Open access journal: http://periodicos.uefs.br/ojs/index.php/sociobiology ISSN: 0361-6525 Sociobiology 60(1): 41-49 (2013) Spatial connectivity of aquatic macrophytes and flood cycle influence species richness of an ant community of a Brazilian floodplain MC Pereira1, JHC Delabie2,3, YR Súarez, WF Antonialli Junior1 Introduction Floodplain environments with flooding regime har- bor a high biological diversity (Welcomme, 1985; Lowe- McConnell, 1999) as a result of the spatial and dynamic complexity existing primarily in habitats with these charac- teristics (Ward et al., 1999; Robinson et al., 2002). Junk et al. (1989) proposed the “flood pulse concept”, which considers the hydrological aspects along with the geomorphological aspects, responsible for spikes in floods and droughts, with different amplitudes and periods along the hydrographic ba- sin and consider this seasonality the greatest driving force of biota composition in the rivers of the floodplain. The initial step of knowledge of available natural resources in a region corresponds to the collection and taxo- nomic identification of species that make up the fauna and flora, in order to facilitate the acquisition of subsidies for more detailed studies of ecological characteristics of their habitats. Furthermore, these studies may eventually lead to Abstract Despite the environmental and economic importance of Pantanal, there are few studies quantifying the influence of sazonality and spatial variation on biological diversity in this ecosystem. In this context, the present work aimed to study the assemblage of ants associated with macrophytes during the flood and dry period of Paraguay river, in marginal environments in the Pantanal of Porto Murtinho, Mato Grosso do Sul. We observed a wide variation in the temporal distribution of the diversity of the assemblages of ants, since from 37 species, 36 occurred in the dry and 20 in the flood period. Of the total of macrophyte species observed, only in 12.5% we found a more specific correlation with ants that were nesting in spaces provided by plants representing a total of 10.52% of the species ana- lyzed. Sociobiology An international journal on social insects 1 - Universidade Estadual de Mato Grosso do Sul, Dourados-MS, Brazil 2- CEPLAC - Itabuna-BA, Brazil 3 - Santa Cruz State University, Ilhéus-BA, Brazil ReSeARCH ARtICle - AntS Article History edited by: Gilberto M.M. Santos, UeFS - Brazil Received 11 October 2012 Inicial acceptance 31 October 2012 Final acceptance 21 november 2012 Keywords Diversity, Flood Pulse, Foraging habits, Formicidae, Pantanal. Corresponding author Márlon César Pereira Centro Integ. de Análise e Monitoramento Ambiental, Universidade estadual de Mato Grosso do Sul Rodovia Dourados/Itahum, Km 12,5 Caixa Postal 351, Dourados-MS, Brazil, 79804-970 e-Mail: marloncesarp@yahoo.com.br a body of knowledge that allows the rational exploitation of biotic resources and adaptation to abiotic conditions of the studied environment (Prado, 1980). According to Lewinsohn et al. (2005) Pantanal is one of the most unknown biomes of Brazil, and the function- al role of invertebrates in this ecosystem is a strong attribute for its conservation. Among these organisms, the ant fauna is one of the most successful group of insects, dominant in number and biomass in several environments (Harada & Adis, 1997; Santos et al., 2003; Battirola et al., 2005), and considered abundant, easy to collect and identify as well as relatively quick to respond to changes in their habitats (Ribas et al., 2003). In addition to its relatively high local abundance, they are especially rich in species and diverse in terms of foraging habits, nesting, among other ecological functions (Blüthgen & Feldhaar, 2010). In tropical ecosystems, ants constitute more than 15% of the total animal biomass (Beattie & Hughes, 2002). Most species are predators and their structuring role in ar- MC Pereira, JHC Delabie, YR Súarez, WF Antonialli Jr - Ant Assemblages respond to Aquatic Macrophytes and Flood Cycle42 thropod community is highlighted in several studies (e.g., Carroll & Janzen, 1973; Jeanne, 1979; Wilson, 1987; Höll- dobler & Wilson, 1990). In addition, when foraging over the vegetation, they can decrease the rate of herbivory and increase the reproductive success of plants (Oliveira et al., 1999). In this