ECOLOGICAL INDICATORS OF RESTORATION SUCCESS: FISH COMMUNITY DISTRIBUTION, COMPOSITION, AND SAMPLING STRATEGIES WITHIN THE PICAYUNE STRAND RESTORATION PROJECT
A Thesis Presented to
The Faculty of the College of Arts and Sciences
Florida Gulf Coast University
In Partial Fulfillment
Of the Requirement for the Degree of
Master of Science in Environmental Science
By
Ryan C. Young
May 2013 ii
APPROVAL SHEET
This thesis is submitted in partial fulfillment of
the requirements for the degree of
Masters of Science
______
Ryan C. Young
Approved: May 2013
______
Edwin M. Everham, III, Ph.D. Committee Chair
______
David W. Ceilley, M.S. Committee Co-chair
______
Michael Duever, Ph.D. Committee Member
The final copy of this thesis has been examined by the signatories, and we find that both the content and the form meet acceptable presentation standards of scholarly work in the above mentioned discipline. iii
ACKNOWLEDGEMENTS
I would like to genuinely thank the Florida Fish and Wildlife Service for granting me access and the necessary permits to conduct this research; the Florida Gulf Coast University and the Office of Research and Sponsored Programs for providing funding to conduct this study; the Inland Ecology Research Group for providing sampling equipment and guidance; my professors who provided me with the core knowledge necessary to understand complex concepts and give me the background knowledge necessary to have a fulfilling graduate experience; Jennifer Nelson for her encouragement; my faithful volunteers including Geoffrey Rosenaw and Garrett Coe who gave their time and energy to help make my field sampling possible; and my loving family Shane, Donna, and Steve Young who came to my rescue whenever I needed assistance, provided endless support and encouragement, and stood behind me every step of the way. This project would not have been possible without these generous aids, tremendous support, and continuous encouragement.
I would also like to express my sincere appreciation and gratitude to my committee chair Dr. Edwin Everham with Florida Gulf Coast University for always giving me the push I needed, the wise words of advice that helped to organize and bring together my thoughts and focus, teaching me to think like a scientist, and sharing with me his passion and love for making a difference in the world; my co-chair David Ceilley with Florida Gulf Coast University for teaching me about experimental design and methodology, helping me to identify my fish samples, teaching me about the Picayune, helping me understand the most current scientific data on inland freshwater fish research in Florida, showing me how to analyze the data I collected, and how fun data analysis can be by letting the information tell a story about the natural world; and Dr. Michael Duever with the South Florida Water Management District for assisting in project design and analysis, critically reviewing my text, sharing his expertise about the area of the Picayune and the great stories about his time in the beautiful area, and providing guidance and encouragement through the project. Their edifying assistance and research experience was the backbone of this study
My heartfelt thanks and endless appreciation goes out to everyone who helped me to complete this project because I could not have done it alone!!
iv
ABSTRACT
Increasing awareness of the damage inflicted upon natural systems by human beings has brought environmental and ecological restoration to the forefront of environmental research and monitoring efforts of the 21st century. Florida leads the country with some of the largest restoration and monitoring projects in our nation’s history. This study was designed to evaluate the success of restoration activities within the Picayune Strand Restoration Project, part of one of the world’s largest restoration efforts the Comprehensive Everglades Restoration Plan. This study was conducted by collecting and analyzing data on fish community structure, species diversity, and species abundance in relation to various restoration phases. These restoration phases included two treatments (impacted unrestored areas, transitional recently restored areas), and reference wetlands (non-impacted natural wetland). Fish community data was collected monthly through the period of inundation with passive sampling using Breder traps as well as active dip net sampling. Based on the abundance and diversity data, results indicated that species richness, abundance, and diversity was lowest in impacted areas, increased in transitional recently restored areas, and was highest in both abundance and diversity in natural reference areas. Fish community data also indicated distinct groupings and similarities within each restoration phase and indicated varied species distribution among sites of different restoration phases. This analysis confirmed that fish community assemblages differed significantly among all three restoration treatments. Several indicator species were identified including Gambusia holbrooki, Jordonella floridae, and Fundulus confluentus which helped to drive the dissimilarity between different phases of restoration. In addition, the majority of species captured were only found in reference wetlands. These findings serve as an indicator that the restoration activities in the Picayune Stand are effective, and that several fish species may be used as indicators of hydrologic restoration success in ephemeral wetlands of Southwest Florida. Further analysis v was conducted to observe patterns in sampling effort and temporal changes in community structure in order to determine the sampling frequency required to obtain a robust signal, the time of year most appropriate for collecting samples of a mature fish community, and patterns of dispersion over multivariate space through the period of inundation. Based on this one-year study, community data suggested that the months of October and November provided the best examples of a mature fish community and that sampling at a frequency of every third month
(September, December, and March) provided sufficient community data to obtain a robust signal. These findings serve as indication that a sampling frequency of every second month is required to obtain the information necessary to make informed decisions about restoration activities, and that the optimal time period for sampling a mature fish community occurs during the months of October and November.
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TABLE OF CONTENTS
Page
Approval Sheet……………………………………………………………………………………ii
Acknowledgements………………………………………………………………………………iii
Abstract…………………………………………………………………………………………...iv
Table of Contents…………………………………………………………………………………vi
List of Figures ...... vii
List of Tables ...... xi
Introduction………………………………………………………………………………………..1 The Florida Everglades……………………………………………………………………1 Study Site – The Picayune Strand Restoration Project…………………………………...3 Ecological Indicators…………………………………………………………………….10 Fish as Indicators………………………………………………………………………...13 Fish Sampling Methods………………………………………………………………….16
Objectives………………………………………………………………………………………..18
Hypothesis……………………………………………………………………………………….19
Methods………………………………………………………………………………………….20 Experimental Design…………………………………………………………………….20 Sample Regions………………………………………………………………………….20 Deep Cypress…………………………………………………………………….21 Cypress Gramminoid…………………………………………………………….21 Gramminoid……………………………………………………………………...22 Restoration Phases……………………………………………………………………….22 Miller Canal……………………………………………………………………...22 Prairie Canal……………………………………………………………………...22 Fakahatchee Strand………………………………………………………………23 Survey Techniques……………………………………………………………………….23 Breder Traps……………………………………………………………………..24 Dip-Netting………………………………………………………………………24 Data Analysis……………………………………………………………………………25
Results……………………………………………………………………………………………28 Fish Community Data…...……………………………………………………………….29 Species Richness…………………………………………………………………30 Relative Abundance……………………………………………………………...33 Potential Indicator Species…………………………………………..…………...38 vii
TABLE OF CONTENTS (Continued) Page
Sampling Frequencies……………………………………………………………………42 Temporal Community Structure…………………………………………………44 Fish Community Structure………………………………………………………….……46 Temporal Dispersion ...... 49 Restoration Success ...... 51
Discussion ...... 53 Potential Indicator Species ...... 59 Sampling Frequency ...... 60 Temporal Sampling ...... 61 Fish Community Structure ...... 61 Temporal Dispersion ...... 62 Restoration Success ...... 63 Future Research Opportunities ...... 65
Conclusions ...... 68
References Cited ...... 69
Appendix A – Fish Sampling Data Form ...... A-1
Appendix B – Fish Sampling Data ...... B-1
Appendix C – Total Fish Abundance...... C-1
Appendix D – Restoration Phases: Cluster Analysis and MDS Ordination ...... D-1
viii
LIST OF FIGURES Page Figure 1. Picayune Strand Restoration Project location map………………………………………..3
Figure 2. Picayune Strand Restoration Project with adjacent restoration lands……………………..3
Figure 3. Aerial photograph of the Picayune Strand Restoration Project boundary………...... 4
Figure 4. Originally constructed roadway network………………………………………...... 5
Figure 5. Plans for the rehydration of the Picayune Strand Restoration Project…………...... 6
Figure 6. Study area with fish sampling locations…………………………………………………20
Figure 7. Average species richness and relative abundance of fishes within each samplable site………………………………………………………………………………………..32
Figure 8. Average diversity metrics including Margalef’s (d), Pielou’s indices (J’), Shannon (H’(loge)), and Simpson (1-Lambda’) for each of the represented sampling sites…...... 33
Figure 9. Percent composition of species documented over the entire study………………………35
Figure 10. Cluster analysis with sample sites representing habitat type and restoration phase using Bray-Curtis similarity from the entire sampling period………………………..….37
Figure 11. MDS ordination with sample sites represented by habitat type and restoration phase using Bray-Curtis similarity from the entire sampling period……...... 38
Figure 12. MDS ordination with superimposed raw abundance data for Gambusia holbrooki…………………………………………………………………………………40
Figure 13. MDS ordination with superimposed raw abundance data for Fundulus confluentus……40
Figure 14. MDS ordination with superimposed raw abundance data for Jordonella floridae……....41
Figure 15. MDS ordination with superimposed raw abundance data for Lepomis sp…………………41
Figure 16. MDS ordination with superimposed raw abundance data for Poecilia latipinna………..42
Figure 17. MDS ordination including composite fish community data from the entire sampling period including the months of September, October, November, December, January, February, and March…………………………………………………………………..…43
Figure 18. MDS ordination including composite fish community data from every other month of sampling including the months of September, November, January, and March…………………………………………………………………………….……...43
Figure 19. MDS ordination including composite fish community data from every third month of sampling including the months of September, December, and March……….44 ix
LIST OF FIGURES (Continued)
Page
Figure 20. MDS ordination including fish community data from the month of October at all samplable sites…………………………………………………………………………...45
Figure 21. MDS ordination including fish community data from the month of November at all samplable sites…………………………………………………………………………...45
Figure 22. MDS ordination representing reference sites of various habitat types sampled each month during the entire sampling period with superimposed relative abundance data for Gambusia holbrooki for each represented month when sampling occurred……………………………………………………………………….47
Figure 23. MDS ordination representing reference sites of various habitat types sampled each month during the entire sampling period with superimposed relative abundance data for Jordonella floridae for each represented month when sampling occurred………………………………………………………………………..48
Figure 24. MDS ordination representing reference sites of various habitat types sampled each month during the entire sampling period with superimposed relative abundance data for Fudulus confluentus for each represented month when sampling occurred……………………………………………………………………….48
Figure 25. MDS ordination of monthly composite fish community data in transitional sites with superimposed trajectory…………………………………………………….....49
Figure 26. MDS ordination of monthly composite fish community data in reference sites with superimposed trajectory……………………………………………………...50
Figure 27. Cluster analysis of Bray-Curtis similarity representing composite fish community data for sampling sites of various restoration phases and habitat types through the entire sampling period……………………………………….51
Figure 28. MDS ordination of composite fish community data for each sampling site representing various restoration phases and habitat types through the entire sampling period using Bray-Curtis similarity from all sampling events……………….52
Figure 29. MDS ordination of composite fish community including superimposed similarity groupings of 40 percent, 50 percent, and 60 percent similarity………….…52
x
LIST OF FIGURES (Continued)
Page
Figure 30. Habitat maps of the Picayune Strand from 1940 and 1995……………………………...57
Figure 31. Principal component analysis including an MDS ordination of all sampling events with superimposed environmental factors including dissolved oxygen, average depth, average conductivity, salinity, and temperature…………………………………………………………………………..58
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LIST OF TABLES
Page
Table 1. Site descriptions with locations and adjacent wells……………………………………...28
Table 2. Presence / absence of water in each sampling site per month…………………………...29
Table 3. Fish species documented via Breder traps and dip-net sampling by scientific name, common name, and species code abbreviations………………….....29
Table 4. Fish species documented in each restoration phase listed by scientific name………..….30
Table 5. Univariate diversity indices including species richness (R) and relative abundance (A), Margalef’s (d), Pelou’s indices (J’), Shannon-Weaver index (H’(loge), and Simpson’s diversity (1-Lambda’) for fish species documented…….…....31
Table 6. Total relative abundance values per restoration phase, total abundance for all species, and total percent composition of fish species documented during the sampling period………………………………………………....34
Table 7. SIMPER analysis of fish species and restoration phases………………………………..39
Table 8 Relative abundance data for Gambusia holbrooki, Jordonella floridae, and Fundulus confluentus over the entire sampling period…………………………………………………………………………………….46
Table 9. Dissimilarity values between consecutive months within various restoration phases and average dissimilarity for each restoration phase through the entire sampling period…………………………………………………………….….50
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INTRODUCTION
The Florida Everglades
The Everglades is a vast wetland comprised of a variety of habitat types including saw grass marshes, wet prairies, sloughs, ponds, tree islands, and mangrove estuaries that dominate
Southern Florida and are characterized by habitat heterogeneity, large spatial extent, and a distinctive hydrologic regime (Dixon 2009; Chimney & Goforth 2006; Davis 1943a,b; Loveless
1959, McCally 1999, Whitney 2004).
Attempts to drain the Everglades for human uses began in the 1850s with the Swamp and
Overflowed Lands Act which allowed for large scale dredging and channelization operations, destroying five million acres of wetlands and creating over 400 miles of drainage canals (Florida
Conservation Foundation (FCF) 1993). Shifting attitudes led to the protection of these wetland systems with the Wetland Protection Act of 1984, which helped to preserve these wetland swamp systems that still cover about 10% of Florida’s land area today (Myers & Ewel 1990).
Nonetheless, this area has also been severely impacted by anthropogenic activities and is severely threatened if it is not protected and restored (COE & SFWMD 1999).
In an effort to protect and restore this landscape, the SFWMD and COE developed a plan titled “Central and Southern Florida Project Comprehensive Review Study Final Integrated
Feasibility Report and Programmatic Environmental Impact Statement” in 1999. The plan, also referred to as the Comprehensive Everglades Restoration Plan (CERP), was designed to create and restore wetlands and reservoirs to increase water storage and water supply (COE & SFWMD
2001). CERP is a multi-billion dollar federal-state partnership that seeks to restore historical hydrologic conditions of the Everglades and encompasses over 200 components addressing water management and ecosystem restoration needs over the southern third of peninsular Florida
(Doren et al. 2009). It was created as, “a framework for modifications and operational changes to -2- the Central and Southern Florida Project that are needed to restore, preserve, and protect the
South Florida ecosystem while providing for other water-related needs of the region, including water supply and flood protection” (COE & SFWMD 2001 p. 1-2). The plan was reviewed and finally approved through the Water Resource Development Act on December 11, 2000. The new plan gave the opportunity to reverse the course of declining health of the Everglades system and leave the irreplaceable system as a legacy for generations to come (COE & SFWMD 1999 p. i).
This document provided a tool to help sustain the environment, economy, and social well-being of South Florida (COE & SFWMD 2004).
Increasing awareness of the damage inflicted upon natural systems by human beings has brought environmental and ecological restoration to the forefront of environmental research and monitoring efforts of the 21st century. The Society for Ecological Restoration International defines ecological restoration as, “the process of assisting the recovery of an ecosystem that has been degraded, damaged, or destroyed.” They also describe the process as an, “activity that initiates or accelerates the recovery of an ecosystem with respect to its health, integrity and sustainability.” (Society for Ecological Restoration International Science & Policy Working
Group 2004.) Restoration projects often require long term commitments by participating parties.
Monitoring is a vital facet in the restoration process as it allows us to assess the progress and effectiveness of restoration efforts, and helps inform the discipline on the relative merits of different approaches to restoration.
