Composition and vegetation structure in a system of coastal dunes of the “de la Plata” river, Uruguay: a comparison with Legrand’s descriptions (1959)

César Fagúndez

In 2008 Acacia longifolia (Andrews) Willd., a coastal shrub native from Australia, was found in the Florianopolis Island, an south brazilian coastal area well knowns for its conservation values and ecological vulnerability. Since then, the surface covered by Acacia longifolia has increased extensively forming thick mats, which can eliminate the local species and threats seriously the coastal systems in this place. This communication shows some preliminary results about the research project on the Acacia sp. in the Florianopolis Island, supported by University of São Paulo.

"ABSTRACT The Brazilian shrublands area (restinga) is composed of marine coast vegetation on Quaternary sandy plains, where the species composition can vary depending on the surrounding ecosystems. Th e aims of this study were to describe the structure and floristic composition of a restinga near the community of Itaúnas, in the northern part of the state of Espírito Santo, to identify any relationships between this restinga and surrounding plant formations, and to determine which are the species that occur preferentially in the coastal forests of Espírito Santo. We sampled woody plants with a diameter at breast height ≥ 2.5 cm, excluding lianas, in 50 plots of 100 m². We selected studies of coastal forests in the states of Espírito Santo and Bahia to prepare a database that would reveal patterns of floristic variation among these areas. We used two-way indicator species analysis for the identification of the species that occur preferentially in the coastal forests of Espírito Santo. We identified 114 species belonging to 38 families. Species richness was greatest for Myrtaceae (26 species), followed by Fabaceae (10). The Shannon index for the study area was 3.96. The estimated total density was 3,330 individuals/ha and basal area was 32.02 m²/ha. Th e highest importance value (IV) was for Protium heptaphyllum(IV, 23.4), indicating that it is characteristic of the Espírito Santo restinga. Th e results of our floristic analysis indicate that the species composition of the Itaúnas restinga is influenced by the so-called tabuleiroforests (coastal lowland forests on Tertiary deposits), which are most common in northern Espírito Santo. This seems to be the main factor responsible for the gradual reduction in floristic similarity between the restingas in the north of Espírito Santo and those in the south of the state, each constituting a distinct floristic block. In addition, we generated lists of species that occur preferentially in the restingaand tabuleiroforests. Th ose lists could inform decisions regarding environmental restoration programs. "

Abstract. Sand dune vegetation along the coast of the Mexican states of Tabasco and Campeche was sampled by means of 1501 releves and the data were analyzed with classification and ordination programs. 36 community types were distinguished, which were grouped into 10 higher-order groups. The communities are described in a sequence reflecting the main vegetation zonation. Beach, embryo dune and foredune community groups include 12 community types, the sheltered zone includes 16 types and the fixed dunes 8 types. The ordination results reveal one main vegetation gradient corresponding to the increase in mean distance from the shoreline and elevation, and being covariant with species richness, mean vegetation cover and mean vegetation height.Additional differences are related to the geographical transition between the Gulf and Caribbean coastal vegetation and environmental conditions, mainly type of sand, occurring in the study area. Siliceous sand and a Gulf climate are characteristic for most of Tabasco, and calcareous sand and a Caribbean-type climate are characteristic for Campeche.Disturbance caused by coconut plantations, cattle grazing and tourism favours the invasion of ruderal species from waste places behind the dunes, which attain local dominance in the dunes described here. Consequently some separate community types had to be distinguished.The distribution of community types is analyzed in the context of the transitions between calcareous and siliceous substrates in the study area.

Composition and vegetation structure in a system of coastal dunes of the “de la Plata” river, Uruguay: a comparison with Legrand’s descriptions (1959) Elena Castiñeira Latorre, Cesar Fagúndez, Edwin da Costa & Andrés Canavero Brazilian Journal of Botany ISSN 0100-8404 Braz. J. Bot DOI 10.1007/s40415-013-0009-2 1 23 Your article is protected by copyright and all rights are held exclusively by Botanical Society of Sao Paulo. This e-offprint is for personal use only and shall not be selfarchived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Braz. J. Bot DOI 10.1007/s40415-013-0009-2 ECOLOGY AND BIOGEOGRAPHY Composition and vegetation structure in a system of coastal dunes of the ‘‘de la Plata’’ river, Uruguay: a comparison with Legrand’s descriptions (1959) Elena Castiñeira Latorre • Cesar Fagúndez Edwin da Costa • Andrés Canavero • Received: 5 October 2011 / Accepted: 26 November 2012 Ó Botanical Society of São Paulo 2013 Abstract Forestation in dune systems blocks the transport of sand, making possible the extension of agriculture or urban development into coastal areas. This process, which has been taking place for a century on the Uruguayan shore, has affected the landscape and the composition and structure of plant communities. In this study, we describe the composition and structure of vegetation stands of a dune system at the seaside resort ‘‘El Pinar,’’ Canelones, Uruguay. In addition, we compare it with a previous survey published by Legrand in Anales del Museo de Historia Natural 6:73, 1959. We recorded 76 species; with Asteraceae, Poaceae, Cyperaceae, and Apiaceae the most represented families. A cluster analysis was used to identify main groups of plant associations. This analysis defined seven groups. The group associated with the foredunes environment exhibits the lowest richness, with