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��ࡱ�>��	�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������{�	��&�bjbjz�z�	B�����?+���������
$$*$*$*$����>$>$>$8v$�*%D>$!U$n%:�%�%�%�%�&>�&�&�T�T�T�T�T�T�T$EW��Y:�T*$'�&�&''�T*$*$�%�%��T747474'\*$�%*$�%�T74'�T7474�I��K�%�����/P�������g2T�J�T�T0!U�J�1Z�3|1Z4�K�K1Z*$�K�''74'''''�T�T74'''!U''''��������������������������������������������������������������������1Z'''''''''� �":	Enhancing Biodiversity by Restoring Wetland Vegetation Communities in Irrigation Ponds 

Darby M. McGrath and Stephen D. Murphy2

1Department of Environment and Resource Studies
 200 University Avenue West 
Waterloo ON Canada
 N2L 3G1
Phone: 1-289-296-7209

7700 Thomas Street 
Niagara Falls ON
L26 6T2
(corresponding author�s permanent address)

2Department of Environment and Resource Studies
 200 University Avenue West 
Waterloo ON Canada
 N2L 3G1
Phone: 1-519-888-4567 x 35616 fax: 1-519-746-0292








Abstract
The purpose of this pilot research is to test the survival and establishment capacities of three different treatments of robust emergent wetland community species in irrigation ponds.  The goal is to determine the most effective way to assemble emergent communities in terms of composition and structure in order design restoration plants that will improve water and biodiversity in irrigation ponds.  A combinatorial repeated measures design was used to test the survival capacity of five different treatments matrices of Scirpus atrovirens, Carex lacustris,and Sagittaria latifolia at two irrigation ponds in the Niagara Region, Ontario Canada.  Growth trait and survival measurements were taken every two weeks in the growing season of 2011.  A vegetation inventory was taken four times throughout the growing season. The findings suggest that S. latifolia is capable of interspecific competition when planted in mixed plot plantings in semi-naturalized ponds.  S. atrovirens was more competitive in monocultural plantings in a pond community comprised of agricultural weed species.  The findings suggest that it is important to determine the type of community that exists in the ponds prior to planting in order to implement the most appropriate plantings for the pond age and structure and the vegetation community composition.  Fragmentation of habitats has reduced the biodiversity of wetland communities at local scales. Incorporating robust emergent species into irrigation ponds offers a way to improve ecosystem functioning for agricultural landowners while providing species recruitment and colonization sites for fragmented metapopulations of wetland species. 

Keywords: ecological assembly, ecosystem functioning, habitat fragmentation, interspecific competition, intraspecific competition, survival, water quality improvements, wetland vegetation
Introduction
Compared to urban sprawl or industry, globally agriculture is the greatest consumer of lands [1]. In North America agriculture and urban development have reduced the gross area of various ecosystem types; reductions in the number of wetlands and associated wetland species are particularly evident [2]. The estimated loss of wetlands is close to one hundred percent in urban and agricultural areas like Southern Ontario [3].  To some degree the loss of wetlands has been mitigated by the creation of farm ponds which now represent a major wetland habitat type in many farming regions [4].   Constructed wetlands are effective for managing agricultural pesticide runoff [5] and when properly vegetated agricultural ponds can ameliorate water quality issues while also increasing biological diversity [6,7 -8, 5].  If irrigation ponds are acting as an analogue to replace lost wetland ecosystems, how can we actively restore wetland communities in irrigation ponds? The potential for irrigation ponds to connect fragmented wetland communities and restore the biodiversity of these ecosystems is significant.  In order to effectively use irrigation ponds as analogues there are important baseline questions that need to be answered which became the purpose of this pilot study.  First we need to understand the dynamics of irrigation ponds in terms of the chemical loads that may be characteristic of ponds that act as collection basins in agricultural landscapes.  Furthermore, before we assemble ecological communities that are characteristic of wetlands, it is important to study the existing vegetative structure and composition in order to determine the potential effects from interspecific competition. It is also important to determine, what if anything is missing in irrigation ponds that one would find in wetlands of a given area.  This pilot study sought to investigate these questions in order to provide basic but significant information and data on these potential ecological analogues.  