Acting as the vanguard for this movement of ecological restoration efforts, Florida leads the country with some of our nation’s largest restoration and monitoring projects. Often noted as the world’s largest restoration effort, the Comprehensive Everglades Restoration Plan (CERP) sets many complex guidelines for evaluating the success of the restoration efforts involving hydrology, water quality analysis, and a variety of other assessment models. The overall CERP -3- project implementation will take over 20 years but, monitoring and evaluation efforts will extend long beyond the project’s completion, well into the future.
Study Site - The Picayune Strand Restoration Project
The Picayune Strand Restoration Project is one of the largest individual CERP projects and as part of the Governor’s ACCELER-8 program in 2004, the first CERP wetland restoration project. Formerly intended for development as a large-scale residential area called the Southern
Golden Gate Estates, Picayune encompasses approximately 85 square miles and is located immediately south of Interstate 75 and extends to US 41 in western Collier County (COE &
SFWMD 2004). Picayune Strand was historically characterized by seasonal flooding and slow- moving sheet flow prior to efforts to develop the area.
Figure 2. This map shows the location of the Picayune Figure 1. This map shows the Picayune Strand Restoration Project Strand Restoration Project in the state of Florida, Collier with adjacent conservation lands including Belle Meade, the County, and the immediate areas surrounding the project Florida Panther National Wildlife Refuge, Fakahatchee Strand (Dixon 2009). Preserve State Park, the Ten Thousand Islands National Wildlife Refuge, Collier-Seminole State Park, Deltona Lands, and the Rookery Bay National Estuarine Research Reserve (Dixon 2009). -4-
These characteristics supported a
typical Southwest Florida wetland
landscape which consisted of cypress
swamps and sloughs, wet prairies,
hardwood hammocks, marshes, pine
and cabbage palm flatwoods, and tidal
areas (Duever & SFWMD, pre-
development vegetation map). The
area is open to the public for various
recreational activities including
hunting, fishing, camping, horseback
riding, hiking, photography, and
wildlife observations and sits in an area
surrounded by additional conservation
Figure 3. This map shows an arial photograph with the Picayune Strand lands including the Fakahatchee Strand Restoration Project boundary. Also visible in the map are the system of 290 miles of road and canal system built during the lands intended Preserve State Park on the east, Belle development (Dixon 2009). Meade to the west. As a result, sheet flow was eliminated, the water table dropped more than 1 meter in some areas, hydroperiod was reduced by 2 to 4 months, drainage rate increased by 16 times, the frequency and intensity of forest fires increased, and the area was invaded by exotic vegetation and other invasive non-native species (SFWMD 2008a). As the water was channelized in the late 1960s by an extensive 48 mile canal system, the overall hydrology of watershed was altered by the discharge of approximately 18 inches of run off annually and the interception of shallow groundwater outflow. This canal system has continually lowered the water table and can still be seen today in the areal image provided in Figure 3. This over- -5-
drainage of the area also resulted in large point source
freshwater discharge into the estuarine system of the 10,000
islands. The decline of ecological function including the
degradation of upland, wetland, and estuarine plant
communities, reduced abundance of native fish, wildlife,
and estuarine shellfish populations, and reduced aquifer
recharge showed even more evidence of the severe negative
impacts that this project had on the natural system. In
Figure 4. This map shows the originally addition to the over-drainage by the canal system, 290 constructed roadway network including 290 miles of paved roads (Dixon 2009). miles of roads were constructed in a grid system throughout the property (Figure 4). The roads were graded and sat approximately 15 centimeters above the surrounding ground elevation, acting as barricades to sheet flow and redirecting any standing water into the nearby canals. Although these modifications severely lowered water levels, they failed to prevent the property from holding water during the wet season, causing the development project to fail (COE & SFWMD 2004, U.S. Department of the Interior (DOI)
2005).
After the company that owned the Southern Golden Gate Estates, the Gulf American
Corporation, went bankrupt concerns over the degradation of environmental quality and water supply potential of the region began to arise. This prompted the State of Florida to include
Picayune Strand in the Save Our Everglades program in 1985 and direct the South Florida Water
Management District (SFWMD) to develop a hydrologic restoration plan to reduce over- drainage, restore historic sheetflow patterns while maintaining the existing levels of flood protection for areas in the north, and enhance the areas environmental value and water resources.
Re-attaining pre-development conditions was the main goal of the proposed restoration project -6-
(COE & SFWMD 2004, Duever 2005) and restoration of the native fish communities was
determined to be an important component of the restoration project (Ceilley 2008).
Prior to the onset of CERP, the need for restoration of this area was recognized. A jointly
funded plan by the SFWMD and the COE was developed and land acquisition of more than
20,000 parcels began with funding assistance from the Department of the Interior in 1984. The
effort extended for over 20 years, continuing through 2005. Restoration efforts began in 2003
after the implementation of CERP,
beginning with the installation of
culverts under US-41 and the
plugging of the northern portion of
Prairie Canal, which lies on the
northeastern most side of the
property. Restoration efforts
continued for four years as the
SFWMD finished the plugging of
Prairie Canal, removed 65 miles of
roads east of Merritt Canal,
demolished 160 structures, and
Figure 5. This map shows the plans for the intended areas of rehydration in conducted soil remediation on the Picayune Strand Restoration Project including canal plugs, road removal, pump station installations, and existing weirs (COE & SFWMD approximately 28 acres of 2004). contaminated soil. Activities halted in 2008 due to lack of funds as a result of the economic
downturn, but have since resumed. The remainder of the project includes continued road
removal to the west of Merritt Canal, plugging of the remaining three canals, and completing the
installation of three pumping stations as well as spreader canals (COE & SFWMD 2004, Duever -7-
2005). Land management activities are being conducted by the Florida Division of Forestry
(DOF), and other cooperating agencies include the U.S. Fish and Wildlife Service, U.S.
Environmental Protection Agency, the National Park Service, and the Florida Department of
Environmental Protection (Dixon 2010, COE & SFWMD 2004).
One of the key restoration components of this project is the plugging of the 48 ± mile canal system. In an effort to restore a more natural hydrologic regime in the area, spoil from the removed roads are being used to plug the canal system. This backfilling technique has been used to return areas to a more natural hydrologic regime and has great potential to improve aquatic wildlife habitat and the area’s hydrologic connectivity. Backfilling is a reasonably simple and short term management strategy which requires simple equipment and no on-site maintenance that has been proven to demonstrate stability over decades (Turner et al 1994). Despite the fact that Prairie Canal was not entirely backfilled, leaving pools of surface water between plugs, the plugs help to prevent water from being released directly into downstream estuaries. The
SFWMD stated that once the water is no longer being conveyed that the canals would no longer over-drain the landscape and surface aquifer resulting in a longer period of inundation for the surrounding area (COE & SFWMD 2004 p. vi). Due to monitoring efforts, SFWMD has already began to document higher water levels near the filled Prairie Canal compared to areas near the unrestored Merritt Canal during the winter of 2006-2007 (Dixon 2010, Ceilley 2008, Chuirazzi
& Duever 2008).
Plugging of the canals in the Picayune Strand, along with the grading of the road system, removal of existing structures, and soil remediation are expected to have significant benefits on the area, as well as adjacent lands, helping to achieve the overall Everglades restoration goal.
“Expected project benefits include restoration of historic wetland communities, restoration of sheetflow towards the coastal estuaries, reduction of harmful surge flows through the Faka Union -8-
Canal into Faka Union Bay, improved freshwater overland flow and seepage into other bays of the Ten Thousand Islands region, improved aquifer recharge, decreased frequency of forest fires, improved habitat for fish and wildlife and threatened and endangered species, reduced invasion of exotic species, and increased spatial extent of wetlands,” (Dixon 2009; Ceilley 2008; COE &
SFWMD 2004; Chuiarazzi & Duever 2008 p.3). These hydrologic improvements are is expected show very positive responses from water dependent animals by increasing their numbers and returning natural distribution patterns once the project is complete (Dixon 2009;
COE & SFWMD 1999).
Although the outcomes of the canal backfilling are expected to be positive, it should be noted that negative impacts of canal backfilling have been documented in the past. A study in the Upper St. John’s River Basin, Florida in 1986 stated that while the plugs slowed canal flow and re-hydrated the marsh, it caused shorter intensity drying events in upstream locations, causing undesirable shifts to flood-tolerant vegetation, and converted areas of marsh to open- water (Chandy, Miller, & Morris 2001). Although some negative impacts of canal plugs have been found in the past, the main objective of the Picayune Strand Restoration Project is to restore the historical hydrology and ecology of the area and surrounding areas. These positive impacts were documented by Baustian and Turner (2006), stating that backfilling restores local hydrologic conditions, creates marsh, and improved ecological processes over a period of 20 years (Baustian & Turner 2006). There may be some negative impacts associated with the canal plugs in the Picayune Strand, but the resulting hydrologic restoration is expected to have a significantly positive impact on the ecological and hydrologic regimes that are currently found there and in the surrounding areas.
Prior to the onset of these restoration efforts, a baseline study was conducted to evaluate the status of Picayune’s habitats and eventually evaluate environmental changes after the PSRP -9- has been completed. The baseline assessment study established reference sites in cypress, wet prairie, hydric pine, and brackish marsh habitats in the Florida Panther and Ten Thousand Islands
National Wildlife Refuges and in Fakahatchee Strand State Preserve and compared them to similar, but impacted wetlands of PSRP (Ceilley 2008, Duever 2005). Aquatic faunal samples were collected from 42 locations six times from August 2005 to February 2007.
For the baseline assessment, fish communities were surveyed at a frequency of three times per year at each location. Wetland ecologists at Florida Gulf Coast University (FGCU) identified a total of 6,381 individual fish collected using Breder traps which consisted of 25 species including nine families, two non-native families, and two non-native species. Eastern mosquito fish was the most abundant species present being found at every sample site. The reference sites had a 58% similarity of fish community structure with seven species contributing to >93% of the similarity, while impacted sites showed an average of 45% similarity with only five species contributing to >93% of the similarity. Reference wetlands showed a higher diversity in fish communities than that of impacted wetlands with a more even distribution of species abundances within and between sites (Ceilley 2008). Abundances of specific fish species will also provide a good indicator of ecosystem health. Deep-water refugia allow certain fish species and higher trophic level fishes to persist in areas where they would not survive during period of frequent dry-downs. Frequent dry-downs may alter the dynamics of native fish communities and may diminish the value of these habitats by not providing access to dry season refugium for aquatic fauna during the dry-down throughout the Picayune. Assessing the change in abundance of specific fishes after the plugging of Prairie and Merritt canal can provide us with another indicator of restoration success.
The baseline study was conducted to eventually evaluate habitat quality and restoration success at the completion of the project. As restoration continues it is expected that fish -10- communities will recover at the impacted wetlands and become more similar to those of reference wetlands. At this point in the restoration process, Phase 1 has been completed, and the continued plugging of additional canals, removal of roads, and the installation of pump stations is yet to occur. We have a unique opportunity, at this time, to measure the improvement of the ecosystems after the completion of Phase 1 of the project, and to examine possible methods for quantifying success.
ECOLOGICAL INDICATORS
Monitoring is required to determine whether the benefits of the project are being achieved (COE & SFWMD 2004). In 2005, the Department of the Interior produced a Science
Plan outlining the importance of monitoring within South Florida to develop and effectively synthesize and communicate the best available science in order to apply it to the adaptive management program for the large-scale restoration project (DIO 2005). The complexity of ecological systems makes it extremely difficult to measure and communicate the effects of such large restoration projects. Despite the difficulty, these monitoring efforts are essential for complex regional restoration programs as stated by Dale & Beyeler et al. (2001), “Ecological systems are complex, and developing effective strategies for measuring and communicating restoration success (or failure) is an extremely difficult, but essential task in any large, complex regional restoration program” (Schiller et al. 2001; Lausch & Herzog 2002; Niemi & McDonald
2004; Ruiz-Jean & Aide 2005; Thomas 2006; Doren et al. 2009 p. 2).
The task of monitoring the entirety of ecological processes within a system would be unmanageable, extremely expensive and time consuming. Ecologists have extensively sought concise and cost-effective measures that can characterize ecosystem conditions (Schiller 2001).
In a large and complex region such as South Florida, there must be means to determine how restoration goals are being met (Doren et al. 2009). -11-
Environmental, biological, and ecological indicators may provide an effective and efficient method of characterizing and assessing ecosystem conditions. The first reference to environmental indicators is attributed to Plato, who cited the impacts of humans on fruit tree harvest (Niemi 2004). The concept of indicators for plant and animal communities can be traced as far back as the 1600s. Examples of early indicators include using migratory bird populations to provide insight into changing environmental conditions, or the use of canaries in coal mines to detect the presence of noxious gasses in the 1920s. One of the more elaborate examples of early biological indicators was the saprobian system developed by Kolkwitz & Marsson in 1908, which used benthic and planktonic plants as indicator species for classifying stream decomposition zones (Niemi 2004). Variations of this system are still used in ecological assessments today in the Florida Department of Environmental Protection’s stream condition index (FL DEP).
Ecological indicators are defined by Niemi (2004) as, “measurable characteristics of the structure (e.g. genetic, population, habitat, and landscape pattern), composition (e.g. genes, species, populations, communities, and landscape types), or function (e.g. genetic, demographic/life history, ecosystem, and landscape disturbance processes) of ecological systems.” Ecological indicators serve several purposes including assessing the condition of the environment, or monitoring trends in condition over time. In addition, these indicators can be used to provide an early warning signal for changes in the environment, or diagnose the cause of an environmental problem (Dale 2001).
The selection of specific indicators for large scale restoration projects comes with many challenges. The selected indicator must be representative of the system as a whole by capturing key pieces of ecosystem function that are tied to the underlying values of the restoration program. The goals and objectives of the monitoring program will help to determine which -12- ecological indicators are used (Doren 2009; Niemi 2004). In addition, the selected indicator should be easily and routinely measurable, be sensitive to stresses on the system, have a known response to disturbances, and can be integrated into management. Indicators can be selected at different hierarchal levels including organism, species, population, ecosystem, and landscape in order to best provide a simple and efficient method to examine the ecological composition, structure, and function of complex ecological systems. This ecological hierarchy includes functional, compositional, and structural elements that can be combined to define ecological systems and provide a means to select indicators representative of the key characteristics of the system (Dale 2001).
Once an indicator is selected, it is necessary to determine the environmental metrics that will be used to most effectively represent the indicator’s response. Indicators previously used for
Florida’s restoration projects have embodied components of the South Florida ecosystem by using the organisms including characteristics distinctive of the Everglades landscape, trophic constituents, biodiversity, physical properties, and associated ecological structure and function
(Doren 2009). These indicators can then be used to reflect the biotic and abiotic state of the environment, reveal evidence for impacts of environmental change, and/or indicate the diversity of other species, taxa, or communities within an area (Niemi 2004).
Although few scientists deny the benefits that indicators provide to research, monitoring, and management efforts, it should be noted that there are three concerns that hinder the use of ecological indicators as a resource management tool. Those concerns include: the use of indicators that may not be able to convey the full complexity of the ecosystem; choosing indicators without defined project goals and objectives, or goals and objectives associated with short-term response rather than long-term maintenance of healthy ecosystems; and failure to establish standard procedures for the selection of indicators to ensure repeatability, avoid bias, -13- impose discipline upon the selection process, and encompass management concerns to ensure scientific rigor (Dale 2001). If these concerns are addressed in the planning process, indicators provide a valuable tool to research, monitoring, and land management activities to conduct concise and cost-effective evaluations of ecosystems (Schiller 2001).
