The selection of priority areas is an enormous challenge for biodiversity conservation. Some biogeographic methods have been used to identify the priority areas to conservation, and panbiogeography is one of them. This study aimed at the utilization of panbiogeographic tools, to identify the distribution patterns of aquatic insect genera, in wetland systems of an extensive area in the Neotropical region (~280 000km2), and to compare the distribution of the biogeographic units identified by the aquatic insects, with the conservation units of Southern Brazil. We analyzed the distribution pattern of 82 genera distributed in four orders of aquatic insects (Diptera, Odonata, Ephemeroptera and Trichoptera) in Southern Brazil wetlands. Therefore, 32 biogeographic nodes corresponded to the priority areas for conservation of the aquatic insect diversity. Among this total, 13 were located in the Atlantic Rainforest, 16 in the Pampa and three amongst both biomes. The distribution of nodes showed that only 15% of the dispersion centers of insects were inserted in conservation units. The four priority areas pointed by node cluster criterion must be considered in further inclusions of areas for biodiversity conservation in Southern Brazil wetlands, since such areas present species from differrent ancestral biota. The inclusion of such areas into the conservation units would be a strong way to conserve the aquatic biodiversity in this region.
La selección de áreas prioritarias es un enorme desafío para la conservación de la biodiversidad. Métodos biogeográficos se han utilizado para identificar áreas prioritarias para la conservación, como la panbiogeografía. Este estudio tuvo como objetivo el empleo de herramientas panbiogeográficas, para identificar los patrones de distribución de los géneros de insectos acuáticos, en los sistemas de humedales de una extensa área de la región Neotropical (~280 000km2), y así comparar la distribución de las unidades biogeográficas identificadas por los insectos acuáticos, con las unidades de conservación del sur de Brasil. Asimismo, se analizaron los patrones de distribución de los 82 géneros de cuatro órdenes de insectos acuáticos (Diptera, Odonata, Ephemeroptera y Trichoptera) en los humedales del sur de Brasil. Ahora bien, 32 nodos biogeográficos correspondieron a las áreas prioritarias para la conservación de la diversidad de insectos acuáticos. Dentro de este total, 13 se encontraban en el Bosque Atlántico, 16 en la Pampa y tres entre los dos biomas. La distribución de nodos mostró que sólo el 15% de los centros de dispersión de los insectos fueron insertados en las unidades de conservación. Las cuatro áreas prioritarias señaladas por criterio de nodo de clúster debe ser considerado en las inclusiones de los diferentes ámbitos para la conservación de la biodiversidad en los humedales del sur de Brasil, debido a que en esas zonas se presentan las 13 especies de la biota ancestrales diferentes. La inclusión de dichas áreas en las unidades de conservación sería una estrategia eficaz para conservar la biodiversidad acuática en la región.
Palabras clave: Panbiogeografía, análisis de trazos, áreas prioritarias, bioma, invertebrados acuáticos. The selection of priority areas is a huge challenge for biodiversity conservation (Sarkar & Margules 2002). Systematic methods for identifying biodiversity priority areas require good data on the distribution and abundance patterns of the ]]>
et al. 2002). Some phylogenetic and biogeographic methods are used for identifying the priority areas for conservation.
Among the biogeographic methods, the panbiogeography identifies, through the recognition of generalized and biogeographic tracks, the centers of origin and the biota evolution ]]>
et al. 1999). The areas selected by panbiogeographic criteria are interesting for the conservation (Morrone 1999) as they represent the true biodiversity hotspots (Crisci et al. 2003).
Biogeographic data for prioritizing the selection of areas for conservation has been used in several countries of South America (Morrone & Lopretto 1994, Menu-Marque et al.. 2000, Contreras-Medina & Eliosa-León 2001, ]]>
et al.. 2003), including Brazil (Franco-Rosselli & Berg 1997, Carvalho et al.. 2003, Lowenberg-Neto & Carvalho 2004, Morrone 2004, Morrone et al. 2004, Prevedello & ]]>
et al.. 2000, Mondragón & Morrone 2004), and gymnosperms in the world (Contreras-Medina et al. 1999, 2001a,b) being ]]>
In wetlands, panbiogeographic methods have been little used for selecting the areas for biodiversity conservation. In such ecosystems, other models, e.g. the species-arearelationship, have been used more in wetland conservation planning and management (Gibbs 2000), including ]]>
et al. 2007, 2008, Rolon et al. 2008). The hydroperiod and habitat heterogeneity also have been used as criteria for conservation of wetland communities (Collinson
et al. 1995, Snodgrass et al. 2000, Babbitt 2005, Van Geest et al. 2005).