The second part of the pilot used the information gathered from answering those basic questions in order to assemble ecological communities in irrigation ponds towards the dual purpose of improving water quality while also connecting fragmented wetland communities. We felt it was important to look at the potential differences in species assemblages and colonization of ponds of different age structure.  The case study ponds were examined as separate case studies but received exactly the same treatments and were compared only using meta-analytical techniques because age and differences in adjacent land-uses can influence wetland species composition. A study that examined 58 wetlands in South-eastern Ontario determined that wetlands are influenced by adjacent land uses [8].  The studied determined that a key component in the maintenance of diverse wetland communities is related to the protection of propagule sources up to 250 meters away from the wetland (in terms of buffering capacity of anthropogenic disturbances and invasive species, e.g. forests) and that vectors of dispersal are requisite for the provision of seed sources. In particular, robust emergent wetland vegetation is an important yet absent functional community in most irrigation ponds. 
Other community guilds i.e. submersed aquatics colonize restored wetlands more readily and rapidly than other community guilds such as robust emergent [9].  Submersed aquatics are distributed by waterfowl and wetland wildlife through ingestion and expulsion of the seeds. However, many emergent species reproduce more frequently by clonal reproduction thereby limiting the dispersal of this guild to other appropriate habitats [9]. As similar habitats become more isolated from one another, as is the case with the wetland loss in Canada species richness decreases [10].  The closer that restored habitats are to other intact habitats and populations the more opportunity there will be for dispersal, colonization, establishment and persistence of native species. Total species richness was higher for non-isolated islands than for isolated island patches [11].  
Assembling a functionally beneficial robust emergent community is an important basis for the restoration experimentation in this study for three main reasons. First, emergent species provide sites for microbes to attach to which increases the nutrient and contaminant breakdown in the water body and also directly uptake nutrients in through their roots [12, 13]. Microbes remove inorganic nutrients, heavy metals, dissolved organic carbon, particulate matter and suspended solids [14]. Wetlands are useful for the treatment of non-point source (NPS) pollution because they can deal with pulses of pesticides from fields in modern agriculture [15].  
Second, rhizomatic vegetation in particular, provides areas for sedimentation, seed collection and seed germination sites while also leaking oxygen from their rhizomes that stimulates aerobic decomposition of organic matter and the growth of nitrifying bacteria (Brix, 1999).  Oxygen leakage from rhizomes serves to oxidize and detoxify potentially harmful reducing substances in the rhizosphere [6]. 
Third, clonally reproducing vegetation is capable of rapid establishment and can function as a nurse crop for other more sensitive species [16].  Planting two grass species in a recently restored depressional wetland can improve native vegetation cover.  By planting species that reproduce clonally, the wetland increased in native coverage, directly from the planted species and also provided safe sites [17] for other native plants to establish. 
In this study, we examined the establishment, survival and competition of a matrix of three robust emergent vegetation species. Specifically, our aims were: (1) to determine the capacity of four different treatments (combinations of planted species) for survival, and (2) to study the relationship between the planted species and agricultural weeds in terms of competition and coverage over time, (3) comparing management approach of different ponds.  Ultimately, the question is whether mixed plantings of robust emergent species will be more successful in establishing and surviving in irrigation ponds as compared to monoculture plantings of the same species. 
Methods 
Site Selection
The Niagara fruit belt is one of three tender fruit producing areas in Canada that can support large-scale commercial production [18].  This area extends about 25 miles along the southern shore of Lake Ontario (between Grimsby and the Niagara River) and extends seven miles inland from the lakeshore overlapping in some areas with the Niagara Escarpment Biosphere Reserve [18].  Many farmers are located in various riverine systems in the Niagara region and either directly pumping water from the streams onto the fruit, or collect water in ponds to ameliorate water scarcity during the growing season.  
Study sites
Two sites were chosen for in the Niagara region both on privately owned farmland. The first site is located on a tender fruit farm that has been in cultivation for at least fifty years in Virgil Ontario (http://goo.gl/LO2LG). The other site is also on a tender fruit farm located in Winger Ontario (http://goo.gl/iCKMA), adjacent to a field crop farm (in 2009 the field was used for corn, soy beans in 2010, and corn in 2011) and also to a bush lot.  The irrigation pond in Virgil (Pond A) was dug in 1980 by a previous owner and is spring fed. The pond in Winger (Pond B) was dug in 2007 by expanding an existing pond on the property.   A third site a provincially significant wetland, the Four Mile Creek Wetland was used as a local reference site to examine the vegetation composition in comparison to the two study sites.  
