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Macrophytes in the upper Parana river floodplain: checklist and comparison with other large South American wetlands.

Wetlands are important sites for biological conservation because they support rich biodiversity and present high productivity (Mitsch & Gosselink 2000). The study of wetland plants has been of interest to botanists for many years, but the effort to identify and understand these plants has increased dramatically since the 1970s, when ecologists began to emphasize the vital role that wetlands play in our landscapes (Cronk & Fennessy 2001).

One of the main ecological characteristics of South America is the existence of large wetlands (Neiff 2001). Inventories of wetlands provide an indication of the sites with the highest biological diversity and productivity (Taylor et al. 1995), and the information collected through inventories is a necessary prerequisite for conservation policies (Pressey & Adam 1995).

Approximately 50% of the inventoried wetlands in South America are located in Brazil (Naranjo 1995). However, specific information related to aquatic macrophytes is extremely scarce. Diegues (1994) performed the first inventory of wetlands in Brazil, and his work provided valuable data for evaluating ecological and economic aspects of these regions. According to Neiff (1978, 1986), aquatic macrophytes are important in shallow ecosystems, such as river-floodplain ecosystems, where they colonize extensive areas and exhibit high rates of primary productivity. In addition, macrophytes are a key component of river-floodplain ecosystems because they enhance nutrient cycling, increase habitat heterogeneity and provide food for a variety of organisms (Esteves 1998).

Floodplains are known as ecosystems with a high diversity of habitats and aquatic and terrestrial species (Junk et al. 2000). Due to their high complexity and seasonal changes in physico-chemistry, these ecosystems are characterized by a variety of assemblages, which differ in richness and composition according to the water level. In the Upper Parana River floodplain, for example, the vegetation is highly conditioned by geomorphology (Souza-Filho 1993); trees dominate the more elevated areas (levees), and shrubs colonize less elevated areas that remain flood-free most of the year, while aquatic macrophytes grow in permanently inundated areas of the wetlands.

Despite the importance of these macrophytes in the Upper Parana River floodplain, a stretch of this river that is key in maintaining the biodiversity of Brazilian inland waters, information about the aquatic vegetation in this region is scattered among several different papers and reports (Bini 1996, Kita & Souza 2003, Thomaz et al. 2004, Thomaz et al. 2009); most of these studies emphasized that the flood pulse and changes in water physico-chemistry are important factors controlling macrophyte populations and communities.

In the present study, we first addressed the number of macrophyte species in the main habitats of the Upper Parana River and its floodplain (herein only Parana floodplain), using records gathered since 1997 and intensive collections performed between 2007 and 2009. Secondly, we used this dataset to compare the species richness and similarity of this area with other South American wetlands. Finally, using species accumulation curves, we examined whether the number of species described in South America is reaching an asymptote, or if more sampling efforts are still necessary to accomplish a comprehensive inventory of the rich aquatic flora of this area.

To accomplish these objectives, we adopted the conceptualization of aquatic macrophytes proposed by Cook (1996), in which the author includes plants which photosynthetically active organs are either permanently, or for several months of the year, total or partially submersed in freshwater or floating in aquatic habitats. More recently, Chambers et al. (2008) also included Charophytes within the definition of macrophytes. To avoid any confusion, we did not use in any part of our text the term "vascular plants" but, instead, consistently used the term "aquatic macrophytes".

MATERIALS AND METHODS

Study area: The floodplain of the Parana River is located downstream from the Porto Primavera Reservoir. This stretch has a length of 160km and is the last region of the river that remains not dammed in Brazilian territory. Thus, it is of key importance to the conservation of the aquatic biodiversity of the Parana Basin (Agostinho & Zalewiski 1995).

According to the Koppen system, the climate in this region is classified as tropical and sub-tropical, with warm summers (mean annual temperature 22[degrees]C) and a mean annual rainfall of 1 500mm (Maack 2002).

The compiled list of taxa was based on records of samplings conducted in the floodplain since 1997. In addition, we utilized several other studies (published and unpublished) that had been conducted in the floodplain. Macrophytes were collected in a variety of habitats, such as the river main channel, lateral channels (anabranches), temporary and permanent lakes, and in the aquatic-terrestrial transition zone (ATTZ, sensu Junk et al. 1989). We also analyzed and revised specimens deposited in the Laboratory of Macrophytes and in the Herbarium (HUEM) of the University of Maringa. To complement the list of species recorded in previous investigations, we carried out additional samplings between 2007 and 2009 in six habitats that are being monitored in the Brazilian Long Term Ecological Research Program (site 6; Thomaz et al. 2009).