The goals of the backfilling efforts in the Picayune Strand encompass the goals of the greater Everglades restoration. Some of the expected benefits surrounding these goals are: the restoration of historic wetland communities; restoration of sheetflow towards coastal estuaries; reduction of harmful surge flows through the Faka Union Canal into Faka Union Bay, improved freshwater overland flow and seepage in to other bays of the Ten Thousand Islands region; improved aquifer recharge; decreased frequency and intensity of forest fires; improved habitat for fish and wildlife and threatened and endangered species; reduced invasion of exotic species; and increased spatial extent of wetland (COE & SFWMD 2004; Chuirazzi & Duever 2008 p. 3).
With these goals and intended benefits in mind, it is imperative to monitor and track changes in environmental and ecological conditions, and the progress and extent of expected benefits as hydrologic restoration advances.
FISH AS INDICATORS
Fish have been identified as key indicators by which to measure restoration success in
South Florida. As essential components to the successful functioning of wetland food webs, changes in fish population sizes and community composition can provide essential information about the structure, composition, and function of an ecosystem (Loftus et al. 2000). Fish are also greatly affected by large scale ecosystem processes, especially in highly variable systems such as the seasonal wetlands of the Picayune Strand. Colonization by fish in these systems depends on the frequency and duration of hydrologic connections, and species specific dispersal abilities.
“The longer and more frequent the connections to other water bodies, the greater the opportunity fish species have to colonize a wetland,” (Barber et al. 2002). Fish assemblages are known to be -14- greatly influenced by these parameters, and it has been noted that higher hydrologic connectivity results in greater richness and diversity. These fundamental effects on fish community structure provide a means for evaluating impacts of changing hydrologic conditions on the health of seasonal wetlands. Monitoring fish assemblages may provide meaningful information on the ecological health of these systems (Main et al. 2007). Fish assemblages have also been noted
(Niemi 2004) as excellent indicators because:
1. They are relatively easy to identify; 2. They are of interest to the public;
3. They are relatively easy to measure;
4. They often contain various locally abundant species with known responses to
disturbance; and
5. They can be monitored at a relatively low cost.
Using fish assemblages as indicators is particularly successful in South Florida due to the topographical characteristics of the area. The lack of topographical relief allows fish species to move throughout the landscape as a function of hydrologic connectivity (Barber et al. 2002). In addition, wetland fish species in south Florida are adapted to deal with historic environmental conditions, which decrease the influence of local abiotic factors on community composition
(Ruetz et al. 2005). Anthropogenic hydrologic alterations such as channelization and compartmentalization of the previously sheetflow-driven system of the Picayune Strand resulted in major changes in the physical and hydrologic processes which has had far reaching effects on the ecological processes and habitat in the area. The physical alterations brought about by canals, impoundments, and roads have resulted in loss of connectivity necessary for sheetflow in this area (Ogden et al. 2005). -15-
The monitoring of ecological responses of fish to hydrologic change within the Picayune
Strand was identified as a necessary science for the south Florida adaptive management process
(DOI 2005). A unique aspect of the current Picayune Strand is that it is in the process of being restored, which allows for immediate biological comparisons to be made through the various stages of hydrologic restoration (Dixon 2009). Monitoring of fish assemblages within various restoration phases would provide excellent biological indication of the hydrologic conditions of these different areas, and the progress of hydrologic restoration over the landscape as a function of connectivity.
Fish community investigations have been conducted in the Picayune Strand since 2001 including pre-restoration wildlife surveys from 2001-2004 (Addison et al. 2006) and continued biological surveys from 2005-2007 (Ceilley 2008). Fish were sampled using Breder traps
(Breder 1960) to develop a baseline for fish communities to be assessed in the future after restoration occurred. Sampling took place over a broad area of the Picayune Strand including the tidally influenced southern portions. During early pre-construction sampling, a total of seven families and 23 species were collected from a variety of habitats (Addison et al. 2006). In the
2005-2007 study, 25 fish species from nine families were collected from a variety of habitat types across Picayune Strand. The study showed that reference wetlands showed a mean species richness of 8.1 species per site, while impacted wetlands showed a mean species richness of 4.5 species per site (Ceilley 2008). This data compared impacted wetlands to reference wetlands, but did not focus on the change in the transitional, recently restored areas of the Picayune Strand.
-16-
FISH SAMPLING METHODS
Many techniques, including active and passive sampling methods, have been used to sample fish communities in wetland habitats. In sampling fish communities, it is important to provide information about the spatial and temporal resolution of fish movement at a low cost and low catch per unit effort, while providing statistical validity through replicable methods (Sargent
& Carlson 1987). Some of the more most commonly used methods are: Breder traps; throw traps; seines; dip-netting; and drop traps. A brief summary of each of these methods advantages and disadvantages are provided below:
1) Breder Traps – This passive sampling method is suitable for use in all types of
vegetation and water levels, replicable with low labor, and inexpensive and
simple to apply. A limitation to this method is the behavior sampling bias against
transient and predatory species, and for active forage fish species (Ceilley et al. in
press, Main et al. 2007, Sargent and Carlson 1987).
2) Throw Traps – This active sampling technique works in herbaceous vegetation,
but can be very destructive to sampling areas and are both labor and time
intensive.
3) Seine Nets – This active sampling method collects large numbers of individuals
and species in the sampled area and provides a largely comprehensive inventory
of fish species in the area. A limitation to this method is that it can only be used
in open water with flat bottoms (Sargent & Carlson 1987).
4) Dip-Netting - This active sampling technique is effective in areas that are difficult
to sample using other active sampling techniques, and are inexpensive and easy to
use. Limitations to this method include disturbance to the sampling area, and -17-
difficulty in standardizing sampling effort and efficiency among individuals and
locations (Main et al. 2007).
5) Drop Traps – This active sampling technique is effective to conduct quantitative
sampling. Some limitations of this method are that it is a destructive method of
sampling, capture is selective due to avoidance behaviors, they are expensive, and
they are complicated to build, set up, and transport (Sargent & Carlson 1987).
There are many fish sampling methods that have been proven effective. “Selection of the appropriate technique or techniques depends on the question being asked, the information required, the nature of the organism(s) or habitat being studied, and the resources available for the project,” (Heyer et al. 1994; Dixon 2009). Since previous research within the Picayune
Strand and Babcock Ranch in Charlotte and Lee Counties has been successfully conducted using a combination of Breder traps and D-frame dip nets (Ceilley 2008 and Ceilley et al. 2013), and because results can be compared to new data collected, these methods were employed during this research project. Breder traps have been identified as one of the most efficient and cost effective techniques for attaining a statistical validity while measuring communities of wetland fish species (Ceilley et al. 2013; Sargent & Carlson 1987). Based on survey methods that have been proven successful within the Picayune Strand, and the identified need for fish community sampling (DOI 2005) in the area, Breder traps in combination with dip-netting appear to be the most effective sampling techniques. By employing these survey methods in the appropriate areas, adequate and appropriate fish community information was expected to be obtained to help determine whether the benefits of the restoration project are being achieved.
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OBJECTIVES
Previous research conducted in the Picayune Strand was intended to eventually evaluate habitat quality and restoration success at the completion of the project. As restoration continues it is expected that fish communities will recover at the impacted wetlands and become more similar to those of reference wetlands. At this point in the restoration process, Phase 1 has been completed, and the continued plugging of additional canals, removal of roads, and the installation of pump stations is in progress. We have a unique opportunity, at this time, to measure the improvement of the ecosystems after the completion of Phase 1 of the project, and to examine possible methods for quantifying success. It is also important to consider sampling cost and efficiency as financial restraints have been a factor in the progress of restoration and monitoring in the Picayune.
Therefore, the objective of this research included the following:
1. Identify specific species to use as performance indicators;
2. Examine the implications of various sampling frequencies to determine the
minimum required to adequately track changes in community structure;
3. Determine the time of year most appropriate to collect samples from a mature fish
community;
4. Determine how fish communities change from the beginning of the wet season
until the dry down;
5. Examine cyclical patterns of temporal dispersion over multivariate space caused
by seasonal hydrologic changes; and
6. Determine whether fish communities at Transitional sites undergoing restoration
are moving from impacted conditions towards minimally impacted Reference
conditions. -19-
HYPOTHESIS
Based on the objectives described above, the hypotheses of this research include the following:
1. There will be significant differences in the distribution of various fish species
relative to the different restoration phases which will lead to the identification
of “indicator species” of restoration success.
2. Monthly sampling of fish communities will provide a similar signal to that of
an abbreviated sampling frequency, identifying the minimum sampling
frequency required to acquire an accurate depiction of fish community
structure among restoration phases.
3. Sampling will be most effective during the months after the start of the dry-
down while the majority of sites are still inundated to allow sampling of fish
communities.
4. Richness, relative abundance, and diversity of fish species will increase
through the dry down as fish communities are concentrated and isolated in
smaller areas.
5. A pattern will emerge from multivariate analysis that can be compared to
additional data in an effort to examine temporal fish community structure
changes, and identify a dispersion pattern through multivariate space
representing fish communities in different phases of restoration.
6. Fish communities in Transitional sites undergoing restoration with show
greater similarity to Reference sites than Impacted sites as hydrologic
restoration will allow movement of fishes across an improved hydrologic
landscape. -20-
METHODS
Experimental Design
There are three general phases specifically related to this study; 1) natural wetland habitats
including deep cypress, cypress gramminoid, and gramminoid marshes (reference); 2) restored canal –
former Prairie Canal including all habitat types (transitional treatment); and 3) un-restored Miller Canal
including all habitat types(impacted treatment). The study area occurred across the center portion of the
Picayune Strand representing the progression of the restoration and hydrologic regimes of the area on a
latitudinal basis.
The study area was divided in to three regions from east to west containing 1) three sample
locations within impacted un-restored sites around Miller Canal; 2) four sample locations within restored
locations around former Prairie Canal; and 3) three sample locations within the natural wetland habitats of
the Fakahatchee Strand. This study looked at these different restoration phases in an attempt to identify
patterns among fish community use and composition in
relation to restoration progress. Survey methods
included using Breder Traps and dip-netting to
incorporate both active and passive sampling techniques.
These surveys were conducted monthly during the end of
the wet season through the dry-down of 2011-2012 (i.e.
September through March).
Sample Regions
The study was divided into three regions with
three habitat types per region to represent major wetland
Figure 6. This map shows the sites selected for this study including three restoration phases with representative habitat types. Sites habitat types across different phases of the overall include two impacted sites (cypress gramminoid and gramminoid habitats), four transitional sites (two deep cypress, and gramminoid restoration project. Ten sample sites were distributed habitats), and three reference sites (deep cypress, cypress gramminoid, and gramminoid habitats). An impacted deep cypress site and a transitional gramminoid site were eliminated due to lack of water and along an East to West transect of the Picayune and samplable fish communities. An additional transitional deep cypress site was added due to its position between recently restored prairie canal and the reference wetlands of the Fakahatchee Strand. -21-
Fakahatchee Strands. Each restoration phase contained 3 to 4 sites representing 3 habitat types including deep cypress, cypress gramminoid, and gramminoid marsh. Therefore, each habitat type could be
analyzed independently and/or together by restoration phase.
Deep Cypress
Deep cypress habitat is characterized by channelized water flow through a “strand” dominated by cypress tree, epiphytes, and a shaded understory. During the dry season, these areas retain water for longer time periods than surrounding wetland areas giving it the longest period of inundation of all the represented habitats in this study
(Whitney, Means, & Rudloe 2004). These habitats may provide dry season refuge for fish species in the area, and contain larger more predatory species in addition to omnivorous forage fish that dominate South Florida wetlands (Odgen 2005).
Cypress Gramminoid
Cypress gramminoid habitat is characterized by a shallower flow way with a less shaded understory, allowing for the growth of herbaceous wetland place such as Jamaica swamp saw grass, ferns, and shrubs (Whitney,
Means, & Rudloe 2004). These habitats are usually ephemeral and provide excellent habitat for colonizer fish species such as Gambusia holbrooki and Jordonella Floridae (Addison et al.
2006). It also provides an advantage to smaller forage fish species that use submerged and emergent
plants as critical habitat and allow protection from predation (Main et al. 2007). -22-
Gramminoid
Gamminoid marshes are ephemeral, broad level plains containing a variety of herbaceous wetland plants and grasses. These habitats become inundated for part of the year, but can remain dry for extended periods of time (Whitney, Means,
& Rudloe 2004). These habitats are also beneficial to colonizer and small forage fish species who take advantage of newly inundated areas for protection from predation and new food sources (Barber 2002).
Restoration Phases Miller Canal
The Miller Canal is the furthest major unrestored drainage canal from the recently restored Prairie
Canal and the natural reference site of the Fakahatchee Strand. This canal runs parallel with the former
Prairie Canal and is located on the far west side of the Picayune Strand. Three sampling sites were selected in the region representing the three habitats of focus in this study. Each of the three habitat sites contained vegetative communities that are characteristic of those habitats. The sites around this canal provided treatment stations for this study which demonstrated pre-restoration characteristics around an existing un-restored canal that were compared against sites around restored sites which should be demonstrating ecological recovery, and un-impacted reference sites located in the Fakahatchee Strand
(Ceilley 2008). All three sites were located around Stewart Boulevard, which is located in the central portion of the Picayune Strand and runs east to west.
Prairie Canal
Restoration on the former Prairie canal was completed in July 2007 which included the installation of 7 miles of canal plugs (Dixon 2009). This former canal is located on the easternmost side -23- of the Picayune Strand running north to south. Three sampling sites were selected in this region representing the three habitats of focus in this study. Each of these habitat sites also contained vegetative communities that are characteristic of those habitats. The sites around this former canal provided treatment stations for this study which demonstrated post-restoration characteristics around a canal that has undergone complete restoration. These sites were compared against sites around the un-restored
Miller canal, and un-impacted reference sites located in the Fakahatchee Strand. All three sites were also located around the east-west transect of Stewart Boulevard.
Fakahatchee Strand
The Fakahathee Strand is a State Park located to the east of the Picayune Strand. The majority of the Picayune Strand has not been negatively impacted by the Southern Golden Gate Estates (SGGE) project, and retains a natural habitats and hydrologic regime. Four sampling sites were selected in this region representing the three habitats of focus in this study, and an additional deep cypress habitat site at a location which had been previously noted as being impacted by the altered hydrology of the Picayune
Strand due to its close proximity to the previously impacted areas (US Army Corps of Engineers 2004).
Each of the selected sites contained a vegetative community that was characteristic of those habitats. The sites within the Fakahatchee Strand provided reference stations for this study which demonstrated natural habitats and hydrology unaffected by the hydrologic alterations in the Picayune Strand, with exception of the additional deep cypress site. These sites were compared against the restored and unrestored sites located in the Picayune Strand. All sites in the Fakahatchee Strand were located along Jane’s Scenic
Drive, which is an extension of Stewart Boulevard bisecting both the Picayune and Fakahatchee Strands on an east to west transect.