Baseline studies in 2010
In 2010 baseline studies were performed at each of the sites to characterize the two study sites in comparison to the reference site.  Water chemistry data was collected at three separate occasions during the growing season from Pond A, Pond B and the reference site to characterize the water quality during the major chemical application times during the agricultural season (May 5th, June 1st, and August 21st ).  In field measurements included water temperature, pH, and conductivity. Grab samples were taken using the protocols established by the Department of Environmental Protection for the State of Maine.  The grab samples were analyzed in the laboratory, analysis included: turbidity, chlorides, nitrates, and total suspended solids.  
Additionally, during the June 1st collection, benthic macroinvertebrates were collected using an adaptation of the protocols used by the Maine Department of Environmental Protection (DEP) which are specifically focused on sampling aquatic macroinvertebrates in freshwater wetlands. Three replicate samples were collected in areas of emergent or aquatic bed vegetation or in other representative habitat areas (replicate samples will be collected in undisturbed locations). The collected samples were preserved in 95% ethyl alcohol for later sorting and taxonomic analysis in the laboratory.  On May 17th six soil samples were collected at Pond A and Pond B (two at each site) to examine the nitrate N and inorganic phosphates at near shore locations. 
The last baseline measurement technique employed in the initial field season was a Wetland Macrophyte Index (WMI) [19]. The sampling of vegetation took place on June 1st (reference site), June 2nd (Pond B) and June 4th (Pond A); all macrophytes were identified to species.
Summary of baseline findings which shaped the experimental design
The soil sample results were insignificant in terms of nutrient levels. The water quality analysis did not reveal any particularly significant changes in water chemistry over the duration of the season. The water quality tests performed were all well within the Provincial standards for recreation, aesthetics and fish and wildlife [20].
The WMI revealed an interesting, albeit not surprising, finding that was consistent with other studies [9].  The irrigation pond in Virgil (Pond A) is devoid of robust emergent wetland species in comparison to the reference site and Pond B which possesses some characteristic wetland species, particularly in the end that was the original pond. Whereas other community guilds like submersed aquatics were present at all three sites because submersed wetland species colonize restored wetlands more readily and rapidly [9].  
Experimental design
For this experiment, plugs were used (as opposed to direct seeding) wetland vegetation direct seeding survival is limited due to the light, heat and water requirements [21].  The planting material was purchased from St Williams Nursery & Ecology Centre: Pterophylla Native Plants and Seeds, Long Point Ontario and transported to both sites in the Niagara Region on the same day (September 22nd 2010). Plant plugs were planted within the plots on September 26th 2010. Monitoring of the plant species were continued biweekly until the end of November 2010. The timing of the plantings was based on the normative seasonal decline in herbicide application on the farms [21].  This was also the time in the growing season when water levels are the lowest in the ponds which will help to protect the plantings because young plants have not yet developed the aerenchymous material necessary for them to survive in anaerobic soils (or standing water) [21]. The water height on the banks provided useful observational data to direct the locations for proper planting for effective establishment [22]. 
Vegetation selection and planting design 
A wide variety of plant species make suitable potential candidates for planting in wetlands, however, the species that were chosen for planting were Scirpus atrovirens Willd., Carex lacustris Willd., Sagittaria latifolia Willd. For consistency sake one of the reasons these species were chosen was because they were absent from the species inventory at both study locations which reduced possible contamination from local seed sources or the rhizomatic spread of the species within the ponds [23] with the potential for considerable vegetative coverage [16].  Other additional considerations included access to planting materials (including the capacity for the nursery to germinate certain species) and precedence from screening tests [24, 25, 15]. The three species were chosen because they reproduce clonally, two of them (Carex lacustris and Scirpus atrovirens) are important contributors to above ground biomass with considerable above ground canopy growth.  Sagittaria latifolia can also reach heights of 80cm providing it a competitive edge for light requirements [26].  All three species are tolerant of variable soil type with the capacity to adapt to degraded wetland habitats [26]. 