indicative species typical of extreme psammophilic environments. Five groups occur in interdune depressions associated with humid sites. The last group was defined in the fixed dunes environment. The species composition similarity was low in comparison with Legrand0 s (1959) survey; furthermore we found a greater presence of nonnative species. We associate this change with the presence of Acacia longifolia, a species with an extremely high invasive potential, considered an ecosystem transformer. Our proposal is the development of an investigation program to assess the effectiveness and challenges of potential management practices. We also suggest applying the tactic of eradication of A. longifolia on the fixed dunes, through different practices of management (e.g., manual control operations, biologic control agents, and the use of fire). E. Castiñeira Latorre E. da Costa Centro Universitario de Rivera, Universidad de la República, Ituzaingó 667, Rivera, Uruguay Introduction E. Castiñeira Latorre (&) Licenciada en Ciencias Biológicas, Facultad de Ciencias, Universidad de la República, Rivera, Uruguay e-mail: elencasti@gmail.com C. Fagúndez Sección Ecologı́a Terrestre, Facultad de Ciencias, Universidad de la República, Iguá 4225, C.P. 11400 Montevideo, Uruguay e-mail: fagundezce@gmail.com A. Canavero Center for Advanced Studies in Ecology & Biodiversity (CASEB), Departamento de Ecologı́a, Facultad de Ciencias Biológicas, Pontificia Universidad Católica de Chile, Santiago, Chile e-mail: acanaver@gmail.com Keywords Acacia longifolia Alien species Floristic vegetation groups Forestation Human pressure Coastal areas around the world are being altered as a consequence of various uses related to an increase in urban development, industrialization, and development of tourism and recreation. The expansion of agricultural and urban areas on coastal systems of dunes has being facilitated by forestation, disrupting sand transportation (Avis and Lubke 1996; Bate and Ferguson 1996; Gadgil and Ede 1998; Kutiel et al. 2004; Carruthers et al. 2011; Kull et al. 2011). These practices have effects at biotic and abiotic levels— plant formations, physical and chemical soil components, and geomorphic changes (Kutiel et al. 2004; Marchante et al. 2008, 2009; Kim and Yu 2009). However, in order to understand the present state of an ecosystem is necessary to know its previous status before modifications—e.g., the 123 Author's personal copy E. Castiñeira Latorre et al. costal dunes of Lake Michigan (Arbogast et al. 2002), southern Israeli coastal dunes (Kutiel et al. 2004) and the Southern Baltic coast (Peyrat et al. 2009). Knowledge about the historical land use allows us to develop an understanding of dune ecology and generates tools for future conservation of this ecosystem and its functions (Antrop 2005; Foster 2006; Rhemtulla and Mladenoff 2007). At the beginning of the 20th century, the global economically driven mindset resulted in a tendency to introduce alien species with the goal of expanding and improving agro-productive and urban areas (Carruthers et al. 2011; Kull et al. 2011). As in the well-reported introduction of Acacia longifolia (Andrews) Willd in Portugal (see Marchante et al. 2008, 2009, 2010, 2011b; Kull et al. 2011), a similar forestry program was performed on the Uruguayan coast introducing this Australian acacia (Delfino and Masciadri 2005; Gutiérrez and Panario 2005; Alonso Paz and Bassagoda 2006). This species consolidates sand dunes, prevents erosion, fertilizes the soil, and provides wind protection for forestation with Pinus spp. (Kull et al. 2011). This mindset of ‘‘national development’’ of the 1900s differs from the ‘‘sustainable development’’ of the 1980s and onwards, with the incorporation of concerns about negative impacts of alien and invasive species on the environment (Carruthers et al. 2011). Acacia longifolia is one of the most successful and prolific invasive species globally, producing landscape modifications and ecosystem changes, and affecting the composition and structure of native vegetation including the loss of biodiversity (Marchante et al. 2010, 2011b, Morris et al. 2011). All of these patterns of change and replacement of native species by invasive exotic species have been reported in the coastal ecosystems of Uruguay (Chebataroff 1952; Legrand 1959; Delfino and Masciadri 2005; Alonso Paz and Bassagoda 2006). Currently, there is a growing interest in the restoration of these modified coastal systems through the elaboration of management programs (Suding 2011). A description of the present spatial patterns of vegetation, in contrast with knowledge of their past status, can be useful for detecting the impacts of human modifications (e.g., introduction of alien species) and contributing to the goals of conservation (Levin 1992; van der Meulen and Salman 1996; Forst 2009). Legrand’s work (1959)—a pioneering description of the floristic composition of the coastal area of ‘‘de la Plata’’ River extending from the eastern border of Montevideo to the Pando stream, Canelones, Uruguay—with botanical records obtained in 1934 allowed us to accomplish a comparison of plant associations with the present state of the ecosystem. These pioneering studies for the area described vegetation communities that develop in extensive fields of dunes, now converted into streets, roads, and 123 housing. Those environments were reduced from 8 km to just 200 m from the shoreline today. Despite the extreme changes observed, you can find primary dunes with an herbaceous stratum composed of Panicum racemosum (P. Beauv.), Senecio crassiflorus (Poir.) DC. and Spartina ciliata Brongn, interdunes depressions with permanent and semi permanent presence of water where you can identify patches of 0.50–2 m high composed of species like Eryngium pandanifolium Cham. & Schltdl., Hydrocotyle bonariensis Lam., Juncus sp. L., Mikania sp. Willd., Paspalum pumilum Ness, Polygonum acuminatum Kunth, Schoenoplectus californicus (C.A. Mey.) Soják, Typha dominguensis Pers. and Typha latifolia L., and a zone of fixed dunes with a 0.50 m high herbaceous stratum of Achyrocline satureioides and Androtrichum