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The search for ecologic criteria for the selection of priority wetlands is important, since these ecosystems are amongst the most affected and degraded ecological systems. Almost half of the wetlands in the world disappeared in the last century (Shine & Klemm 1999). In Southern Brazil, conservative data indicate that 90% of the wetlands have disappeared. Maltchik (2003) consider that approximately 72% of the remaining wetlands are smaller than 1km2. This pattern is a consequence of a severe habitat fragmentation due to agricultural expansion, especially rice ]]>
Southern Brazil is in an extensive area of Neotropical region (~280 000km2), represented by two large ]]>
2). The specific aims of this study were to: (1) recognize the common patterns (generalized tracks) of distributions of the aquatic insects in Southern Brazil wetlands; (2) recognize the centers of origin of aquatic insects (biogeographic nodes) in Southern Brazil; (3) compare the distribution of the biogeographic units identified by the aquatic insects with the conservation units of Southern Brazil and (4) propose the prioritization of areas for aquatic insects conservation in the Neotropical region.
This survey was developed in a large number of wetland systems, ranging a wide gradient of altitude and latitude. Aquatic insects are an important trophic level in wetland ]]>
et al. 2004).
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Materials and methods
Study area: The state of Rio Grande do Sul (RS) is located in Southern Brazil (27º04’ - 33º45’S - 49º42’ - 57º38’ W), has an area of 282 184km2 and presents Humid Subtropical Mid-Latitude Climate. The annual precipitation varies between 1 200 and 1 800mm, and it is relatively well distributed along the year without the existence of a dry period (Cfclimate classification of Köpen). The mean temperature varies between 15oC and 18oC, with minimum temperature bellow 10oC in the winter and maximum temperature above 32oC in the summer (RADAMBRASIL 1986).
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The state of Rio Grande do Sul has approximately 3 441 wetlands, presenting a total flood area of approximately 30 332km2 (Maltchik 2003). For this study, a total of 146 wetlands (Fig. 1) were ]]>
Aquatic insect sampling: Aquatic insect collections were carried using a kick net (D-shaped, 30cm in width, 400mm-opening mesh). Sampling was limited to the littoral zone of wetlands (water depths of less than 50cm), kicking up the substrate and then sweeping above the disturbed area to capture dislodged or escaping aquatic insects (Rosenberg et al. 1997). The sampling effort was the same for all wetlands, represented by 25 sweeps of 1m over several habitats of the littoral zone (detritus, rooted macrophytes and other dominant vegetation). Sweeps were pooled into one sample per wetland (3.5-l plastic bucket) and preserved in situ with 10% formaldehyde.
In laboratory, each sample was washed through a 400mm sieve, being the leaves, the stems, and other debris removed. The remaining material was preserved with 80% ethanol. Aquatic insects were separated and identified up to genus level under 7X magnification, according to Lopretto & Tell (1995), Trivinho- Strixino & Strixino (1995), Merritt & Cummins (1996), Fernández & Domínguez (2001) and the aid of specialists.
The data regarding the distribution of the genera of aquatic insects from four orders were considered for the analysis: Diptera (Chironomidae), Ephemeroptera, Odonata and Trichoptera. The criterion used for selecting the genera was the presence of more than one wetland in Southern Brazil.
The panbiogeographic approach basically consists of plotting distributions of different taxon on maps, connecting their separate localities with lines called individual tracks (Fig. 2A). When different individual tracks are superposed, the resulting summary lines are considered generalized tracks, which indicate the pre-existence of ancestral biotic components that were fragmented in the past due to tectonic and/or climatic changes. When two or more generalized tracks converge in a given area, they determine a node, which represents a complex area where different ancestral geological and biotic components interrelate in time and space (Morrone & Crisci 1995).
The analysis of the generalized tracks and biogeographic nodes was carried individually for each order and for all orders together. First, individual tracks were built for ]]>
Since each wetland was sampled once from March-October (eight months), the influence of the sampling period (seasonal element) on the insect distribution patterns ]]>
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Results
Insect distribution: The data regarding the distribution in Southern Brazil wetlands revealed 82 genera -37 of Chironomidae (Diptera), 29 of Odonata, eight of Ephemeroptera and eight of Trichoptera from 13 families. Based on the ]]>
Table 1).
The genera of Chironomidae (Diptera) defined 17 generalized tracks and 11 ]]>
Fig. 2B). The most important generalized tracks were g and o, created from six and five individual tracks, respectively. The track g is located in both biomes, whereas the track o is located in the Pampa (Fig. 2B). The most important nodes were 6 and 11 -both supported by ]]>
6 -located in the Pampa- was defined by the tracks g, n, and o, and the node 11 -located in the Atlantic Rainforest- was represented by the tracks c, e and g (Table 2,
Fig. 2B).