We sought to test the effects that the three treatments had on growth, regeneration, abundance of planted species as well as species diversity and abundance and community composition (wetland species guilds, natives versus non-native) at the two different ponds.  Of interest was the potential variability of species competition and establishment between the two different age structures [27, 28].  A combinatorial approach to planting was chosen because the experiment�s purpose was to test the capacity for establishment of the three species (Scirpus atrovirens, Carex lacustris, Sagittaria latifolia) in terms of intraspecific competition (monocultures of each species), interspecific competition (the polyculture plots included one plug of each species) as well as the influence that the monocultures and polycultures have on other naturally occurring species in the robust emergent wetland vegetation zone [29].  
In total 156 plots of 50cm2 plots were established (seventy-eight at each of the ponds). Sixty of the plots received one of three combinatorial planting treatments; eighteen plots at each site were left alone and used as a control.  On August 23rd, 2010 a plot-based vegetation inventory was taken prior to hand-weeding during site preparation for planting to outline the characteristic community members [22].  The treatment plots were not weeded and received no plants. The planting design used was based on four treatments and the pattern is consistent around the perimeter of each pond.  The planting density for each 50cm2 plot is three plugs per plot. 
The five planting treatments created at each site are as follows:
1) Mixed Plot Treatment � includes one plant each of Sagittaria latifolia, Scirpus atrovirens and Carex lacustris. 
2) Monoculture: Sagittaria latifolia � Each plot contained three plant plugs of S. latifolia (plugs were planted approximately 5 cm from one another). 
3) Monoculture: Scirpus atrovirens � Each plot contained three plant plugs of S. atrovirens (plugs were planted approximately 5 cm from one another). 
4) Monoculture: Carex lacustris � Each plot contained three plant plugs of C. lacustris (plugs were planted approximately 5 cm from one another).  
5) Treatment Monitor � No plant plugs were planted in these plots
This planting pattern is repeated twelve times at each site location, resulting in a total of 15 replicates of each treatment at each site. In total 180 plugs were planted at each site, 60 of each species.  
Variables measured
Two types of sampling took place during the growing season of 2011.  A monthly vegetation inventory was executed in May, June, July and August to gather information on plant abundance and diversity. During these inventories the maximum height of the canopy with the associated species was recorded, in addition to the percent cover of plants, water, rocks, detritus, bryophytes, and soil for each plot (including the control plots).  The second set of measurements that were executed every two weeks beginning when the planted species were first visible (June 13th) and are classified as survival and establishment measurements [31, 32].  
The variables measured included presence and absence of the planted species, diameter of the stem at the base of the plant (three measurements were taken and averaged) using a digital caliper, diameter of the entire canopy (taken in two measurements as North to South and East to West using a ruler), maximum plant height, leaf length (three measurements taken and averaged) and leaf width (taken as three measurements and averaged), the total number of leaves per plot (this includes the mother plant and the clones that establish from that original plant), and the total number of ramets per mother plant (i.e. the number of clones that occur).  For the plants that survived the first winter, these measurements were taken seven times during the growing season in 2011 at two week intervals.
Statistical analysis
The purpose of the analysis was to examine the interaction of time and treatments, which was why repeated measures design was suitable for statistical analysis.  A repeated measure, or within-subject design, studies the same subjects over time to assess the influence of change temporally.  Repeated measures design reduces the variance of estimates of treatment-effects which allows for statistical inferences to be made using fewer subjects [32].
            The data was analyzed using the program SYSTAT 12�. Following the Shapiro-Wilks test for normality, it was determined that the survivorship data sets were normally distributed.  Since the data from this study was normally distributed, it has multiple response variables and it violates the rule of compound symmetry and sphericity the Repeated Measures MANOVA test was used to analyze the data. Six treatments were considered during the analysis: Treatment One: Sagittaria latifolia in the mixed plot, Treatment Two: Scirpus atrovirens in the mixed plot, Treatment Three: Carex lacustris in the mixed plot, Treatment Four: Sagittaria latifolia monoculture planting, Treatment Five: Scirpus atrovirens monoculture planting, and Treatment Six: Carex lacustris monoculture planting.  
