LAKES AND PONDS OF THE WORCESTER-MODNADNOCK PLATEAUECOREGION IN MASSACHUSETTS: COMPARATIVE LIMNOLOGY AND PATTERNS OF AQUATICPLANT DIVERSITY Matthew Hickler and Stuart Ludlam The Millers River Environmental Center Athol, Massachusetts With Support From The Massachusetts Environmental Trust 2006 CONTENTS FORWARD..........................................................................................................................v ACKNOWLEDGEMENTS...............................................................................................vi CHAPTERS INTRODUCTION...............................................................................................................1 Setting......................................................................................................................2 METHODS..........................................................................................................................3 Pond Selection.........................................................................................................3 Species Inventories..................................................................................................3 Comparative Limnology..........................................................................................4 DEFINITIONS.....................................................................................................................4 SECTION 1: BIODIVERSITY AND SPECIES DISTRIBUTIONS..................................6 GENERAL PATTERNS..........................................................................................6 Species Richness..........................................................................................6 Species Presence..........................................................................................7 Rare Species Non-Native Species.....................................................................................8 ALPHA, BETA ANDGAMMADIVERSITY..........................................................9 Diversity Estimators.....................................................................................9 Biodiversity..................................................................................................9 GammaDiversity.........................................................................................9 Alpha Diversity............................................................................................9 Beta Diversity............................................................................................10 Species Accumulation Curves.......................................................12 AssessingBeta Contribution Using Presence Values....................14 Summary of Beta Diversity...........................................................15 SECTION 2: ANALYSES OF DIVERSITY PATTERNS..............................................17 ARTIFICIAL VS. NATURAL PONDS................................................................17 Background................................................................................................17 Pond Origins..............................................................................................17 Biodiversity and Pond Origins...................................................................18 Alpha Diversity..............................................................................18 State-listed Species........................................................................18 Beta Contribution...........................................................................19 Species Composition......................................................................19 ii Indicator Species............................................................................19 Environmental Differences – Artificial vs. Natural Ponds........................20 SUMMARY...............................................................................................21 EXPLORATORY ANALYSES OF BIOLOGICAL AND ENVIRONMENTAL VARIABLES.........................................................................................................23 Background................................................................................................23 PCA Analyses............................................................................................23 Results........................................................................................................27 Species Richness Correlations.......................................................27 Summary of Species Richness Correlations......................30 Beta Diversity................................................................................31 Summary and Discussion...................................................32 State-listed Species........................................................................33 FINAL SUMMARY..........................................................................................................34 CONSERVATION ISSUES..............................................................................................35 Artificial Ponds: Qualities and Threats..................................................................36 LITERATURECITED......................................................................................................37 SECTION 3: COMPARATIVE LIMNOLOGY............................................................L-1 INTRODUCTION...............................................................................................L-1 