Survey Techniques
Fish were sampled using both active and passive capture methods to inventory present fish species and determine species richness and relative abundance. Each sample location included Breder trap sampling, and dip-netting. The survey data was recorded on a pre-prepared observation form
(Appendix A). Environmental data was obtained for each sampling event using a YSI Model 85 handheld oxygen, conductivity, salinity, and temperature system (YSI 1998). -24-
Breder Traps
Many fish sampling techniques can be both destructive to habitat and difficult to conduct in areas with thick wetland vegetation (Main et al. 2007;
Sargent and Carlson 1987). As Breder traps have been shown to be a successful technique in measuring absolute or relative densities of wetland fish species (Ceilley et al. in press, Sargent and Carlson 1987), and they were used in previous studies in the Picayune Strand (Ceilley et al. 2007, Ceilley 2008), this method was also selected for use in this study. For sampling, ten of these plastic fish traps (Breder 1960) were deployed at each sampling location when depths were sufficient to permit effective sampling (>2cm). Breder traps were placed in a stratified pattern throughout the aquatic habitat to sample available fish habitat at each location and to maximize capture efficiency (Sargent and Carlson 1987; Main et al. 1997; Ceilley et al.
1999). Following the protocol employed for the baseline assessment (Ceilley 2008), traps were submerged for a period of one hour and then retrieved. The fish collected were field identified, enumerated, and recorded on field sheets before being released live back into the water. Voucher specimens for each fish species, along with any unidentified species in the field were anesthetized using a solution of benzocaine and then preserved in 10% formalin and labeled by site and date to be brought back to the FGCU lab for positive identification.
Dip Netting
Concurrent active dip net sampling was conducted by 2 persons with standard D-frame, 500 micron mesh dip nets in all of the major and minor micro-habitats (e.g. emergent and submerge vegetation, snags, roots, algal mats, benthos, rocks, etc.) available at each site (Ross 1990; FDEP 1993).
Dip nets were selected for they were used in previous studies conducted in the Picayune Strand (Main et al. 2007) and their relative ease of use and versatility in diverse habitats. “There are no definitive rules about the number of sweeps needed to sample a habitat adequately…a reasonable procedure is to survey -25- each aquatic habitat for an equal period of with an equal number of sweeps,” (Heyer et al. 1994). Dip netting was standardized by sampling each represented habitat with three sweeps within each sampling location. The net was vigorously worked through vegetation, open water, and/or surficial bottom sediments. The net contents were then placed in a white pan and sorted with forceps. Fish were identified, enumerated, and recorded on field sheets before being released live back into the water. Any unidentified species in the field were anesthetized using a solution of benzocaine and then preserved in 10% formalin and labeled by site and date to be brought back to the FGCU lab for positive identification.
Data Analysis
Plymouth Routines in Multivariate Ecological Research, Volume 6 (PRIMER v6) was utilized to perform analyses for the fish community data collected in this study (Clarke & Gorley 2006). Univariate analysis was used for a comparison of treatments (reference, transitional, natural), for mean diversity using several diversity indices, richness, and abundance. Non-parametric multivariate techniques were also used to analyze the community data. All fish abundance data for this analysis were forth-root transformed prior to analysis to down-weight the influence of extremely abundant species, (Clarke and
Warwick 2001; Ceilley 2008) which was consistent with previous studies. “Standardization is appropriate for comparing data sets with varying levels of effort to obtain percent composition values for individual taxa within a community (by samples) or percent composition of individual taxa between communities (by variable) to identify relative distribution among habitat types and endemism where an individual species or taxa is found only at one site or habitat type,” (Ceilley 2008). Differences in water levels between reference, transitional, and impacted sites that allow for, or preclude the development of -26- aquatic communities was considered as a primary variable to be monitored. Therefore, by weighting the samples by averaging abundance and/or emphasizing percent composition between sites may mask differences that may be ecologically important for measuring restoration of hydrology and sheetflow.
During scheduled wet season sampling events when dry conditions were observed (and therefore no fishes were present) data were recorded as zero (0) fishes collected. Due to the small number of samples collected at each site, data were pooled for each site for all sampling events for the overall analysis of site conditions (Ceilley 2008). Data were also analyzed by sampling events to determine temporal trends, and indicate adequate sampling frequencies over the sampling period.
Hierarchical agglomerative cluster analysis (using group average linking) based on Bray-Curtis similarities was employed to examine natural groupings among samples and sampling locations. The groupings produced in the cluster diagram were further evaluated for statistically significant evidence
(p<0.05) of genuine clusters in sample assemblages using a series of similarity profile (SIMPROF) random permutation tests, a feature incorporated in PRIMER v6 (Clarke & Gorley 2006). Because cluster analysis is best applied to communities responding to distinct environmental, non-metric multidimensional scaling (MDS) was also employed. This statistical tool can also be based on Bray-
Curtis similarity matrices and has been recommended for use in examining communities structured by environmental gradients (Clarke & Warwick 2001; Clarke & Gorley 2006). In the case of MDS, plots may be considered useful for interpretation if associated stress values are >0.2 (Kruskal 1964).
Analysis of similarity (ANOSIM), a non-parametric analog of analysis of variance was used to test for a priori groupings of samples. When using ANOSIM, global tests were considered significant at p<0.05 and pair-wise tests were interpreted using the absolute value of the associated R statistics (Clarke
& Warwick 2001). Relative contributions, in terms of relative abundance were graphically displayed as overlays on MDS plots to illustrate relative importance of ordination.
Additional statistical analysis was conducted on the data set using trajectories to observe temporal fluctuations in species assemblages within and between sites through multivariate space. Given the cyclical nature of inundation in south Florida landscape, cyclic season patterns in fish community were observed in sites that provided a period of inundation long enough for analysis. The sampling frequency -27- of this study allowed for the detection of changes in community structure through the period of inundation as well as the presence/absence of individual taxa known to be indicators of wetland condition.
This information allowed for the observations of patterns to inform wetland restoration science of what time of year was the most appropriate for collecting a mature wetland fish community, how wetland fish communities change from the beginning of the wet season through the dry-down, and what sampling frequency was necessary for obtaining an accurate representation of the temporal changes in community structure and diversity metric s.
Baseline studies show high dissimilarity in fish communities at impacted sites and high similarity between reference sites, suggesting that disturbance of sheetflow causes dispersion, or high dissimilarity of fish communities even from like habitats. This dispersion observed in the baseline study at most of the impacted wetland areas should show a decrease over time in transitional areas as wetlands are restored.
The analysis in this study allows for the observation of this trend at a critical time in the Picayune Strand
Restoration Project process that is currently not being evaluated.
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RESULTS
A total of 8 sites were successfully sampled for fish including two impacted sites, three transitional sites, and three reference sites (Table 1). Two sites, impacted cypress and transitional gramminoid, did not have a long enough period of inundation, or enough hydrologic connectivity to support a fish community, and therefore did not provide fish community data. These sites were eliminated from analysis as they did not provide sufficient data for this study. A total of 18 sampling events were conducted from September 2011 through March 2012 where adequate standing water was available. All 8 sites were successfully sampled a minimum of 1 time during the sampling period as water levels allowed (Table 2). For potential use in future research projects, a raw data set for all sampling events are provided in Appendix B. A total of 15 species were documented within the study area by use of the sampling methods described.
Table 1. Site descriptions with locations and adjacent wells. Sites highlighted in red have been eliminated from the study due to lack of water and a samplable fish community.
Adjacent Well Restoration Phase Habitat Type Location T3W1 Impacted Gramminoid Picayune T3W2 Impacted Cypress Gramminoid Picayune T3W2 Impacted Deep Cypress Picayune T3W5 Transitional Gramminoid Picayune T3W5 Transitional Cypress Gramminoid Picayune T3W5 Transitional Deep Cypress Picayune T3W7 Transitional Deep Cypress Fakahatchee - Reference Gramminoid Fakahatchee - Reference Cypress Gramminoid Fakahatchee - Reference Deep Cypress Fakahatchee
-29-
Table 2. This table shows the presence/absence of water in each sampling site per month. Cells highlighted in blue represent inundated sites that held standing water and a samplable fish community during the represented month. Blank cells represent dry sites without a samplable fish community.
Site September October November December January February March Impacted Cg X X Impacted C X Impacted G X X Transitional C X X X Transitional Cg X X Transitional G X X Transitional C2 X X X X X X Reference C X X X X X X X Reference Cg X X X X X Reference X X X X
Fish Community Data
Table 3. This table shows fish species documented via Breder traps and dip-net sampling by scientific name, common name, and species code abbreviations.
Species Capture Scientific Name Common Name Code Method Belonesox belizanus Pike killifish PKKF Breder Cichlasoma urophthalmus Mayan cichlid CIUR Breder Elassoma evergladei Everglades pygmy sunfish ELEV Dip Net Fundulus chrysotus Golden topminnow FUCR Breder Fundulus confluentus Marsh killifish FUCO Both Gambusia holbrooki Mosquito fish GAHO Both Hemichromis letourneuxi Jewelfish cichlid HELI Breder Heterandria formosa Least killifish HEFO Breder Jordonella floridae Flagfish JOFL Both Lepomis macrochirus Bluegill LEMAC Breder Lepomis marginatus Dollar sunfish LEMAR Breder Lepomis sp. (juv) Sunfish juvenile LESP Both Lucania goodei Bluefin killifish LUGO Breder Notropis petersoni Coastal shiner NOPE Breder Poecilia latipinna Sailfin molly POLA Breder
A total of 15 species were documented through this study using Breder traps (Breder 1960) and dip-netting. Each species was identified in the field and released alive unless positive identification was required, upon which they would be preserved and brought to the FGCU laboratory for further investigation. The fish species abbreviations for each fish species were based on the first two letters of -30- the Genus and species in each scientific name. These species abbreviation are used in various graphs and figures throughout the remainder of this report (Table 3).
Species Richness
To begin evaluating status and trends of fish communities, species richness was determined.
Table 4 summarizes fish species that were documented at least once in each of the represented restoration phases. Impacted study sites contained a total of two species, the Transitional study sites contained a total of 6 species, and reference sites contained a total of 15 species. A total of 15 fish species were collected representing 7 families including 1 non-native family and 3 non-native species. The most abundant species by far was the eastern mosquito fish (Gambusia holbrooki) (n = 2965) which was found at all locations where fish were collected. Also abundant in some locations was the marsh killifish (Fundulus confluentus) (n = 208), Florida flagfish (Jordonella floridae) (n = 107), and juvenile sunfish (Lepomis sp.
(juv)) (n = 30).
Table 4. This table shows fish species documented in each restoration phase listed by scientific name. An ‘X’ shows that a fish species was present at a site at some time within the period of this study.
Scientific Name Impacted Transitional Reference Belonesox belizanus X X Cichlasoma urophthalmus X Elassoma evergladei X Fundulus chrysotus X Fundulus confluentus X X X Gambusia holbrooki X X X Hemichromis letourneuxi X Heterandria formosa X Jordanella floridae X X Lepomis macrochirus X Lepomis marginatus X Lepomis sp. (juv) X X Lucania goodei X X Notropis petersoni X Poecilia latipinna X Total Documented Species 2 6 15
Univariate diversity metrics were conducted for each of the sampling locations in all restoration phases. The metrics applied to this analysis included six different indices: richness (species richness (R) -31- and relative abundance (A)), evenness (Margalef’s (d) and Pielou’s indices (J’)), and heterogeneous measures (Shannon-Weaver index (H’(loge)) and Simpson’s diversity index (1-Lambda’). Total documented species for each site ranged from a low of 1 species at the impacted gramminoid site, to a high of 13 species at the reference cypress gramminoid site (Table 5). The mean species richness was 1.5 for impacted sites, 3.3 for transitional sites, and 8.0 for reference sites. In general, the greatest diversity values were reported in reference sites. Figure 7 illustrates the average species richness and relative abundance of fishes within each restoration phase calculated from the values described in Table 5. The lowest species richness and relative abundance values were documented in impacted sites. Relative abundance was the greatest within transitional sites while diversity was greatest within reference sites
Table 5. : Univariate diversity indices including species richness (R) and relative abundance (A), Margalef’s (d), Pielou’s indices (J’), Shannon-Weaver index (H’(loge), and Simpson’s diversity (1-Lambda’) for fish documented with highest values for each region in bold and lowest value underlined are represented in this table. The highest diversity values were found in reference sites and the highest abundance values were found in the transitional cypress site ‘C2’. The lowest abundance and diversity values were found in impacted sites.
Diversity Metrics Sample Species Abundance Margelef Pielou Shannon Simpson 1- S N d J' H'(loge) Lambda' Impacted Cg 2.0 28.0 0.3 0.4 0.3 0.1
Impacted G 1.0 14.0 0.0 **** 0.0 0.0 Transitional C 3.0 376.0 0.3 0.0 0.0 0.0 Transitional Cg 2.0 108.0 0.2 0.1 0.1 0.0 Transitional C2 5.0 1574.0 0.5 0.1 0.1 0.0 Reference C 7.0 962.0 0.9 0.3 0.5 0.2 Reference Cg 13.0 442.0 2.0 0.5 1.4 0.7 Reference G 4.0 183.0 0.6 0.5 0.7 0.4
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Total Abundance (N) Species Richness (S)
1600.0 14.0
1400.0 12.0
1200.0
10.0
1000.0 8.0 800.0 6.0
600.0 RichnessSpecies Relative Abundance Relative 4.0 400.0
200.0 2.0
0.0 0.0
Figure 7. This is a graphic representation of average species richness and relative abundance of fishes within each restoration phase documented during sampling. Species richness was highest in Reference sites, and relative abundance was highest at transitional site ‘C2’, followed by reference sites and transitional site ‘C’. Lowest values were observed in impacted sites.
Figure 8 illustrates the average diversity metric values (Margalef’s (d), Pielou’s indices (J’),
Shannon-Weiver (H’(loge)), and Simpson’s diversity (1-Lambda’)) of fishes within each restoration phase calculated from the values described in Table 5. Diversity metrics were generally highest in reference sites with the exception of the impacted cypress gramminoid site. Transitional sites showed a general increase in diversity metric values from transitional sites, and reference sites showed a general increase from transitional sites. Elevated diversity metrics in the impacted gramminoid site may have been driven by the low relative abundance of species. -33-
2.0 1.8
1.6 1.4 1.2 1.0 0.8 0.6
Diversity Diversity MetricValues 0.4 0.2 0.0
Margelef (d) Pielou (J') Shannon (H'(loge)) Simpson (1-Lambda') Sample Sites
Figure 8. Average diversity metrics including Margalef’s (d), Pielou’s indices (J’), Shannon (H’(loge)), and Simpson (1- Lambda’) for each of the represented sampling sites are shown in this graph. The highest values were found in reference sites.