Relevant to the MANOVAR model, the measures were repeated over time on a per-plot basis, and interactions were expected to occur on an individual basis.  However, the response variables were expected to be non-independent as well. For all the repeated measures, there was one within-subjects factor (time, measured in weeks; measurements were taken every two weeks from June until September).  Seven sets of measurements were collected and recorded.  There were six between-subjects factors for the survivorship analysis (six different treatments) and five between-subjects factors for the vegetation inventory analysis (five different treatments). 
Multivariate and univariate F values are representative of the degree of difference in the dependent variable created by the independent variable; in this study the F value refers to the influence that treatment and/or time have on the response variables (e.g. plant height, number of ramets, stem diameter). 
The Pillai�s Trace Test was used to assess the statistical significance between the groups of independent variables.  Data are reported using F, p, and Pillai�s trace because Pillai�s trace is the actual test for significant differences of the repeated factor of time (and time X treatment) within subjects [30]. 
Results
Survivorship analysis
Pond A: Virgil, Ontario
The means for all of the survival traits measured the means Treatment Five, the monoculture of Scripus atrovirens differed significantly as compared to the other treatments. The second most successful treatment consistently for each measured trait was Treatment Two, which refers to the Scirpus atrovirens in the mixed plot. For Mean Leaf Surface Area, Treatment One, Sagittaria latifolia in the mixed plot was not different than Treatment Two (Scirpus atrovirens in the mixed plot). 
[INSERT TABLE 1 HERE]
Pond B: Winger Ontario
Of the means for each of the survival traits measured, Treatment One, S. latifolia in a mixed plot, differed significantly than of the other treatments including the monoculture planting of S. latifolia; although S. latifolia as a monoculture was second to Sagittaria latifolia in the mixed plot.  Carex lacustris had a zero percent survival rate after winter at this pond.  
[INSERT TABLE 2 HERE]
Pond A compared to Pond B
Flowering only occurred at Pond B for one species; Sagittaria latifolia.  The measurements taken for the flowers included number of bolts, mean flower height, and mean number of flowers.  For the traits pertaining to flowering, the means of Treatment One, S. latifolia in a mixed plot, differed significantly than all of the other treatments. Since S. latifolia was the only species that flowered, Treatment Four (the monoculture of S. latifolia) was second in trait means. 
[INSERT FIG. 1 HERE]
[INSERT FIG. 2 HERE]
[INSERT FIG. 3 HERE]
[INSERT FIG. 4 HERE]
[INSERT FIG. 5 HERE]
Vegetation inventory 	
The vegetation inventory data was analysed using five specific treatments. Treatment One: is a mixed plot (one plant each of S. latifolia, S. atrovirens and C. lacustris); Treatment Two: S. latifolia monoculture; Treatment Three: S. atrovirens monoculture; Treatment Four: C. lacustris monoculture; and last, Treatment Five: a control (no plants were planted). The analysis was concerned with examining whether or not time and/or treatment influences the seven particular response variables.  The response variables are: total number of species, number of native species, number of exotic species, canopy height, percent cover of the treatment, percent cover of native species and percent cover of non-native species. 
Pond A
The analyses revealed that Treatment One (the mixed plot treatment) was consistently the most influential treatment for all seven response variables over time for Pond A.  Treatment Three was the next most influential treatment on the response variables over time. This is consistent with the survivorship analyses where Treatment Three was the most influential Treatment for survivorship. 
[INSERT TABLE 3 HERE]
Pond B 
The analyses revealed that Treatment One (the mixed plot treatment) was consistently the most influential treatment for all seven response variables over time for Pond B.  Treatment Two was the next most influential treatment on the response variables over time. This is consistent with the survivorship analyses where S. latifolia was most successful in the mixed plot planting. Treatment Two was the second most influential treatment on the seven response variables. This is also consistent with the survivorship findings where S. latifolia in a monoculture was the second most influential treatment for survivorship in Pond B. 
[INSERT TABLE 4 HERE]
[INSERT FIG. 6 HERE]
[INSERT FIG. 7 HERE]
[INSERT FIG. 8 HERE]
Discussion
For ecological restoration on agricultural lands, the results suggest that planting plugs of S. latifolia (at a spacing of 50 cm) is an effective strategy for establishing a dominant plant community in semi-naturalized irrigation ponds.  The findings suggest that S. latifolia is better suited to competing with existing agricultural weeds and wetland species (interspecific competition) rather than intraspecific competition. S. latifolia provides important benefits to organisms dependent on wetlands for food including waterfowl, muskrats, porcupines and habitat including usage as cover and for fish and macroinvertebrates.