sense, there are mutualistic interactions be- tween ants and plants in which the latest provide nesting site and food for ants in exchange for its defensive activity (Beattie, 1985; Styrsky & Eubanks, 2010). Despite the environmental importance of Pantanal, which plays an important role in biological diversity due to the variety of natural habitats, opportunities for feeding and reproductive niches, and essential ecosystem services including carbon storage, flooding control, fish production, and aquifer recharge (Alho, 2005; Alho & Gonçalves, 2005; Alho et al., 1988; Keddy & Fraser, 2005), we can still say that there are few studies quantifying the influence of sea- sonality and spatial variation on its biological diversity. The few ecological studies are concentrated in some places, like the plain of Cuiabá River, Paraguay River in the region of Corumbá, Miranda River and Negro River and refer mainly to communities of aquatic plants, fish, phyto and zooplankton, except for the work of Raizer and Amaral (2001) on spiders associated with aquatic macrophytes and Alves-dos-Santos (1999) on the pollination of Pontederiace- ae. In this context, this study aimed to evaluate the ant as- semblages that occur during the change in the disposition of macrophytes during the season of flood and drought in mar- ginal environments of the Paraguay River in Porto Murtinho Pantanal , Mato Grosso do Sul , Brazil. Material and Methods Study area The samples were collected during the day pe- riod from March/2009 to March/2010 in Porto Murtin- ho, state of Mato Grosso do Sul, Brazil (Fig. 1), in Para- guay (21°41’40.86”S and 57°53’10”W) and Amonguijá Rivers (21°41’10.39” S and 57°52’51.81”W), Crimi- nosa (21°40’27”S and 57°53’30.6”W) and Flores Lakes (21°45’56”S and 57°54’58.67”W). The altitude of these ar- eas is around 75 m, and the vegetation ranges from grassy to sparse shrub areas. Data sampling The sampling sites, in each water body described above, were randomly chosen. In each sampling site, the ants were searched and collected in macrophyte beds at littoral zones. We developed our ant samplings at the emerged por- tions of aquatic macrophytes, using entomological nets and tweezers. The ant samplings were carried out on a monthly basis, at least in five plants in each sampling site throughout the study. The sampled ants were fixed at 70% ethanol in labeled vials, and identified by Dr. Jacques C. H. Delabie, using by reference the Coleção do Laboratório de Mirme- cologia do Centro de Pesquisas do Cacau (CEPEC), follow- ing the nomenclature adopted by Bolton (1994; 1995b and 2003). Voucher specimens were deposited at number 5602 in the collection previously cited. The ants were collected with MMA – SISBIO permission (17487 – 1). Sampled macrophytes were collected and pressed to posterior identification at Herbário da UFMS in Campo Grande/MS by Dr. Arnildo Pott, Dr. Vali Joana Pott and Gabriela Serra, with taxonomic keys proposed by Scremin- Dias (1999), Pott &Pott (2000) and Amaral et al. (2008) and voucher specimens were deposited in Herbário da UFMS (CGMS) in Campo Grande/MS. To facilitate the visualization of the dynamics of ant assemblages that occurred in macrophytes, we divided the year of collection in steps of dry season (September-Febru- ary) and flood season (March-August), based on the level of the Paraguay River every month (Fig. 2). The total species richness was estimated using the bootstrap method (Smith & van Belle, 1984), with a confi- dence interval (α = 0.05), using presence/absence data for all samples. This method was selected for its robustness, with relatively large sample sizes (Hellmann & Fowler, 1999). We used Pearson`s correlation between ant richness and environmental variables obtained from the meteorological station of Porto Murtinho (rainfall) and from field data (tem- perature and river level) (Fig. 3). To assess whether or not there was specificity between species of ants and macrophytes, the data referring to all ants Figure 1. Map of the study area. Porto Murtinho, state of Mato Gros- so do Sul, Brazil. Sociobiology 60(1): 41-49 (2013) 43 and macrophytes were submitted to cluster analysis, through the methods of Jaccard and UPGMA as linkage method, us- ing the statistical program R (R Development Core Team, 2011). To assess whether the generated dendrogram reflects the similarity