Relative Abundance
Fish species documented in this study are classified as small omnivorous fishes that move through the hydrologic landscape via active dispersal (Trexler 2010; Main et al. 2007). Hydrologic connectivity and distance shape the patterns of fish dispersal over the south Florida landscape which impacts the relative abundance of species at locations of varying distance from source populations (Ruetz et al. 2005). The relative abundance of fish species documented via sampling in this study was analyzed -34-
Relative Abundance Scientific Name Species Impacted Transitional Reference Total Belonesox belizanus PKFF 0 3 2 5 Cichlasoma urophthalmus CIUR 0 0 1 1 Elassoma evergladei ELEV 0 0 2 2 Fundulus chrysotus FUCR 0 0 2 2 Fundulus confluentus FUCO 2 20 186 208 Gambusia holbrooki GAHO 31 2027 907 2965 Hemichromis letourneuxi HELI 0 0 15 15 Heterandria formosa HEFO 0 0 6 6 Jordanella floridae JOFL 0 6 101 107 Lepomis macrochirus LEMAC 0 0 2 2 Lepomis marginatus LEMAR 0 0 4 4 Lepomis sp. (juv) LESP 0 1 29 30 Lucania goodei LUGO 0 1 1 2 Notropis petersoni NOPE 0 0 1 1 Poecilia latipinna POLA 0 0 13 13
Total Documented Species 33 2058 1272 3363 Percent Composition Scientific Name Species Impacted Transitional Reference Belonesox belizanus PKFF 0.00% 0.15% 0.16% Cichlasoma urophthalmus CIUR 0.00% 0.00% 0.08% Elassoma evergladei ELEV 0.00% 0.00% 0.16% Fundulus chrysotus FUCR 0.00% 0.00% 0.16% Fundulus confluentus FUCO 6.06% 0.97% 14.62% Gambusia holbrooki GAHO 93.94% 98.49% 71.31% Hemichromis letourneuxi HELI 0.00% 0.00% 1.18% Heterandria formosa HEFO 0.00% 0.00% 0.47% Jordonella floridae JOFL 0.00% 0.29% 7.94% Lepomis macrochirus LEMAC 0.00% 0.00% 0.16% Lepomis marginatus LEMAR 0.00% 0.00% 0.31% Lepomis sp. (juv) LESP 0.00% 0.05% 2.28% Lucania goodei LUGO 0.00% 0.05% 0.08% Notropis petersoni NOPE 0.00% 0.00% 0.08% Poecilia latipinna POLA 0.00% 0.00% 1.02% Table6. The above table shows the total relative abundance values per restoration phase, total abundance for all species, and total percent composition of fish species documented during the sampling period. The most abundant species included Gambusia holbrooki and Fundulus confluentus, and the least abundant species included Cichlasoma urophthalmus and Notropis petersoni. Gambusia holbrooki composed the majority of all species observed.
-35- in an attempt to identify dominant fish species, species composition similarities among restoration phases, and potential indicator species. Table 6 shows the total abundance values for each individual fish species documented during this study. This table shows the total relative abundance for each species per restoration phase, as well a total relative abundance for all sampling events and total percent composition of each species per restoration phase.
Figure 9 illustrates the total percent composition for each fish species over the entire sampling period. This was determined by totaling the relative abundance of each fish species over the entire sampling period and calculating the overall percentage.
The fish species abbreviations in Figure 8 are described in Table 2 and Table 6. Gambusia holbrooki (88 percent) comprised the overwhelming majority of species collected during this study and was the most abundant of documented species. Other abundant species observed included Fundulus confluentus (6 percent) and Jordonella floridae (3 percent). All other documented species comprised less than 1 percent of the total collected specimen.
CIUR HELI 0.03% FUCR0.45% 0.06% FUCO6.18% LUGO0.06% 0.15% PKKF 88 88.17% 11.83% GAHO0.18% 0.39% HEFO3.18% POLA0.89% 0.12% JOFL0.03% LESP0.06% 0.06% LEMAC LEMAR
Figure 9. The above figure shows percent composition of species documented via sampling efforts based on overall relative abundance in the form of a pie chart. Gambusia holbrooki makes up the majority of all fish collected during the sampling period.
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Abundance data for fish species was used to observe similarities and difference between and among sites in the various restoration phases. Similarity percentage tests (SIMPER) for reference, transitional, and impacted sites are included in Table 7. Reference sites averaged 58% similarity (32% dissimilarity) of fish community structure, with 4 species contributing >91% of the similarity between reference sites. Transitional sites averaged 63% similarity (37% dissimilarity) of fish community structure, with 2 species contributing to 100% of the similarity between transitional sites. Of the two sampled impacted sites, there was an average of 72% similarity (28% dissimilarity) of fish community structure, with 1 species contributing to 100% of the similarity between impacted sites. Of the 12 species that contributed to the >90% dissimilarity between sites of different restoration phases, the average abundances of 10 species were higher in reference wetlands than in transitional wetlands including many considered as indicators of wetland hydrology (Main et al. 2008), and the lowest average abundance of various species was found in impacted wetland. These indicators include the Florida flagfish, marsh killifish, mosquito fish, and sailfin molly which made the largest contribution to dissimilarity between sites of varying restoration phases.
Using this data, a cluster analysis dendrogram was prepared of all sample locations with data from all sampling events to illustrate natural groupings between fish communities based on Bray-Curtis
Similarity of fish abundance data (Figure 10). The restoration phase is provided by the symbol used to represent each site and each habitat type is represented by the letters beneath. The cluster analysis identified two main clusters. Reference and transitional sites grouped together in clusters that were greater than 50 percent similar and impacted sites grouped together in a cluster that was greater than 70 percent similar. The less distinct groupings of transitional sites and reference sites may be a function of other influences including the advanced restoration and geographic location of transitional site C2. The measure of relative abundance were also assessed for each of the sampling sites using composite data from all sampling events by compiling non-metric MDS ordination to illustrate the relative similarity between fish communities based on restoration phase and habitat type (Figure 11).
The restoration phase is provided by the symbols used to represent each site, and the habitat type is represented by the habitat letter codes assigned to each symbol. The results of the MDS ordination -37- show distinct groupings of the sample locations among like restoration phases where impacted sites are in one group and transitional and reference sites group together in another distinct group.
Figure 11 is a 2-dimensional (2D) ordination that was reduced from a 3-dimensional (3D) ordination for incorporation into this document. The 2D stress value was 0.03 which provides a useful image (Clarke & Warwich 2001). The 3D stress value was 0.01, which provides an even more support for the flattened 2D image.
Figure 10. The above figure is a cluster analysis showing restoration phase and habitat type using Bray-Curtis similarity from all samples. The cluster analysis shows two distinct groupings of sites at 95% confidence (p<0.05) with three additional sub-groupings. Major groupings include a cluster of impacted sites, transitional sites ‘C’ and ‘Cg’, and reference site ‘G’ at about 50% similarity. The second major cluster shows reference sites ‘C’ and ‘Cg’ as well as transitional site ‘C2’ at >50% similarity. Sub groupings include impacted sites at >70% similarity in one group, transitional sites ‘C’ and ‘Cg’ with reference site ‘G’ at >60% similarity in the second group, and transitional site ‘C2’ with reference sites ‘C’ and ‘Cg’ in a third sub grouping. -38-
Figure 11. The above figure is an MDS ordination with restoration phase and habitat type for each sampling site using Bray-Curtis similarity from all sampling events. The ordination through multivariate space is based on similarity values and places points in a three-dimensional multivariate space. The three dimensional image has been compressed into a 2D representation in order to represent the multivariate ordination effectively.
Potential Indicator Species
The MDS ordination was overlaid with superimposed total abundance data for each species that had significant impact on the on the similarities and differences among and between sampling sites in an attempt to identify “indicator species” of restoration success (Table 7). The scales for the MDS ordinations with superimposed fourth root abundance figures represent 5 species including Gambusia holbrooki, Jordonella floridae, Fundulus confluentus, Lepomis sp., and Poecilia latipinna. These species made up most of the dissimilarity among various restoration phases in the SIMPER test due to their high occurrence in reference sites, moderate occurrence in transitional sites, and low occurrence and/or absence in impacted sites. Figures 12-16 are MDS ordination figures with superimposed fourth root abundances for the 5 species attributing the most dissimilarity between sites of different restoration phases. The highest overall abundances for all species except Gambusia holbrooki were documented in reference sites, while G. holbrooki was most abundant in transitional sites.
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Table 7. SIMPER analysis showing the species contributing to similarity within groups (A) and the dissimilarity between groups (B). Impacted sites: T3W1-Cg, T3W1-G. Transitional Sites: T3WS-C, T3WS-Cg, FS-C2. Reference sites: FS-C, FS-Cg, FS-G. Cumulative species contributions cut off at 90%. Species codes shown in Table 3. The information shown in this table was used for identifying potential indicator species. A Impacted Sites: Average Similarity = 71.89 Species Average Abundance Average Similarity Similarity/SD Percent Cumulative Similarity Percent GAHO 2.1 71.89 - 100 100
Transitional Sites: Average Similarity = 63.39 Species Average Abundance Average Similarity Similarity/SD Percent Cumulative Similarity Percent GAHO 4.63 48.76 4.77 76.93 76.93 JOFL 1.17 14.62 3.81 23.07 100.00
Reference Sites: Average Similarity = 58.30 Species Average Abundance Average Similarity Similarity/SD Percent Cumulative Similarity Percent GAHO 4.19 23.43 3.35 38.69 38.59 JOFL 2.40 12.34 3.14 25.33 64.02 FUCO 2.74 8.06 1.14 23.08 87.10 LESP 1.28 3.69 0.64 5.01 92.12 B Impacted and Transitional Sites: Average Dissimilarity = 52.79 Species Impacted Transitional Average Dissimilarity/ Percent Cumulative Abundance Abundance Dissimilarity SD Contribution Percent GAHO 2.1 4.63 23.43 3.35 44.38 44.38 JOFL 0 1.17 12.34 3.14 23.38 67.76 FUCO 0.59 0.70 8.06 1.14 15.27 83.03 LUGO 0 0.33 3.69 0.64 7.00 90.03
Impacted and Reference Sites: Average Dissimilarity = 68.56 Species Impacted Transitional Average Dissimilarity/ Percent Cumulative Abundance Abundance Dissimilarity SD Contribution Percent JOFL 0 2.4 14.02 2.98 20.45 20.45 GAHO 2.1 4.19 11.86 2.23 17.30 37.75 FUCO 0.59 2.74 11.48 2.45 16.75 54.49 POLA 0 1.05 6.27 1.15 9.15 63.64 LESP 0 1.28 5.97 1.18 8.71 72.35 FUCR 0 0.67 3.06 1.24 4.46 76.81 HELI 0 0.66 2.51 0.65 3.67 80.48 LEMAC 0 0.40 2.12 0.64 3.09 83.56 ELEV 0 0.40 2.12 0.64 3.09 86.65 HEFO 0 0.52 2.00 0.65 2.92 89.56 LEMAR 0 0.47 1.81 0.65 2.63 92.20
Transitional and Reference Sites: Average Dissimilarity = 47.77 Species Impacted Transitional Average Dissimilarity/ Percent Cumulative Abundance Abundance Dissimilarity SD Contribution Percent JOFL 0.70 2.74 9.42 1.83 19.72 19.72 GAHO 4.63 4.19 5.65 1.36 11.82 31.54 FUCO 1.17 2.40 5.58 3.15 11.69 43.23 POLA 0 1.05 4.83 1.20 10.10 53.33 LESP 0.33 1.28 4.73 1.28 9.90 63.23 FUCR 0 0.67 2.52 1.27 5.27 68.50 PKKF 0.44 0.40 2.17 0.85 4.55 73.05 HELI 0 0.66 2.14 0.66 4.47 77.52 LUGO 0.33 0.33 1.93 0.78 4.04 81.56 LEMAC 0 0.40 1.70 0.66 3.57 85.12 ELEV 0 0.40 1.70 0.66 3.57 88.69 HEFO 0 0.32 1.70 0.66 3.56 92.24 -40-
Figure12. The above figure is an MDS ordination with superimposed raw abundance for Gambusia holbrooki. Abundance for Gambusia holbrooki was highest in the transitional ‘C2’ site, followed by the reference sites.
Figure 13. The above figure is an MDS ordination with superimposed raw abundance data for Fundulus confluentus. Abundance data for Fundulus confluentus was highest in reference sites followed by the transitional site ‘C2’.
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Figure 14. The above figure is an MDS ordination with superimposed raw abundance data for Jordonella floridae. Abundance data for Jordonella floridae was highest at reference sites, followed by transitional sites. Jordonella floridae was absent in impacted sites.
Figure15. The above figure is an MDS ordination with superimposed raw abundance data for Lepomis spp. This species was most abundance in reference sites ‘C’ and ‘Cg’, followed by transitional site ‘C2’. Lepomis spp. were absent in all other sites during the sampling period.
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Figure16. The above figure is an MDS ordination with superimposed raw abundance data for Poecilia latipinna. This species was only found in reference sites ‘C’ and ‘G’ and was absent in all other sites through the sampling period.
Sampling Frequencies
Sampling data used to create the MDS ordination was then observed at various levels of organization in an attempt to identify sampling frequencies that most effectively measure fish communities in comparison to monthly sampling. Data was divided into three subsets using composite data from various months in order to represent different sampling frequencies over the sampling period.
Subsets included every month (September- March), every other month (September, November, January,
March), and every third month (September, December, March) for comparison with the MDS ordination representing the complete set of composite fish community data.
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Figure 17. The above figure is an MDS ordination including composite fish community data from the entire sampling period including the months of September, October, November, December, January, February, and March. The MDS ordination shows groupings of sites of various restoration condition and habitat type when sampled every month during the period of inundation.
Figure 18. The above figure is an MDS ordination including composite fish community data from every other month of sampling including the months of September, November, January, and March The MDS ordination shows groupings of sites of various restoration condition and habitat types when sampled every other month during the period of inundation. This ordination was observed in comparison to an ordination representing a sampling frequency of every month through the period of inundation.
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Figure 19. The above figure is an MDS ordination including composite fish community data from every third month of sampling including the months of September, December, and March. The MDS ordination shows groupings of sites of various restoration phases and habitat types when sampled every third month during the period of inundation. This ordination was observed in comparison to an ordination representing a sampling frequency of every month during the period of inundation.
Temporal Community Structure
MDS ordinations were produced from subsets of sampling data that were broken down
further into monthly composites of fish community data in an attempt to observe the time of year
most appropriate for collecting samples from a mature fish community. Monthly MDS
ordinations were observed in comparison with the composite fish community MDS ordination
over the entire sampling period for this study. Monthly MDS ordinations that most closely
resemble that of the MDS ordinations from the complete data set would provide an example of a
mature fish community from which to sample. It should be noted that the only months during
which every samplable site held enough water to support a fish community were the months of
October and November. All other months contained at least one site that did not hold enough
water to support a samplable fish community. The months of October and November are
represented in the MDS ordinations pictured below (Figure 20, Figure 21). Monthly MDS
ordinations for all other months and restoration phases may be found in Appendix D.
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Figure 20. The above figure is an MDS ordination including fish community data from the month of October at all samplable sites. The MDS ordination shows groupings of sites of various restoration phases and habitat types during the month of October. The ordination was then observed in comparison to an MDS ordination representing all sampling sites sampled through the entire sampling period.
Figure21. The above figure is an MDS ordination including fish community data from the month of November at all samplable sites. The MDS ordination shows groupings of sites of various restoration phases and habitat types during the month of October. The ordination was then observed in comparison to an MDS ordination representing all sampling sites sampled through the entire sampling period.
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Fish Community Structure
To observe patterns of fish community change throughout the period of inundation and among sites of different restoration phases, abundance data was analyzed in a visual representation using
Microsoft Excel (Table 7). Community data was also represented in MDS ordinations showing the relative abundance of the three species having the greatest impact on differences between sites of different restoration phases (Gambusia holbrooki, Jordonella floridae, Fundulus confluentus) over the sampling period. MDS ordinations for reference sites are pictured below (Figure 22, Figure 23, Figure
24).