trigynum (Spreng.) H. Pfeiff. that grows mixed with an arboreal stratum of A. longifolia. The aim of this study is to describe the present composition and structure of the vegetation stands and their association with a series of environments—foredunes, interdune depressions, and fixed dunes—registered in a system of coastal dunes at the seaside resort ‘‘El Pinar,’’ Canelones, Uruguay. We also compare our data with those presented in Legrand0 s pioneering descriptions of this flora. This study allows us to reflect upon the effects of the introduction of A. longifolia and the processes that operated on the Uruguayan coastal ecosystem given their known characteristics before their urbanization. This knowledge can be useful in generating management strategies for restoration. Material and methods Study area Uruguay is located in the southeast of South America (30°350 S, 53°580 W) with a surface of 176,215 km2 and approximately 670 km of coast line (Chebataroff 1973). It is included in the phytogeographic unit Pampeana Province, which is a transition zone between Paranaense Province and the Atlantic forest (Cabrera and Willink 1973; Brussa and Grela 2007). The climate is temperate and rainy, with irregular precipitation all year round and with a mean yearly temperature of 17.5° C. The Uruguayan territory is ‘‘Cfa’’ according to Koeppen’s climatic classification (Bidegain and Caffera 1997). The study area occupies a surface of 125,560 m2 on the seashore of ‘‘de la Plata’’ River in the seaside resort ‘‘El Pinar,’’ Canelones, Uruguay (34°480 S, 55°540 W) (Figs. 1, 2). It comprises a coastal system of transgressive dunes originated by variations in the sea level during the Holocene (Panario and Piñeiro 1997; Cavallotto et al. 2004, 2005). The beach has Author's personal copy Vegetation of a coastal dune system white and fine sands with a non-continuous fore dune which shows erosive forms (Gutiérrez and Panario 2005). The ‘‘Carrasco’’ stream constitutes the western limit of the study area and toward the east the ‘‘Pando’’ stream, both with great extensions of wetlands (Fig. 1). Plant survey The sampling sites were assigned using an aerial photograph of the study area (taken on 2001 by the ‘‘Servicio de Sensores Remotos Aeroespaciales’’ of Uruguayan Air Force). We determined three environment types in the study area: foredunes, interdune depressions, and fixed dunes (Fig. 2). After an entitation carried out by visual inspection of the whole study area we considered nine vegetation stands (according to the dominant species, e.g., Acacia stand; or according to a combination of floristic characteristics and the availability of water, e.g., psammophilic grassland): AS = Acacia stand, ES = Eryngium stand, HPS = hydrophilic pasture stand, PGS = psammophilic grassland stand, PPS = psammophilic pasture stand, PSS = psammophilic steppe stand, SS = Schoenoplectus stand, ThS = Thelypteris stand, and TyS = Typha stand. 22 transects (sample units) were placed in the previously chosen vegetation stands: three in AS, two in ES, three in HPS, two in PGS, one in PPS, two in PSS, three in SS, one in ThS, and five in TyS; each transect was located randomly within each vegetation stand (Fig. 2). In order to survey the species composition we performed a point-survey (0.5 mm diameter needle) over a transect in each vegetation stand, registering the species in contact with ten needles (each separated by 2 m) per transect (Brown 1954; Kent and Coker 1994). Species were determined in the field and in the laboratory using taxonomic keys (Rosengurtt et al. 1970; Lombardo 1982, 1983, 1984; Alonso Paz 1997; Izaguirre and Beyhaut 2003a, b) and consulting national collections (MVJB-‘‘Jardı́n Botánico de Montevideo,’’ MVFA-‘‘Facultad de Agronomı́a, UDELAR, Montevideo’’) (Table 1). We deposited vouchers of reference at the national collection MVJB-‘‘Jardı́n Botánico de Montevideo.’’ The scientific names of the species list present in this work and that of Legrand0 s work follow Instituto de Botánica Darwinion (2011), Royal Botanic Gardens (2011) and W3TROPICOS (2009). In order to compare the composition of present-day vegetation at the study area with observations accomplished by Legrand, we used zones defined by him at ‘‘Cantón C’’ (the ‘‘Canton C’’ is a larger area than the study area but it includes it) where zones I and II are comparable to foredunes, zone III to interdune depressions, and zone IV to fixed dunes. Legrand used occasional surveys during 20 years to describe the floristic characteristics of the study area (see Legrand 1959).To Fig. 1 Map of Uruguay showing the location of the study area at ‘‘El Pinar,’’ Canelones, Uruguay 123 Author's personal copy E. Castiñeira Latorre et al. Fig. 2 Aerial photo of the study area (year 2001) at the seaside resort ‘‘El Pinar,’’ Canelones, Uruguay. (292 9 430 m parallel to the coastal shore, 34°480 38.600 S, 55°540 39.400 W; 34°480 46.400 S, 55°540 32.900 W; 34°480 54.200 S, 55°540 47.000 W; 34°480 46.600 S, 55°540 53.300 W). We indicate the three environments (fixed dunes, interdune depressions, and foredunes), the sampling sites and the vegetation stands. AS Acacia stand, ES Eryngium stand, HPS hydrophilic pasture stand, PGS psammophilic grassland stand, PPS psammophilic pasture stand, PSS psammophilic steppe stand, SS Schoenoplectus stand, ThS Thelypteris stand, and TyS Typha stand determine the status of conservation of species we used the data base of the ‘‘IUCN Red List of Threatened Species’’ (IUCN 2009), a local classification generated at the ‘‘Laboratorio de Botánica, Facultad de Agronomı́a, Universidad de la República, Uruguay’’ (Marchesi et al. 2008), and the species conservation priority report for Uruguay conceived by a group of experts for the SNAP (National System of Protected Areas) (Alonso Paz et al. 2009). Data were collected in three field works between the months of January and March 2009. Statistical analysis We developed an agglomerative cluster analysis using the Jaccard index as a distance metric and the Ward’s method as the fusion algorithm. We incorporated an indicator species analysis (ISA) to detect the most informative level