The genera Ephemeroptera defined three generalized tracks and three biogeographic nodes in Southern Brazil wetlands (Fig. 2C). The generalized track c was defined by three individual tracks of the genera Americabaetis, ]]>
and Caenis. The other two generalized tracks -a and b- were presented by two individual tracks each (Fig. 2C). The generalized tracks a and c are located in the Pampa; the track b is located in the transition of both biomes. The ]]>
Table 2, Fig. 2C). The genera of Odonata identified 12 generalized tracks and nine biogeographic nodesseven nodes in the Atlantic Rainforest and two in the Pampa (Fig. 2D). The most important generalized tracks were h (represented by seven individual tracks), and f (represented by ]]>
Table 2). The track h was defined by the genera Coryphaeschna, Acanthagrion, Cyanallagma, Homeoura, Oxyagrion, Micrathyria, and Tramea, and was mainly associated with the biome Atlantic Rainforest. The track f was represented by Cyanallagma, Homeoura, Leptobasis, Oxyagrion, Lestes, Micrathyria, Tramea, and Telebasis, and was located in ]]>
Table 2, Fig. 2D). The genera of Trichoptera defined only one generalized track, but no biogeographic node (Table 2, Fig. 2E). The track identified by the genera Oecetis and Oxyethira link both biomes. ]]>
The superposition of all 63 individual tracks defined 16 generalized tracks and seven biogeographic nodes represented by different orders of aquatic insects (Table 2, Fig. 2F). The generalized track h was defined by eight genera of Chironomidae, Odonata, Ephemeroptera, and Trichoptera. Other important generalized tracks were c, f, and n -all represented by seven individual tracks each (Fig. 2F). The most important biogeographic nodes defined by three generalized tracks, at least, were: one, represented by the generalized tracks i, p, and o and all located in the Atlantic Rainforest; two, represented by the generalized tracks f, g, i, and m and all located in the Atlantic Rainforest; and five, represented by the generalized tracks c, e, and j and all located in the Pampa (Table 3, Fig. 2F). The other four biogeographic nodes were located in the Pampa (Fig. 2F).
Implications for conservation: All generalized tracks and biogeographic nodes identified in Southern Brazil wetlands were superposed in a summary map (Fig. 3A). Therefore, 32 biogeographic nodes corresponded to the priority areas for conservation of the aquatic insect diversity in Southern Brazil wetlands. Among this total, 13 were located in the Atlantic Rainforest, 16 in the Pampa, and three amongst both biomes (Fig. 3A). According to the node cluster criterion, the priority regions for conservation were the following: (a) Site one (~9 000km²), located in the Western region in the Pampas, presenting two main areas that totalize ten nodes; (b) Site two (~1 000km²), located in the South-eastern region, in the Pampa and represented by the three nodes; (c) Site three (~7 000km²), located in the Eastern region, in the Atlantic Rainforest, and represented by four nodes; and (d) Site four (~6 000km²), located in the central region, in the Atlantic Rainforest and presented by three nodes (Fig. 3B).
The comparison between the distribution of the biogeographic nodes and the distribution of the conservation units in Southern Brazil, showed that several priority areas for the conservation of the diversity in aquatic insects are found outside the conservation units (Fig. 4). Only five out of 32 biogeographic nodes (approximately 15%) were partially within the conservation units (three nodes in the Pampa, one node in the Atlantic Rainforest, and one node between both biomes). The corresponding areas to the other biogeographic nodes were not inserted in the conservation units (Fig. 4). All genera composition matrix was correlated with geographic distance matrix (r=0.118, p=0.001). When we analyzed each invertebrate group separately, Trichoptera and Chironomidae genera were correlated with geographic distance matrix (r=0.094, p=0.016 and r=0.072, p=0.001, respectively). Odonata and Ephemeroptera genera were not correlated with geographic distance matrix (r=0.037, p=0.138 and r=0.057, p=0.085, respectively), nor with sampling seasonality matrix (r=0.025, p=0.201 and r=0.015, p=0.34, respectively). These results showed that the distribution patterns of aquatic insects genera in Southern Brazil wetlands were not influenced by the sampling period.
On the other hand, the number of nodes was loosely correlated with the number of genera collected in the studied wetlands (R²=0.267, F1,40=13.862, p<0.001). In this sense, the number of nodes cannot be highly predicted by insect richness because the major portion of variation in the number of nodes was not explained by insect richness, given that many areas fall far from the regression line (coefficient of determination of only 0.267) (Fig. 5).