Breder Dip Net Impacted Transitional Reference No Water Mosquito Fish (Gambusia holbroki) Site September October November December January February March Site September October November December January February March T3W1-Cg 7 16 T3W1-Cg 1 2 T3W2-C T3W2-C T3W1-G 1 3 T3W1-G 1 T3W5-C 4 325 T3W5-C 45 T3W5-Cg 7 96 T3W5-Cg 3 T3W5-G T3W5-G FS-C2 8 4 16 679 752 FS-C2 51 37 FS-C 7 111 23 413 FS-C 3 4 3 95 FS-Cg 2 23 9 4 95 16 FS-Cg 3 8 2 FS-G 2 36 34 6 FS-G 1 1 6 Flagfish (Jordanella floridae) Site September October November December January February March Site September October November December January February March T3W1-Cg T3W1-Cg T3W2-C T3W2-C T3W1-G T3W1-G T3W5-C 1 T3W5-C T3W5-Cg 2 T3W5-Cg T3W5-G T3W5-G FS-C2 1 FS-C2 2 FS-C 1 3 8 11 4 FS-C 1 1 FS-Cg 13 27 FS-Cg 4 FS-G 1 11 16 FS-G Marsh Killifish (Fundulus confluentus) Site September October November December January February March Site September October November December January February March T3W1-Cg 2 T3W1-Cg T3W2-C T3W2-C T3W1-G T3W1-G T3W5-C T3W5-C T3W5-Cg T3W5-Cg T3W5-G T3W5-G FS-C2 6 11 FS-C2 3 FS-C 1 25 18 24 FS-C 1 FS-Cg 1 1 3 93 5 FS-Cg 3 FS-G 1 1 9 FS-G Table 8. The above table shows relative abundance data for Gambusia holbrooki, Jordonella floridae, and Fundulus confluentus over the entire sampling period with dry months represented in yellow. The table is divided into two sampling methods including Breder traps and dip netting. The table was used to observe changes in abundance and community structure throughout the entire sampling period from month to month.
Monthly relative abundance data for these three species having the greatest impact on
dispersion patterns through multivariate space were observed to identify temporal patterns of -47-
changes in fish community structure through the dry down. Species abundance data for
Gambusia holbrooki, Jordonella floridae, and Fundulus confluentus showed general patterns of
increased abundance from inundation through the dry down. Gambusia holbrooki was the only
species ubiquitous to all sites where fish were captured. Earlier colonizers in reference and
transitional sites included Gambusia holbrooki and Jordonella floridae. Total abundance of these
species also varied among sites of varying restoration phases. Reference sites had a higher
abundance of all represented species throughout the period of inundation and especially during
sampling events occurring during the dry down.
Figure 22. The above figure is an MDS ordination representing reference sites of various habitat types sampled each month (indicated by month codes 9,10,11,12,1,2,and 3) during the entire sampling period with superimposed relative abundance data for Gambusia holbrooki for each represented month when sampling occurred. -48-
Figure23. The above figure is an MDS ordination representing reference sites of various habitat types sampled each month during the entire sampling period with superimposed relative abundance data for Jordonella floridae for each represented month when sampling occurred.
Figure24. The above figure is an MDS ordination representing reference sites of various habitat types sampled each month during the entire sampling period with superimposed relative abundance data for Fundulus confluentus for each represented month when sampling occurred.
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Temporal Dispersion
Temporal trajectories were superimposed over monthly MDS ordinations of fish community data for each restoration phase in order to observe temporal dispersion patterns through multivariate space
(Figure 25, Figure 26). Trajectories for restoration phases with sufficient data follow a similar dispersion pattern through multivariate space over the period of inundation. Dissimilarity between consecutive months for each restoration phase was observed through SIMPER analysis which is displayed in Table 8.
Reference sites show a decrease in dissimilarity (increase in similarity) from September until January when the dissimilarity slightly increases. Impacted and Transitional sites do not show apparent patterns of changing dissimilarity between consecutive months. Average dissimilarity between months was lowest in impacted sites, increased in transitional sites, and was highest in reference sites. These differences are displayed graphically on the MDS ordination trajectories and with a table of SIMPER analysis dissimilarity values from month to month.
Figure25. The above figure is an MDS ordination of monthly composite fish community data in transitional sites with superimposed trajectory to observe movement through multidimensional space and changes in community structure through the period of inundation. -50-
Figure26. The above figure is an MDS ordination of monthly composite fish community data in reference sites with superimposed trajectory to observe movement through multidimensional space and changes in community structure through the period of inundation.
Table 9. The table below shows dissimilarity values between consecutive months within various restoration phases and average dissimilarity for each restoration phase through the entire sampling period to observe changes in community structure through the period of inundation. An ‘X’ represents a lack of sufficient data to calculate dissimilarity values due to the dry down of sampling locations.
Monthly Dissimilarity
Impacted Transitional Reference September x x 62.67 October 22.93 36.06 43.18 November x 20.35 31.75 December x 33.77 18.41 January x 18.04 21.05 February x x 19.49 March
Average Dissimilarity 22.93 27.06 32.76
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Restoration Success
Temporal composite data for each of the sampled sites was analyzed using an MDS ordination and a cluster analysis with SIMPER in order to observe fish communities at sites in each conditional replicate and whether transitional sites undergoing restoration are moving from impacted conditions towards minimally impacted reference conditions. The MDS ordination shows three groupings of sites of the similar restoration condition. All sites displayed had <42% similarity overall. Using SIMPER analysis impacted sites showed 72% similarity, transitional sites showed 63% similarity, and reference sites showed 58% similarity. Cluster analysis showed impacted sites in a distinct grouping. A second grouping contained two transitional sites and one reference site all having >60% similarity. The third apparent grouping contained one transitional site and two reference sites that show >50% similarity with sites of other conditions.
Figure 27. The above figure is a cluster analysis showing restoration phase and habitat type using Bray-Curtis similarity from all samples. The cluster analysis shows two distinct groupings of sites with three additional sub-groupings. Major groupings include a cluster of impacted sites, transitional sites ‘C’ and ‘Cg’, and reference site ‘G’ at about 50% similarity. The second major cluster shows reference sites ‘C’ and ‘Cg’ as well as transitional site ‘C2’ at >50% similarity. Sub groupings include impacted sites at >70% similarity in one group, transitional sites ‘C’ and ‘Cg’ with reference site ‘G’ at >60% similarity in the second group, and transitional site ‘C2’ with reference sites ‘C’ and ‘Cg’ in a third sub grouping. -52-
Figure28. The above figure is an MDS ordination of composite fish community data for each sampling site representing various restoration phases and habitat types through the entire sampling period using Bray-Curtis similarity from all sampling events. The ordination through multivariate space is based on similarity values and places points in a three- dimensional multivariate space. The three dimensional image has been compressed into a 2D representation in order to represent the multivariate ordination effectively.
Figure29. The above figure is an MDS ordination of composite fish community data for each sampling site representing various restoration phases and habitat types through the entire sampling period including superimposed similarity groupings of 40 percent, 50 percent, and 60 percent similarity. Impacted sites show a distinct grouping at 50 percent and all transitional and reference sites group together at 40 percent similarity.
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DISCUSSION
The primary focus of this study was to evaluate the success of the restoration activities in the
Picayune Strand by obtaining and analyzing fish species richness and relative abundance in relation to various phases of restoration. Fish communities provide a means for evaluating the effects of changing hydrologic conditions on the ecological health of wetlands (Main et al.
2007). Therefore, the biological health of the various restoration phases in the Picayune Strand was measured by accessing the status and trends of fish populations.
The fish species richness and relative abundance data collected via active and passive sampling techniques varied among sites of different restoration phases. Total abundance was highest in transitional sites and lowest in impacted sites, while species richness was highest in reference wetlands and lowest at impacted sites. The higher species richness values within reference wetlands were expected because the associated hydrology and habitat quality are greater and more natural than that of the impacted and transitional sites. It should also be noted that 12 of the 15 species identified in this study had a higher relative abundance than in sites of both impacted and transitional restoration phases. These findings support previous research that fish can serve as performance indicators to monitor hydrologic restoration (Main et al. 2007).
The results of the cluster analysis showed two major clusters with one cluster containing two groups of <50% similarity (Figure 9). Of the two main clusters, the most similar cluster was comprised of two reference sites (cypress and cypress gramminoid) and one transitional site
(cypress). The overall cluster analysis showed that the transitional restored sites grouped primarily with natural reference sites, but shares <50% similarity with impacted un-restored sites. It appears that restored sites are transitioning between impacted un-restored sites towards a more natural reference condition. -54-
The non-metric MDS ordination showed distinct groupings of the sample locations within natural reference wetlands, transitional recently restored areas, and impacted un-restored areas as each restoration phase appeared close together in 2D space (Figure 29). SIMPER analysis showed that Reference sites were the least similar (58 percent), transitional sites were slightly more similar (63 percent), and impacted sites were the most similar (72 percent). This may be a result of the low species richness found at impacted sites, and increasing species richness in transitional and reference sites. SIMPER analysis also showed that impacted and transitional sites were 53 percent dissimilar, impacted and reference sites were 69 percent dissimilar, and transitional and reference sites were 48 percent dissimilar. This indicated that transitional sites are more similar to reference sites than they are with impacted sites. This can be observed on the MDS ordination as impacted sites tightly group in the upper left, transitional sites group in the center, and reference sites group in the upper right of the multidimensional space. This may indicate that transitional sites are moving towards reference conditions, at least within multidimensional ecological space.
Breder trap and dip-net sampling provided data on the fish communities found in the
Picayune Strand which was identified as necessary data by the DOI (2004) Science Plan.
Sampling identified a total of 15 fish species throughout the Picayune and Fakahatchee Strands.
All 15 species were found in reference wetlands, six species were found within transitional restored sites, and two species were found within impacted un-restored sites. Native species found only in reference wetlands included the Everglades pygmy sunfish (Elassoma evergladei), golden topminnow (Fundulus chrysotus), least killifish (Heterandria formosa), spotted sunfish
(Lepomis macrochirus), dollar sunfish (Lepomis marginatus), coastal shiner (Notropis petersoni), and sailfin molly (Poecilia latipinna). Native species also found in transitional sites included marsh killifish (Fundulus confluentus), mosquito fish (Gambusia holbrooki), Florida -55- flagfish (Jordonella floridae), juvenile sunfish species (Lepomis sp. juv), and bluefin killifish
(Lucania goodei) of which only mosquito fish and marsh killifish were also found in impacted un-restored sites. Exotic species found in reference sites included the pike killifish (Belonesox belizanus) which was also found in transitional sites, Mayan cichlid (Cichlasoma urophthalmus), and jewelfish cichlids (Hemichromis letourneuxi). Fish populations in the Everglades are driven mostly by hydroperiod (Ruetz 2005). Two exceptions to this general pattern are Jordonella floridae and Gambusia holbrooki as their primary mechanism for synchronous population dynamics is dispersal due to their higher tolerance for low dissolved oxygen and harsh conditions. Poecilia latipinna is another species that can tolerate low dissolved oxygen and harsh water conditions, though they are not known for using active dispersal as a primary means of recolonization. The majority of species rely on dry season refugia to repopulate inundated areas through the wet season. It should be noted that Fundulus confluentus is also a rapid colonizer of newly inundated areas through active dispersal (Trexler 2010).
Species found in sites of varying restoration phases can be associated with these species specific dispersal patterns. Impacted sites contained only species that use active dispersal as a recolonization strategy, indicating a lack of connected deep-water refugia to newly inundated areas of the represented natural habitats (cypress, cypress gramminoid, gramminoid).
Transitional sites contained these species as well as species that require dry season refugia in order to recolonize inundated areas including Lepomis species and Lucania goodei which may indicate greater connectivity in the region resulting from recent restoration activities. Figure 30 shows habitat maps of the Picayune Strand from 1940 and from 1995 showing the changes in connectivity and habitat composition as a result of the original development project (CERP
2004). Many of the areas that may have provided dry season refugia to fish species prior to the original construction have been converted to areas that may not provide the same functions to -56- fish communities as a result of altered hydrology. For example, one of the sites intended for this study in an impacted un-restored area contained a vegetative community characteristic of a deep cypress habitat, but did not retain enough water during the wet season to support a fish community.
With these altered conditions, it is also important to consider environmental factors that area associated with various restoration phases which could influence fish species richness and relative abundance values. Differences in water quality parameters, hydroperiods, predator prevalence, and water flows vary among restoration phases which may have contributed to the fish community composition within phases. For example, the population density, size structure, and relative abundance of fishes in South Florida wetlands are regulated by the annual duration of uninterrupted flooding according to Ogden et al. (2005). Ogden goes one to explain that fishes such as sunfishes, golden topminnows, and bluefin killifish are less represented in areas with shorter periods of inundation. Loss of connectivity and sheetflow in the Picayune Strand may explain the differences in fish community composition within sites of different restoration phases. “Channelization and compartmentalization of the previous sheet-flow-driven system have resulted in major changes in physical and hydrologic processes, resulting in far-reaching effects on ecological processes and habitat.” (Ogden et al. 2005).
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Figure30. The figure above shows habitat maps of the Picayune Strand from 1940 and 1995 in order to observe the changes that have occurred over time as a result of hydrologic alterations. Various habitat types are represented using colors and patterns which have been superimposed onto a map of the Picayune Strand Restoration Project (COE & SFWMD 2004). -58-
To further examine some of the environmental factors driving change in fish
community structure and composition, a principle component analysis (PCA) was
conducted using environmental data collected during sampling events. Figure 31 shows
an MDS ordination of all sampling events for sites of each habitat and restoration phase
representing environmental data collected for each event. Variables included dissolved
oxygen, temperature, average depth, salinity, and average conductivity. Distributions of
sampling events on the MDS ordination appear to be separated along the axes of both
average depth and average conductivity. Impacted sites appear to group as a result of a
low average depth, while transitional and reference sites appear to group on either side of
both the average depth and average conductivity axes throughout the sampling period.
This ordination may indicate that average depth within sites may be the driving factor
influencing species abundance and relative abundance variations among site of various
restoration phases.
Figure31. The above figure is a principal component analysis including an MDS ordination of all sampling events of various habitats and restoration phases with a superimposed representation of environmental factors including dissolved oxygen, average depth, average conductivity, salinity, and temperature. Location of points in comparison to the various environmental factors represented by lines allows for the observation of which factors are driving the differences between sampling events.
-59-
Potential Indicator Species
As hydrologic restoration continues in the Picayune Strand, fish can provide useful information to monitor the changes in the ecological health of the areas affected (Main et al.