of cluster and indicative species of each group (Dufrene and Legendre 1997). The species indicator value (IVkj = RAkj 9 RFkj 9 100, where IVkj is the indicator value for the species j in group k, RAkj is the relative abundance of species j in group k, and RFkj is the relative frequency of species j in group k.) combines information on the concentration of species abundance in a particular group and the faithfulness of occurrence in a particular group (Dufrene and Legendre 1997). In each group it varies between 0 % (no indication) and 100 % (perfect indication) (Table 1). Statistical significance of the IV was tested using a Monte Carlo permutation test (10,000 replication) (McCune and Mefford 1999). The most informative level 123 of the cluster was assigned calculating the maximum value of the sum of the IV of each species greater than 70 % (McGeoch et al. 2002; Gautreau and Lezama 2009). ISA was run on PC-ORD program (McCune and Mefford 1999). We used the software PAST. v. 1.24 to estimate the rarefaction curves for each plant group (Magurran 1988; Gotelli and Graves 1996; Hammer et al. 2001). We used the richness estimator Chao 2 computed with log-linear 95 % confidence intervals (Chao 1987) to estimate the species richness of total sample with the program estimates 8.2 (Colwell 2006). We also calculated the Jaccard Index to evaluate the degree of similarity between the species list of our study with that of Legrand (1959) [J = (shared species)/(exclusive species of Legrand study ? exclusive species of current study ? shared species)], and to evaluate the degree of similarity between the species of the three principal different environments observed in the study area (foredunes vs. zone I ? II of Legrand; interdunes depressions vs. zone III of Legrand; fixed dunes vs. zone IV of Legrand) [J = (shared species)/(exclusive species of Legrand0 s zone ? exclusive species of the environment types of current study ? shared species)]. Since Legrand used occasional surveys along 20 years to describe the floristic characteristics of the study area (see Legrand 1959), to make his survey comparable with our data, not only do we included the transect survey in this analysis but we also included species registered off of transects. This exception was made only for the calculation of the Jaccard Index to compare the Legrand0 s survey with data presented in this study. Species Carpobrotus edulis (L.) N.E. Br. PS Transects Family 1 * Aizoaceae * Apiaceae Sagittaria montevidensis Cham. & Schltdl. Centella asiatica (L.) Urb. Vegetation stands Eryngium serra Cham. & Schltdl. Apiaceae Hydrocotyle bonariensis Lam. Apiaceae Achyrocline satureioides (Lam.) DC. Asteraceae Baccharis articulata (Lam.) Pers. Asteraceae Baccharis dracunculifolia DC. Asteraceae Baccharis gnaphalioides Spreng. Asteraceae Baccharis spicata (Lam.) Baill. Asteraceae Baccharis trimera (Less.) DC. Asteraceae Chevreulia sarmentosa (Pers.) S.F. Blake Asteraceae * 1 2 3 1 TyU 2 3 1 EU 2 3 4 5 1 2 10 10 ThU PG 1 1 AU 2 1 IV 2 3 Asteraceae Enhydra sessilis (Sw.) DC. Asteraceae Eupatorium buniifolium Hook. Arn. Asteraceae Eupatorium tremulum Hook. & Arn. Asteraceae Gamochaeta coarctata Cabrera Asteraceae * 2 5 1 3 1 3 5 (97.2) 1 1 3 1 3 (54.8) 1 2 7 (60) 1 1 2 1 5 1 1 Asteraceae Mikania micrantha Kunth Asteraceae Mikania sp. Willd. Asteraceae Pluchea sagittalis (Lam.) Cabrera Asteraceae Pterocaulon lorentzii Malme Asteraceae Senecio crassiflorus (Poir.) DC. Asteraceae Senecio selloi (Spreng.) DC. Asteraceae * 1 1 Asteraceae Conyza bonariensis (L.) Cronquist Sonchus asper (L.) Hill 1 HP Author's personal copy Apiaceae Hypochoeris radicata L. SU Alismataceae Eryngium pandanifolium Cham. & Schltdl. Cirsium sp. Mill. 2 PP 2 6 1 2 2 1 9 1 1 5 1 1 9 3 (90.9) 2 1 1 (66.7) Asteraceae Androtrichum trigynum (Spreng.) H. Pfeiff. Cyperaceae Carex riparia Curtis Cyperaceae 123 Eleocharis sellowiana Kunth Cyperaceae Eleocharis sp. R. Br. Cyperaceae Schoenoplectus californicus (C.A. Mey.) Soják Cyperaceae Scirpus giganteus Kunth Cyperaceae Scirpus olneyi A. Gray Cyperaceae Eriocaulon sp. L. Eriocaulaceae 1 1 1 6 9 7 4 9 3 7 (54.9) 6 1 1 9 2 1 10 2 (75) 1 3 4 2 1 1 Vegetation of a coastal dune system Table 1 List of species, family, vegetation stands where it was registered (AS Acacia stand, ES Eryngium stand, HPS hydrophilic pasture stand, PGS psammophilic grassland stand, PPS psammophilic pasture stand, PSS psammophilic steppe stand, SS Schoenoplectus stand, ThS Thelypteris stand, and TyS Typha stand), and the indicator value (IV) are shown between brackets and outside of the brackets is the number of the groups, 1 to 7, of the agglomerative analysis that the species is the indicator of (only taxa with significant values, P \ 0.05) 123 Table 1 continued Species Acacia longifolia (Andrews) Willd. * Vegetation stands PS Transects Family 1 Haloragaceae Juncus maritimus Lam. Juncaceae Juncus sp. L. Juncaceae Hyptis floribunda Briq. Ex Micheli Lamiaceae Eugenia uniflora L. Myrtaceae Ludwigia hookeri (Micheli) H. Hara Onagraceae Ludwigia peploides (Kunth) P.H. Raven Onagraceae 2 3 1 2 3 1 EU 2 3 4 5 1 2 ThU PG 1 1 AU IV 2 1 2 3 1 9 10 9 7 (80) 1 4 2 1 4 (92.3) 7 2 Pinaceae 1 1 Plantaginaceae Andropogon arenarius Hack. Poaceae Andropogon sp. L. Poaceae Chascolytrum subaristatum (Lam.) Desv. Poaceae Cortaderia selloana (Schult. & Schult. f.) Asch. & Graebn. Poaceae Panicum schwackeanum Mez 1 TyU Onagraceae * Plantago sp. L. Cynodon dactylon (L.) Pers. 1 HP Author's personal copy Pinus maritima Mill. SU Fabaceae Myriophyllum aquaticum (Vell.) Verdc. Oenothera affinis Cambess. 2 PP * 1 1 1 Poaceae Poaceae Paspalum notatum Flüggé Poaceae Paspalum pumilum Ness Poaceae Spartina ciliata Brongn. Poaceae Sporobolus indicus (L.) R. Br. Poaceae Polygala leptocaulis Torr. & A. Gray Polygalaceae Polygonum acuminatum Kunth Polygalaceae Polygonum punctatum Elliott Polygalaceae Rhynchospora sp. Vahl. Potamogetonaceae Margyricarpus pinnatus (Lam.) Kuntze Rosaceae Rubiaceae Richardia brasiliensis Gomes Rubiaceae Salvinia auriculata Aubl. Salviniaceae Dodonaea viscosa Jacq. Sapindaceae Agalinis communis (Cham. & Schltdl.) D’Arcy Scrophulariaceae Bacopa monnieri (L.) Wettst. Scrophulariaceae Solanum americanum Mill Solanaceae Solanum sisymbrifolium Lam. Solanaceae Sphagnum sp. (Herb Linn) Sphagnaceae 3 6 1 1 1 7 4 1 (100) 6 6 3 9 9 4 1 7 2 4 (66.2) 1 7 1 (62.9) 1 5 6 7 4 9 1 6 1 1 2 (58.5) 3 1 1 8 1 1 1 1 4 E. Castiñeira Latorre et al. Cephalanthus glabratus (Spreng.) K. Schum. 