonomically motivated character of most studies, (Burseraceae; Silva & Oliveira 1989, Assumpção & there may be more cases of overlooked or incorrectly Nascimento 2000, Pereira & Alvez 2006, Leite et al. identified species. For example, Clusia palmicida 2007, Pimentel et al. 2007, Sacramento et al. 2007). Rich. ex Planch. & Triana (Clusiaceae) has so far These species are, however, not restricted to restinga been reported only from Bonifácio (Silva 2008); it habitats. Other common woody restinga species in Pará is probably more widespread but may not have been are Chrysobalanus icaco (Chrysobalanaceae), Humiria noted as it is easily mistaken for C. grandiflora Splitg. balsamifera Aubl. (Humiriaceae) and Byrsonima (J. B. F. Silva, unpublished data). crassifolia (Malpighiaceae). B. crassifolia is not reported Non-metric multidimensional scaling with from southeast and southern Brazil, where B. sericea DC. presence-absence data of plant species did not yield a is common (Assumpção & Nascimento 2000, Menezes distinct grouping of sites (figure 2); possible divisions & Araujo 2004, Matallana et al. 2005, Pereira & Alvez exist among sites of the Bragança-Viseu basin (sites 2006, Sacramento et al. 2007). 7-10 in figure 1) on one hand, localities around Marajó An analysis of the João Murça Pires Herbarium Bay (sites 1-3 in figure 1) on the other and then the specimen data base shows that in Pará the only woody remaining, loosely associated sites Salinópolis, Crispim restinga species restricted to the coastal zone are and Algodoal (4-6 in figure 1). Additional field work Coccoloba ramosissima Wedd. (Polygonaceae), the is needed to confirm the existence of such a regional aforementioned Myrcia cuprea (Myrtaceae) and the pattern. Hierarchical clustering based on a Bray-Curtis (= Sapindaceae Dodonaea viscosa Jacq. The latter species Czekanowski/Sørensen) dissimilarity matrix for a similar is, however, widespread and not restricted to the coastal data set presented by Amaral et al. (2008) shows a close region elsewhere. D. viscosa is not a typical lowland association between the well-sampled sites Algodoal rainforest tree, and it may be that collectors in Pará so far and Crispim and non-conclusive grouping of the other neglected, e.g., open and anthropogenically influenced localities. areas, resulting in a biased view of D. viscosa’s distribution It can be expected that the seasonal precipitation in this region. It proved difficult to ascertain whether regime of the study region induces dominance shifts species were restricted to inland sites e.g. to sandy soils, as in plant communities. Bastos (1996) documented by such information is often omitted from herbarium labels. means of wells that the ground water table at Algodoal However, occurrence on sandy soils and in campina or sunk during the dry season by 2.6, 2.9, 1.5 and 1.4 m campinarana (open shrub on poor sandy soils and white in comparison to the local wet season maximum in sand forest, respectively) environments was mentioned HAD, PWM, HDV and HS formations, respectively; for a number of species (3-23% of non-restinga vouchers her HDV and HS wells were covered by 0.1 and 0.7 m for the woody species listed in table 1). of surface water during the rainy season, respectively Among the herbaceous plants, Ambrosia (included in the figures given above). Bastos (1996) microcephala DC. (Asteraceae) is cited as the only noted corresponding changes in the composition of the Revista Brasil. Bot., V.33, n.4, p.563-573, out.-dec. 2010 569

Bastos (1996), Amaral (1997) and Silva (2008) provided data on structural properties of woody vegetation at Algodoal, Crispim and Bonifácio, respectively. Bastos (1996) considered trees and shrubs with breast height stem diameters ≥ 1.6 cm. Amaral (1997) and Silva (2008) measured the stem diameter at ground level, Amaral (1997) including all woody plants and Silva (2008) only those ≥ 2.5 cm (measured within the cylindrical stem section next to the soil surface). Coverage/basal area of woody vegetation was reported by all authors as projected stem area (at breast height or basal) per unit of ground area. Amaral (1997) and Silva (2008) observed that a great number of stems show prostrate growth, and that, as a consequence, stem length often exceeds actual crown height. Pouteria ramiflora, Anacardium occidentale and Humiria balsamifera were among the largest trees at Bonifácio, with maximum/average Figure 2. Non-metric multidimensional scaling (NMDS) basal stem diameters of 32/8, 25/12 and 29/16 cm, of restinga sites along the coast of Pará (Raup-Crick respectively (Silva 2008); even larger specimens of H. dissimilarity matrix based on occurrence of plant species; R2 balsamifera with basal diameter > 80 cm were observed for non-metric fit between observed rank dissimilarities and at Bonifácio outside Silva’s sampling plots (R. M. Silva, ordination distance: 0.99). Dotted lines show the minimum U. Mehlig and L. O. Santos, unpublished data). However, spanning tree among sites derived from NMDS results. the latter species lagged behind the more abundant P. ramiflora, A. occidentale and Byrsonima crassifolia in herbaceous vegetation, resulting in a reduced number terms of average basal area (3.3, 1.8, 1.2 and 1.1 m2 ha‑1, of species during the dry season. The same author respectively; Silva 2008). Amaral (1997) and Bastos observed a reduction of vegetation cover during the dry (1996) do not report average and maximum diameters season along with leaf fall in the persisting plants, both of single species; at Crispim, H. balsamifera is the indicators of reduced water availability. Elevation profiles species with the highest basal area (5.7 m2 ha-1) within for Algodoal and Bonifácio (Bastos 1996, Silva 2008) the larger patches of woody vegetation. Interestingly, H. show that most woody constituents of restinga vegetation balsamifera was not found by Bastos (1996) at Algodoal, were restricted to stabilised, elevated dune regions not where Byrsonima crassifolia (in PWM) and Tapirira reached by tidal inundation; structural development and guianensis (in RF) reach the highest basal areas (0.9 and species richness increased with distance from the sea 2.0 m2 ha-1, respectively). A summary of forest structure (Bastos 1996). data is given in table 2.