2007; Ceilley 2008). Hydrology is a major component in fish species abundance and relative abundance as indicated by abundance data and principle component analysis. To further link the effects of hydrologic restoration to fish community data, several potential indicator species have been identified. Species that contributed the most to the dissimilarity between impacted, transitional, and reference sites included Gambusia holbrooki, Fundulus confluentus, and
Jordonella floridae. All of these species are known to use active dispersal as a method of recolonizing newly inundated areas (Trexler 2010; Main et al. 2007) and are often pioneer species as they are more tolerant of harsh water conditions than many other species of fish found in this study (Ruetz et al. 2005). Active dispersal requires connectivity between habitats and the presence of these species may indicate that they are connected to adjacent areas containing source populations. It should be noted that these species do not need a long period of inundation in order to colonize a newly inundated area due to their tolerance and active method of recolonization. Therefore, additional indicator species may be required in order to observe trends in both connectivity and period of inundation in various habitats and restoration phases as restoration continues. Species such as Fundulus chrysotus, Lucania goodei, Lepomis species, and Elassoma evergladei have been identified as indicators of longer periods of inundation as frequent dry downs appear to limit the density of these species (Ruetz et al. 2005; Ceilley 2008).
Heterandria formosa has also been noted for having a response to drought similar to that of
Fundulus chrysotus and Lucania goodei, but was found to have a weaker association with period of inundation than the other species (Ruetz et al. 2005).
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Sampling Frequencies
When conducting ecological monitoring, it is important to consider sampling effort in order to determine the resources necessary to effectively carry out monitoring activities.
“Ecologists have long sought out concise and cost-effective measures that can characterize ecosystem conditions,” (Schiller 2001). Data collected via monthly monitoring during this study was observed in various subsets in order to identify sampling frequencies that most effectively and efficiently measure fish communities in comparison to monthly sampling. The data was divided into subsets representing every other month (September, November, January, March), every second month (October, December, February), and every third month (November,
February) and analyzed using non-metric MDS ordination plots. MDS ordinations were observed against the composite data including all data collected from monthly fish sampling in order to identify ordinations that most closely represented that of the complete data set. The sampling frequency that most closely resembled the MDS ordination of the complete data set was the subset including data from every second month (October, December, February) followed by the subset that included data from every other month (September, November, January,
March). During the 2011/2012 wet season a sampling frequency of every other month most closely resembled the complete data set as the MDS ordination groupings were the most similar to that of the original. It should be noted that additional sampling would be necessary to establish trend data that supports this sampling frequency as the most effective over several seasons due to hydrologic pattern variations from season to season. An established sampling frequency containing a smaller number of sampling events would significantly lower sampling effort and provide a more concise and cost-effective means to measure ecosystem conditions using fish communities.
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Temporal Sampling
In addition to identifying a sampling frequency that best represents the fish community through the entire period of inundation, it is also important to identify the time of year when sampling efforts most effectively measure a mature fish community. Subsets of fish data collected during each sampling month were analyzed with non-metric MDS ordinations in order to identify months that most closely resembled the MDS ordination including the entire data set.
Due to the short periods of inundation in impacted sites, the only month that contained fish data from all sites was November. The month of November was recommended for mature sampling by Ceilley (2008) and Duever (personal communication). The month of October contained data from all sites except for the transitional cypress #2 site as no fish were trapped during that sampling event. When compared with the MDS ordination containing data from all sampling events, the ordination that most closely resembles the complete ordination groupings is the month of October. Even though there was lack of complete data for the month of October, it was determined that both October and November would provide fish community data that best represents a mature fish community. Additional sampling would be required to establish these months as the most effective for sampling a mature fish community as hydrologic patterns vary from year to year.
Fish Community Structure
Fish community data was also used to observe how fish communities change through the period of inundation. Monthly sampling data for each site and restoration phase was analyzed with non-metric MDS ordinations and superimposed relative abundance data, and a visual representation of total abundance data for the three fish species (Gambusia holbrooki, Fundulus confluentus, Jordonella floridae) that had the greatest impact on ordinations identified using -62-
SIMPER analysis. It should be noted that data from impacted sites was not used in this analysis as total abundance for each species was too small to observe change. Abundance data showed a general pattern of increasing total abundance for all three species from inundation through the dry-down. This pattern was also observed by DeAngelis et al. (2010), who stated that seasonal fluctuations of water level in the Everglades allows for the expansion of small fishes over an extensive flooded area during the wet season, and surviving small fishes then tend to be concentrated in smaller water bodies when water levels fall during the dry season, making fish biomass available to higher trophic levels such as wading birds. Abundance data also showed that Gambusia holbrooki and Jordonella floridae were early colonizers of newly inundated areas as they were documented early in the sampling period.
Temporal dispersion
To further examine patterns of fish community changes through the period of inundation, temporal trajectories were superimposed over monthly MDS ordinations of fish community data for transitional and reference restoration phases in order to observe temporal dispersion patterns through multivariate space, and community changes over time (Figures 25-26). It should be noted that impacted sites could not be shown with a trajectory due to lack of data resulting from the dry-down. The trajectory for reference and transitional sites are used to create a visual of month to month variation within sites over the entire sampling period and can be used in future studies to observe trajectory patterns in multidimensional ecological space. To further examine this pattern, SIMPER analysis was conducted to observe dissimilarity from month to month in each restoration phase (Table 6). Average month-to-month dissimilarity between sites that provided sufficient data was lowest within impacted sites (22.93 percent) and increased as restoration progressed to transitional sites (27.06 percent), and finally reference sites (32.76 percent). Reference sites also show a pattern of decreasing dissimilarity from month-to-month -63- through the dry-down with the exception of January-February. The pattern shows that the fish community is becoming more similar among reference sites as fish species become concentrated into smaller areas with decreasing water levels. The jump in dissimilarity may be a function of this concentration of species into smaller areas with the dry-down, which resulted in the capture of species that were previously un-documented. Transitional sites showed a similar pattern of decreasing dissimilarity with an increase in dissimilarity occurring in December-January. This difference may be a result of a shorter overall period of inundation than that of reference sites.
Impacted sites provided data for only two months during the sampling period which only allowed for a single dissimilarity value between the months of October and November.
Increasing average dissimilarity values from impacted sites through reference sites may be a result of a more diverse fish community that can be found in reference sites, and a recovering fish community in transitional sites. This pattern may provide a reference for future studies on changes in fish community structure though the dry-down, and can be used to assess restoration activities in ephemeral wetlands.
Restoration Success
Fish species abundance and relative abundance data for all sites of each restoration phase was analyzed to observe the effects of hydrologic restoration in the Picayune Strand after the initial phases of restoration were completed. Cluster analysis and MDS ordinations were created for composite fish data from sites of different habitats though each of the impacted un-restored sites, transitional recently restored sites, and natural reference wetlands to observe whether fish communities in areas undergoing restoration are moving from impacted conditions towards minimally impacted reference conditions. The resulting figures (Figures 27-29) showed distinct groupings among sites of similar restoration phases. It is important to note that transitional sites and reference sites also grouped together in the MDS ordination at a minimum of 40 percent -64- similarity and a maximum of 60 percent similarity. Impacted sites grouped together at 40 percent similarity, and only showed 20 percent similarity to groups of other restoration phases.
Groupings of transitional and reference sites show a shift from impacted conditions prior to restoration towards a more natural condition, confirming that the hydrology of transitional sites is successfully being restored as restoration efforts continue.
It should be noted that transitional site C2 showed fish community data that very closely resembled reference wetland sites through the period of inundation. Transitional site C2 was selected based on its proximity to Prairie Canal and known hydrologic impacts resulting from the large development activities in the area. The site is located slightly south of the additional transitional sites about one mile east of Prairie Canal. Transitional C2 exhibited a longer period of inundation than other sites of the transitional restoration phase, which may be a result of its location further downstream. Downstream areas tend to flood sooner and retain water longer through the dry season due to water’s stacking effect moving upstream and being situated slightly closer to sea level, which may have given rise to a more mature fish community and a more rapid overall restoration of the site.
Cluster analysis shows transitional cypress gramminoid and cypress sites grouping with the gramminoid reference site. Gramminoid habitats are generally known to have a shorter period of inundation than that of both cypress gramminoid and cypress habitats, which may indicate that transitional sites continue to show impairment from the original hydrologic alterations in the Picayune Strand. Another potential reason for this could be the unrestored areas to the west of the transitional sites that may be continuing to impact the hydrology of the newly restored areas. Additionally, this finding may be a result of restoration “time lag”, which is the time between restoration and recovery when newer restored areas may be more susceptible to disturbance that can hinder the overall goals of restoration. We must recognize that -65- restoration itself is a disturbance and the newly establishing systems need time to organize and move toward more complex, stable, native communities. It is expected that through further restoration of impacted areas to the west in addition to increased recovery time, transitional habitat areas will continue to move towards more natural reference conditions. Continued fish sampling will be required to observe these changes as restoration continues.
Future Research Opportunities
A multi-agency Restoration Coordination and Verification Team (RECOVER) was established to support the implementation of CERP with science that can be integrated into management components through research, monitoring, modeling, planning, assessment, and adaptive management (Dixon 2009; DOI 2005). Fish community monitoring is a critical component of RECOVER and was identified to be an important aspect of the adaptive assessment process set forth in the goals of CERP (USGS 2004).
A major problem that conservation biology faces is the lack of baseline data that exists which is needed to measure and assess population changes (Dixon 2009; Busby & Parmelee
1996). However, baseline fish data is available within the Picayune and Fakahatchee Strand
Preserve is available (Addison et al. 2006). Additional fish data was also collected during the first phase of restoration (Ceilley 2008). Experimental design used during baseline studies and follow up studies during restoration was replicated in this study, allowing for the comparative studies within the Picayune and surrounding areas. By, using this data with additional monitoring, temporal trends in species richness and abundance can be evaluated (Dixon 2009).
The previous studies within the Picayune Strand help to support the results of this study. When reviewed in combination, these studies can make future fish studies stronger by evaluating trends in fish community composition over time. Therefore, fish monitoring within the Picayune -66-
Strand should be continued in the future to further observe and document ecological responses to the hydrologic changes (DOI 2005).
Future research can be improved by adapting techniques and sampling methods based on experiences from this study and previous research. If additional fish surveys are conducted, different sampling frequencies may be employed in maximize data quality and minimize sampling effort by utilizing data from this study. Future restrictions in monetary support and man-power may result in a minimal sampling opportunity which can be optimized by sampling during the recommended time periods identified in this study.
Additional sampling sites are recommended for future sampling efforts in order to minimize and offset the amount of sites chosen that may retain enough water to support a samplable fish community in a given year. Further additional sites may also be added to examine the effects of hydrologic restoration on fish communities in additional habitat types such as hydric pine flatwoods and downstream habitats. The addition of downstream habitats would provide an opportunity to observe fish community changes as a result of additional environmental factors such as conductivity, salinity, and water flow. It is also recommended that sampling is continued for the duration of inundation to provide a complete data set that includes the entire period of inundation through dry-down. Sampling of source populations and dry season refugia may also provide valuable data on hydrologic connectivity and dispersion of fish species across the landscape post inundation. Additional research on indicator species is also recommended to determine if dispersion is directly linked to hydrologic connectivity, restoration, and/or dry season refugia in the Picayune Strand.
Access to sampling sites at the height of the wet season was difficult due to high water and lack of a sufficient 4WD vehicle. It is recommended that a swamp buggy or access to -67- additional forms of off road transportation is arranged prior to the onset of the wet season to ensure access to sampling sites.
Since fish are only one part of the overall aquatic fauna affected by the restoration in the
Picayune Strand, future research should incorporate the collection of other aquatic fauna including macroinvertebrates and anurans. Long-term monitoring is needed for fish and other taxa to determine if a continued display of positive responses to hydrologic restoration over time persists. Aquatic faunal research should continue through the process of restoration and after the restoration is complete in the areas highlighted in this study as well as additional areas undergoing hydrologic restoration in the Picayune. Future studies, complimented by this and previous studies, will continue to document restoration progress and feasibility to improve future monitoring efforts and inform decision making. This document could potentially aid in justification for continued funding, support for funding and management decisions, and assist with acquiring new funding sources within the Picayune Strand.
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CONCLUSIONS
After careful analysis and review of the fish data collected as part of this research project, the following conclusions can be made:
1) Restoring hydrology within the Picayune by the plugging of Prairie Canal recovered
fish habitat and hydrologic connectivity which resulted in increased species richness
and relative abundance within restored areas.
2) There were significant differences in the distribution of fish species relative to the
restoration phases: specifically Gambusia holbrooki, Fundulus confluentus, and
Jordonella floridae appear to be common indicator species of hydrologic
connectivity. Fundulus chrysotus, Lucania goodei, and Elassoma evergladei were
also unique to reference sites and may also be used as potential indicator species.
3) Sampling frequencies of every other month provide a similar signal to that of monthly
sampling, and may be adequate for collection of fish community data which has
implications for sampling effort.
4) The months of October and November provided samples of a mature fish community
which indicates the most effective time of year for sampling events.
5) Fish community structure and abundance changes throughout the period of inundation
as fish are concentrated into smaller areas by the dry-down. These patterns may
provide a snapshot of overall abundance and identify pioneer species that are early
colonizers of newly inundated areas.
6) Reference wetlands become more similar through the period of inundation and dry-
down while average month to month similarity increases as restoration progresses.