Discussion
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Insect distribution: Congruences of biogeographic patterns of the phylogenetically related groups may be more associated to a common natural history than to independent dispersion events. Independent dispersion events occur as a reaction from each group to different environmental conditions (Gullan & Cranston 2008). The coinciding patterns of individual tracks of genera analyzed (e.g. generalized track) indicated a narrow relationship between biota and studied region history. In this sense, we can infer ]]>
The faunal similarity observed was not influenced by the sampling period, but by the natural history of aquatic insects. Since the studied groups present ephemeral adults ]]>
The distribution patterns found varied between the studied biomes. Most of the tracks and nodes found for different orders of aquatic insects were within the Pampa, excepting for Odonata. For Chironomidae, ten out of 17 generalized tracks were located in the Pampa. According Ferrington (2008), intermittent and ephemeral aquatic ecosystems, like Brazilian lowlands, ]]>
Most of tracks of the Odonata genera were distributed in higher altitude of the Atlantic Rainforest. According to Kalkman et al. (2008), significant regions of the Odonata diversification include the regions with higher altitude, such as the ]]>
Odonata species observed belong to these two families, occurring in lentic water habitats of savannas, such as the wetlands in the Pampa. The occurrence of two families of ]]>
et al. (2004), these families were found in the extratropical, warm temperate latitudes of Laurasia (England and Siberia), and they dispersed in the Early Cretaceous across other landmasses including Gondwana (Brazil). A limited dispersal ability of both families of Trichoptera reflects the current distribution restriction of the genera found in Southern Brazil wetlands, since such order of aquatic insects was presented by a unique generalized track and by no biogeographic node at the region studied.
Despite the peculiarities in the distribution of each group, the general patterns of distribution of aquatic insects in Southern Brazil wetlands are related to the formation of the Atlantic Rainforest and the Pampa. The three most important biogeographic nodes supported by many generalized tracks, reflect a huge vicariant event at the studied region and represent large areas of aquatic biodiversity. The two largest nodes present in ]]>
distributed in areas located at lower altitudes and with restricted dispersal ability such as Ephemeroptera and Trichoptera. Such relation shows the importance of the landscape on the diversity and distribution of aquatic insects at the Neotropical region.
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Implications for conservation: The biogeographic nodes guide historically the generalized regional tracks (Craw et al.1999), since the patterns of distribution of the biota are influenced by the events that occur in each region. Such centers reflect indirectly the long-term maintenance of the biodiversity and spatial structure of the biota. They also include areas of high ]]>
Our results showed that the conservation unit system in Southern Brazil do not contemplate several priority areas for the historical and biogeographic conservation of wetland aquatic insects. Several areas important for ]]>
Brazil (Carvalho & Ozório 2007). The priority areas pointed by node cluster criterion must be considered in further inclusions of areas for biodiversity conservation in Southern Brazil wetlands, since such areas ]]>
Acknowledgments
This research was supported by funds from Universidade do Vale do Rio dos Sinos – NISINOS (02.00.023/00-0), and Conselho Nacional de Desenvolvimento Científico e Tecnológico -CNPq (52370695.2). Leonardo Maltchik holds a Brazilian Research Council -CNPq Research Productivity grant. We declare that the data collection complied with the Brazilian current laws. ]]>
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*Correspondencia: Leonardo Maltchik: Laboratory of Ecology and Conservation of Aquatic Ecosystems, Av. Unisinos, 950 CEP 93.022-000, UNISINOS, São Leopoldo, RS, Brazil; UNISINOS, São Leopoldo, Brazil; maltchik@unisinos.br Marina Schmidt Dalzochiov: Laboratory of Ecology and Conservation of Aquatic Ecosystems, Av. Unisinos, 950 CEP 93.022-000, UNISINOS, São Leopoldo, RS, Brazil; UNISINOS, São Leopoldo, Brazil; mahsdalzochio@gmail.com Cristina Stenert: Laboratory of Ecology and Conservation of Aquatic Ecosystems, Av. Unisinos, 950 CEP 93.022-000, UNISINOS, São Leopoldo, RS, Brazil; UNISINOS, São Leopoldo, Brazil; cstenert@unisinos.br Ana Silvia Rolon: Laboratory of Ecology and Conservation of Aquatic Ecosystems, Av. Unisinos, 950 CEP 93.022-000, UNISINOS, São Leopoldo, RS, Brazil; UNISINOS, São Leopoldo, Brazil; asrolon@gmail.com
Received 22-III-2011. Corrected 30-VI-2011. Accepted 28-VII-2011.
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