Table 2. Forest structure data from Algodoal, Crispim, and Bonifácio, coast of Pará State, Brazil. Sampling area refers only to the plots where forest structure data were collected; floristic data were also recorded outside of these plots. Bastos (1996) used a minimum breast height diameter of 1.6 cm, Silva (2008) a minimum ground level diameter of 2.5 cm; to facilitate comparison, data of Amaral (1997; no minimum diameter) are presented for basal diameters ≥ 2.5 cm (obtained by interpolation from his histograms).

Sampling área Density Basal área Height max. Diameter max. Diameter mean Site (ha) (indiv. ha-1) (m2 ha-1) (m) (cm) (cm)

Algodoal (Bastos 1996) 0.50 1,669** 16.2 10 > 24.5 4.6 0.05 6,060** Crispim (Amaral 1997) 0.32 1,581* to 6,023 15.4 08 > 45.3 – Bonifácio (Silva 2008) 0.46 2,293** 09.9 08 > 25.0 5.5

* = calculated for patches of woody vegetation, excluding open areas; ** = restinga forest. 570 R. M. Silva et al.: The coastal restinga vegetation of Pará

The restinga vegetation of Pará is, as far as is Apg II. 2003. An update of the Angiosperm Phylogeny Group known, rather poor compared to the more abundant flora classification for the orders and families of flowering of the sandy coastal plains of southeastern Brazil. This plants: APG II. Botanical Journal of the Linnean Society is probably due to the less extensive restinga areas in 141:399-436. Amazonia. While a few species are specialists for coastal AraUjo, D.S.D. 2000. Análise florística e fitogeográfica environments, the majority of taxa also occurs at sites das restingas do Estado do Rio de Janeiro. Tese de doutorado, Universidade Federal do Rio de Janeiro, distant from the sea; however, a number of species seem Rio de Janeiro. to prefer sandy soils in the hinterland, as well. No endemic AraUjo, D.S.D. & Henriques, R.P.B. 1984. Análise species are known and no clear pattern emerges in respect florística das restingas do Rio de Janeiro. In Restingas: to the grouping of restinga plant communities along the origem, estruturas e processos (L.D.L. Lacerda, D.S.D. coast of Pará. The currently applied terminology for plant Araujo, R. Cerqueira & B. Turcq, eds.). CEUFF, Niterói, formations is not satisfactory; transitions between the p.159-193. different environments have to be analysed ever more AraUjo, D.S.D., Pereira, M.C.A. & Pimentel, carefully, and drastic seasonal changes in the species M.C.P. 2004. Flora e estrutura de comunidades na composition of herbaceous communities are not covered restinga de Jurubatiba – Síntese dos conhecimentos com by the superficial classification “formations” provide. enfoque especial para a formação aberta de Clusia. In A more precise description of plant communities is Pesquisas de longa duração na restinga de Jurubatiba: necessary as a starting point for further ecological studies ecologia, história natural e conservação (C.F.D. Rocha, F.A. Esteves & F.R. Scarano, eds.). RiMa, São Carlos, as well as for specifying conservation and management p.59-76. strategies. Future research should not only try to identify Assis, A.M., Thomaz, L.D. & Pereira, O.J. 2004a. the plant associations and their set of characteristic species Florística de um trecho de floresta de restinga no within the different restinga habitats, but also focus on município de Guarapari, Espírito Santo, Brasil. Acta the environmental parameters that may determine the Botanica Brasilica 18:191-201. occurrence of each plant association. Assis, A.M., Pereira, O.J. & Thomaz, L.D. 2004b. Fitossociologia de uma floresta de restinga no Parque Acknowledgements – R. M. Silva is grateful for the fellowship Estadual Paulo César Vinha, Setiba, município de granted by the Conselho Nacional de Desenvolvimento Guarapari (ES). Revista Brasileira de Botânica 27:349- Científico e Tecnológico (CNPq). 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