[FIGURE 1 OMITTED]

In each lake, the aquatic macrophytes were analyzed by boat at a slow speed along the entire shoreline. In the ATTZ, samplings were carried out on foot. We used a grapple attached to a line to record submersed species. Because ponds and lakes have small areas (from 0.006 to 113.8ha) and samplings were carried out on the entire shores, we considered the recorded species as the actual richness of these habitats, and did not correct the results to account for sampling effort (rarefaction curves, for example).

Identification followed comparative morphology and a specialized bibliography (e.g., Hoehne 1948, Cook 1996, Pott & Pott 2000, Amaral et al. 2008, Bove & Paz 2009). The list of taxa contains families and genera according to the "Angiosperm Phylogeny Group-APG II" (2003) for Magnoliophyta (Angiospermae), Willis (1973) for Pteridophyta, and Crandall-Stotler (1980) for Hepatophyta.

Plant life forms were chosen according to Pedralli (1990), and we followed Tur (1972) for epiphytic forms. Plants growing in wet soils (marshes locally known as "varjao" or "varzea") were included in the category of amphibious (Irgang & Gastal 1996).

In order to make comparisons among South American wetlands, we used the following lists of macrophytes obtained in long-term surveys (Table 1).

All of the investigations and the lists of species that we used were carried out by specialists and included several types of habitats (Fig. 1). Although there are several other papers describing single habitats, we did not use these studies. It is difficult to guarantee that all studies follow the same methodology, but we believe that they are similar enough to at least contribute a first tentative of comparison of Neotropical wetlands to make inferences about the richness of macrophytes in this region.

To find similarity among these surveys, we first converted all data into a large matrix containing species occurrence presence/absence. A matrix of similarity was built using the Bray-Curtis distance coefficient (Krebs 1999). To compare all surveys, we used the method of complete linkage (Sneath & Sokal 1973). A dendrogram of similarities was built using the PRIMER v. 6 software, Plymouth Routines in Multivariate Ecological Research (Clarke & Gorley 2006).

Using the entire dataset, which included all wetlands, we assessed whether the richness of macrophytes in South America reaches an asymptote, or if there are many species yet to be found. The expected species accumulation curve was calculated according to a Mao Tau function. Using an accumulative curve with "studies" as units of sampling effort, an asymptote would indicate whether almost all species have already been recorded; however, the lack of an asymptote would indicate that the number of aquatic macrophyte species found until now is still far from the real total of these species. We also estimated the richness of macrophytes using a first-order Jacknife estimator (Jack1) with the objective to assess the extent to which the number of macrophyte species in South America remains underestimated. Accumulation and estimation curves were constructed using the EstimateS program (Colwell 2009).

RESULTS

A total of 153 species of macrophytes was recorded in the Upper Parana River floodplain. These species were distributed in 100 genera and 37 families (Appendix 1), representing a variety of taxonomic groups (Charophyta, Bryophyta, Pteridophyta, Basal Angiospermae and Angiospermae).

Sixteen of the recorded species are cryptogams and are classified as follows: two charophytes, two hepatophytes and 12 pteridophytes. Of the Angiospermae, Poales exhibited the highest number of taxa (40), followed by Alismatales (17), Myrtales and Lamiales (12 species each). The families with the highest numbers of species were Poaceae (21), Cyperaceae (17), Pontederiaceae (8), Hydrocharitaceae (7), Polygonaceae and Onagraceae (6) and Fabaceae (5). Araceae, Alismataceae, Commelinaceae, Amaranthaceae and Plantaginaceae were represented by four species each and the other families by three or fewer species.

All life forms were found in the area, and emergent and amphibious types were the most representative macrophytes, contributing 45% and 26% of the species, respectively (Fig. 2). Rooted submersed (11%) and free-floating species (9%) were also important, while the lowest numbers of species were found for rooted floating and free-submersed types (2%) and epiphytes (1%).

The number of species recorded in the Parana River and its floodplain consistently increased over time (Fig. 3). The greatest increase occurred between 2007 and 2009, due to intensified sampling efforts and refined taxonomic identification, which led to the addition of 105 species to the recorded flora of the Upper Parana River floodplain.