matrix in species distribution in the plants ana- lyzed we used the cophenetic correlation coefficient, defin- ing the minimum value of 0.75 as a measure of dendrogram adjustment quality. Figure 2. Accumulated of rainfall and level of the Paraguay River during the year, in the region of Porto Murtinho – MS. Results and Discussion We collected 582 ant individuals, belonging to six sub-families, 17 genera and 37 species that occurred in 36 species of aquatic and paludal macrophytes belonging to 14 orders and 18 families (Table 1). Through the boostrap method we estimate that 41 species of ants occur associated with these species of macrophytes (IC = 37 to 46) so, this result suggests that 92.7% of species of ants were sampled, indicating good sample size sufficiency. The higher proportion of ant species was repre- sented by the subfamily Formicinae with thirteen species (35.1%), followed by Myrmicinae with ten species (27%) and Dolichoderinae and Pseudomyrmicinae with five spe- cies (13.5%) each. These results are not consistent with other studies about ant fauna (Longino & Nadkarni, 1990; Johnson & Ward, 2002; Diehl et al., 2005) which the authors found more species of the subfamily Myrmicinae in differ- ent biomes. Moreover, the results differ from the findings of Corrêa et al. (2006) “Capões” of Pantanal - MS, which fol- lows the pattern described above. However, the great propor- tion of Formicinae found in our survey could be explained by the fact that our samples were accomplished in open ar- eas, as in the investigations of Marinho et al. (2002), Leal (2002, 2003). The natural habitat of Formicinae is vegeta- tion (Hölldobler & Wilson, 1990; Brühl et al., 1998), there- Figure 3. Person`s correlation analyses (a) ants species richness and temperature, (b) ants species richness and rainfall, (c) ants species richness and river level. fore they can be frequently found in this place (Marques & Del-Claro, 2006). As for the correlation analyses, rainfall (Fig. 3a) was the variable that most explained the ant’s richness, followed by temperature (Fig. 3b) and river level (Fig. 3c). The high- MC Pereira, JHC Delabie, YR Súarez, WF Antonialli Jr - Ant Assemblages respond to Aquatic Macrophytes and Flood Cycle44 Macrophyte Species Code Ant Species Visited Macrophytes Species Order Alismatales Azteca sp (15) Family Alismataceae Dorymyrmex sp. 1 (4 - 33 Echinodorus tenellus 1 Dorymyrmex sp. 2 (40) Order Asterales Dorymyrmex sp. 3 (330 Family Asteraceae Linepithema humile (1 - 2 - 3 - 4 - 6 - 7 - 8 - 9 - 10 - 11 - 12 - 13 - 14 - 15 - 16 - 17 - 18 - 19 - 20 - 21 - 22 - 23 - 24 - 25 - 28 - 29 - 30 - 31 - 32 - 33 - 34 - 35 - 36) Enydra radicans 2 Subfamily Ectatomminae Melanthera latifolia 3 Ectatomma brunneum (8 - 11 - 12 - 14 - 15 - 22 - 28 - 33) Pacourina edulis 4 Subfamily Formicinae Order Caryophyllales Brachymyrmex patagonicus (15) Family Amaranthaceae Camponotus (Myrmaephaenus) sp 1 (11) Alternanthera aquática 5 Camponotus (Myrmaephaenus) sp 2 (5 - 8 - 14 - 15 - 21 - 33) Family Polygonaceae Camponotus (Myrmaephaenus) sp 3 (2 - 14 - 22 - 28 - 33) Polygonum acuminatum 6 Camponotus crassus (2 - 3 - 7 - 8 - 10 - 11 - 14 - 15 - 19 - 22 - 23 - 28 - 29 - 32 - 33 - 35) Polygonum ferrugineum 7 Camponotus leydigi (2 - 22 - 28) Polygonum punctatum 8 Camponotus melanoticus (14 - 22 - 30 - 33) Family Portulacaceae Camponotus novogranadensis (4 - 8 - 10 - 11 - 12 - 14 - 22 - 32 - 33) Portulaca grandiflora 9 Camponotus rufipes (4 - 11 - 14 - 16 - 22 - 33) Order Commelinales Camponotus sexguttatus (4 - 14 - 32 - 33 - 35) Family Pontederiaceae Camponotus vittatus (14) Eichhornia azurea 10 Nylanderia sp1 (4 - 5 - 7 - 14 - 29 - 33 - 36) Eichhornia crassipes 11 Nylanderia sp2 (33 - 35) Pontederia rotundifolia 12 Subfamily Myrmicinae Order Curcubitales Acromyrmex balzani (4 - 8 - 10 - 11 - 14 - 30 - 33) Family Curcubitaceae Atta sexdens rubropilosa (2 - 4 - 10 - 11 - 22 - 35) Cyclanthera hystrix 13 Cephalotes minutus (4 - 10 - 11 - 20 - 33 - 35) Order Fabales Cephalotes pavonii (4 - 14 - 15 - 28 - 33) Family Fabaceae Crematogaster sp (8 - 10 - 11 - 12 - 35) Aeschynomene sensitiva 14 Crematogaster victima (33) Bauhinia bauhinioides 15 Cyphomyrmex transversus (33) Mimosa pigra 16 Pheidole obscurithorax (4 - 6 - 8 - 10 - 11 - 14 - 16 - 29 - 30 - 33 - 35) Neptunia plena 17 Solenopsis invicta (3 - 7 - 11 - 14 - 15 - 19 - 22 - 28 - 29 - 32 - 33 - 35 - 36) Senna aculeata 18 Solenopsis sp (4 - 36) Senna alata 10 Subfamily Ponerinae Senna occidentalis 20 Hypoponera opaciceps (30) Sesbania virgata 21 Hypoponera sp. (11 - 32) Vigna lasiocarpa 22 Odontomachus haematodus (10 - 16 - 33 - 35) Order Malpighiales Subfamily Pseudomyrmecinae Family Euphorbiaceae Pseudomyrmex denticollis (11 - 9 - 29 - 30 - 32) Caperonia castaneifolia 23 Pseudomyrmex gracilis (3 - 21 - 22 - 29 - 33 - 35) Order Malvales Pseudomyrmex simplex (11 - 28 - 29 - 33 - 34) Family Malvaceae Pseudomyrmex sp (pr. palidus) (19 - 26 - 27 - 33) Hibiscus striatus 24 Pseudomyrmex termitarius (2 - 8 - 15 - 22 - 33) Pavonia laetevirens 25 Order Myrtales Family Onagraceae Ludwigia grandiflora 26 Ludwigia sedoides 27 Order Poales Family Poaceae Setaria paucifolia 28 Urochloa subquadripara 29 Order Salviniales Family Azollaceae Azolla filiculoides 30 Family Marsileaceae Marsilea crotophora 31 Family Salviniaceae Salvinia auriculata 32 Order Solanales Family Convolvulaceae Ipomoea alba 33 Ipomoea chiliantha 34 Order Vitales Family Vitaceae Cissus spinosa 35 Order Zingiberales Family Marantaceae Thalia geniculata 36 Table 1. List of ant species and macrophytes species sampled in marginal environments of the Paraguay and Amonguijá Rivers and Criminosa and Flores Lakes, in the region of Porto Murtinho, Mato Grosso do Sul, Brazil, from March/2009 to March/2010. Sociobiology 60(1): 41-49 (2013) 45 est richness was observed in July, probably because in this month there is the flood peak of the region, which makes the ants that are foraging to look for fixation in the substrate, since the macrophytes that are more inside the river become subject to being dragged by the flood. As for the cluster analysis performed, in Figure 4 we can observe 6 distinct groups. Group 1 (Fig. 4) in which the plants were used by the ants as a means of connection to other plants, such as Azolla filiculoides (Lam.), or the plants presented little floral cycles during the work, Mimosa pigra (Willd). Group 2 (Fig. 4) less common plants or plants with well-defined cycles of flowering, so that ants were only found in them when flowering occurred. The macrophytes of Group 3 , all showing in common the fact of being with- out connection with other plants during the full sampling period, becoming “islands”, in which some species of ants were found isolated from other areas and/or species, as in the case of Senna alata (Roxb.) in which we found, in some instances, entire colonies of Solenopsis invicta (Buren 1972) adhered to the plant, possibly reaching it by boating floating (Tschinkel, 1988). In Group 4, are the macrophytes specie in which occurred the higher frequency of ant species through- out the study period. In common, they have as features, be- sides being abundant, the provision of important resources such as flowers, domatias and nectaries during most part of the year (Pott & Pott, 2000). In group 5 is the macrophyte Pontederia rotundifolia (L. F.), which occurred usually in the center of large groups of macrophytes of Pontederiaceae family, which hindered the access of ants to these plants. And the Group 6 with a single member, Salvinia auriculata (Aubl.) which is used as a means of connection, as A. filicu- loides, but presented large distribution during samplings and possesses bristles that hinders the passage of small ants. The plant species with the highest number of visit- ing ants species was Ipomoea alba (L.), in which 25 spe- cies were recorded, probably due to the fact that this plant ia shrubby and amphibious, providing attractiveness to ants throughout the year. Eichhornia crassipes (Mart Solms) was another species with a high relative number of visits, with 15 species recorded, and high abundance throughout the study period. For the same reason, and also because during the entire year the plant species Pacourina edulis (Aubl.), Cissus spinosa (Cambess.) and Vigna lasiocarpa (Benth. Verdcourt) flourish, in these species we recorded 13, 11 and 11 species, respectively. Figure 4. Cluster analysis of the species of macrophytes according to the species of visitor ants. G1= macrophytes used as a means of transi- tion by the ants, G2 = Macrophytes that flourished during a time of year, G3 = islands macrophytes, G4 = macrophytes with high frequency of visits, G5 = connection