METHODS..........................................................................................................L-2 RESULTS............................................................................................................L-3 Ranges of Water Quality Measures.........................................................L-3 Graphic Distribution of Water Quality....................................................L-5 Statistical Analysis of Water Quality and Morphometric Measures .......L-9 Differences Between Artificial and Natural Ponds................................L-11 Water Quality Profiles, Results..............................................................L-12 DISCUSSION....................................................................................................L-13 Water Quality.........................................................................................L-13 Graphic Distribution of Water Quality Variables..................................L-17 PCA and Factor Analysis of Water Quality...........................................L-18 PCA and Factor Analysis of Water Morphometry................................L-19 Morphometry and Water Quality...........................................................L-20 Differences Between Artificial and Natural Ponds................................L-21 iii Water Quality Profiles...........................................................................L-22 Defining Stratification...............................................................L-22 Non Stratified Ponds..................................................................L-22 Stratified Ponds..........................................................................L-23 SUMMARIES Water Quality.........................................................................................L-24 Influence of Watersheds on Water Quality............................................L-24 PCA Analysis.........................................................................................L-25 Lake Morphometry................................................................................L-25 Artificial Versus Natural Ponds.............................................................L-25 Profiles and Stratification......................................................................L-25 Limnology Literature Cited...............................................................................L-25 APPENDICES................................................................................................................A-1 Appendix A: Individual Pond Descriptions and Summaries..............................A-1 Appendix B: Flora.............................................................................................A-27 Composite Species List.........................................................................A-27 Species Presence Values Table.............................................................A-28 State-Listed Species Occurrences Table...............................................A-29 Individual Pond Floras..........................................................................A-30 Flora Notes............................................................................................A-35 Appendix C: Species Accumulation Curves.....................................................A-36 Appendix D: Tables of Lake Chemical and Physical Variables.......................A-40 Water Quality........................................................................................A-40 Physical Variables.................................................................................A-41 Appendix E: General Notes on Ponds and Their Origins ................................A-42 iv FORWARD This project was a collaboration between the two authors. Stuart Ludlam took the lead on the physical/chemical limnological components and Matthew Hickler coordinated the work and handled the botanical/ecological portions. Data collection, analyses, and writing were, for the most part, handled independently (but with frequent discourse). We have organized background information, ecological/botanical analyses, and limnological components in separate, stand-alone, chapters. Summaries of major findings appear at the end of most sections. A wealth of additional information including data tables, summary information on individual ponds, species lists, etc. may be found in the appendices. ACKNOWLEDGEMENTS The project was funded by a grant from the Massachusetts Environmental Trust, Ecosystem Health and Biodiversity Program. The Millers River Environmental Center (and Director, Sue Cloutier in particular) administered the grant, and provided much appreciated equipment and support. Our conversations with lake shore property owners, recreational users, and knowledgeable “locals” were invaluable. We are particularly grateful to those who shared data with us: MelissaCullina, The Massachusetts Division of Fisheries and Wildlife, Natural Heritage and Endangered Species Program. Robert Bertin, Holy Cross College. Jeff Mangum, Army Corps of Engineers, Tully Lake Flood Control Project. James West, Army Corps of Engineers, Tully Lake Flood Control Project. Rick McVoy, Massachusetts DEM, Division of Watershed Management. Scott Ryan, Massachusetts Division of Conservation and Recreation, Dam Safety Program. Leo Colette, Upper Naukeag Lake, Ashburnham. Matthew LaPointe, Ashburnham water filtration plant. Aaron Ellison and Albert Belanger (independently) helped