[FIGURE 2 OMITTED]

[FIGURE 3 OMITTED]

The Parana floodplain exhibits the third highest richness of macrophytes (153 species) of the 12 areas for which we have data in South America. The coastal area of the State of Rio Grande do Sul (Brazil) ranked first (321 species), and the Pantanal Matogrossense, one of the largest wetlands in the world, ranked second (247 species).

A dendrogram built using the Bray-Curtis coefficient of distance showed that South American wetlands are dissimilar with respect to macrophyte assemblages (Fig. 4). We can roughly recognize three groups of wetlands. The first includes the seasonal ponds (Northeast Brazil), coastal lagoons of the State of Rio de Janeiro, Pantanal, Parana floodplain in Brazil, the coastal plain in the State of Rio Grande do Sul and the Parana floodplain in Argentina, with a similarity of 11.7% (Fig. 4). These areas share only two species in common, Polygonum ferrugineum and Nymphoides indica, both of which are widespread hydrophytes. In this first group, the most similar areas were the Parana floodplain and the Pantanal Matogrossense with 40.2% similarity and 79 species in common (9% of the total species)

The second group included the Amazon basin of Ecuador, Amazon River floodplain and Amapa wetlands, with a similarity of 13% and sharing four species, Eichhornia azurea, Hymenachne amplexicaulis, Salvinia auriculata and Utricularia foliosa. Within this group, the aquatic flora of the Amazon River floodplain and the Amazon Basin of Ecuador share 13 species (2% of the total species) and exhibit 35.1% similarity. Finally, group three was formed by Peru, Ecuador and Bodoquena (Mato Grosso do Sul), with a similarity of 18% and sharing 11 species; within this group, Peru and Ecuador had the highest similarity of 33.6% and sharing 35 species (4% of all species).

Considering all of the surveys that we found for South America together with the survey we conducted in the Parana floodplain, a total of 854 species of macrophytes was compiled. However, the species accumulation curve produced using these studies as a surrogate of sampling effort, did not reach an asymptote (Fig. 5). In fact, the number of species estimated through Jack1 was 1 388, indicating an approximate underestimation of 534 species.

[FIGURE 4 OMITTED]

[FIGURE 5 OMITTED]

DISCUSSION

According to Chambers et al. (2008), the Neotropical region has the highest number of macrophyte species in the world (984 species). The number of species of macrophytes recorded in the Upper Parana River and its floodplain (153 species; 16% of the Neotropical region) can be considered high due to the small relative area of this ecosystem (2 500km2) compared to other aquatic areas from this region, such as the Amazon and the Pantanal. Even considering the higher species richness found in the Pantanal Matogrossensse (247 species), our survey still indicates that the Parana floodplain is highly diverse because the Pantanal is 55 times larger in area, extending over approximately 138 183[km.sup.2]. Thus, the conservation units contained inside this stretch of the Parana River can be considered important for the conservation of aquatic macrophyte diversity. Again, we emphasize that the number of species we recorded does not represent the real species richness because there are a number of habitats not yet investigated in this stretch.

The comparison of the region investigated in this study with other dammed stretches suggests the importance of the Parana floodplain as a hotspot of macrophyte species diversity in this basin. In a survey of 18 reservoirs of the Parana River and some of its main tributaries, Martins et al. (2008) found only 39 species of macrophytes. Even in the Itaipu Reservoir, which is dendritic (and thus, favorable for macrophyte colonization), a long-term dataset showed a total of 110 species (Mormul et al. 2010). The same conclusion can be made when we compare our data with reservoirs from other basins; for example, only 23 species of macrophytes were recorded in the Guri Reservoir, Venezuela (Vilarrubia & Cova 1993). The great variety of habitats found in the Parana floodplain, together with the natural disturbance caused by seasonal oscillations in the water level, might explain these differences in relation to different reservoirs. On the other hand, the sampling effort was not controlled in these different surveys, and thus the results should be viewed with caution. However, this pitfall might be minimized because all of the investigations that we included in this report were floristic surveys, which tend to maximize the sampling within a region. In addition, there may be differences in the definition of macrophytes used in different surveys. Considering species composition, the assemblages of the Parana floodplain can be considered as a sample of the aquatic flora from the Pantanal, as was previously pointed out by Thomaz et al. (2009). In fact, the most representative families in number of species are largely the same for both ecosystems (Poaceae and Cyperaceae, Onagraceae, Pontederiaceae, Plantaginaceae and Fabaceae). The families with few species in both ecosystems are also the same (Typhaceae, Cucurbitaceae, Maranthaceae, Haloragaceae, Solanaceae and Orchidaceae). Our cluster analysis also indicated that these three wetlands are the most similar amongst all ecosystems in our dataset, which could be due to their geographical proximity and to hydrological similarities (all areas are subjected to seasonal variation in the water level and also include a great variety of habitats). Furthermore, both wetlands belong to the same larger Parana basin.