bridges abundant at sampling area, and G6 = macrophyte isolated in Pontederiaceae group. MC Pereira, JHC Delabie, YR Súarez, WF Antonialli Jr - Ant Assemblages respond to Aquatic Macrophytes and Flood Cycle46 The ant species Linepithema humile (Mayr 1868) is an indicator of 4 groups of plants (Fig. 4) and occurred in 91.6% of plant species. However, it was more frequent in Ipomoea chiliantha (Hallier). This ant species is invasive in several environments, mainly in Europe, United States, Aus- tralia and Brazil and as such, end up being opportunistic in environments newly colonized (Suarez et al., 2001; Holway et al., 2002). Because they are aggressive in environments in which they settle, they can monopolize food sources of many native ants by the high recruitment capacity that this species presents (Majer, 1994; Gómez & Oliveiras, 2003; Touyama et al., 2003). In addition to this, we found colonies of L. humile nesting inside P. edulis and indeed the doma- tias offered by this macrophyte have already been described (Pott & Pott, 2000). The genus Camponotus (Emery 1889) was the most rich , with 11 ant species (29.7% of species richness), prob- ably because it is one of the most abundant genera world- wide, with varied habits and an excellent foraging system (Holldobler & Wilson, 1990), reflecting the prevalence de- scribed by Wilson (1976) and Bolton (1995a). According to them, Camponotus, Pheidole (Westwood 1839), Solenopsis (Smith 1858) and Crematogaster (Lund 1831) are the genera with the highest diversity of species and adaptations, great- est geographical distribution extension and greatest local abundance, and, therefore, are considered the most preva- lent genera on a global scale. In fact, this genus is always very frequent in inventories of ants as can be seen in works such as those of Majer and Delabie (1994) studying the com- munity of ants of the Amazon, Soares et al. (1998) in euca- lyptus plantations and secondary Atlantic forests, Verhaagh and Rosciszewski (1994) in different biomes in Bolivia, and in open areas such as restinga (Gonçalves & Nunes, 1984), cerrado (Marinho et al., 2002), caatinga (Leal, 2002, 2003) and also associated with vegetation (Wilson, 1976). In this group of ants is the species Camponotus novogranadensis (Mayr 1870) which has been shown to be an indicator of 3 groups of plants (Fig. 4): Group 2, probably because it is an abundant group and occurs throughout the year; Group 5, due to the fact that the plant is difficult to reach and the ant is an opportunistic forager; and Group 6, because the plant is widely distributed throughout the year as the Group 2. The most common species of the genus Camponotus was C. crassus (Mayr 1862), occurring in 16 macrophyte species. Only two species of the genus Solenopsis were found (Table 1), however, occurred in 14 species of macrophytes, this is probably related to physiological and behavioral as- pects of these species, which may spend long periods of food shortage and compete with other species of ants or other groups of animals since they present an efficient bulk recruitment strategy (Fowler et al., 1991). In both species the boating floating behavior was observed, this behavior (Tschinkel, 1988) probably indicates that such colonies built their nests on the banks of rivers and streams during the dry season, and when those areas were flooded, they floated, waiting for the opportunity to restore their colony in some other appropriate place. Those ants demonstrate the strong association of biological traits of such ant species with hy- drological systems in general. The genus Pseudomyrmex (Lund 1831), frequently described as arboreal (Ward, 1989), was found in 55.55% of the plants near the river bank, probably using the plants as substrate in search for prey, presented five species and one of them registered for the first time in Brazil (Pseudomyrmex denticollis Emery 1890). However, its distribution was not- ed at the central region of Paraguay, approximately 350km far from our sampling area in Porto Murtinho. Considering, that the region where P. denticollis was registered in Para- guay has the same vegetal formation of Porto Murtinho (dry Chaco). The least frequent species of ants were found, most part of the time foraging, in the leaves, trunk or flowers, or even using the plant as a means of connecting to