us with statistical approaches for working with our data. However, the actual analyses and interpretations are our own. Margaret Ludlam provided field assistance on many trips. Contact Information Downloadable copies of the report are available from the Millers River Environmental Center: www.millersriver.net. Or, for paper copies contact: Millers River Environmental Center 100 Main Street Athol MA 01331 978-248-9491 For data used in the report, in electronic form, or other business, contact: Matthew Hickler 52 Butterworth Rd. Orange, MA 01364 978 249-5517 [email protected] vi INRODUCTION The contribution made by freshwater habitats to biodiversity is known to be exceptionally large relative to the portion of the landscape occupied. University of New Hampshire botanist Garrett Crow has called New England “the Amazonia of aquatic plant diversity” (NEBC Meeting Notes, 1998). In recognition of the unique contribution aquatic systems make to biodiversity, they have justifiably become a focus for biologists and conservation specialists (Natural Heritage and Endangered Species Program, 2003; Weiher et al., 1994; Rörslett, 1991). However, biodiversity patterns (and the ecological factors from which they arise) are complex, and shortcomings in our understanding hamper the design of effective conservation and management strategies. In particular, large, landscape-scale studies are rare and, thus, information on how biodiversity accumulates and how it is maintained at this scale is scarce. Parts 1 and 2 of this report document patterns of aquatic plant diversity in lakes and ponds of the Worcester Plateau Ecoregion in Massachusetts. Part 3 covers the physical and chemical limnology of waterbodies. The project was designed with two principal goals in mind: (1) determine how site variables associated with individual lakes and ponds correlate with key biological attributes (species richness, presence of rare species, and presence of regionally uncommon species); and (2) quantify how variation in species composition among lakes and ponds (beta diversity) contributes to the overall (gamma)diversity of the regional aquatic flora. More generally, the objective was to elucidatehow patterns of species distribution among ponds promote and constrain the overall biodiversity of the habitat type, and to search for site variables associated with observed patterns. Conservation and management efforts typically take place at large, landscape scales, whereas most ecological data are collected at small plot scales. Translating small- grain biological and environmental data to larger scales is fraught with pitfalls because many important ecological processes are scale dependant (Reed et al., 1993; Poiani, 2000; Jonsson and Moen, 1998). To ensure that data wereappropriate to the scale of interest, our approach was to collect biological data at the whole pond scale (i.e., whole- pond floras) and site variables at commensurate scales. In addition to exploring general patterns of biodiversity, we identified a number of specific questions to be addressed by the study: Do natural ponds differ biologically from man-made ponds? If so, how? Is it reasonable to presume that natural ponds, as a group, have higher quality biological attributes than artificial ponds? Do ponds that support rare species differ biologically from those that do not? Is it reasonable to presume that by focusing protection efforts on rare species habitats, the overall regional diversity can be captured and protected? Do shoreline and watershed development correlate with species richness or other biological qualities of ponds? How many ponds are needed to capture most of the aquatic species in the Ecoregion? What specific subset of ponds yields maximum diversity in minimum area? 1 Note: Our original scope-of-work included addressing questions about patterns of non- native invasive plant distributions. However, we were forced to abandon this plan because too few occurrences (three) were found. Setting The Worcester Monadnock Plateau Ecoregion (Figure 1) covers 1,136 km2 (438 square miles) in North-Central Massachusetts. It is characterized bymoderatelyhigh- elevation (500 to 1,400 feet), hilly terrain, poor acidic soils, and granitic and high-grade metamorphic bedrock. Population density is low relative to surrounding regions and the vast majorityof the land remains undeveloped. The 1999 landuse statistics show 8% of the land developed, 3% in agriculture, and the remaining 89% in natural vegetation - mostlyforest land. Figure 1 The Worcester-Monadnock Plateau Ecoregionof Massachusetts Worcester Plateau WorcesterPlateauEcoregion SurveyedPonds BERNARDSTON LEYDEN NORTHFIELD WARWICK ROYALSTON WINCHENDON ASHBY ASHBURNHAM ORANGE ERVING ATHOL FITCHBURG GARDNER PHILLIPSTON TEMPLETON WESTMINSTER PETERSHAM HUBBARDSTON PRINCETON BARRE N 10 0 10 Kilometers 2 METHODS POND SELECTION We used GIS (ARCVIEW) and data layers from Massachusetts Geographic Information System (MASSGIS) to identify waterbodies for potential inclusion in the study. The 1:5000 scale DEP Wetlands data layer was used to identify candidate waterbodies, and all polygons coded as “open water” were extracted in the initial step. The Open water polygons were clipped by the Upper Worcester Plateau Ecoregion polygon in the Ecoregions data layer to produce a new layer consisting of open water bodies contained within the Ecoregion. Water bodies that spanned the New Hampshire state line or were partially within other ecoregions were excluded. Finally, the surface area