However, some families differ considerably in the number of species between these two wetlands. For example, there are only two species of Nymphaeaceae in the Parana floodplain, whereas there are eight in the Pantanal that occur mainly in rain-fed shallow ponds and seasonal standing or slow flowing water in addition to in the river floodplain, except for Victoria amazonica, which grows in oxbow lakes. Similarly, 10 species of Characeae were found in the Pantanal, which is attributed to the alkaline and brackish waters in the Southwestern Pantanal (Bueno 1993, Pott & Pott 1997), but only two were identified in the Parana floodplain, where acid soils predominate and charophytes do not thrive. The importance of the type of habitat in determining species composition can also be observed if we compare a survey carried out on lakes, reservoirs and wetlands in the Southern Parana State (Cervi et al. 2009), only 200km away, that shares only 23% of macrophytes with the Parana floodplain.

The low richness of aquatic epiphytes reported in the Parana floodplain is related to the small sampling effort that has been carried out on floating meadows. Epiphytes usually colonize advanced stages of aquatic succession (Pott & Pott 2003). For example, surveys carried out by Tur (1972) and Neiff (1982) in the Middle Parana (Argentina) identified 70 species of epiphytes. Even though both regions are on the Parana River, the flood pulses differ between the Middle and the Upper Parana basins, which may influence the accumulation of organic matter and, thus, the formation of floating-substrates, as well as the displacement of these islands, and this may explain the differences in the richness of epiphytes found in these wetlands.

The species number increase over time reported for the Parana floodplain can be mainly attributed to the refinement of the taxonomic searches and identification carried out. In addition to this effect, we also considered a higher sampling effort in the Parana floodplain, with collections made in habitats not previously investigated (e.g., rocks in the Parana channel) and the arrival of new species (e.g. Hydrilla verticillata). The high level of species richness that we found indicates that the Parana floodplain is still in a good conservation state, despite the strong anthropogenic pressures in the region related to changes in hydrometric levels, nutrient cycling and suspended solid loadings (Souza-Filho 2009).

However, despite the good status of conservation with respect to the aquatic flora, we contend that there is a concern related to the presence of two invasive species, H. verticillata and Urochloa subquadripara. The first is a submersed species native to Asia and the North of Africa, that colonizes the Parana main channel and has a high competitive ability, threatening native species due its rapid regeneration following hydrological disturbances (Sousa et al. 2009, Thomaz et al. 2009). Its success in the Parana main channel is associated with the same effect leading to an increase in the colonization by submersed species, i.e., the increase in water transparency and propagule pressures originating in the upstream reservoirs (Thomaz et al. 2009). Hydrilla verticillata has not yet colonized lakes in either the Baia or Ivinhema river habitats (Sousa et al. 2009). The second species, U. subquadripara, belongs to the family Poaceae, which contributes with several invasive species (Petenon & Pivello 2008). Although U. subquadripara has been rarely recorded in the Parana floodplain, it reduced significantly the diversity of macrophytes in a lake close to the Baia River, the only place where it occurs with high biomass in this floodplain (Michelan et al. 2010). Disturbances associated with the oscillation in water levels may explain why this species is so rare in most habitats in the floodplain, but in light of its severe threat to macrophyte diversity, its monitoring is a priority, especially in the best preserved areas of this region.

The results of our cluster analysis indicate that Neotropical wetlands are different regarding macrophyte composition. Thus, we infer that such differences may be due to multiple factors, such as climate, flood regime and geography.