7) Transitional recently restored sites are moving towards more natural reference
conditions as restoration continues and hydrology/connectivity improves. -69-
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A-1
APPENDIX A
FISH SAMPLING DATA FORM
Picayune Strand Aquatic Fauna Field Data Sheet
Site ID______Sky Condition______Date______Air Temp:______Field Crew______Wind:______Inverts Collected: Yes No Sampling Start Time______Invert Start______Breder Start______Sampling Finish Time______Invert Finish______Breder Finish______DO:______Salinity:______pH:______SI:______Temp:______Trap No. Depth(cm) Species Count Total
Dipnet & Visuals for fish/amphibians Species Number Comments
Comments:______B-1
APPENDIX B
FISH SAMPLING DATA
Appendix B-1. Species table with species codes
Family Genus Species Common Name Species Code Lepisosteidae Lepisosteus platyrhincus Florida gar LEPL Ictaluridae Ameiurus natalis Yellow bullhead AMNA Callichthyidae Hoplosternum littorale Brown hoplo HOLI Cichlidae Cichlasoma bimaculatum Black acara CIBI Cichlidae Cichlasoma urophthalmus Mayan cichlid CIUR Cichlidae Hemichromis letourneuxi Jewelfish Cichlid HELI Cichlidae Oreochromis aureus Blue tilapia ORAU Fundulidae Fundulus chrysotus Golden Topminnow FUCR Fundulidae Fundulus confluentus Marsh Killifish FUCO Fundulidae Lucania goodei Bluefin killifish LUGO Poeciliidae Belonesox belizanus Pike Killifish PKKF Poeciliidae Gambusia holbrooki Mosquitofish GAHO Poeciliidae Heterandria formosa Least killifish HEFO Poeciliidae Poecilia latipinna Sailfin molly POLA Cyprinodontidae Jordanella floridae Flagfish JOFL Centrarchidae Lepomis sp. (juv) Sunfish juv. LESP Centrarchidae Lepomis gulosus Warmouth LEGU Centrarchidae Lepomis macrochirus Bluegill LEMAC Centrarchidae Lepomis marginatus Dollar sunfish LEMAR Centrarchidae Lepomis microlophus Redear sunfish LEMI Centrarchidae Lepomis punctatus Spotted sunfish LEPU Centrarchidae Micropterus salmoides Largemouth bass MISA Percidae Etheostoma fusiforme Swamp darter ETFU Clariidae Clarias batrachus Walking catfish CLBA Soleidae Trinectes maculatus Hogchoker TRMA Cyprinidae Notropis petersoni Coastal shiner NOPE Elassomatidae Elassoma evergladei Everglades pygmy sunfish ELEV Loricariidae Pterygoplichthys sp. (juv) Ptero PTSP
B-2
Appendix B-2. Fish sampling data – Impacted Cypress Gramminoid (9/21/2011 – 3/13/2011)
Species Impacted- Impacted- Impacted- Impacted- Impacted- Impacted- Impacted- Code Cg- Cg- Cg- Cg- Cg- Cg- Cg- 9-21-11 10-24-11 11-10-11 12-12-11 1-13-12 2-14-12 3-13-11 LEPL 0 0 0 0 0 0 0 AMNA 0 0 0 0 0 0 0 HOLI 0 0 0 0 0 0 0 CIBI 0 0 0 0 0 0 0 CIUR 0 0 0 0 0 0 0 HELI 0 0 0 0 0 0 0 ORAU 0 0 0 0 0 0 0 FUCR 0 0 0 0 0 0 0 FUCO 0 0 2 0 0 0 0 LUGO 0 0 0 0 0 0 0 PKKF 0 0 0 0 0 0 0 GAHO 0 8 18 0 0 0 0 HEFO 0 0 0 0 0 0 0 POLA 0 0 0 0 0 0 0 JOFL 0 0 0 0 0 0 0 LESP 0 0 0 0 0 0 0 LEGU 0 0 0 0 0 0 0 LEMAC 0 0 0 0 0 0 0 LEMAR 0 0 0 0 0 0 0 LEMI 0 0 0 0 0 0 0 LEPU 0 0 0 0 0 0 0 MISA 0 0 0 0 0 0 0 ETFU 0 0 0 0 0 0 0 CLBA 0 0 0 0 0 0 0 TRMA 0 0 0 0 0 0 0 NOPE 0 0 0 0 0 0 0 ELEV 0 0 0 0 0 0 0 PTSP 0 0 0 0 0 0 0
B-3
Appendix B-3. Fish sampling data – Impacted Gramminoid (9/21/2011 – 3/17/2011)
Species Impacted- Impacted- Impacted- Impacted- Impacted- Impacted- Impacted- Code G-9-21-11 G-10-24- G-11-10- G-12-12- G-1-13-12 G-2-14-12 G-3-17-12 11 11 11 LEPL 0 0 0 0 0 0 0 AMNA 0 0 0 0 0 0 0 HOLI 0 0 0 0 0 0 0 CIBI 0 0 0 0 0 0 0 CIUR 0 0 0 0 0 0 0 HELI 0 0 0 0 0 0 0 ORAU 0 0 0 0 0 0 0 FUCR 0 0 0 0 0 0 0 FUCO 0 0 0 0 0 0 0 LUGO 0 0 0 0 0 0 0 PKKF 0 0 0 0 0 0 0 GAHO 0 11 3 0 0 0 0 HEFO 0 0 0 0 0 0 0 POLA 0 0 0 0 0 0 0 JOFL 0 0 0 0 0 0 0 LESP 0 0 0 0 0 0 0 LEGU 0 0 0 0 0 0 0 LEMAC 0 0 0 0 0 0 0 LEMAR 0 0 0 0 0 0 0 LEMI 0 0 0 0 0 0 0 LEPU 0 0 0 0 0 0 0 MISA 0 0 0 0 0 0 0 ETFU 0 0 0 0 0 0 0 CLBA 0 0 0 0 0 0 0 TRMA 0 0 0 0 0 0 0 NOPE 0 0 0 0 0 0 0 ELEV 0 0 0 0 0 0 0 PTSP 0 0 0 0 0 0 0
B-4
Appendix B-4. Fish sampling data – Transitional Cypress Gramminoid (9/21/2011 – 3/17/2011)
Species Transitional Transitional Transitional Transitional Transitional Transitional Transitional Code -Cg- -Cg- -Cg- -Cg- -Cg- -Cg- -Cg- 9-21-11 10-25-11 11-17-11 12-12-11 1-13-12 2-14-12 3-17-12 LEPL 0 0 0 0 0 0 0 AMNA 0 0 0 0 0 0 0 HOLI 0 0 0 0 0 0 0 CIBI 0 0 0 0 0 0 0 CIUR 0 0 0 0 0 0 0 HELI 0 0 0 0 0 0 0 ORAU 0 0 0 0 0 0 0 FUCR 0 0 0 0 0 0 0 FUCO 0 0 0 0 0 0 0 LUGO 0 0 0 0 0 0 0 PKKF 0 0 0 0 0 0 0 GAHO 0 10 96 0 0 0 0 HEFO 0 0 0 0 0 0 0 POLA 0 0 0 0 0 0 0 JOFL 0 0 2 0 0 0 0 LESP 0 0 0 0 0 0 0 LEGU 0 0 0 0 0 0 0 LEMAC 0 0 0 0 0 0 0 LEMAR 0 0 0 0 0 0 0 LEMI 0 0 0 0 0 0 0 LEPU 0 0 0 0 0 0 0 MISA 0 0 0 0 0 0 0 ETFU 0 0 0 0 0 0 0 CLBA 0 0 0 0 0 0 0 TRMA 0 0 0 0 0 0 0 NOPE 0 0 0 0 0 0 0 ELEV 0 0 0 0 0 0 0 PTSP 0 0 0 0 0 0 0
B-5
Appendix B-5. Fish sampling data – Transitional Cypress (9/21/2011 – 3/17/2011)
Transitional Transitional Transitional Transitional Transitional Transitional Transitional Species -C- -C- -C- -C- -C- -C- -C- Code 9-21-11 10-25-11 11-17-11 12-12-11 1-13-12 2-14-12 3-17-12 LEPL 0 0 0 0 0 0 0 AMNA 0 0 0 0 0 0 0 HOLI 0 0 0 0 0 0 0 CIBI 0 0 0 0 0 0 0 CIUR 0 0 0 0 0 0 0 HELI 0 0 0 0 0 0 0 ORAU 0 0 0 0 0 0 0 FUCR 0 0 0 0 0 0 0 FUCO 0 0 0 0 0 0 0 LUGO 0 0 0 1 0 0 0 PKKF 0 0 0 0 0 0 0 GAHO 0 0 4 370 0 0 0 HEFO 0 0 0 0 0 0 0 POLA 0 0 0 0 0 0 0 JOFL 0 0 0 1 0 0 0 LESP 0 0 0 0 0 0 0 LEGU 0 0 0 0 0 0 0 LEMAC 0 0 0 0 0 0 0 LEMAR 0 0 0 0 0 0 0 LEMI 0 0 0 0 0 0 0 LEPU 0 0 0 0 0 0 0 MISA 0 0 0 0 0 0 0 ETFU 0 0 0 0 0 0 0 CLBA 0 0 0 0 0 0 0 TRMA 0 0 0 0 0 0 0 NOPE 0 0 0 0 0 0 0 ELEV 0 0 0 0 0 0 0 PTSP 0 0 0 0 0 0 0
B-6
Appendix B-6. Fish sampling data – Transitional Cypress 2 (9/30/2011 – 3/17/2011)
Transitional Transitional Transitional Transitional Transitional Transitional Transitional Species -C2- -C2- -C2- -C2- -C2- -C2- -C2- Code 9-30-11 10-27-11 11-14-11 12-7-11 1-27-12 2-14-12 3-17-12 LEPL 0 0 0 0 0 0 0 AMNA 0 0 0 0 0 0 0 HOLI 0 0 0 0 0 0 0 CIBI 0 0 0 0 0 0 0 CIUR 0 0 0 0 0 0 0 HELI 0 0 0 0 0 0 0 ORAU 0 0 0 0 0 0 0 FUCR 0 0 0 0 0 0 0 FUCO 0 0 0 0 6 14 0 LUGO 0 0 0 0 0 0 0 PKKF 0 0 0 0 0 3 0 GAHO 0 8 4 16 730 789 0 HEFO 0 0 0 0 0 0 0 POLA 0 0 0 0 0 0 0 JOFL 0 0 0 1 0 2 0 LESP 1 0 0 0 0 0 0 LEGU 0 0 0 0 0 0 0 LEMAC 0 0 0 0 0 0 0 LEMAR 0 0 0 0 0 0 0 LEMI 0 0 0 0 0 0 0 LEPU 0 0 0 0 0 0 0 MISA 0 0 0 0 0 0 0 ETFU 0 0 0 0 0 0 0 CLBA 0 0 0 0 0 0 0 TRMA 0 0 0 0 0 0 0 NOPE 0 0 0 0 0 0 0 ELEV 0 0 0 0 0 0 0 PTSP 0 0 0 0 0 0 0
B-7
Appendix B-7. Fish sampling data – Reference Cypress Gramminoid (9/15/2011 – 3/17/2011)
Reference Reference Reference Reference Reference Reference Reference Species -Cg- -Cg- -Cg- -Cg- -Cg- -Cg- -Cg- Code 9-15-11 10-13-11 11-7-11 12-5-11 1-13-12 2-7-12 3-17-12 LEPL 0 0 0 0 0 0 0 AMNA 0 0 0 0 0 0 0 HOLI 0 0 0 0 0 0 0 CIBI 0 0 0 0 0 0 0 CIUR 0 1 0 0 0 0 0 HELI 0 14 0 1 0 0 0 ORAU 0 0 0 0 0 0 0 FUCR 0 0 0 1 0 0 0 FUCO 0 1 1 3 96 50 0 LUGO 0 1 0 0 0 0 0 PKKF 0 2 0 0 0 0 0 GAHO 2 23 9 43 103 18 0 HEFO 0 2 1 2 1 0 0 POLA 0 0 0 0 9 0 0 JOFL 0 0 0 13 31 0 0 LESP 0 4 3 2 0 0 0 LEGU 0 0 0 0 0 0 0 LEMAC 0 0 0 0 0 0 0 LEMAR 4 0 0 0 0 0 0 LEMI 0 0 0 0 0 0 0 LEPU 0 0 0 0 0 0 0 MISA 0 0 0 0 0 0 0 ETFU 0 0 0 0 0 0 0 CLBA 0 0 0 0 0 0 0 TRMA 0 0 0 0 0 0 0 NOPE 0 1 0 0 0 0 0 ELEV 0 0 0 0 0 0 0 PTSP 0 0 0 0 0 0 0
B-8
Appendix B-8. Fish sampling data – Reference Gramminoid (9/15/2011 – 3/17/2011)
Reference Reference Reference Reference Reference Reference Reference Species -G- -G- -G- -G- -G- -G- -G- Code 9-15-11 10-13-11 11-7-11 12-5-11 1-13-12 2-14-12 3-17-12 LEPL 0 0 0 0 0 0 0 AMNA 0 0 0 0 0 0 0 HOLI 0 0 0 0 0 0 0 CIBI 0 0 0 0 0 0 0 CIUR 0 0 0 0 0 0 0 HELI 0 0 0 0 0 0 0 ORAU 0 0 0 0 0 0 0 FUCR 0 0 0 0 0 0 0 FUCO 0 1 1 9 0 0 0 LUGO 0 0 0 0 0 0 0 PKKF 0 0 0 0 0 0 0 GAHO 3 37 34 66 0 0 0 HEFO 0 0 0 0 0 0 0 POLA 0 0 1 3 0 0 0 JOFL 1 11 0 16 0 0 0 LESP 0 0 0 0 0 0 0 LEGU 0 0 0 0 0 0 0 LEMAC 0 0 0 0 0 0 0 LEMAR 0 0 0 0 0 0 0 LEMI 0 0 0 0 0 0 0 LEPU 0 0 0 0 0 0 0 MISA 0 0 0 0 0 0 0 ETFU 0 0 0 0 0 0 0 CLBA 0 0 0 0 0 0 0 TRMA 0 0 0 0 0 0 0 NOPE 0 0 0 0 0 0 0 ELEV 0 0 0 0 0 0 0 PTSP 0 0 0 0 0 0 0
B-9
Appendix B-9. Fish sampling data – Reference Cypress (9/30/2011 – 3/13/2011)
Reference Reference Reference Reference Reference Reference Reference Species -C- -C- -C- -C- -C- -C- -C- Code 9-30-11 10-27-11 11-14-11 12-7-11 1-13-12 2-7-12 3-13-12 LEPL 0 0 0 0 0 0 0 AMNA 0 0 0 0 0 0 0 HOLI 0 0 0 0 0 0 0 CIBI 0 0 0 0 0 0 0 CIUR 0 0 0 0 0 0 0 HELI 0 0 0 0 0 0 0 ORAU 0 0 0 0 0 0 0 FUCR 0 0 0 0 0 0 1 FUCO 0 0 0 1 25 18 25 LUGO 0 0 0 0 0 0 0 PKKF 0 0 0 0 0 0 0 GAHO 0 0 3 7 115 206 508 HEFO 0 0 0 0 0 0 0 POLA 0 0 0 0 0 0 0 JOFL 0 1 0 3 9 11 5 LESP 2 1 1 0 1 2 13 LEGU 0 0 0 0 0 0 0 LEMAC 0 0 0 0 0 1 1 LEMAR 0 0 0 0 0 0 0 LEMI 0 0 0 0 0 0 0 LEPU 0 0 0 0 0 0 0 MISA 0 0 0 0 0 0 0 ETFU 0 0 0 0 0 0 0 CLBA 0 0 0 0 0 0 0 TRMA 0 0 0 0 0 0 0 NOPE 0 0 0 0 0 0 0 ELEV 0 0 0 0 0 0 2 PTSP 0 0 0 0 0 0 0
C-1
APPENDIX C
TOTAL FISH ABUNDANCE
Total Numbers of Fish - Breder Traps
Site September October November December January February March Impacted Cg 7 18 Impacted Cg 0 Impacted G 1 3 Transitional C 0 4 327 Transitional Cg 7 98 Transitional G 0 0 Transitional C2 1 8 4 17 685 766 Reference C 2 2 1 11 145 55 455 Reference Cg 6 49 14 26 225 21 Reference G 3 48 36 34 Total Numbers of Fish - Dip Netting
Site September October November December January February March Impacted Cg 1 2 Impacted Cg 0 Impacted G 1 0 Transitional C 0 0 45 Transitional Cg 3 0 Transitional G 0 0 Transitional C2 0 0 0 0 51 42 Reference C 0 0 3 0 5 3 100 Reference Cg 0 0 0 3 15 2 Reference G 1 1 0 6 Total Numbers of Fish - Composite
Site September October November December January February March Impacted Cg 0 8 20 0 0 0 0 Impacted Cg 0 0 0 0 0 0 0 Impacted G 0 2 3 0 0 0 0 Transitional C 0 0 4 372 0 0 0 Transitional Cg 0 10 98 0 0 0 0 Transitional G 0 0 0 0 0 0 0 Transitional C2 1 8 4 17 736 808 0 Reference C 2 2 4 11 150 58 555 Reference Cg 6 49 14 29 240 23 0 Reference G 4 49 36 40 0 0 0
D-1
APPENDIX D
RESTORATION PHASES:
CLUSTER ANALYSIS AND MDS ORDINATIONS
Appendix D-1 Impacted Cluster Analysis and MDS Ordination
D-3
Appendix D-2 Transitional Cluster Analysis and MDS Ordination
D-3
Appendix D-3 Reference Cluster Analysis and MDS Ordination