1 1 Poaceae Panicum racemosum (P. Beauv.) Spreng. 1 Author's personal copy Vegetation of a coastal dune system 1 1 20 1 1 1 10 1 Typhaceae Vitaceae Typha latifolia L. Cissus striata Ruiz & Pav. * Nonnative species Thelypteridaceae Typhaceae Thelypteris rivularioides (Fée) Abbiatti Typha dominguensis Pers. 1 Transects Family The scientific names follow Instituto de Botánica Darwinion (2011), Royal Botanic Gardens (2011) and W3TROPICOS (2009) 10 1 1 1 1 PS Vegetation stands Species Table 1 continued 2 PP SU 2 3 HP 2 3 TyU 2 10 3 10 4 8 5 EU 2 ThU PG 2 AU 2 3 IV 6 (92.7) Results We registered the presence of 75 species from 27 families (Table 1). The families with the greatest number of species were Asteraceae (S = 21), Poaceae (S = 11), Cyperaceae (S = 7), and Apiaceae (S = 4) (Fig. 3). Of 75 species, 50 were registered with point sampling and 25 observed occasionally outside of the transects. Estimates of total expected species diversity of plants at 22 sites (with the point-survey) predicted that nearly 23 additional species are still remaining to be found in addition to the 50 already recorded (Chao 2 ± 1 SD; 73.2 ± 14.3) (Fig. 4). Considering the species recorded in the study area (S = 75), we are close to the expected richness predicted by the estimation analysis. From the 80 species registered by Legrand (1959) (for the same study area), 64 species were absent in our samples (Table 2). In contrast, we registered 59 species not registered by Legrand and we share 16 species (Table 2). The Jaccard index (J = 0.16) confirms a low degree of similarity between both species lists. The agglomerative classification analysis and the sum of significant IV jointly define seven groups for the plant association (Fig. 5). These were grouped in three higher groups that are associated with the following environments: foredunes, interdune depressions, and fixed dunes (Fig. 2). Group 1, Psammophilic steppe stands and Psammophilic pasture stand (PSS1, PSS2, PPS) (Fig. 2). This is an herbaceous stratum that grows on fine sands on foredunes. The indicative species is P. racemosum (P. Beauv.) Spreng. (IV = 100), but other species presented high IV: S. crassiflorus (Poir.) DC. (IV = 66.7) and S. ciliata Brongn. (IV = 62.9) (Table 1). This group presented the lowest observed richness (S = 5). This group is located in areas equivalent to zone I and II in Legrand0 s work (1959) where the most abundant species were P. racemosum and S. crassiflorus (Table 2). The degree of similarity between the species of the foredunes (present data) versus zone I ? II of Legrand report a low degree of similarity (J = 0.19). Group 2, Schoenoplectus stand and Thelypteris stand (SS1, SS2, SS3, ThS) (Fig. 2). This is an herbaceous stratum 2-meters high over a peaty ground flooded permanently or semipermanently located in the interdune depressions. The indicative species is S. californicus (C.A. Mey.) Soják (IV = 75) followed by P. acuminatum Kunth (IV = 58.5) (Table 1). This group presented an observed richness of S = 17. Group 3, Hydrophilic pasture stand and Typha stand (HPS1, TyS3) (Fig. 2). This is a dense pasture located in the interdune depressions. The Typha-stand of T. latifolia L. grows in the lowest zone of the study area. The indicative species is Mikania sp. Wild. (IV = 90.9) (Table 1) followed by H. bonariensis Lam. (IV = 54.8) (Table 1). This group presented an observed richness of 123 Author's personal copy E. Castiñeira Latorre et al. Fig. 3 Frequencies diagram: number of species per family registered in the study area S = 10. Group 4, Hydrophilic pasture stands (HPS2, HPS3) (Fig. 2). This is a dense pasture located in the interdune depressions which only reaches 0.5-meters in height. The indicative species is Juncus sp. L. (IV = 92.3) followed by P. pumilum Ness (IV = 66.2) (Table 1). This group presented an observed richness of S = 7. Group 5, Eryngium stands (ES1, ES2) (Fig. 2). It is an herbaceous stratum continuum of 1.5-meters in height that grows in a semipermanent flooded area located in the interdune depressions. The indicative species is E. pandanifolium Cham. & Schltdl. (IV = 97.6) (Table 1). This group presented an observed richness of S = 12. Group 6, Typha stands (TyS1, TyS2, TyS4, TyS5) (Fig. 2). It is a gramineous stratum continuum of 2 m in height that grows in a permanent flooded area located in the interdune depressions. The indicative species is T. dominguensis Pers. (IV = 92.7) (Table 1). This group presented the highest observed richness (S = 19). Groups 2–6 are located in humid areas equivalent to zone III in Legrand’s work where the most abundant species were: Cyperus prolixus Kunth, Juncus microcephalus Kunth, Leiothrix arechavaletae (Körn.) Ruhland, Lycopodiella alopecuroides (L.) Cranfill, Scleria distans Poir., Typha domingensis and Xyris jupicai Rich (Table 2). The comparison between interdunes depressions (present data) versus zone III of Legrand also report a low degree of similarity (J = 0.22). Group 7, Acacias stands and psammophilic grassland stands (PGS1, PGS2, AS1, AS2, AS3) (Fig. 2). It is a 0.50 m high herbaceous stratum that grows on fixed dunes mixed with an arboreal stratum of A. longifolia. The indicative species is A. longifolia (IV = 80.0) followed by A. satureioides (Lam.) DC. (IV = 60.0) and A. trigynum 123 (Spreng.) H. Pfeiff. (IV = 54.9) (Table 1). This group had the same values of observed richness as group 6. This group is located in areas equivalent to zone IV in Legrand0 s work (1959) where the most abundant species were A. satureioides, Andropogon selloanus (Hack.) Hack., Noticastrum diffusum (Pers.) Cabrera, Dichanthelium sabulorum (Lam.) Gould & C.A. Clark, Eragrostis bahiensis Roem. & Schult., Eragrostis trichocolea Arechav., Gamochaeta stachydifolium (Lam.) Cabrera, Helianthemum brasiliense (Lam.) Pers., H. bonariensis Lam., P. racemosum (P. Beauv.) Spreng., S. crassiflorus (Poir.) DC., and Sisyrinchium vaginatum Spreng. (Table 2). The Fig. 4 Measure of species richness (lower continuous curve), performance of diversity estimator Chao 2 (continuous curve with dots) and its standard deviation (dotted curves) for the 22 samples of vegetation stands. Data from Estimates using 1,000 replications Author's personal copy Vegetation of a coastal dune system Table 2 Legrand0 s (1959) list of species for the study area Species Family Zone Abundance category Iresine portulacoides (A. St.