In fact, the most similar areas (Pantanal and Parana floodplain) share many similarities: they are both large floodplains located from 80-160 m.a.s.l., have a great variety of habitats and are subjected to seasonal water level fluctuations (Agostinho & Zalewiski 1995, Vila da Silva 1995). However, the cluster analysis also shows that South American wetlands are diverse regarding macrophyte assemblages, and even ecosystems located in the same basin may differ considerably (e.g., the upper and middle/lower Parana floodplains). The differences observed for these two floodplains may be accounted for by differences in their nutrient regimes (the Argentinean floodplains receive high phosphorus inputs from the Andes tributaries) and also to the types of habitats investigated (e.g., the widespread occurrence of floating meadows in Argentina with a high richness of epiphytes). As previously mentioned, the groups formed by the cluster analysis suggest that, though they represent geographically distant environments, such as the Amazonian floodplains and the Argentina plain, the sampled landscapes are determinant in forming groupings.

The accumulation curve reflects the differences found in the cluster analysis. In other words, the great differences among the South American surveys included in the cluster analysis indicate a high beta-diversity, leading to a lack of an asymptote in the accumulation curve. The total number of species found in all 12 surveys represents 87% of the number found by Chambers et al. (2008) for the Neotropical region. Despite the fact that our findings are close to the total number of Neotropical macrophytes, the lack of an asymptote, together with the high underestimation of true richness suggested by the Jack1 estimator, indicates that we are still far from describing the actual richness for this region. The number of plants to be described in Brazil, what may reflect the situation of South America, is considered very high (Pimm et al. 2010). In fact, there has been a clear lack of investigations conducted in pristine habitats in South America, such as in parts of the Amazon and the Andes, which are areas of high biodiversity. Future investigations at these sites, together with the description of new species (e.g., Bove et al. 2006, Amaral & Bittrich 2008), will certainly increase the number of species of Neotropical macrophytes recorded and give a better idea of the biodiversity provided by the great variety of ecosystems of this biogeographical region.

Our results reinforce the hypothesis of Irgang & Gastal (1996) that Uruguay, North Argentina, Paraguay and South Brazil form a phytogeographic unit, and therefore, the sampled number of species does not closely correlate with other evaluated areas. There are many other large wetlands in South America that should be included in this analysis but that were not included because of insufficient floristic inventories, such as Guapore and Ilha do Bananal.

In summary, this report highlights the flora of different wetlands of South America and indicates that the actual species richness of macrophytes of this continent is far from being well understood. Our hypothesis sustains that macrophyte records, together with existing surveys, indicate a continuous need for carrying out increasing numbers of collections in new areas in the upper Parana river-floodplain system and in other South American wetlands, as the number of species so far reported remains far from the predicted total. The checklist generated in this study is intended to support other research in wetlands and, in particular, to assure the continuity of ongoing long-term ecological programs, and it reveals a rich flora that is practically unknown to botanists and ecologists.
APPENDIX I

List of taxa recorded in the Parana River floodplain between
the years 1996 and 2009

                       TAXA                          L.F.