another which was providing resources at that moment. A. filicu- loides and S. auriculata, for example, are species associated with floating macrophytes abundant in the region, which are commonly used as a means of connection so that these ants can forage in other plants. Most species of ants visited more than one species of macrophyte, suggesting that in general they do not establish dependency relationship with these plants, what is probably related to the fact that this environ- ment is very unstable throughout the year, mainly, due to the fast change in the level of the river during flood season (Fig. 2). We can observe a large variation in the temporal dis- tribution of diversity in the studied ant assemblage. From 37 species observed, 36 occurred in the dry season and 20 in the flood. This variation occurs probably because there is a vari- ation in the availability of macrophyte banks throughout the year since, during the flood, the greater volume of water that generates greater transport capacity, carries along its course the majority of macrophytes in which the ants were foraging in the previous season. A greater diversity of species during the dry season, probably because during the months of flood, macrophytes decrease in number as they will gradually fall- ing off, being dragged and changing position by the force of the currents and wind (Tur, 1972). This dynamics, for sure, causes a considerable decrease in the foraging substrate for ants during this period. In the analysis of similarity among the ant species (Fig. 5) we can see three main groups: Group 1 contains the species that were uncommon in a few species of mac- rophytes. This group fits, for example, Camponotus vittatus (Forel 1904), which was isolated by having occurred only in Aeschynomene sensitiva (Sw.) during the full season (Fig. 5). The low frequency is probably explained by the com- monness of this species in urban areas (Soares et al., 2006), not in rural areas. Sociobiology 60(1): 41-49 (2013) 47 The species Hypoponera opaciceps (Mayr 1887), Hypoponera sp. and P. denticollis foraged only in A. filicu- loides, S. auriculata and E. crassipes. In these cases, the low frequency is probably explained by their trophic nature as generalist predators (Santschi, 1938), specially Hypoponera (Taylor 1967). These ants must use these plants as substrate to capture prey that forage in search of different resources in these plants. In Group 2, we also found species that occupied spaces offered by the plants, i. e. domatias (Holldobler & Wilson, 1990). This was the case of Dorymyrmex sp 2, Cre- matogaster victima (Smith 1858) and Cyphomyrmex trans- versus (Emery 1894) found nesting in P. edulis and in young plants of I. alba. Finally, belonging to Group 3, are the ants that oc- curred in most plants and sampling months, which we con- sidered opportunistic foragers of several resources. S. in- victa represents such an example of taking advantage of plants, using them as resources when water flooded their nests and they used the mechanism of boating floating (Tschinkel, 1988) to reach the closest plants, as discussed earlier. In plants where we found S. invicta no other ant spe- cies was found. Most likely, this occurs because of the size and aggressiveness of S. invicta colonies (Delabie & Fowler, 1995). Another species similar in occurrence was C. cras- sus, the most common ant of the genus Camponotus, and L. humile, the Argentine invasive ant (Majer, 1994; Suarez et al., 2001). Thus, we can conclude that the several species of macrophytes are important resources for ants in Pantanal, either as substrate where foragers find preys, nectaries, or other resources or as places to establish their colonies, find- ing therefore, besides food resources, shelter. Acknowledgments The authors thank to Pantanal Research Center, Uni- versidade Estadual de Mato Grosso do Sul and Fundect for financial support. We thank Gabriela Serra do Vale Duarte for the plant species identification and Ingrid de Carvalho Guimarães for the revision of the English text. We thank two anonymous reviewers for valuable comments that improved this article. References Alho, C.J.R. (2005). The Pantanal. In: L.H. Fraser & P.A. Keddy (Eds.), The World’s Largest Wetlands - Ecology and Conservation (pp. 203-271). 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