For ecological restoration of irrigation ponds on agricultural lands devoid of facultative wetland species planting S. atrovirens in monocultures (at a density of three plugs in a fifty by fifty centimetre arranagement) is a good strategy in establishing a dominant emergent vegetation community presumably capable of managing field run-off [15].  Intraspecific competition is not as of much an issue for S. atrovirens� survival and dominance; rather interspecific competition is improved with at least three plants per fifty cm2 plot.  
At both sites the survivorship of C. lacustris was limited. Although plugs were used for planting, this research may confirm findings from Yetka and Galatoswitsch which suggest that   rhizome planting is most successful if executed in the spring [33] .  More likely, however, is that the low seasonal water levels influenced survival.  Budelsky and Galatowitsch observed that seedling survival of Carex lacustris was highly dependent on water levels during the first growing season (precipitation was extremely low during the growing of 2011 which required the landowners to irrigate regularly [34].  Percent cover from the vegetation surveys indicate that by June some plots at both sites were entirely dry and by July all of them were devoid of contact with water). 
Our study has also determined that mixed plots of facultative wetland species, in this study, S. atrovirens, S. latifolia, and C. lacustris is the most effective way to influence the native and exotic species community composition of agricultural irrigation ponds.  Land managers must clearly identify their particular goal for restoration to determine the most appropriate type of planting to execute.  S. atrovirens in a monoculture differs significantly in its capability to establish and dominate the planted plots adjacent to weedy agricultural fields.  Therefore, a monoculture planting of this species may be preferable for some land managers if the goal is to create a dominate community capable of managing agricultural pollutants. If the goal is to establish a more biodiverse and �natural� ecosystem in the irrigation pond, planting a mixed plot would help to influence the community by increasing the total number of species colonizing each plot.
The vegetation survey indicated many more agricultural weeds in the plots in Pond A.  It may be more difficult for native introduced plants to establish into the communities of highly resistant agricultural weeds. For the duration of the vegetation inventory, Pond B supported thirteen facultative wetlands species whereas Pond A supported only four before planting.  
Conclusions
Although this study was not focused on the spatial linkages between scales explicitly, there is an implied recognition that improving the delivery of ecosystem services at the farm-scale has implications for enhancing biodiversity at the regional scale.  The question then becomes one of trying to determine how best to introduce the suitable species.  One major component of this study looked at the importance of functional dominance in introducing native species to irrigation ponds. 
The findings suggest that community establishment and dominance is influenced by the pre-existing vegetative community, the density of the planted species, and the capacity for clonal reproduction.  The species chosen for the study have the ability to reproduce clonally which offers them a chance to maximize their spread and coverage within the first growing season.  
Due to the potential constraints of competing with the pre-existing community (interspecific) and within species (intraspecific) this research demonstrates that landowners/ land managers looking to improve water quality and habitat in irrigation ponds need to investigate species exist in/around the community prior to plant and density selection. Additionally one needs to consider the age of the pond and think about other challenges (e.g. habitat fragmentation, species dispersal/ isolation, deer densities, deer densities and existing community) to maximize the success of plant survival and dominance.  This study is the initial phase in determining how best to assemble ecological communities in irrigation ponds; based on these findings a full-scale restoration study should first determine whether the site is well-suited to monoculture or polyculture plantings.  The next phase of this study should look at incorporating more species particularly rarer or more sensitive species that are largely absent from regional landscapes. 
Acknowledgments 
The authors would like to thank Jennifer Baldson, Simon Green and Virginia McGrath for their dedicated work in the field. The authors are also grateful for the funding through the Ontario Graduate Scholarship provided by the Ontario Ministry of Training, Colleges, and Universities.
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 Darby M. McGrath; E-mail: dmcgrath@uwaterloo.ca
 When establishing the sampling protocols for the field and laboratory components of the wetland analysis many sources were consulted, however, the EPA for the State of Maine offers one of the most comprehensive and well-researched manuals for sampling and analysis.









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