of each waterbody was calculated using ARCVIEW and lakes less than 5 hectares in surface area were removed. The final tally was 71 waterbodies greater than five hectares in surface area and totally contained within the Upper Worcester Plateau Ecoregion of Massachusetts. The distribution of size classes in the data set was heavily skewed towards small ponds. To ensure a relatively even distribution of size classes of lakes chosen for the study, we first stratified the 71 ponds into four size classes (5-10 ha., 10-25 ha, 25-50 ha and > 50 ha) then randomly selected five water bodies from each class for inclusion in the study, for a total of 20 ponds.The lower size limit of 5 hectares was set arbitrarily, but with the intent of avoiding, to the extent possible, small beaver impoundments, farm ponds, seasonal pools, and other small open-water habitats that are likely to have ecological qualities that are distinct from larger, more permanent ponds. Following the first season of field work and preliminary data analyses, we discovered that the data set was dominated by man-made ponds (19’th century mill ponds, water supply reservoirs, flood control impoundments, etc.) It turns out that artificial water bodies far outnumber natural ponds in the region. The study was designed, among other things, to evaluate differences between natural and artificial ponds. To ensure the dataset had enough natural ponds to do the necessary analyses, we added five natural ponds to the dataset using stratified random selection as described above, after the first year of the study was completed. The final dataset, therefore, consists of 25 waterbodies. Data on flora and summer water chemistry for the final five ponds were collected in 2005 (the year following data collection for the original 20 ponds). No spring water chemistry data were collected on the final five ponds. SPECIES INVENTORIES Species tallied for pond floras were limited to those perceived to be “true aquatics”, i.e. species that occur in areas that are routinely flooded for the entire growing season (although they may be rarely exposed during unusually dry periods). For most species, there was little ambiguityas to whether they qualified as true aquatic species or not. Some wetland species, however, occasionally find a foothold in shallow shoreline areas, and a few amphibious species have both aquatic and terrestrial forms that can occupy either habitat. Species with population centers in bordering wetlands with only 3 rare, errant individuals found growing in shallow water were excluded from the floras. Examples include: Lythrum salicaria, Decodon verticillata, Triadenum virginiana, and someCarexspecies. Amphibious species were counted as present when they had aquatic populations on a pond, but not if they were limited to bordering wetlands. Examples include: Proserpinaca palustris, Gratiola aurea, Juncus pelocarpus, Eleocharis acicularis, and Hypericum boreale. Thus, some species such as Typha latifolia were counted as present on ponds if found growing as an emergent aquatic, but not if limited to seasonally emersed bordering wetlands. The frequency of occurrence of a small number of amphibious species in the data set may not be representative of their actual regional presence; for instance Carex stricta, aubiquitouswetland species in the region, is rare in the dataset, found growing as a true aquatic on only a single pond. All inventories were completed between August 1, and September 15. Twenty ponds were inventoried in 2004; five additional ponds were added to the dataset and inventoried in 2005. Floras were compiled by thoroughly canvassing each lake by boat. The entire perimeter of each lake was explored in a zigzag pattern from shoreline to the maximum depth where visual observations were possible (usually about 2 meters). Frequent grab samples of vegetation were taken using a three-tined garden rake attached to a pole. The intent was always to find and identify every resident species of vascular plant, no matter how uncommon or limited in distribution. In practice, even with great vigilance, some species were undoubtedly overlooked. On all except the very smallest ponds, an entire field day was devoted to each pond’s inventory; however theactual field time spent on an inventory varied with size and complexityof vegetation. Collections were routinelymade of difficult-to-identify taxa for later confirmation. Voucher specimens were prepared for selected taxa and are on file at the University of Massachusetts herbarium. Nomenclature follows Sorrie and Somers, 1999. COMPARATIVE LIMNOLOGY Methods used in collecting lake physical and chemical data are presented in Section 3; Comparative Limnology. DEFINITIONS Some terms that are used throughout this report have been used inconsistently by various authors, or have meanings that vary depending on context. Others may be unfamiliar to some readers. The following definitions are provided to clarify how these terms are used in this report. Diversity or biodiversity: If not qualified (e.g. alpha diversity or beta diversity) means species richness, which is a simple count of species. Alpha diversity: When applied to a specific pond, alpha is the count of species documented during that pond’s inventory. It is the same as species richness. When applied to a group of ponds, or the dataset as-a-whole, it is the mean species richness of ponds in the group. Our alpha diversity estimates are based on intensive, whole-pond 4
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