-Hil.) Moq. Amaranthaceae I Eryngium pandanifolium Cham. & Schltdl. Apiaceae III nc Hydrocotyle bonariensis Lam. Apiaceae I, II, III, IV Abundant Acanthospermum australe (Loefl.) Kuntze Asteraceae IV Achyrocline satureioides (Lam.) DC. Asteraceae IV Baccharis genistifolia DC. Asteraceae IV Baccharis gnaphalioides Spreng. Asteraceae IV Baccharis microcephala Baker Asteraceae Baccharis rufescens Spreng. Asteraceae ? ? Sparse Frequent Abundant ? Frequent III, IV ? nc IV ? Frequent Frequent Baccharis spicata (Lam.) Baill. Asteraceae III Berroa gnaphalioides (Less.) Beauverd Conyza blakei (Cabrera) Cabrera Asteraceae Asteraceae IV II ? ? Frequent nc nc Conyza sp. Less. Asteraceae III, IV ? nc Gamochaeta calviceps (Fernald) Cabrera Asteraceae II ? nc Gamochaeta falcata (Lam.) Cabrera Asteraceae II ? nc Gamochaeta stachydifolium (Lam.) Cabrera Asteraceae I, II, IV ? Abundant Pluchea sagittalis (Lam.) Cabrera Asteraceae III nc Senecio crassiflorus (Poir.) DC. Asteraceae II, IV Very frequent Cakile maritima Scop. Brassicaceae I *? Sparse Calycera crassifolia (Miers) Hicken Calyceraceae I, II ? Abundant Pratia hederacea (Cham.) G. Don Campanulaceae III ? Abundant Helianthemum brasiliense (Lam.) Pers. Cistaceae IV ? Abundant Noticastrum diffusum (Pers.) Cabrera . Compositae IV ? Abundant ? Rare Calystegia soldanella (L.) Roem. & Schult. Convolvulaceae I Androtrichum trigynum (Spreng.) H. Pfeiff. Cyperaceae III Abundant Cyperus prolixus Kunth Cyperaceae III ? Cyperus reflexus Vahl Cyperaceae III ? Abundant nc Cyperus rigens J. Presl & C. Presl Cyperaceae III, IV ? nc Eleocharis sp. R. Br. Cyperaceae III Kyllinga vaginata Lam. Cyperaceae III ? nc nc Rhynchospora brownii Roem. & Schult. Cyperaceae III ? nc Rhynchospora microcarpa Baldwin ex A. Gray Cyperaceae III ? nc Schoenoplectus californicus (C.A. Mey.) Soják Cyperaceae III Scleria distans Poir. Cyperaceae III ? Abundant Drosera brevifolia Pursh Droseraceae III ? nc nc Eriocaulon modestum Kunth Eriocaulaceae IV ? nc Leiothrix arechavaletae (Körn.) Ruhland Eriocaulaceae III ? Abundant Syngonanthus gracilis (Bong.) Ruhland Eriocaulaceae III ? Sparse Stylosanthes leiocarpa Vogel Fabaceae IV ? Frequent Laurembergia tetrandra (Schott ex Spreng.) Kanitz Haloragaceae III ? nc Sisyrinchium vaginatum Spreng. Iridaceae IV ? Abundant Juncus acutus L. Juncus microcephalus Kunth Juncaccea Juncaceae III III ? nc Abundant ? nc Juncus scirpoides Lam. Juncaceae III Juncus sp. L. Juncaccea III Utricularia gibba L. Lentibulariaceae III ? nc Utricularia tridentata Sylvén Lentibulariaceae III ? nc Lycopodiella alopecuroides (L.) Cranfill Lycopodiaceae III ? Abundant nc 123 Author's personal copy E. Castiñeira Latorre et al. Table 2 continued Species Family Zone Abundance category Lycopodiella caroliniana (L.) Pic. Serm. Lycopodiaceae III ? Scarce Oenothera mollissima L. Onagraceae II, IV ? Frequent Ludwigia hookeri (Micheli) H. Hara Onagraceae III nc Plantago brasiliensis Sims Plantaginaceae IV ? Frequent Andropogon selloanus (Hack.) Hack. Poaceae IV ? Abundant ? Frequent Chascolytrum erectum (Lam.) Desv. Poaceae IV Cortaderia selloana (Schult. & Schult. f.) Asch. & Graebn. Poaceae III Dichanthelium sabulorum (Lam.) Gould & C.A. Clark Poaceae IV ? Abundant Eragrostis bahiensis Schrad. ex Schult. Poaceae IV ? Abundant Eragrostis trichocolea Arechav. Poaceae IV ? Abundant Ischaemum minus J. Presl Poaceae III ? Panicum racemosum (P. Beauv.) Spreng. Poaceae I, II, IV Abundant Paspalum pumilum Nees Paspalum vaginatum Sw. Poaceae Poaceae III I ? nc Rare Piptochaetium panicoides (Lam.) E. Desv. Poaceae IV ? Frequent Poa lanuginosa Poir. Poaceae II, IV ? Frequent Poidium uniolae (Nees) Matthei Poaceae III ? nc Polypogon chilensis (Kunth) Pilg. Poaceae III ? nc Schizachyrium condensatum (Kunth) Nees Poaceae III ? nc ? Frequent Spartina ciliata Brongn. Poaceae I, II Polygala cyparissias A. St.-Hil. & Moq. Polygalaceae II, IV nc nc Scarce Polygala tenuis DC. Polygalaceae III ? nc Rumex cuneifolius Campd. Polygalaceae II ? nc Anagallis filiformis Cham. & Schltdl. Primulaceae III ? nc Margyricarpus pinnatus (Lam.) Kuntze Rosaceae II, IV Mitracarpus megapotamicus (Spreng.) Kuntze Rubiaceae IV ? nc Richardia brasiliensis Gomes Rubiaceae IV ? Dodonaea viscosa Jacq. Sapindaceae IV nc Bacopa monnieri (L.) Wettst. Scrophulariaceae III nc Scoparia montevidensis (Spreng.) R.E. Fr. Scrophulariaceae IV Typha domingensis Pers. Typhaceae III Xyris jupicai Rich. Xyridaceae III ? nc nc Frequent Abundant ? Abundant 0 Zone (I, II, III and IV) and abundance category correspond to Legrand s observations. The scientific names follow Instituto de Botánica Darwinion (2011), Royal Botanic Gardens (2011) and (W3TROPICOS 2009) nc not categorized by Legrand, sparse, frequent, very frequent, and abundant, ? exclusive species of Legrand0 s survey, * non-native species lowest similarity is reported for the comparison of the fixed dunes (present data) versus zone IV of Legrand (J = 0.08). Rarefaction curves showed that group 1 (psammophilic steppe stands and psammophilic pasture stand in foredunes) had the lowest observed and expected richness (Fig. 6). Interdune depressions had a wide range of richness values (Fig. 6). Group 4 (hydrophilic pasture stands) had the lowest richness, followed by group 3 (hydrophilic pasture stand and Typha stand of T. latifolia) (Fig. 6). Group 5 (Eryngium stand) had a higher richness than group 3 and, although it has a lower observed richness than group 2 (Schoenoplectus stands and Thelypteris stand), they did not differ significantly in their expected richness (Fig. 6). 