  Characeae -- Charophyta
    Chara guairensis R. Bicudo                       Rs
      Nitella furcata (Roxb. ex Bruz.)               Rs
        Ag. emend. R.D. Wood
  Ricciaceae -- Hepatophyta (Bryophyta)
    Riccia sp.                                       Em
    Ricciocarpus natans L. (Corda)                   Ff
  Azollaceae -- Pteridophyta
    Azolla filiculoides Lam.                         Ff
    A. microphylla Kaulf.                            Ff
  Blechnaceae -- Pteridophyta
    Blechnum brasiliense Desv.                       Em
    B. serrulatum Rich.                              Em
  Pteridaceae -- Pteridophyta
    Ceratopteris pteridoides (Hook.) Hieron.         Ff
    Pityrogramma calomelanos (L.)                    Em
      Link var. calomelanus
    P. trifoliata (L.) R. Tryon                      Em
  Salviniaceae -- Pteridophyta
    Salvinia auriculata Aubl.                        Ff
    S. biloba Raddi emend de la Sota                 Ff
    S. minima Baker                                  Ff
  Thelypteridaceae -- Pteridophyta
    Thelypteris interrupta (Willd.) K. Iwats.        Em
    T. serrata (Cav.) Alston                         Em
    Nymphaeaceae -- Basal Angiospermae
    Cabomba furcata Schult. & Schult. f.             Rs
    Nymphaea amazonum Mart.                          Rf
      ex Zucc. subsp. amazonum
Alismatales -- Monocots
  Araceae
    Lemna valdiviana Phil.                           Ff
    Pistia stratiotes L.                             Ff
    Wolffiella lingulata (Hegelm.) Hegelm.           Ff
    W. oblonga (Phil.) Hegelm.                       Ff
  Hydrocharitaceae
    Egeria densa Planch.                             Rs
    E. najas Planch.                                 Rs
    Hydrilla verticillata (L.f.) Royle               Rs
    Limnobium laevigatum (Humb. & Blonpl.            Ff
      ex Willd.) Heine
    Najas conferta (A. Braun) A. Braun               Rs
    N. microcarpa K. Schum.                          Rs
  Alismataceae
    Echinodorus grandiflorus                         Em
      (Cham. & Schltdl) Micheli
    E. tenellus (Mart. ex Schult. &                  Rs
      Schult. f.) Buchenau
    Sagittaria montevidensis Cham. & Schltdl.        Em
    S. rhombifolia Cham.                             Em
  Limnocharitaceae
Hydrocleys nymphoides (Willd.) Buchenau              Rf
  Potamogetonaceae
Potamogeton pusillus L. ssp. pusillus                Rs
Asparagales
  Orchidaceae
    Habenaria repens Nutt.                           Ep
    Habenaria sp.                                    Ep
Poales
  Typhaceae
    Typha domingensis Pers.                          Em
  Xyridaceae
    Xyris jupicai Rich.                              Em
  Cyperaceae
    Cyperus diffusus Vahl                            Em
    C. digitatus Roxb.                               Am
    C. ferax Benth.                                  Em
    C. giganteus Vahl                                Em
    C. haspan L.                                     Em
    C. surinamensis Rottb.                           Em
    Cyperus sp.                                      Am
    Eleocharis elegans (Kunth) Roem. & Schult.       Em
    E. filiculmis Kunth                              Em
    E. geniculata (L.) Roem. & Schult.               Em
    E. minima Kunth                                  Em
    E. montana (Kunth) Roem. & Schult.               Em
    Frimbristylis autumnalis L.                      Am
    Oxycaryum cubense (Poepp. & Kunth) Palla         Ep
    Rhynchospora corymbosa (L.) Britton              Am
    Scleria melaleuca Rchb. ex Schltr. & Cham.       Am
    S. pterota C. Presl                              Am
  Poaceae
    Acroceras zizanioides (Kunth) Dandy              Am
    Echinochloa polystachya (Kunth) Hitchc.          Am
    E. crus-pavonis (Kunth) Schult.                  Em
    Eragrostis bahiensis (Schrad.                    Am
      ex Schult.) Schult.
    Eragrostis hypnoides (Lam.) Britton,             Em
      Sterns & Poggenb.
    Hymenachne amplexicaulis (Rudge) Nees            Em
    Leersia hexandra Sw.                             Am
    Megathyrsus maximus (Jacq.) B. K.                Am
      Simon & S. W. L. Jacobs.
    Panicum dichotomiflorum Michx.                   Em
    P. mertensii Roth                                Am
    P. pernambuncense (Spreng.) Mez ex Pilg.         Em
    P. prionitis Nees                                Am
    P. rivulare Trin.                                Am
    P. sabulorum Lam.                                Am
    Paspalum conspersum Schrad.                      Am
    P. millegrana Schrad.                            Em
    P. repens P.J. Bergius                           Em
    Setaria pauciflora Linden ex Herrm               Am
    Steinchisma laxa (Sw.) Zuloaga                   Am
    Urochloa humidicola (Rendle)                     Am
      Morrone & Zuloaga
    Urochloa subquadripara                           Em
      (Trin.) R.D. Webster
Commelinales
  Commelinaceae
    Commelina diffusa Burm. f.                       Am
    C. nudiflora L.                                  Em
    C. schomburgkiana var. brasiliensis Seub.        Em
    Floscopa glabrata (Kunth) Hassk.                 Em
  Pontederiaceae
    Eichhornia azurea (Sw.) Kunth                    Rf
    E. crassipes (Mart.) Solms                       Ff
    Heteranthera reniformis Ruiz & Pav.              Em
    H. seubertiana Solms                             Em
    Heteranthera sp.                                 Em
    Pontederia cordata L.                            Em
    P. parviflora Alexander                          Em
    P. triflora (Seub.) G. Agostini,                 Em
      D. Velasquez & Velasquez
Zingiberales
  Maranthaceae
    Thalia geniculata L.                             Em
Ceratophyllales -- Eudicotyledoneae
  Ceratophyllaceae
    Ceratophyllum demersum L.                        Fs
Caryophyllales -- Core Eudicotyledoneae
  Polygonaceae
    Polygonum acuminatum Kunth                       Em
    P. ferrugineum Wedd.                             Em
    P. hydropiperoides Michx.                        Em
    P. meisnerianum Cham. & Schltdl.                 Em
    P. punctatum Elliot                              Em
    P. stelligerum Cham.                             Em
  Amaranthaceae
    Alternanthera philoxeroides (Mart.) Griseb.      Am
    Gomphrena elegans Mart.                          Em
    Pfaffia glomerata (Spreng.) Pedersen             Am
    P. iresinoides (Kunth) Spreng.                   Am
  Haloragaceae
    Myriophyllum aquaticum (Vell.) Verdc.            Rs
Myrtales -- Rosidea
  Onagraceae
    Ludwigia grandiflora (Michx.)                    Em
      Greuter & Burdet
    L. helminthorrhiza (Mart.) H. Hara               Rf
    L. lagunae (Morong) H. Hara                      Em
    L. leptocarpa (Nutt.) H. Hara                    Em
    L. octovalvis (Jacq.) P.H. Raven                 Am
    L. peruviana (L.) H. Hara                        Em
  Lythraceae
    Cuphea melvilla Lindl.                           Em
    C. sessiliflora A. St.-Hil.                      Am
    Rotala mexicana Schltdl. & Cham.                 Rs
  Melastomataceae
    Acisanthera sp.                                  Em
Malpighiales -- Eurosideae I
  Podostemaceae
    Crenias sp.                                      Rs
    Podostemum rutifoliumWarming var. rutifolium     Rs
  Euphorbiaceae
    Caperonia castaneifolia (L.) A. St.-Hil.         Em
  Phyllanthaceae
    Phyllanthus niruri L.                            Am
Fabales
  Fabaceae
    Aeschynomene montevidensis Vogel                 Em
    A. sensitiva Sw.                                 Em
    A. virginica (L.) Britton, Sterns & Poggenb.     Am
    Sesbania cf. exasperata Kunth                    Am
  Fabaceae
    Vigna lasiocarpa (Mart.ex Benth.) Verdc.         Em
Cucurbitales
  Cucurbitaceae
    Cyclanthera hystrix (Gillies) Arn.               Am
  Begoniaceae
    Begonia cucullata Willd.                         Am
Malvales -- Eurosideae II
  Malvaceae
    Byttneria scabra L.                              Am
    Hibiscus sororius L.                             Em
    Melochia arenosa Benth.                          Am
Gentianales -- Euasterideae I
  Rubiaceae
    Diodia brasiliensis Spreng.                      Am
  Apocynaceae
    Oxypetalum sp. 1                                 Am
    Oxypetalum sp. 2                                 Am
    Rhabdanenia pohlii Mull. Arg.                    Em
Lamiales
  Plantaginaceae
    Bacopa salzmannii Wettst. ex Edwall              Rs
    Mecardonia procumbens (Mill.) Small              Em
    Scoparia dulcis L.                               Em
    S. montevidensis (Kuntze) R.E. Fr.               Em
  Linderniaceae
    Lindernia sp. 1                                  Rs
    Lindernia sp. 2                                  Rs
  Acanthaceae
    Hygrophila costata Nees                          Em
    H. guianensis Nees                               Em
    Justicia comata (L.) Lam.                        Am
  Lentibulariaceae
    Utricularia foliosa L.                           Fs
    U. gibba L.                                      Fs
    U. nigrescens Sylven                             Fs
Solanales
  Solanaceae
    Schwenckia americana L.                          Am
    Solanum glaucophyllum Desf.                      Am
  Convolvulaceae
    Ipomoea alba L.                                  Em
  Hydroleaceae
    Hydrolea spinosa L.                              Em
Apiales -- Euasterideae II
  Apiaceae
    Eryngium ebracteatum Lam.                        Em
    E. ekmanii H. Wolff                              Em
    Hydrocotyle ranunculoides L.f.                   Rf
Asterales
  Menyanthaceae
    Nymphoides indica (L.) Kuntze                   Rf
  Asteraceae
    Ecliptaprostrata (L.) L. (=alba) (L.) Hassk.)   Am
    Mikania cordifolia (L. f.) Willd.               Am
    Pluchea sagittalis (Lam.) Cabrera               Em