123 Group 6 (Typha stands of T. dominguensis) showed the highest observed richness of all groups of the interdune depressions and had similar values with group 7 (Fig. 6). None of the species registered at the study area, in ours and Legrand’s survey, has been evaluated and included at the ‘‘IUCN Red List of Threatened Species’’ (IUCN 2009), at local or global level. However, local evaluations using IUCN criteria (see Marchesi et al. 2008; Alonso Paz et al. 2009) considered Eriocaulon modestum Kunth, Anagallis filiformis Cham. & Schltdl. and Syngonanthus gracilis (Bong.) Ruhland as species with restricted distributions and in a retraction process by the action of human impact, and Laurembergia tetrandra (Schott) Kanitz a species in a Author's personal copy Vegetation of a coastal dune system Fig. 5 Cluster analysis of the plant associations based on Jaccard0 s distance and Ward0 s method retraction process with a socioeconomic value. All these species were registered in Legrand0 s survey. We report here Eleocharis sellowiana Kunth, which is a species in a retraction process due to human impacts, and Paspalum notatum Flüggé, a species that has a restricted distribution but high socioeconomic value (Alonso Paz et al. 2009). We also reported the following non-native species: Carpobrotus edulis (L.) N.E. Br., Centella asiatica (L.) Urb., Cirsium sp. Mill, Hypochoeris radicata L., Sonchus asper (L.) Hill, A. longifolia, Pinus maritima Mill., Cynodon dactylon (L.) Pers. (Table 1). Legrand (1959) registered Cakile maritima Scop. as the only non-native species (Table 2). Discussion The global tendency of the introduction of Australian acacias to fix dune systems at the beginning of the 20th century, with the goal of expanding and improving agroproductive and urban areas, was also imposed in Uruguayan coastal ecosystems (Carruthers et al. 2011; Kull et al. 2011). The study area at the shore of the ‘‘de la Plata’’ River in the seaside resort ‘‘El Pinar,’’ Canelones, Uruguay, corresponds to a relic dune system of a larger ecosystem that was modified with the introduction of A. longifolia and Pinus spp. for urban development (Legrand 1959; Gutiérrez and Panario 2005). This ecosystem management has produced large landscape modifications, affecting the soil, fire regimes, water availability, and the composition, structure, and dynamics of native vegetation in our study area, as has been reported in other studied dunes systems (Marchante et al. 2010, 2011b; Morris et al. 2011). Representativeness of families of plant species at the study area agrees with the research of Delfino and Masciadri (2005) for the oceanic Uruguayan seashore, where Asteraceae and Poaceae were the prevailing families. Fontana (2005), at ‘‘Bahı́a Blanca,’’ Buenos Aires, Argentina, found these two families to be the most abundant, but with Poaceae more common than Asteraceae. When comparing the description of plant species composition of each environment (i.e., foredunes, interdune depressions, and fixed dunes) written by Legrand (1959) with our results, we found similar indicator species. Legrand recognized this species as indicative based on their experience and by not using a statistical index like the indicator value (Dufrene and Legendre 1997). In spite of this similarity in the indicator species, one major result from the historical perspective is the low similarity of species composition between the present and past communities. Recognizing the causes behind this pattern is a major concern for ecological science and in particular for restoration ecology (Suding 2011). In this sense, the existence of local evaluations of conservation status of species (see Marchesi et al. 2008; Alonso Paz et al. 2009), allowed us to verify that some species registered by Legrand had conservation problems and disappeared from the study area. Nevertheless, the magnitude of this change is due to the loss of these 123 Author's personal copy E. Castiñeira Latorre et al. Fig. 6 Rarefaction curves showing mean expected richness and 95 % confidence intervals for each of the seven groups of the cluster analysis threatened species. We observed a great change in the floristic composition and vegetation structure in less than 100 years in this system of coastal dunes, accompanied by the presence of non-native and in some cases invasive species (C. edulis, C. asiatica, Cirsium sp., H. radicata, S. asper, A. longifolia, P. maritima, C. dactylon). The agglomerative cluster analysis allowed us to identify seven plant groups which were associated with three coastal environments: foredunes, interdune depressions, and fixed dunes. The foredunes harbor communities of pioneer psammophilic plants (Hesp 2002). In this extreme environment, the action of the wind triggers mechanical and physiological effects in vegetation (e.g., abrasive effect, increase of transpiration rate, homeostatic changes) leading to a reduced number of species becoming established (Herrmann et al. 2008; Judd et al. 2008). As it was reported by Cordazzo and Seeliger (1993) for the Atlantic coast of ‘‘Rı́o Grande do Sul,’’ Brazil and by Fontana (2005) for ‘‘Bahı́a Blanca,’’ Buenos Aires, Argentina, the least diverse plant association was located in the foredunes, where P. racemosum, S. crassiflorus, and S. ciliata were indicators of this psammophilic extreme environment. These species have special importance for conservation of dune systems because they contribute to the regeneration of foredunes after storms (Cordazzo and Davy 1999; Hesp 2002). From a conservation perspective, we witnessed an extreme retraction of a shore environment, because Legrand (1959) described that this plant association extended for 200 m from the beginning of the foredunes (i.e., zone I and II) and currently it only reaches about 20 m. Behind foredunes we found interdune depressions. Here, the outcrop of the water table generates a moisture increase leaving the ground permanently or semi-permanently 123 flooded. At interdune depressions wind and abrasive effects of transportation of sand decrease, hydrophilic plant associations become established and the specific diversity as well as the number of plant groups increases (Cordazzo and Seeliger 1993; Fontana 2005). The indicative species are the highest emergent herbs adapted to flooded environments: E. pandanifolium, S. californicus, Juncus sp. and T. dominguensis (Grace and Wetzel 1981; Grace 1989; Newman et al. 1996; Costa et al. 2003; Calviño and Martı́nez 2007). Mikania sp. was abundant in humid environments climbing on stems of T. latifolia (Lombardo 1983). L. arechavaletae is a perennial grass considered endemic for Uruguay with records from the Departments of Montevideo and Rivera. This species has been observed and categorized as ‘‘abundant’’ by Legrand (1959); however it has not been reported at the present study. One of the most diverse environments was the fixed dunes where the indicator species were common in coastal dunes: A. satureioides, A. trigynum and an alien species A. longifolia (Legrand 1959; Lombardo 1983, 1984). This environment presents the lowest similarity between present and past. We associate this striking change with the presence of A. longifolia, as has been noted in several other places (e.g., Marchante et al. 2008, 2009, 2010, 2011b; Morris et al. 2011). This species presents ecophysiological traits associated to the resources acquisition that confers competitive advantages over native species (e.g., initial high relative growth rates, N2 fixing symbioses and the development of ectomycorhitic symbiosis), and generates modifications in the composition of nitrogen and carbon, nutrients cycling, and in microbial soil processes (Smith and Read 2008; Morris et al. 2011). A. longifolia is considered a species with an extremely high invasive potential Author's personal copy Vegetation of a coastal dune system and an ecosystem transformer species (sensu Richardson et al. 2000; Marchante et al. 2008, see also Table 1 in Wilson et al. 2011). When it appears it modifies the community composition, species richness, and soil ecology (Marchante et al. 2008, 2009; Marchante 2011; Echeverrı́a et al. 2009; Wilson et al. 2011). In terms of the environmental change, the Acacia stands can be interpreted as the psammophilic grassland stands invaded and modified by A. longifolia which gives a colonization opportunity to other non-native species (C. asiatica (L.) Urb., C. dactylon (L.) Pers.) (Marchante et al. 2011b). In this context of extreme change of coastal plant communities, we consider the necessity of implementation of management plans for the restoration at least for the fixed dunes of the costal ecosystems of Canelones, Uruguay. Now we have a theoretical and empirical framework generated by the experience of almost 200 years of introduction of Australian acacias in many ecosystems around the world (see special issue: ‘‘human-mediated introductions of Australian acacias—a global experiment in biogeography’’, Diversity and Distribution 2011). This knowledge has led to the proposal of a set of management strategies for A. longifolia (van Wilgen et al. 2011; Wilson et al. 2011). Given the scarcity of local knowledge about the naturalization, invasion, and effects on native communities of A. longifolia in Uruguay, we encourage the development of an investigation program to assess the effectiveness and challenges of the potential management practices. To achieve the greatest benefit with the management strategy, we suggest prioritizing the fixed dunes as the spatial areas of action. The population of A. longifolia in this environment has a low socioeconomic value [in this area it has not been considered a timber resource like in other places such as South Africa (see van Wilgen et al. 2011; Wilson et al. 2011) and it has invaded a low area. Although the species is invasive in all dune systems of the Uruguayan coast, we consider that the area invaded the study area is low due it being confined to a narrow strip of relict dunes. In this scenario, van Wilgen et al. (2011) suggest applying the tactic of eradication including a combination of different practices of management (see Fig. 3 in van Wilgen et al. 2011): manual control operations, biologic control agents [e.g., bud-galling wasp Trichilogaster acaciaelongifoliae (Hymenoptera: Pteromalidae) and seed-feeding weevil Melanterius ventralis (Coleoptera: Curculionidae)] and the use of fire. These management practices will have effects on the seed bank, buds, juvenile, and adult plants (Le Maitre et al. 2011, Marchante et al. 2010, 2011a, b, c, Wilson et al. 2011). Despite the importance of the conservation of coastal environments, this area has been impacted for a century by human activities, primarily recreation and urbanization. In Uruguay, as in many other coastal areas around the world, plantations, dune fixation, and high human pressure have produced important modifications in the health of coastal dunes. The historical knowledge compared with the present status of the coastal ecosystems has allowed us to understand and disentangle causes and processes behind the ecological change. In consequence, we encourage progress in the design of a management program to control the A. longifolia invasion and the restoration of Uruguayan coastal dune systems. Acknowledgments In the memory of my mother Elena Latorre. We thank A. Camargo, J.M. Piñeiro and J.A. Simonetti for comments on data interpretation and for their constructive comments on the manuscript. We thank Meredith Root-Bernstein, A. Camargo and G. Franco Castiñeira for contributions on English. AC is grateful for the support of FONDECYT-FONDAP Grant 1501-0001 and received a fellowship from the ‘‘Vicerrectorı́a Adjunta de Investigación y Doctorado-PUC, Chile’’. We are also very grateful to two anonymous reviewers and editors of the Brazilian Journal of Botany for their comments on an earlier version of this manuscript. 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Composition and vegetation structure in a system of coastal dunes of the “de la Plata” river, Uruguay: a comparison with Legrand’s descriptions (1959)

Composition and vegetation structure in a system of coastal dunes of the “de la Plata” river, Uruguay: a comparison with Legrand’s descriptions (1959)

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