(L.F. = life forms; Em = emergent, Rs = rooted submerged,
Ff = free-floating, Am = amphibious, Rf = rooted floating,
Fs = free submerged and Ep = epiphyte).


ACKNOWLEDGMENTS

This study was funded by the Brazilian Council of Research (CNPq) through a Long Term Ecological Program (site number 6). S. M. Thomaz received a CNPq Productivity Research Grant and acknowledges this agency for long-term funding. A. Pott received a CNPq and CAPES Productivity Research. Additionally, F. A. Ferreira and R. P. Mormul acknowledge CNPq and CAPES (Coordenacao de Aperfeicoamento de Pessoal de Nivel Superior) for furnishing Ph.D. fellowships, respectively.

Received 04-VI-2010.

Corrected 30-XI-2010.

Accepted 07-I-2011.

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Fernando Alves Ferreira (1), Roger Paulo Mormul (1), Sidinei Magela Thomaz (2) *, Arnildo Pott (3) & Vali Joana Pott (3)

(1.) Postgraduate Program in Ecology of Continental Aquatic Environments, Universidade Estadual de Maringa- UEM, Av. Colombo 5790, bloco H-90, CEP 87020-900, Maringa, Parana, Brazil; ferreirabot@gmail.com, roger.mormul@gmail.com

(2.) Research Group in Limnology, Ichthyology and Aquaculture-Nupelia, Biological Science Department, Universidade Estadual de Maringa-UEM, Av. Colombo 5790, bloco H-90, CEP 87020-900, Maringa, Parana, Brazil; smthomaz@nupelia.uem.br

(3.) Biological Science Department, Universidade Federal do Mato Grosso do Sul--UFMS CEP 79.070-900, Caixa Postal 549, Campo Grande, MS, Brazil; arnildo.pott@gmail.com, vali.pott@gmail.com

* Corresponding author
TABLE 1
List of reports used to compare South American wetlands. The terms
in bold represent the corresponding analysis cluster

         Author(s)     Study region        Type of environment

1    Thomaz et        Upper Parana      River channels, secondary
     al. (2009)       River             channels, lagoons, swamps
                      floodplain,
                      Brazil (PR#)
2    Irgang &         Coastal plain     Swamps, saltmarshes,
     Gastal (1996)    of Rio Grande     rivers, lakes,
                      do Sul,           temporary ponds
                      Brazil (CP#)
3    Pott &           Pantanal          Shallow lakes, rivers,
     Pott (2000)      Matogrossense,    swamps, floodplains,
                      Brazil (Pan#)     meandering  ponds meander,
                                         "corixos", "vazantes",
                                        borrow pits, temporary
                                        ponds, permanent ponds
4    Bove et al.      State of Rio      Coastal lagoons, lakes,
     (2003)           de Janeiro,       permanent and temporary
                      Brazil (RJ#)      swamps, floodplains
5    Franca et        Brazilian         Artificial ponds
     al. (2003)       semiarid
                      Northeast
                      region (SA#)
6    Thomaz et        State of Amapa,   "Ressaca", environments
     al. (2003)       Brazil (AMA#)     influenced by tidal
                                        water regimes
7    Kahn &           Peru (Pe#)        Brackish ponds, mangroves,
     Leon (1993)                        rivers, lakes, wetlands
8    Crow (1993)      Ecuador (Ec#)     Lacustric systems
9    Terneus (2007)   The Amazon        Lakes, streams, rivers
                      basin of
                      Ecuador (AmEc#)
10   Scremin-Dias     Bodoquena in      Limestone springs and
     et al. (1999)    the State of      streams
                      Mato Grosso do
                      Sul,
                      Central-West,
                      Brazil (Bo#)
11   Junk &           Amazon River      Floodplain, "varzea"
     Piedade (1993)   near Manaus,      lakes, floating islands,
                      Brazil (Am#)      low-lying flats,
                                        low-lying swales,
                                        river shores, lake shores
12   Neiff (1986)     Middle Parana     Rivers, swamps, washways,
                      River             permanent ponds,
                      floodplain,       flooded ponds
                      Argentina
                      (PRA#)

Note: Bold character are indicated with #.
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Author:Alves Ferreira, Fernando; Mormul, Roger Paulo; Magela Thomaz, Sidinei; Pott, Arnildo; Pott, Vali Joa
Publication:Revista de Biologia Tropical
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Date:Jun 1, 2011
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